Mangeenee thine TUT HME 0 whet vee d ” u ‘ * "I wae Sita ROR i on sacar ee ea chee : SEE RN Th EN aol eee MEU RETIN FID De ie nite hs tenmal RENE NM TH Pate ayes ss “ ’ Stee meet witae SE rene ae ote meee ot ehit het aeh sino Dp ets cece teen A. e Pte Bh Aah a an oe Fa, Sere ad ofa CRA peel oe Hear ot fe ne pee wed Ws oF teat e Pen enema o © Aram be tert “ LOS Mal ntind oP Putin a Pe Prats» eee : ~ “ near pete et tet No — ‘ ey ek SY aes * - ai Bulge aol orale atic gee eeyauavtt Colt bapocen sem hctee Porcartineubefgartegeh HARVARD UNIVERSITY ELE AS LIBRARY OF THE Museum of Comparative Zoology Me i - T va i TULANE STUIDINES UN AZOOLOGY AND BOTANY VOLUME 15 1968-1969 TULANE UNIVERSITY NEW ORLEANS TULANE STUDIES IN ZOOLOGY AND BOTANY is devoted primarily to the biology of the waters and adjacent land areas of the Gulf of Mexico and the Caribbean Sea, but manuscripts on organisms outside this geographic area will be considered. Each number is issued separately and contains an individual monographic study, or several minor studies. As volumes are completed, title pages and tables of contents are distributed to institutions receiving the entire series. Manuscripts submitted for publication are evaluated by the editor or associate editor and by an editorial committee selected for each paper. Contributors need not be members of the Tulane University faculty. The editors of Tulane Studies in Zoology and Botany recommend conformance with the principles stated in chapters I and II (only) of the Style Manual for Biological Journals, 2nd ed., published in 1964 by the American Institute of Biological Sciences, Washington, DE. Manuscripts should be submitted on good paper, as original typewritten copy, double- spaced, and carefully corrected. Two copies, carbon or other suitable reproduction, must accompany the original to expedite editing and assure more rapid publication. Legends for figures should be prepared on a separate page. Illustrations should be proportioned for one or two column width reproductions and should allow for insertion of legend if occupying a whole page. An abstract not exceeding three percent of the length of the original article must ac- company each manuscript submitted. This will be transmitted to Biological Abstracts and any other abstracting journal specified by the writer. The editors also recognize the policy adopted by the Federal Council for Science and Technology, and endorsed by the Conference of Biological Editors, that page charges for publication of scientific research results in scientific journals will be budgeted for and paid as a necessary part of research costs under Federal grants and contracts. Accord- ingly, writers crediting research grant support in their contributions will be requested to defray publication costs if allowable under the terms of their specific awards. Illustrations and tabular matter in excess of 20 percent of the total number of pages may be charged to the author; this charge is subject to negotiation. Exchanges are invited from institutions publishing comparable series but subscriptions are available if no exchange agreement can be effected. Separate numbers or volumes can be purchased by individuals and subscriptions are accepted. Remittance should accompany orders from individuals. Authors may obtain separates of their articles at cost. Address all communications concerning manuscripts and editorial matters to the editor; communications concerning exchanges, and orders for individual numbers to Dr. Royal D. Suttkus, Department of Biology, Tulane University, New Orleans, Louisiana 70118. When citing this series authors are requested to use the following abbreviations: Tzlane Stud. Zool. and Bot. Price per number (flat rate): $1.50 Price per volume (flat rate): $5.00 Gerald E. Gunning, Editor Muse Alfred E. Smalley, Associate Editor 2 = COM Pz Arthur L. Welden Associate Editor LIBRARy COL, Department of Biology Tulane University JA New Orleans, Louisiana 70118, U.S.A. Meade Natural History Libra NV 9) 19 Ar stony. J70 Tulane University Riverside Research Laboratoftés VAR Route 1, Box 46-B Niven o Belle Chasse, Louisiana 70037 SITy, Vickie L. Haddenhorst Assistant to the Editors NUMBER Ih CONTENTS OF VOLUME 15 SPEOCIROLANA THERMYDRONIS (CRUSTACEA: ISOPODA) FROM NORTH- EAST MEXICO, RE-DISCOVERY, HABITAT, AND SUPPLEMENTAL DESCRIP- UPON sega 2 A Dees ee cp gi ee Bn RN a W. L. Minckley and Gerald A. Cole A NEW CRAWFISH OF THE GENUS PROCAMBARUS FROM MISSISSIPPI (DECAPODA, ASTACIDAE) __ Joe B. Black OBSERVATIONS ON THE BEHAVIOR AND ECOLOGY OF THE NUTRIA IN Ts ESIC IN Ap oe ees Marilyn J. Warkentin NOTROPIS EDWARDRANEYI, A NEW CYPRINID FISH FROM THE ALABAMA AND TOMBIGBEE RIVER SYSTEMS AND A DISCUSSION OF RELATED SPECIES Royal D. Suttkus and Glenn H. Clemmer AGGREGATIVE BEHAVIOR AND HABITAT CONDITIONING BY THE PRAIRIE RINGNECK SNAKE, DIADOPHIS PUNCTATUS ARNYI _. eee Harold A. Dundee and M. Clinton Miller III A NEW TURTLE SPECIES OF THE GENUS MACROCLEMYS (CHELYDRIDAE ) FROM THE FLORIDA PLIOCENE James L. Dobie STUDIES ON FROG TRYPANOSOMIASIS II. SEASONAL VARIATIONS IN THE PARASITEMIA LEVELS OF TRYPANOSOMA ROTATORIUM IN RANA CLA- MITPTCAN SMER© Miclt OUTSTAN A 2 oe ee ee Ralph R. Bollinger, John Richard Seed and Albert A. Gam STUDIES ON AMERICAN PARAGONIMIASIS VI. ANTIBODY RESPONSE IN THREE DOMESTIC CATS INFECTED WITH PARAGONIMUS KELLICOTTI___ John Richard Seed, Franklin Sogandares-Bernal, and Albert A. Gam PLANOPHILA TERRESTRIS, A NEW GREEN ALGA FROM TENNESSEE SOIL Robert D. Groover and Sister Adrian Marie Hofstetter, O.P. LIFE CYCLES OF CARNEOPHALLUS CHOANOPHALLUS N. SP. AND C. BASODACTYLOPHALLUS N. SP. (TREMATODA: MICROPHALLIDAE) ________ John F. Bridgman ETHEOSTOMA COLLETTEI, A NEW DARTER OF THE SUBGENUS OLIGO- CEETALUSSHROM EOUISTANA AND ARKANSAS 2 a ee Ray S. Birdsong and Leslie W. Knapp MYSIDOPSIS BAHIA, A NEW SPECIES OF MYSID 2 eRe MYSIDACEA ) FROM GALVESTON BAY, TEXAS - _ we ~Joane “Molenock DIGENETIC TREMATODES OF MARINE TELEOST FISHES FROM BISCAYNE BAY, FLORIDA Robin M. Overstreet PAGE 10 41 59 64 70 75 81 106 119 aT 7 | cr | | WT Sere eee i - ra Ot ae » ee eo a ee « ee; TULANS STUDIES IN ZOOLOGY AND BOTANY 3 \ ys —— & NOV 1 1968 Volume 15, Number 1 HOOPER LO, 1968 UNIVERSITY. SPEOCIROLANA THERMYDRONIS (CRUSTACEA: ISOPODA) FROM NORTH- EAST MEXICO, RE-DISCOVERY, HABITAT, AND SUPPLEMENTAL DESCRIPTION W. L. MINCKLEY AND GERALD A. COLE Department of Zoology, Arizona State University, Tempe, Arizona 85281 1, A NEW CRAWFISH OF THE GENUS PROCAMBARUS FROM MISSISSIPPI (DECAPODA, ASTACIDAE ) JOE B. BLACK Department of Biological Sciences McNeese State College Lake Charles, Louisiana 70601 p. 5 OBSERVATIONS ON THE BEHAVIOR AND ECOLOGY OF THE NUTRIA IN LOUISIANA MARILYN J. WARKENTIN Department of Biology, Tulane University New Orleans, Louisiana oy KO) NOTROPIS EDWARDRANEYI, A NEW CYPRINID FISH FROM THE ALABAMA AND TOMBIGBEE RIVER SYSTEMS AND A DISCUSSION OF RELATED SPECIES ROYAL D. SUTTKUS and GLENN H. CLEMMER Department of Biology, Tulane University New Orleans, Louisiana p. 18 TULANE UNIVERSITY NEW ORLEANS TULANE STUDIES IN ZOOLOGY AND BOTANY is devoted primarily to the biology of the waters and adjacent land areas of the Gulf of Mexico and the Caribbean Sea, but manuscripts on organisms outside this geographic area will be considered. Each number is issued separately and contains an individual monographic study, or several minor studies, As volumes are completed, title pages and tables of contents are distributed to institutions receiving the entire series. Manuscripts submitted for publication are evaluated by the editor or associate editor and by an editorial committee selected for each paper. Contributors need not be members of the Tulane University faculty. The editors of Tulane Studies in Zoology and Botany recommend conformance with the principles stated in chapters I and II (only) of the Style Manual for Biological Journals, 2nd ed., published in 1964 by the American Institute of Biological Sciences, Washington, De. Manuscripts should be submitted on good paper, as original typewritten copy, double- spaced, and carefully corrected. Two copies, carbon or other suitable reproduction, must accompany the original to expedite editing and assure more rapid publication. Legends for figures should be prepared on a separate page. Illustrations should be proportioned for one or two column width reproductions and should allow for insertion of legend if occupying a whole page. An abstract not exceeding three percent of the length of the original article must ac- company each manuscript submitted. This will be transmitted to Biological Abstracts and any other abstracting journal specified by the writer. The editors also recognize the policy adopted by the Federal Council for Science and Technology, and endorsed by the Conference of Biological Editors, that page charges for publication of scientific research results in scientific journals will be budgeted for and paid as a necessary part of research costs under Federal grants and contracts. Accord- ingly, writers crediting research grant support in their contributions will be requested to defray publication costs if allowable under the terms of their specific awards. Illustrations and tabular matter in excess of 20 percent of the total number of pages may be charged to the author; this charge is subject to negotiation. Exchanges are invited from institutions publishing comparable series but subscriptions are available if no exchange agreement can be effected. Separate numbers or volumes can be purchased by individuals, but subscriptions are not accepted. Remittance should accompany orders from individuals. Authors may obtain separates of their articles at cost. Address all communications concerning manuscripts and editorial matters to the editor; communications concerning exchanges, and orders for individual numbers to the Meade Natural History Library. When citing this series authors are requested to use the following abbreviations: Tulane Stud. Zool. and Bot. Price per number (flat rate): $1.50 Price per volume (flat rate): $5.00 Gerald E. Gunning, Editor Arthur L. Welden Associate Editor Department of Biology Tulane University New Orleans, Louisiana 70118, U.S.A. Meade Natural History Library Tulane University New Orleans, Louisiana 70118, U.S.A. Tim M. Berra Assistant to the Editors TULANE STUDIES IN ZOOLOGY AND BOTANY Volume 15, Number 1 Ge Xi Py October 16, 1968 MUS. COMP. ZUOL. LIBRARY NOV 1 1968 HARVARD Eprrors’ NOTE UNIVERSITY. Effective with Volume 15, Number 1, the name of the biological journal published by Tulane University will be Twlane Studies in Zoology and Botany instead of Tulane Studies in Zoology. The Departments of Zoology and Botany were united into a single Department of Biology some time ago. The name change for the journal reflects a de- sire of the biology faculty to include botanical papers. Policies with regard to editorial review and exchanges remain the same. SPEOCIROLANA THERMYDRONIS (CRUSTACEA: ISOPODA) FROM NORTH- EAST MEXICO, RE-DISCOVERY, HABITAT, AND SUPPLEMENTAL DESCRIPTION W. L. MINCKLEY AND GERALD A. COLE Department of Zoology, Arizona State University, Tempe, Arizona 85281 ABSTRACT Speocirolana thermydronis, previously known from a single specimen, was re- discovered in interstices of travertines that rim large spring-fed pools near Cuatro Ciénegas Coahuila, México. The 29 new specimens differ little from the original de- scription, but some variations are described. Speocirolana thermydronis Cole and Minckley (1966) was described from a single, large female collected 12 April 1964 near Posos de la Becerra, 13.7 km southwest of Cuatro Ciénegas, central Coahuila, México. In August 1967, 29 additional speci- mens were caught, 22 from an unnamed poso (sinkhole) ca. 12 km southwest of Cuatro Ciénegas, and 7 specimens from the west laguna of El Mojarral, 11 km southwest of that town. Considerable time spent at a number of other places, including the area of original discovery, failed to obtain the isopod. Habitats —The unnamed poso is a spring tributary to the west laguna of EI Mojarral. Its dimensions are about 10 by 12 m, by about 2 m deep. There was about 0.85 m of water in the pit, over a dark, flocculent bot- tom of silt, shells of the snails Paludiscala caramba Taylor and Durangonella sp., and travertine blocks and fragments. The banks are of gypsum, matted roots of desert plants, and travertine. The inflowing water was 34 C. This is the type-locality of P. caramba, the most distinctive of the freshwater snails of the Cuatro Ciénegas basin (Taylor, 1966: 207-8). The second locality for S$. thermydronis, the western limnocrene in the marshy area called El Mojarral, has been described by Taylor (1966:163, pls. 9-10). It is a large, clear pool, about 25 by 100 m and up to 5 m deep. Large springs enter at the north- west end and water leaves by a subsurface channel and by small surface outflows at the southeast. Water temperatures at the inflow are about 33° C, but some variations occur in places remote from the source. Much of the bottom is soft, gray, flocculent silt, stabilized locally by beds of waterlily, Nymphaea sp. Banks are undercut and mostly of gypsum and _ travertines. All our new specimens of S. thermydronts were collected by manually breaking large, loosely-cemented, porous travertine blocks that were abundant around the margins of the pools. In some instances, individual isopods moved from interstices of a block as water drained from it, and could be picked from the surface. Minckley (1961:454) described similar habitats for a troglobitic asellid, Asellus stygius (Packard), in porous marls of Doe Run, Meade County, Ken- tucky. S. thermydronis was accompanied in its habitat by an undescribed species of aquatic isopod, by an aquatic mite, and by an oligochaete. The last three animals also lived in the softer sediments; however, S. thermydronis was consistently absent from such places. It seems unlikely that S. thermy- dronis moves extensively within the sedi- ments that characterize most lagunas of the Cuatro Ciénegas area. Living specimens re- tained in the field soon became entangled with loose sediment particles on the bottoms of their containers, and were distinctly in- capacitated. A. stygivs is similarly affected under certain conditions (Minckley, 1961: 454). Life on the surface of the undisturbed sediments also is most likely limited to places where fishes are absent; as many as 12 species of fish, many of which are highly predaceous, occur in larger springs of the area. EDITORIAL COMMITTEE FOR THIS PAPER: Dr. THOMAS E. BOWMAN, Curator, Division of Crustacea, The Smithsonian Institu- tion, Washington, D. C. Dr. ALFRED E. SMALLEY, Associate Professor of Biology, Tulane University, New Orleans, Louisiana No. 1 Neil F. Hadley (personal communica- tion), now of Arizona State University, was with Mary L. Allesio when she collected the holotype in 1964, and has provided addi- tional data on the type-locality. The holo- type was one of three, similarly-sized in- dividuals observed in a small pit, about 0.5 m in diameter and one m deep, ca. 0.6 km south of the head-spring of Posos de la Becerra. The animals were in the shade and were moving about on the surface of gray, flocculent sediment beneath about 0.25 m of water. The pit was near extensive marshes that existed downflow from La Becerra prior to diversion of water by canalization in late 1964 (Cole and Minckley, 1966). The en- tire marsh-river-poso complex south and west of La Becerra now is dry, and during core-drilling operations in 1967 the water table was encountered about 3 m below the land surface—there is little doubt that the point of original capture also is destroyed. Our failure to find S. thermydronis in the remaining habitats of Posos de la Becerra may be attributed, in part, to destruction by desiccation of most marginal deposits of suitable travertine. The sudden drying of the laguna decimated fish and endemic snail populations (Minckley, in Taylor, 1966: 162-3). In 1967, some of the snails were becoming re-established, as may S. thermy- dronis, as the habitat stabilizes. On the other hand, absence of the species from ap- parently suitable habitat in a number of other springs diminishes somewhat the im- portance of its apparent absence in La Becerra. This may result from the well- known vagaries of collecting such secretive organisms. Much of the basin floor has water within a meter of its surface; numer- ous large, subterranean channels are known, and interstitial habitat for S. thermydronis, as well as for other animals, must be exceed- ingly abundant. We have little doubt that additional specimens and habitats will be found. Supplemental description —Those features of S. thermydronis adequately described in our original paper are not elaborated here. We find that part of the description was in error, however, and reproduce that section (Cole and Minckley, 1966:18), with cor- rections in boldface type, as a footnote.! 1In S. thermydronis the mandibles (Fig. 6) are asymmetrical, with the left incisive process Cirolanid Isopod 3 In addition, the second and third pereonites of S. thermydronis bear epimera, as do other species of the genus—their absence in the holotype was due to an artifact of prepara- tion, as was suspected. Body shape and proportions in S. thermy- dronis are quite constant. The telson, broad and acutely tipped in both sexes, seems a good character separating the Coahuilan form from S. pelaezt (telson broadly rounded ) and S. bolivari (telson truncate). It appears that length and number of segments in the antennal flagella in the genus Speocirolana may vary considerably with size, with larger individuals tending to add length and segments. All specimens of S. thermydronis are small (the holotype at 15 mm is largest), whereas S. pelaezi ranges from 19 to 26 mm and S. bolivari from 15 to 35 mm (Bolivar y Peltain, 1950; Rioja, 1953). The second antenna of the 15- mm holotype of S. thermydronts reaches pos- teriad to the seventh pereonite (as in S. pelaezi). In the smaller, new specimens of S. thermydronis, the maximal extension is to the fifth pereonite. And, in the large S. bolivari, its extension is far past the seventh. Articles in the first antennal flagellum of S. thermydronis range from 6 to 14 (including the holotype, at the maximum). The second antenna is similarly variable, ranging from 25 to 35 (with the large holotype again bearing the maximum). S. bolivari has 22 to 28 seg- ments in the first antennae, and 48 to 82 in the second, and S. pelaezi has 20 and 30 segments, respectively. The relative elonga- tion of terminal segments in the second an- tennal flagellum is variable—the holotype has long terminal segments (as in S. pelaezt), yet some smaller specimens have shortened segments indistinguishable from those of S. bolivart. The inner plate of the first maxilla of S. thermydronis is invariable in bearing three sparsely-plumose spines and two delicate setae. The outer plate, however, often has as many as 12 strong, distal spines (10 in the holotype). The maxilliped of S. thermydronis seems highly diagnostic. Penultimate and ante- overlapping the right ventrally. The pars molaris of the mandible is sub-triangular, bear- ing on its margin about 34 short, cone-shaped teeth. 4 Tulane Studies in Zoology and Botany penultimate segments of the palp have only fine, hairlike setae on their outer surfaces, except for a strong spine at each of their distal corners. The ultimate segment is some- times naked on its outer surface, as in the holotype, but in some specimens there are fine hairlike setae on the proximal half. The outer surfaces of the last three palp segments in S. bolivari are strongly setaceous, and such a condition also is found on the penultimate and antepenultimate segments in S. pelaezi. These species, however, lack the strong, isolated spines at the distal, outer corners of the last two segments. Such a spine is present only on the first segment of their palps (Bolivar y Peltain, 1950:Fig. 5; Rioja, 1953:Fig. 18). All pereopods of S. thermydronis are es- sentially as described, except for variable armament on the subcheliform, first three pairs. The holotype has the palmar margins of the propodus armed with 2, 3, and 4 stout spines, respectively; many of the new specimens have 2, 4, and 4 such spines. The exopod of the fifth pleopod of the new material (rather than the fourth, as erroneously given for the holotype) is only sparsely invested with plumose setae. We also find that both sexes of S. thermydronis lack distal setae on the endopodites of all pleopods. Presence of such setae on the first two pleopods, and their absence on the suc- ceeding three, as in S. bolivari and S. pelaezi, is therefore not a character of generic rank as was tentatively suggested by Bowman (1964:233-4). Distribution of setae on the pleopods does, however, provide good specific characterization—S. bolivari has setae on endopods of at least the first and second pleopods in males (Rioja, 1953: Figs. 28, 30); S. pelaezi has setae on the endopod of the first pleopod in females and at least the second in males (Bolivar y Peltain, 1950:Figs. 9, 11); and, as given before, all pleopodal endopods of both sexes of S. thermydronts lack setae. Except for the modified second pleopods of the male, no sexual dimorphism is obvious in the genus Speocirolana. The copulatory organ of S. thermydronts rises from the base Noles of the pleopodal endopod. The organ 1s variable in length, ranging from slightly shorter to slightly longer than the endopod. Living specimens of S. thermydronis have ivory-white exoskeletons, with dark-brown or black masticatory surfaces on the man- dibles and terminal claws of the pereopods. In addition, the spines on the pereopods are straw-yellow. There is a variable amount of red-to-brownish pigment associated with internal organs, and this is visible through the dorsum of living animals, but fades rapidly to light-brown or yellow in alcoholic material. Specimens of S. thermydronis from the 1967 collections are housed at the U. S. National Museum, Washington, D. C., the Instituto Nacional de Investigaciones Bio- l6gico Pesqueros, México, D. F., and the National Museum of Canada, Ottawa, On- tario. A permit for collection of aquatic ani- mals in México was supplied by the Direc- cion General de Pesca e Industrias Conexas, México, D. F. Numerous personnel of Arizona State University deserve thanks for help in the field and in the laboratory. Studies were supported, in part, by Grants GB-2461 and GB-6477X from the National Science Foundation. LITERATURE CITED Bourvar y PELTAIN, C. 1950. Estudio de una Cirolana cavernicola nueva de la region de Valles, San Luis Potesi, México. Ciencia, 10( 11-12) :211-218. Bowman, T. E. 1964. Antrolana lira, a new genus and species of troglobitic cirolanid isopod from Madison Cave, Virginia. In- ternat. J. Speleol., 1( 1-2) :229-236, pls. 50-57. Coir, G. A., and W. L. Muinckiey. 1966. Speocirolana thermydronis, a new species of cirolanid isopod crustacean from central Coa- huila, México. Tulane Stud. Zool., 13(1): a2" Mrincxtey, W. L. 1961. Occurrence of subter- ranean isopods in epigean waters. Amer. Midl. Nat., 66(2):452-455. Rroya, E. 1953. Estudios carcinologicos. XXX. Observaciones sobre los cirolanidos cavernico- los de México (crustaceos, isopodos). An. Inst. Biol., Mexico, 24(1):141-170, lam. 1-6. Taytor, D. W. 1966. A remarkable snail fauna from Coahuila, México. Veliger, 9(2):152- 228, pls. 8-19. October 16, 1968 A NEW CRAWFISH OF THE GENUS PROCAMBARUS FROM MISSISSIPPI (DECAPODA, ASTACIDAE ) JOE B. BLACK Department of Biological Sciences McNeese State College Lake Charles, Louisiana 70601 ABSTRACT A new crawfish species of the Spiculifer Group, Blandingii Section, of the genus Procambarus is described from tributaries of the Tombigbee River in eastern Missis- sippi. P. lagniappe, new species, is related to P. penni and P. spiculifer. The first specimens of the species de- scribed here were collected on June 16, 1955 by the late George H. Penn and me from two creeks in Kemper Co., Mississippi (Tombigbee River drainage). Dr. Penn first identified these specimens as Procambarus penni Hobbs (1951:273). Later he changed the labels to read “atypical P. versutus” (Hagen, 1870:51). In August, 1965, I col- lected in this area again, finding additional similar specimens recognizing them as an undescribed species. This species belongs to the Spiculifer Group of the Blandingii Section of the genus Procambarus and is the fourth species, to date, of this group to be recorded from Mississippi. Other species are P. penni from the Pearl and Pascagoula River drainages, P. ablusus Penn (1963:121) from the Hatchie River drainage, and P. viosca: Penn (1946:27) from the Pearl, Amite, Tangipa- hoa, Homochitto, Big Black, Yazoo, Wolf, and Tombigbee River drainages. I wish to express my gratitude to Dr. Horton H. Hobbs, Jr., Senior Zoologist, Smithsonian Institution, for examination of specimens and confirmation of identifica- tion. Thanks are also due Dr. Alfred _E. Smalley, Tulane University for the loan of specimens of this species from the Tulane University collections. -y Procambarus \agniappe,! new species Diagnosis —Body pigmented; eyes nor- mal. Rostrum with prominent marginal spines; acumen attenuate, constituting 43.3- 44.5 per cent of total length of rostrum; postorbital ridges terminating in spines; suborbital angle much reduced; lateral sur- face of carapace with two cervical spines. Areola 5.0-6.2 times longer than wide, con- stituting 25.3-27.7 per cent of entire length of carapace. Simple hooks present on ischio- podites of third and fourth pereiopods; basipodites of fourth pereiopods bear an op- posable tubercle on cephalic surface. First pleopod of first form male (Figs. 1, 6, 12) without shoulder on cephalic margin, termi- nating distally in three distinct elements; mesial process subspiculiform and directed caudodistally with only tip corneous; cephalic process lacking; caudal process represented by low hemi-disc arising from much reduced caudal knob; central projection corneous, subacute, extending caudodistad at about 60 degree angle to axis of pleopod shaft; fusion lines of its component elements clearly marked. Annulus ventralis as figured (Fig. Dye Holotypic male, Form I1—Carapace (Figs. 3,7) subovate, slightly compressed laterally; abdomen slightly longer than carapace (39.0 and 38.2 mm, respectively). Maximum height of carapace slightly more than maxi- mum width (16.8, 15.4 mm). Areola rela- tively broad and short (constituting 25.6 per cent of entire length of carapace), about 1 Creole French: in Louisiana, a little some- thing extra given to customers by tradesmen. EDITORIAL COMMITTEE FOR THIS PAPER: Dr. Horton H. Hoss, JR., Senior Zoologist, Department of Invertebrate Zoology, Smithsonian Institution, Washington, D. C. Dr. JOSEPH F. FITZPATRICK, JR., Department of Zoology, Mississippi State Univer- sity, State College, Mississippi 6 Tulane Studies in Zoology and Botany Vol. 15 Figures 1-12. Procambarus lagniappe, new species; 1. Mesial view of first pleopod of holo- type; 2. Mesial view of first pleopod of morphotype; 3. Dorsal view of carapace of holotype; 4. Upper view of carpus and chela of holotype; 5. Lateral view of first pleopod of morphotype; 6. Lateral view of first pleopod of holotype; 7. Lateral view of carapace of holotype; 8. Epistome of holotype; 9. Annulus ventralis of allotype; 10. Basipodites and ischigpodites of third (upper) and fourth (lower) pereiopods of holotype; 11. Antennal scale of holotype; 12. Caudal view of first pleopod of holotype (pubescence removed from all structures illustrated). No. 1 five times as long as wide, with five fine punctations in narrowest part. Cephalic por- tion of carapace nearly three times as long as areola. Rostrum long, acumen extending cephalad beyond peduncle of antennule, excavate; with acute marginal spines. Rostrum widest at base, margins elevated and converging anteriorly; no median carina. Acumen long and thin. Postorbital ridges well developed, each terminating cephalad in acute spine; punctations along mesial sur- faces of ridges, tubercles along lateral sur- faces. Brachiostegal spine well developed. Suborbital angle much reduced. Cervical groove interrupted on each side by two conspicuous acute spines. Upper two-thirds of carapace punctate, lower one-third strongly granulate. Epistome (Fig. 8) with raised lateral margins converging toward small cephalo- median spine. Cephalic section of telson with three spines in each corner. Antennules of usual form, with prominent acute spine on ventral side of basal segment. Antennae not quite reaching tip of abdomen. Antennal scale (Fig. 11) long, tip extend- ing beyond tip of acumen, widest slightly proximal of midlength, outer distal margin with strong spine. Chela (Fig. 4) ovoid in cross-section, long, slender; single row of seven tubercles along inner margin of palm; smaller tuber- cles in six irregular longitudinal rows on upper surface of palm. Both fingers terminat- ing in short, corneous tips bent toward each other. Movable finger with one prominent tooth on opposable margin near base. Im- movable finger with two prominent, widely spaced teeth on opposable margin. Fingers long; dactyl about 53 per cent of total length of outer margin of chela. Carpus (Fig. 4) ovoid in cross-section; with two prominent, acute tubercles in distal half of inner margin; few squamous tubercles arranged in two rows between inner margin and longitudinal groove. Basipodite of first pereiopod with- out spines. Ischiopodites of first pereiopods each with four spines on inner margin. Hooks on ischiopodites (Fig. 10) of third and fourth pereiopods simple, subequal in length; basipodites of fourth pereiopods each bearing small tubercle on distal cephalic surface opposite hook. Apophyses on coxo- podites of fourth and fifth pereiopods with caudomesial projections, that on fourth Procambarus lagniappe y knoblike, that on fifth compressed and acute. First pleopod (Figs. 1, 6, 12) extending to middle of coxopodite of third pereiopod when abdomen flexed. Pleopod essentially straight; terminating in three distinct ele- ments; mesial process subspiculiform and directed caudodistally at about 55 degree angle to axis of main shaft, tip corneous; cephalic process lacking; caudal process low, flattened corneous hemi-disc; caudal knob low, reduced; central projection corneous, subacute, extending caudodistad at 60 de- gree angle to axis of main shaft, fusion lines of component parts clearly defined. Pleo- pods asymmetrical with base of left dis- placed in plane posterior to right. Allotypic female—Differs from holotype in following respects: rostrum more slender, acumen proportionately longer; areola pro- portionately narrower; rostrum widest slightly cephalad of base; tip of antennal scale reaching tip of acumen; chela_pro- portionately wider, thicker; antennae extend caudad to middle of fifth abdominal seg- ment. Annulus ventralis (Fig. 9) movable; spindle-shaped, widest in transverse axis; cephalic margin partially obscured by sev- eral tuberculate prominences extending cau- dad from sternum immediately cephalic to it; sinus originating near midcephalic mar- gin and forming sinuous line to near mid- caudal margin of annulus. Area adjacent to sinus raised, smooth. Morphotypic male, Form Il.—Differs from holotype in following respects; carapace less compressed laterally; height and width of carapace subequal; areola proportionately wider (4.5 times as long as wide); an- tennae extending caudad to caudal margin of telson; chela and hooks on ischiopodites reduced; caudomesial processes on COxo- podites of fourth and fifth pereiopods less well developed; ischiopodites of each first pereiopod with three spines. First pleopods (Figs. 2, 5) asymmetrical, right pleopod reaching almost to anterior edge of coxo- podite of third pereiopod when abdomen flexed, left displaced posteriorly reaching to middle of coxa of third pereiopod. All termi- nal elements present but less acute and non- corneous; caudal knob more prominent. Type Locality—Pawticfaw Creek (tribu- tary to Sucarnoochee Creek, in turn a tribu- tary of Tombigbee River) 6.0 miles south of DeKalb, Kemper County, Mississippi, at 8 Tulane Studies in Zoology and Botany Measurements (in mm) — Mor- Holo- Allo- pho- type type type Carapace: length 38.2 48.0 34.8 width (max. ) 15.4 19.6 14.5 height (max. ) 16.8 20.6 14.5 Abdomen length 39.0 49.7 36.5 Areola: length 9.8 OR 9.0 width (min. ) 1.8 2.0 1.8 Rostrum: length 14.8 19.0 13.3 width 5.2 6.8 4.9 length of acumen 6.6 9.1 6.1 Antennal scale: length 13:3 15.5 Hig) width 4] 5.0 3.8 First pleopod length 9.5 -- 7.9 Right chela: length, outer margin 30.3 31.5 21.0 length, dactyl 15.8 16.8 11.8 width, palm TA 8.9 al thickness of palm (max. ) 5.6 6.2 3.5 Mississippi Highway 39. Here the stream varies in width from 10 to 30 feet, with a bottom of white sand and plant debris. The water is Clear, with good current, suggestive of a spring-fed stream. Other crawfishes present include an unidentified species of Orconectes (no first form males collected ) , which was predominant and was only found in piles of plant debris. All specimens of P. lagniappe were taken from scattered growths of Vallisneria in the swifter parts of the stream. Disposition of types—tThe holotypic male, Form I, the allotypic female and the morpho- typic male, Form II, all from the type lo- cality, are deposited in the United States National Museum (nos. 119088, 119089, 119090, respectively). The following para- types are retained in my personal collection at McNeese State College: JBB 108 (type locality)—one ¢ I, five ¢¢ II, two 2 2; JBB 206 (type locality)—one ¢ II. Para- types in the Tulane University Collections are: TU 3174, Blackwater Creek, 7.8 miles south of DeKalb—one 4 I, three ? 2, GHP and JBB coll.; TU 3613 (type locality )— one 6 I, one’ o: Il, one 2, JBB coll: Color notes——Background color of most of the dorsal part of the carapace is medium brown. The sides of the cephalic and thoracic Voli region are marked by an incomplete (in the mid-dorsal area of the areola) U-shaped yoke of olive brown. The yoke increases in width caudally and is widest at caudal edge of carapace where the base of the U almost meets in the median line. A large cream colored area is below the “yoke” on the cephalic portion, which area darkens to light olive-tan on the thoracic portion of the carapace. The mid-dorsal portion of the abdomen is marked by a lightly-defined stripe of medium brown. On either side of this stripe, the background color is cream. The caudal portion of each abdominal seg- ment is deep olive brown. A deep, olive brown, lateral “V” connects the caudal por- tions of adjacent segments. A pair of small, rectangular, deep olive brown blotches mark the cephalolateral portion of each segment. Between the transverse caudal stripes and above the “V” on either side is a rust-colored blotch on the second through fifth abdominal segments. The cephalic sec- tion of the telson is marked by an inverted cream colored crescent surrounded by olive brown, the caudal section is uniformly medium brown. The bases of the uropods are cream, with mesial rectangular stripes of dark brown surrounded by a large area of rust brown. The palms of the chelae are medium brown with dark olive brown tubercles. The dactyl and immovable finger are dark olive brown, except for cream colored tips. Variations —There are few variations of import in observed specimens. The annuli ventrales of smaller adult females are sig- nificantly different. Smaller females do not possess tuberculate prominences on the sternum just cephalic to the annulus ven- tralis. Also the median sinus on the annulus ventralis of smaller females is less deeply sculptured and shows less lateral twisting. Relationships——On the basis of the struc- tures of the first pleopod of Form I males, P. lagniappe is most closely related to P. spiculifer (LeConte, 1856:401), resembling it in lacking a distinct cephalic process, but differing from it in having a much reduced caudal knob. This resemblance may be more superficial than actual. Geographically, its closest Spiculifer Group relative is P. penni Hobbs (1951:273). Second form males show a closer resemblance to P. penni than No. 1 Procambarus lagniappe 9 to P. spiculifer; also the annuli ventrales of females of P. penni and P. lagniappe are markedly similar. LITERATURE CITED Hacen, H. A. 1870. Monograph of the North American Astacidae. Illus. Cat. Mus. Comp. Zool., Harvard Coll., 3:1-109. Hosss, H. H., Jr. 1951. A new crayfish of the genus Procambarus from Louisiana, with a key to the species of the Spiculifer Group. J. Washington Acad. Sci., 41:272-276. LEConreE, J. 1856. Descriptions of new species of Astacus from Georgia. Proc. Acad. Nat. Sci. Philadelphia, 7:400-402. PENN, G. H. 1946. A new crawfish of the genus Procambarus from Louisiana. J. Washington Acad. Sci., 36:27-29. see (Decapoda, Astacidae). Proc. Biol. Soc. Washington, 76:121-125. October 16, 1968 OBSERVATIONS ON THE BEHAVIOR AND ECOLOGY OF THE NUTRIA IN LOUISIANA MARILYN J. WARKENTIN!} Department of Biology, Tulane University New Orleans, Louisiana ABSTRACT A population of feral nutria (Myocastor coypus) on Tulane University’s Riverside Campus near Belle Chase, Plaquemines Parish, Louisiana, was observed at 2-3 day intervals from January 29, 1967, through July 11, 1967. Twenty-eight nutria were trapped, marked with permanent numbered tags and with distinctive field markers, and released. There were 21 recaptures. Ac- tivity peaked at noon, late afternoon, and late evening. No observations were made of predawn activity. The major type of activity changed from sunning to constant grooming and feeding with increasing ambient temperatures. Solitary nutria were rarely observed. Daily cruising ranges of nutria were usually less than 200 yds. Preference was shown for areas of shade and areas covered by duckweed. Nutria lived colonially in large dens built into the banks of ponds. Each colony lived in a separate den; a typical colony consisted of 15-20 nutria. No nests or platforms were built; instead, floating objects or exposed roots of willow trees were utilized for sun- ning and feeding. The main diet was duck- weed (Lemna minor, Wolfia sp., and Wolfiella floridana) and white clover ( Tri- folium sp.). Within each social group, there was one alpha female and one alpha male; all other animals were equally subordinate. The alpha female dominated the alpha male. Aggression consisted of warning sounds (“mooing”) and chasing subor- dinate animals. Fighting was ritualistic in the water. There was little care of young by the mother and no attempt was made to protect them from predators. Garfish, snakes, and red-shouldered hawks took young nutria. Man was the major pred- ator on adult nutria. When alarmed, nutria assumed a “watching” position in the water, or dived underwater, slapping the water with the hind feet to produce an alarm sound. The South American beaver or nutria, (Myocastor coypus), is a large hystrico- morph rodent which was first successfully introduced into Louisiana from Argentina by E. A. MclIlhenny in March, 1937, as a fur animal to be raised in captivity. The subspecies found in Louisiana and Texas, according to Lowery (1943), is Myocastor coypus bonariensis, a form native to north- ern Argentina, Uruguay, Paraguay, and southern Brazil. Many nutria which escaped or were released from fur ranches have es- tablished feral colonies throughout Louisiana (Harris, 1956). These colonies have done great damage in agricultural areas, especially to rice and sugar cane crops (Evans, pers. comm. ). In an effort to develop some method of control, much interest has arisen in the behavior and ecology of nutria. Tulane University’s Riverside Campus near Belle Chase, Plaquemines Parish, Louisi- ana, supports a moderately large, relatively isolated population of nutria. The purpose of this paper is to report on a study of the habits and habitat of this population of feral nutria, Description of Study Area Three interconnected ponds (comprising an area of 130,000 sq ft) were chosen as the study site. Two biotic communities were found around these ponds. Disturbed areas, which had been cleared of trees and shrubs, were in closely-cropped grass and white clover, Trifolium sp. Undisturbed areas, which surrounded the ponds on three sides, were typical lowland wooded habitat. The dominant vegetation was Acer rubrum, Ul- mus americana, Populus deltoides, Acer negundo, Salix nigra, Myrica cerifera, Quer- 1 Present address: Department of Zoology, University of Missouri, Columbia, Missouri. EDITORIAL COMMITTEE FOR THIS PAPER: Dr. JAMes N. Layne, Director, Archbold Biological Station, Lake Placid, Florida Mr. JAMrEs Evans, Leader, Jackrabbit Research Station, U. $. Department of the Interior, Twin Falls, Idaho 10 No. 1 cus virginiana, Corundus drummondt, Celtis laevigata, and Sambucus canadensis, Several species of the mustard family grew along the water’s edge wherever there was suf- ficient sunlight. During this study, the ponds were always covered to some extent by a mixture of Lemna minor, Wolfia sp., and Wolfiella floridana (all referred to as “duckweed” in the remainder of the paper). During February and early March, this cover was relatively sparse, but by April, a thick blanket of aquatics covered the ponds. All ponds con- tained numerous underwater fallen logs on which the nutria sat while eating. Many of these logs were later exposed when water levels in the ponds dropped. METHODS AND MATERIALS Nutria were observed in the study area at regular (2-3 day) intervals from Janu- ary 29, 1967, through July 11, 1967. At- tempts were made to observe nutria activity at all hours, but due to lack of adequate night-viewing equipment, most observations were made during daylight hours. Observa- tions usually lasted 3-4 hours (for a mini- mum total of 200 hrs), and whenever pos- sible, air and water temperatures, as well as current water conditions, were recorded for each group of observations. Most ob- servations (75%) were made from a boat in the center of the first pond. Nutria were trapped, marked, and released during three different periods. On March 1, 1967, ten traps were placed around the edges of two ponds in areas showing definite signs of nutria activity. Traps used were double-door treadle traps, 10” X 10” X 32”, baited with carrot cubes. Traps were set with both doors open. These traps were removed on March 10, 1967. On April 19, 1967, one 4’ X 4 raft was placed in the center of the first pond. During the second period of trapping, four traps of the same size as above, baited with carrots, were placed on this raft and trapping continued for six days (April 24-30). On the raft, only one door of each trap was opened and the traps were arranged so that each one had a “lead-in” of carrot bits. On May 19, 1967, a second raft, 4.5’ x 6.5’, was placed on the third pond. During this period of trapping, six traps were placed on this raft and four traps were placed on the raft in Nutria Behavior and Ecology 1] the first pond. Trapping started on June 26, 1967; all traps were removed on July 2, IGA Each captured nutria was removed from the trap by means of a handling bar, mea- sured, weighed, and then marked in three ways: by a National #3 metal numbered tag through the cartilage of the left ear, by a correspondingly numbered #3 tag in the webbing of the left hind foot, and by a distinctively colored plastic flag pinned by a stainless steel safety pin through the skin of the nape of the neck. The numbered tags were for long-term identification; the nape marker was for field identification and was generally short-term due to sloughing. Each animal was released at the site of capture. If a nutria was recaptured, the location of the trap and the animal's number were recorded. The nutria was then measured and weighed, and if necessary, the nape marker was replaced. Samples of the vegetation around the study ponds were collected regularly from February through July, 1967. Specimens were pressed, dried, identified, and placed in the Environmental Biology Herbarium of Tulane University to be mounted at a later date. RESULTS AND DISCUSSION In the first ten days of trapping, 19 nutria were marked and 3 were recaptured. Marked animals included 8 adult females, 5 juvenile females, and 5 juvenile males. One adult male was taken. Two juvenile females and 1 juvenile male were recaptured. In the second period of trapping, 5 nutria (3 juve- nile males, 1 juvenile female, and 1 adult female) were recaptured a total of 11 times, but only 1 new animal, a juvenile male, was marked. During the last trapping pe- riod, 8 nutria (4 juvenile females and 4 adult males) were marked and 4 animals (2 juvenile females, 1 juvenile male, and 1 adult female) were recaptured a total of 7 times. Age determination was in ac- cordance with the findings of Atwood (1950): all animals with a total length greater than 29.5 inches were considered to be adults. Total lengths at initial marking varied from 23.3 inches (a juvenile male) to 36.1 inches (an adult female); 54% were less than 29.5 inches, that is, were juveniles. Insufficient data were obtained 12 Tulane Studies in Zoology and Botany Mean Number Nutria Observed Ol 6am Time Figure 1. from recaptures to determine any growth increments. Recaptures were low. The nutria became trap-shy very rapidly, particularly if the traps were placed on land. One individual was recaptured 7 times, but 7 other nutria were retrapped less than 4 times each. Only observations on nutria with field markers were recorded. Of the 28 nutria marked, 5 were never seen again, 13 were seen only once, and 7 others were seen on 5 or more separate occasions. The low num- ber of sightings may have been due to the sloughing of field markers during the study. Thus, many of the animals originally marked may actually have been present but not recognized. Activity Nutria were active periodically through- out the day and night. Activity (based on the number of animals seen at any given time) peaked at noon, late afternoon, and late evening (Fig. 1). No observations were made of predawn activity. Chabreck (1962) found that activity was highest during the night, peaking just before sunrise. On the other hand, Norris (1967) reported greatest activity between 1800 hrs and sunset, when nutria were grazing. He observed only a 12noon Volsals Mean number of nutria observed at different hours of the day. limited amount of activity at night. In- creased activity during dawn hours was be- low the peak amount seen prior to sunset. Davis and Jenson (1960) reported that in England in cold weather, more nutria were seen in the daytime than usual, This did not appear to be the case at Riverside. The number of animals observed at any given time did not change with temperature, but the type of activity did change. At tem- peratures up to approximately 27° C, most nutria were seen lying in groups of three or four, sunning or sleeping. The areas chosen for this were generally the same (Fig. 2), but at ambient temperatures above 15.5° C, only those not in bright sunlight were used. At night the major activity changed from sleeping to eating; grooming was rarely observed. At temperatures above approximately 28° C, most nutria were seen eating, with frequent grooming; few were seen sleeping. When the temperature ex- ceeded 34° C, no animals were seen sunning. The nutria spent most of the time groom- ing, or swimming about, frequently stop- ping to eat for a short period before swim- ming again. Single nutria were rarely observed. Gen- erally, two Or more nutria were in the same area; the maximum number seen together G Seay G a B<—_RAFT 53) O Q Q O Ci Key: e- entrance cays G- grooming area i -restin First Road > ae 30 feet Figure 2. Map of one of the ponds studied showing locations of den entrances, grooming stations, and resting areas. Nutria were seen eating only to the left of the dotted line during the day, but were seen on both sides of the line at night. was 13. The only exception to this was in the case of several adult males, which were always seen alone. Movements Most nutria remained in the study area. Of animals marked, six were observed in the pond adjacent to the original trap site, and three were seen two ponds away (approxi- mately 300 yds) from the original trap site. Only one marked nutria, a large female, was seen further from the study area, at a dis- tance of 600 yds. Adams (1956) and Kays (1956) found that in southwestern Louisi- Nutria Behavior and Ecology 13 ana, daily cruising ranges of nutria rarely exceeded 200 yds and that nutria usually re- mained in one general area throughout their life. Norris (1967) observed that males ranged farther and most movements out of the home area were made to exploit tempo- rary food supplies. At Riverside, although the only marked animal found outside the study area was a female, a few males were observed wandering alone, at distances up to 200 yds from a pond. These animals were generally grazing on clover. Movements of nutria within the ponds were closely correlated with the location of duckweed (when the ponds were only par- tially covered) and with amounts of shade. A preference was always shown for areas of the ponds covered with duckweed, even if the animal was not feeding at the time. At high ambient temperatures, a preference was also shown for shady spots, both in ponds and on land. Nutria suffer from heat prostration if exposed to hot sun for any length of time (Evans, pers. comm.) and were rarely seen in brightly sunlit areas ex- cept when moving from one section of a pond to another or when sunning at ambient temperatures below 15.5° C. Burrows Nutria lived in large dens in the banks of ponds. A den usually had 3-5 above-water entrances and at least two below-water en- trances. The surface of the den was free of vegetation and slickened from the con- stant activity of household members. Each den complex covered an area 13-18 ft in width, extending into the bank to a depth of 10-13 ft; the height of the top of the den above water varied from 4 ft to 10 ft. Each underwater opening was 8-10 inches in diameter, but above-water entrances were larger (20-30 inches) and often opened into two or more tunnels extending back into the den. Water levels in the ponds dropped 21 inches during May and June, so that many underwater entrances were re- vealed. In addition to large dens, there were also many simple burrows with 1-2 openings into the bank close to water level. Ashbrook (1948) reported that each pair of nutria builds its own burrow. They work in and up until they are well above water level, then a space is cleared and grasses de- posited. As the family grows, the burrow 14 Tulane Studies in Zoology and Botany is enlarged, since the offspring from the original pair continue to live in the same burrow. Eventually the burrow, now a den, becomes the home of a large colony. As long as the water supply is sufficient and food plentiful, the colony will remain in the same locality. Observations of colonies at Riverside agreed with the findings of Ashbrook. Each colony lived in a separate den and spent most of the time in the “home” pond. The number of active dens per pond and the number of individuals in each colony varied. A typical colony consisted of 15-20 nutria and was composed of approximately equal numbers of adults and juveniles and of males and females. No nests of the type reported by Laurie (1946) were observed, nor did nutria build platforms for sunning, as reported by At- wood (1950). Instead of platforms, nutria utilized floating logs or exposed roots of willow trees located in the water near the pond’s edge. The size of the areas varied from 2-3 ft X 6-10 ft and were 2-17 inches above water. Sticks and bits of bark were carried onto these constantly. Most sunning was done in these areas; nutria usually lay in close contact, often partly on top of each other. These areas were also used to some extent as grooming areas (Fig. 2). Feeding Feeding appeared to be a major activity of nutria when they were not in the dens. Shady areas were usually chosen for feeding locations. The main diet consisted of duck- weed and white clover, but occasionally red maple leaves or bark of willow trees were also eaten. Swank and Petrides (1954) and Atwood (1950) listed many foods con- sumed by nutria, but neither included duck- weed or clover as a major component of the diet. Laurie (1946) noted that in England, nutria barked trees only in captivity. When eating a maple leaf or a piece of carrot, the nutria sat up on its hind feet and held the food item in its forepaws. Duck- weed was eaten with a shovelling motion, both forepaws scooping it into the mouth with a rapid, coordinated motion. Young nutria ate duckweed in the same manner as adults, except that motions were less well coordinated, much more rapid, and frequently resulted in the young animal get- Vol ting a face-full, rather than a mouthful, of the aquatics. Nutria did not form feeding platforms from non-eaten material, as mentioned by Swank and Petrides (1954), nor was food carried to specific feeding stations, as ob- served by Atwood (1950). Both previous studies were in a marshy habitat, where cat- tail, bulrush, and other coarse emergents were utilized to a great extent. At River- side, the ponds contained no such vegeta- tion; the only food carried was an occasional leaf or piece of bark. As reported by Atwood (1950), when- ever possible, nutria utilized floating ob- jects capable of supporting their weight, such as large branches, planks, or logs. If these were not available, shallow or exposed areas along banks were used. In either case, forepaws were left free to hold and manipu- late food. Occasionally a nutria would swim around slowly as it ate, paddling with the hind feet, using the tail as a rudder, and scooping duckweed with the forefeet. Although duckweed was the major com- ponent of the diet, clover was also eaten, mainly during hours of darkness. Nutria appeared to select primarily the clover blos- soms, leaving leaves and stems, as well as surrounding grasses, untouched. Grooming Periodically a feeding nutria would stop and groom itself thoroughly. Young ani- mals stopped to groom more often than adults, but they appeared to be less efficient at removing duckweed from the fur. Groom- ing always started with much scratching of the sides and stomach by the hind feet. Then the fur was groomed, using the forefeet. The grooming motion is back and forth, rapidly, with the apparent intention of roughing the fur rather than cleaning it; the forefeet were never licked to remove the dirt and duckweed gathered from the fur. Forefeet were used together whenever pos- sible, especially on the head, stomach, and chest. They were used out of phase or only one at a time when grooming the flanks, axilla, and back. Occasionally the teeth were also used, especially on the flanks. Most at- tention was paid to the abdomen, insides of the flanks, and genitalia. After these areas were thoroughly groomed, the head was cleaned and attention was given to the rest of the body. No. 1 Grooming motions appeared to be some- what stereotyped. Nutria were observed to continue grooming even in a pouring rain, just as if they were dry. A young nutria which was kept in captivity exhibited ex- actly the same pattern of grooming as ani- mals in the wild, even though it was neither wet nor covered with duckweed. There was never any evidence of mutual grooming; each individual groomed only itself. Dominance Relationships A simple hierarchy existed among nutria in the ponds. Within each social group, there was one alpha female and one alpha male. All other animals were equally sub- ordinate; that is, there was no dominance among subordinate members of the group. The alpha female exhibited aggressive be- havior towards the alpha male and all sub- ordinate females. The alpha male exhibited aggressive behavior towards all subordinate males and females. Only when the alpha fe- male was in estrus was she submissive to the alpha male. Dominance was expressed by the alpha animal chasing the subordinate individual. The dominant nutria would swim up very rapidly to a subordinate nutria, sometimes “mooing” several times either before or dur- ing the approach. Response in the sub- ordinate animal was generally flight. When the aggressive nutria approached within 5- 10 ft, the subordinate animal would dive with a loud splash, swim underwater for a considerable distance (20-100 ft), and re- emerge. The dominant animal usually turned 180°at the spot where the other had dived and would begin to feed or groom, paying no attention to any other nutria. During the course of the study, the domi- nant female in one of the social groups was displaced by a subordinate female. While the subordinate female was establishing her superiority, she continuously chased all other females and occasionally also chased the males. None of the nutria actually fought with her, although the original alpha female sometimes waited until the last moment before retreating. Dominance was estab- lished in 2-3 weeks, but the exact time is not known. Actual fighting was observed in only four instances. In each case, a male was per- sistently sexually aggressive toward the al- Nutria Behavior and Ecology 15 pha female when she was not in estrus. When the male ignored warning sounds (low-pitched “mooing”) from the female, she turned on him and they fought literally tooth-to-tooth. During this battle, the only sounds were the gnashing of teeth as the nutria lunged at one another with mouths open, pushing incisors against incisors. Al- though there appeared to be some attempt to bite the opponent, the main effort was directed at the opponent’s teeth. Most fight- ing took place with heads above water; oc- casionally the animals rolled underwater and re-emerged splashing as each tried to over- power the other by pushing its head under- water. In the four fights observed, after less than 3 minutes the male exhibited the typi- cal subordinate flight pattern described above. Whenever a male was rebuffed in his ef- forts, he spent much time grooming himself and marking the area. Marking consisted of dragging the genitalia (penis extruded) over logs, rocks and any other objects which might be encountered. The male also would raise his tail, back toward the object to be marked or just back up the bank, then uri- nate backwards in spurts carrying 2-3 ft behind him. Reproductive Behavior When the dominant female approached estrus, as judged by the increasingly swollen condition of the external genitalia, both she and the adult males spent more time grooming. Frequently they alternated areas where they sat to groom, so that first a male, then the female, sat on a certain log or part of the bank. Each spent much time sniffing spots where the other had been. But until the female was actually receptive, she continued to chase any aggressive males that came closer than 10-15 ft. Copulation was not observed but prob- ably took place in the woods. After much vocalization and grooming, the female would follow the male into the woods. Several minutes later when the pair emerged, the female was completely submissive to the male, following him in the pond and being chased when she approached nearer than 3-4 ft to him. There was more vocalization (“mawing”) between male and female dur- ing this time, always soft and in an alternat- ing chorus. 16 Tulane Studies in Zoology and Botan Receptivity is reported to last 2-3 days (Ashbrook, 1948). At the end of this time, the female again became the dominant mem- ber in the relationship. Care of the Young There was little care of the young by the mother. All nursing took place inside the den. Frequently when the mother sat in one of the den openings, young crawled over her head and back. Occasionally, the mother nuzzled them, but she never groomed them. When the young were out of the den, the mother did not stay near them. If one of the young called to her, she answered and some- times went to it. Always, when the mother came up to one of the young, she first touched noses with it, then either swam on by it or stopped to eat. If a young nutria swam up to the mother, again, touching of noses occurred first. Frequently a young nutria swam around the mother, under her chin, nuzzling her, before it stopped to feed. Other members of the group also looked after the young. As the young grew more adventurous and became better swimmers, they swam farther away from the den. The young always stayed together or with an- other member of the colony. Generally a young nutria swam out to a juvenile that was feeding nearby. After touching noses, the young nutria would begin to feed, facing away from the juvenile but still touching its body. Often the juvenile would swim away without the young noticing. The young nutria would continue to eat for a few minutes, then suddenly realize it was alone, and with a startled cry, head back to the den as quickly as possible. Juveniles often played with young nutria and generally spent much more time with them than did the mother. This may have been because the adult nutria were never observed to play or do much adventuring, whereas young nutria were continually chas- ing each other, splashing in the water, and investigating everything. There appeared to be no attempt to pro- tect young from predators. When the mother was frightened, she would run away and leave the young to fend for themselves. De- pending on how badly frightened she was, the mother might or might not return for Vol. 15 them. It was by frightening away the mother of a litter of five that I managed to catch a baby nutria, which was kept in captivity for four months. Predation In the United States, as in England, nutria still have few natural enemies. Man and severe winter weather take the largest toll of nutria in England (Newson, 1966). In the western United States, bobcats and coyotes prey to some extent on nutria, but in the southern United States, predatory birds are the only major predators besides man (Harris, 1956). In the study area at Riverside, garfish (Lepisosteus sp.), cotton- mouths (Ancistrodon piscitvorous), and red- shouldered hawks (Bwteo lineatus) were ob- served to take young nutria. Only one adult was found killed by another animal, pos- sibly by a bobcat (Lynx rufus). Although at Riverside nutria had few predators except man, they were extremely wary and could disappear very quickly. When mildly alarmed, a nutria would swim behind a log or a branch sticking out of the water. Only its head and back would be above the surface; this was the “watching” position. If a nutria was on land when frightened, it would dash for the pond, enter with a splash, dive, and when it re-emerged, assume the “watching” position described above. As long as the animal was alarmed, it remained completely motionless and in this position easily passed as a log. When a nutria was frightened to a greater degree, it would dive underwater, slapping the water with its hind feet as it did so. The resounding sound of this splash alerted any other nutria in the area and these also disappeared. Generally the dive lasted 30 sec, and the nutria again emerged at some distant, hidden spot. But occasionally when badly frightened, a nutria would remain submerged for a much greater length of time. Sooter (1943) reported a nutria re- mained underwater for 3 min 15 sec, but the longest dive observed at Riverside lasted less than 2 min. No attempt by one animal to protect an- other was ever observed. Each nutria had to fend for itself, including the young, as dis- cussed earlier. No. 1 Acknowledgments The writer wishes to thank Mr. James Evans, Director of the Nutria Research Sta- tion, U. S. Department of Interior, Houma, Louisiana, for his advice on trapping and marking methods and for his generous as- sistance in the loaning of materials used in the study. Thanks are also due Frank Thomas for aid in collection of data. The study was supported in part by an Environmental Bi- ology Traineeship under the direction of Dr. R. D. Suttkus at Tulane University, New Orleans, Louisiana. LITERATURE CITED Apams, W. H. 1956. Ecological studies of coastal marsh in the vicinity of Price Lake, Rockefeller Refuge, Cameron Parish, La. Un- published M. S. Thesis, La. St. Univ., Baton Rouge. 118 pp. AsHBROOK, FRANK G. 1948. Nutria grow in the United States. J. Wildl. Mgt., 12(1):87-95. Atwoop, EArt L. 1950. Life history studies of nutria or coypu in coastal Louisiana. J. Wildl. Met., 14(3):249-265. Cuasreck, Ropert H. 1962. Daily activity of nutria in Louisiana. J. Mamm., 43(3):337- 344, Nutria Behavior and Ecology 17 Davis, R. A., and A. G. JENson. 1960. A note on the distribution of the coypu (Myocastor coupusy in Great Britain. J. Anim. Ecol., 29: Harris, VAN T. 1956. The nutria as a wild fur mammal in Louisiana. Trans. 21st N. Am. Wildl. Conf., pp. 474-486. Kays, Caritos E. 1956. An ecological study with emphasis on nutria (Myocastor coypus ) in the vicinity of Price Lake, Rockefeller Refuge, Cameron Parish, La. Unpublished M. S. Thesis, La. St. Univ., Baton Rouge. 145 pp. LaurRIE, E. M. C. 1946. The coypu in Great Britain. J. Anim. Ecol., 15(1):22-34. Lowery, GrorcE H., Jr. 1943. Check-list of mammals of Louisiana and adjacent waters. Occ. Papers, Mus. Zool., La. St. Univ., No. 13> pp. 23-25% Newson, R. M. 1966. Reproduction in the feral coypu (Myocastor coypus). Comp. Biol. Reprod. Mamm., Symposia Zool. Soc. Lond., No. 15, Academic Press, London. Norris, J. D. 1967. The control of coypus (Myocastor coypus Molina) by cage trapping. J. Appl. Ecol., 4(1):167-190. SooTER, CLARENCE A. 1943. Nutria a good diver. J. Mamm., 24(4):503-504. Swank, WENDELL G., and GrEorGE A. PETRIDEs. 1954. Establishment and food habits of the nutria in Texas. Ecology, 35(2):172-176. October 16, 1968 NOTROPIS EDWARDRANEYI, A NEW CYPRINID FISH FROM THE ALABAMA AND TOMBIGBEE RIVER SYSTEMS AND A DISCUSSION OF RELATED SPECIES ROYAL D. SUTTKUS and GLENN H. CLEMMER Department of Biology, Tulane University New Orleans, Louisiana ABSTRACT Notropis edwardraneyi, an_ endemic eyprinid of the Alabama — Tombigbee River system, is described from 32,493 specimens. A close relative of N. blennius, it is restricted to the main channels and larger tributaries below the Fall Line where it is the dominant minnow. N. ed- wardraneyi is compared to N. blennius and N. potteri, and differs in having a larger eye and in pigmentation. Additional data are given for N. blennius and N. potteri, and their interrelationship is discussed. This new species of Notropis is of mod- erate size and presumably is a close relative of Notropis blennius (Girard), 1856, and Notropis pottert Hubbs and Bonham, 1951. Probably, Hubbs and Bonham’s (1951:103 ) reference to Notropis blennius in the Tom- bigbee River was based on specimens of this new form. The senior author collected the new species for the first time on 1 August 1957 from below Lock No. 2 on the Tombigbee River. Five years later Meredith May (Black- well), then a student of Dr. Herbert T. Bos- chung, requested identification of some speci- mens taken from the Cahaba River and these proved to be the same as the Tombigbee specimens. An intensive survey of the fishes of the main channel of the Alabama River was started in 1964 and the new form was common, if not the most abundant cyprinid as indicated by the numbers of specimens in the series listed below. The bulk of the specimens used in this study was obtained by the authors and Dr. Gerald E. Gunning. Some of the remainder were collected by the senior author with the aid of the Environmental Biology Training Program! students. We wish to acknowledge assistance in collecting by Dr. C. Robert Shoop, Dr. Sylvia Earle, and Mr. Armand Kurtis. We are indebted to Mr. Ben Stimpson for his generous hospitality. He provided us with a campsite near Choctaw Lake and use of a private boat ramp into the Alabama River for three summers, 1964-1966. This enabled us to obtain samples from the lower part of the Alabama River, a few miles above its junction with the Tombigbee River. During our stay on the Stimpson property, Mr. Elwood Overstreet gave us much valuable assistance. We are grateful to Mr. B. Frank Wilson, Sr., Executive Vice President of Peoples Bank in Selma, Alabama who made arrange- ments for our use of the old Crocheron house and property in Cahaba, Alabama as a camp- site during the summer of 1964. Mr. J. Campbell Banks of Columbus, Mis- sissippi made available for our use an area along the Tombigbee River for which we are most grateful. We would like to take this opportunity to extend our thanks to the biologists and en- forcement personnel of the Alabama Con- servation Department for their cooperation in all of our biological studies in the State of Alabama. Mr. Carlyle Suttle’s cooperation and help deserve special mention. For permission to examine material housed in the University of Alabama collection, we wish to thank Dr. Herbert T. Boschung. We 1Assistance was made through the National Institutes of Health Grants 3-T1-ES-27-02S1, 03S1, 0481, 0581, and WP-00082-04, 05. EDITORIAL COMMITTEE FOR THIS PAPER: Dr. GEORGE A. Moore, Professor Emeritus, Department of Zoology, Oklahoma State University, Stillwater, Oklahoma Dr. PHILIP W. SMITH, Taxonomist, Illinois Natural History Survey, Urbana, Illinois 18 No. 1 extend our gratitude to Dr. Ernest A. Lach- ner for making arrangements to examine type-material of Notropis blennius in 1958, and other materials during subsequent years; Drs. Reeve M. Bailey and Robert R. Miller for personal efforts to locate series of No- tropis blennius in the University of Michi- gan, Museum of Zoology collection for our use as comparative material and Dr. Neil H. Douglas of Northeast Louisiana State College for the loan of some fine series of Notropis blennius and Notropis potteri from the lower Mississippi River and Red River (in Louisiana) respectively. We wish to acknowledge our appreciation for the pen and ink illustrations by Miss Betsy Grover and for the photographs by Dr. Clyde Barbour. Notropis edwardraneyi new species Fluvial Shiner GEios 14. 7,9) The description is based on thousands of specimens (32,493), all of which were taken from the Alabama and Tombigbee drainages. Material. The holotype, Tulane University number 49485, an adult 57.6 mm in standard length, was collected from the Alabama River at Yellow Jacket Bar, River Mile 129.8 (U.S. Corps of Engineers Navigation Chart, 1958), 1.2 miles down-river from Holly Ferry crossing or 12.5 miles east of Pine Hill, Wilcox County, Alabama, on 8 March 1967, at 2245 to 2345 hours, RDS 4097, by R. D. Suttkus and G. E. Gunning. Paratypes: Taken with the holotype were 8,224 paratypes (22.1-56.0 mm in standard length) which were distributed as follows: TU 44028 (7,324 specimens); United States National Museum, USNM 202435 (100); Cornell University, CU 52941 (100); Uni- versity of Michigan, Museum of Zoology, UMMZ 187475 (100); Academy of Natural Sciences of Philadelphia, ANSP 109424 (100); Museum of Comparative Zoology, MCZ 45878 (100); Stanford University, SU 66551 (100); University of Kansas, Mu- seum of Natural History, KU 12674 (100); University of Alabama, UAIC 2791 (100); and Field Museum of Natural History, FMNH 74294 (100). Other paratopotypes (7,551) were taken 7-8 April 1966 at 2340- 2422 hr, (TU 40303, 1020: 19-62); 28 Notropis edwardraneyt Lo June 1966 at 2227-2308 hr, (TU 40925, 85: 24-52); 1 July 1966 at 1945-2100 hr, (TU 41400, 390: 23-59); 5 August 1966 at 1645-17 10° hr, (TU 41726. 16-12-29. 4 October 1966 at 2200-2235 hr, (TU 41618, 33: 19-40): 19 December 1966 at 235-2215 hr GW) 427.37, 5630") 19 Se 31 May 1967 at 2150-2230 hr, (TU 46802, 223: 24-55); 7 August 1967 at 2110-2215 hr, (TU 47361, 49: 18-46); and 26 Sep- tember 1967 at 2155-2250 hr, (TU 47924, LOS S=55))e Other paratypes (11,773 specimens), all taken from the main channel of the Ala- bama River from Watts Bar, 3.5 mi above Cahaba, River Mile 204.5, downstream to the Choctaw Bluff area, River Mile 45, are as follows: TU 33381 (1613, 27-65), Dal- las Co., Watts Bar, 29 June 1964, Royal D. Suttkus 3515; TU 35243 (236, 24-52). Dal- las Co., old ferry landing across river from Cahaba, 27-28 June 1964, RDS 3508; TU 35269 (314, 25-55), Dallas Co., old ferry landing across river from Cahaba, 28 June 1964, RDS 3512; TU 47822 (85, 16-47), Wilcox Co., Hurricane Island, River Mile 166.5, 19 Aug. 1967, RDS 4193; TU 47838 (75, 18-46), Wilcox Co., St. Johns Bar, River Mile 164.8, 19 Aug. 1967, RDS 4192: TU 47762 (143, 20-50), Wilcox ’Go., Lower Canton Bar, west bank, River Mile 156.7, 18 Aug. 1967, RDS 4190; TU 47781 (9, 16-43), Wilcox Co., Lower Canton Bar, east bank, River Mile 156.7, 18 Aug. 1967, RDS 4191; TU 47515 (306, 17-50), Wil- cox Co., Hobbs Bar, River Mile 149.5, 18 Aug. 1967, RDS 4189; TW 40295" (297, 16-54), 7 Apr. 1966, RDS 3857; TU 40900 (289, 24-54), 28 June 1966, RDS 3918: TU 41695 (83, 8-24), 5 Aug. 1966, RDS 3945, TU 41608 (284, 18-56), 4 Oct. 1966, RDS 401i LO) 42741 (4705 19250) et Dec. 1966, RDS 4065; TU 44011 (175, 17- 62), 8 Mar. 1967, RDS 4096; TU 46783 (202, 16-41), 31 May 1967, RDS 4143; TU 47346 (64, 21-51), 7 Aug. 19675200 47909 (152, 18-47), 26 Sept. 1967; RDS 4202, all from Wilcox Co., Evans Upper Bar, River Mile 135.8; TU 41711 (29, 10- 17), 5 Aug. 1966, RDS 3946; TU 47477 (89, 22-45), 9 Aug. 1967, RDS 4186, from Wilcox Co., Evans Lower Bar, River Mile 133. TU 46796 (395, 26-54), Wilcox Co., new bar above Yellow Jacket Bar, River Mile 130.1, 31 May 1967, RDS 4144; TU 20 Tulane Studies in Zoology and Botan) Voll Figures 1 and 2. 1 S.L. 54.8 mm, T.L. 71.0 mm. 2 (bottom) Notropis blennius: ISG) Silos 4-20. mms, Tele. 69-4) mm. 41732 (19, 15-22), 5 Aug. 1966; RDS 3949: TU 41632 (92, 20-46), 4 Oct. 1966, RDS 4013; TU 47374 (86, 19-42), 7 Aug. 1967, RDS 4179; TU 47936 (39, 13-58), 26 Sept. 1967, RDS 4204; all from Wilcox Co., Reeves Bar, River Mile 128.5. TU 40320 (491, 18-52), 8 Apt. 1966, RDS 3859; TU 40940 (226, 27-57), 28 June 1966, RDS 3920: TU 41745. (63, 14-46), 5 Aug. 1966, RDS 3950; TU 41639 (187, 18-47), 4 Oct. 1966, RDS 4014; TU 42746 (687, 21-56), 19 Dec. 1966, RDS 4067; TU 44029 (94, 21-53), 9 Mar. 1967, RDS 1098: TU 46819 (43, 15-42), 31 May 1967, RDS 4146; TU 47387 (168, 21-36), 7-8 Aug. 1967, RDS 4180; TU 47968 (230, 20-51), 27 Sept. 1967, RDS 4207; all from Wilcox Co., Tait Bar, River Mile 122.4; UW 40355 “(243. -22-61)). 8 Apr. 1966, RDS 3860; TU 40950 (109, 25-63), 29 June 1966, RDS 3921; TU 41755 (12, 20- 18), 5 Aug. 1966, RDS 3951; TU 41655 (119, 19740), 5 Oct 71966." RDS: 4015: (top) Notropis edwardraneyi: lateral view of a paratype (TU 44028) lateral view of topotype (TU TU 42759 (900, 16-54), 19-20 Dec. 1966, RDS A0683 SW 44045, SAGs 845) Mar. 1967, RDS 4099; TU 46830 (32, 29-38), 31 May - 1 June 1967, RDS 4147; TU 47396 (33, 20-44), 8 Aug. 1967, RDS 4181; TU 47980 (108, 15-48), 27 Sept. 1967, RDS 4208; all from Wilcox Co., Wil- cox Bar, River Mile 120.3. TU 41761 (9, 28-45), 5 Aug. 1967, RDS 4182: TU 41666 (147, 19-44), 5 Oct. 1966, RDS 4016: TU 47408) (45> 16-48). 6 Aug. 1967, RDS 4182: TU 47994 (65, 19- 46), 27 Sept. 1967, RDS 4209, all from Wilcox Co., Ohio Bar, River Mile 111.6; LUA 72 Giada 55)) 16 Aug. 1966, RDS 3953: TU 41670 (301, 16-41), 5 Oct. 1966, RDS 4017: TU 47420, (173, 18-49), 8 Aug. 1967, RDS 4183; TU 48012 (232, 17-48), 27 Sept. 1967, RDS 4210, all from Monroe Co., Stein Island, River Mile 107.5; TUAW C2 ie) Aue: 1966, RDS 3954; TU 47491 (24, 24-47), 10 Aug. 1967, RDS 4187, from Monroe Co., St. James No. 1 Notropis edwardraneyt 21 Figures 3-5. 3 (top) Notropis potteri (NLSC 5357) S.L. 46.7 mm, T.L. 60.1 mm. 4 No- tropis edwardraneyi (TU 44028) S.L. 48.9 mm, T.L. 62.3 mm. 5 Notropis blennius (TU 43167) S.L. 48.8 mm, T.L. 61.6 mm. i) bo Tulane Studies in Zoology and Botan) Woll IS Figures 6-8. t raneyi (TU 44028) S.L. 48.9 mm, T.L. 62.3 mm. 8 Notropis blennius (TU 43167) S.L. RL. 61.6 mm: Bar, River Mile 104: TU 41797 (4, 18-27), 5 Aug. 1966, RDS 3955; TU 47435 (229, 15-48), 8 Aug. 1967, RDS 4184, from Mon- roe Co., Bates Bar, River Mile 99; TU 47452 (134, 22-55), 8 Aug. 1967, RDS 4185, Monroe Co., Haines Island, River Mile 96; TU 41813 (10, 18-41), 6 Aug. 1966, RDS 3957; TU 47499 (16, 30-47), 17 Aug. 1967, RDS 4188, from Monroe Co., Silver Creek Bar, River Mile 87.7; TU 41823 (1, 22), 6 Aug. 1966, RDS 3958, Monroe Co., Mouth of Limestone Cr., River Mile 80.1; TU 35323 (28, 25-33), 2 July 1964, RDS 3519, Clarke Co., Choctaw Bluff, River Mile 45. Other material, 1297 specimens. Tombig- bee R.: TU 40497 (7, 40-48) 10 Apr. 1966, 6 Notropis potteri (NLSC 5357) S.L. 46.7 mm, T.L. 60.1 mm. 7 Notropis edward- 48.8 mm, RDS 3865, Miss., Monroe Co., 3.9 mi W of Amory, Hwy 278; TU 34614 (32, 35- 58), 9 Sept. 1964, RDS 3583; TU 37585 (25, 42-60), 6 May 1965, RDS 3663; TU 39420 (15, 26-43); TU 40162 (4, 25-43); TU 40217 125 20-45) NU 40489) (49: 26-60); TU 48758 (127, 26-53), all from Miss., Lowndes Co., 9.3 mi NW of Co- lumbus, 4 mi above Hwy 50; TU 48837 (30, 30-49), Miss., Lowndes Co., 9.3 mi NW of Columbus, 42 mi below Hwy 50; TU 48867 (758, 20-36), Ala., Pickens Co., 0.7 mi W of Pickensville; TU 16124 (6, 34-39), Ala. Choctaw Co., 5.5 mi SE of Pennington, below Lock No. 2; UAIC 2593 (145, 30-62), Pickens Co. at Vienna; UAIC 1470 (4, 31-46), Sumpter-Greene No. 87° Notropis edwardraneyt nN Oo TENNESSEE ALABAMA MISSISSIPP | GEORGIA 325 1020 _30__40 mies ee 10 0 10 20 3040 50 Kilometers | 86° 85° Figure 9. Distribution of collection sites of Notropis edwardraneyi. Triangle symbol indicates type-locality. Cos., Noxube Cr., 1 mi N of mouth; UAIC 2475 (12, 34-42), Clarke Co., 300 yds below Hwy 43 at Jackson. Black Warrior River: TU 43332 (14, 23-34), 22 Dec. 1966, Glenn H. Clemmer 518, Ala., Tuscaloosa Co., 0.5 mi S of North- port, below dam; UAIC 889 (346, 29-51); UAIC 1056 (217, 31-62); UAIC 1060 C1I56,531-46)- WAIC 1264 (2. 42-43); WAIG 1502 (235.720-50) | UAIG 1594nG;, 35-44) UAIG 1595 1(77, 23-54); UAIC 1608 (36, 28-57); UAIC 1647 (184, 29- 56); UAIC 1648 (58, 29-54); UAIC 1694 G93. (28255) UAIC W719 420; 30-5207 WAIG 1935) (287, 26-54 = UAIG, 2052 (ZiGy 25-54 )2 WAIC, 2033) (OZ, Sze UAIC 2515 (678, 25-55), all from Tusca- loosa Co., below Oliver Lock and Dam at 24 Tulane Studies in Zoology and Botan Vols E | See x & e Z w 3 _ = a o- N.blennius S 2b e@- N.potteri l L jae Ne fie L ae es) 6 8 10 12 14 16 18 20 22 HEAD LENGTH (mm) Figure 10. Relation of orbit length to head length in Notropis blennius (TU 22422, TU 43167, TU 43174) and Notropis potteri (TU 4936, TU 42395). Tuscaloosa: WAIG 1570055. 17-47) salus- caloosa Co., bar above mouth of Big Sandy Creek. Cahaba River: TU 35134 (127, 39-45), 26 June 1964, RDS 3505; TU 37698 (13, 45-54), 11 May 1965, RDS 3671, all from Perry Co., 1 mi W of Sprott, off Hwy 14. TU 34055 (14, 39-46), Bibb Co., 8 mi N of Centreville, Hwy 27; TU 35090 (5, 40- 44), Bibb Co., 2.1 mi N of Centreville, off Hwy 5; TU) 29893) (3; 5051), TU 30099 (9940-50) UAIC 96251, 37-52) Ee WAIe 1437 (24, 36-47), 1 mi W of Sprott. Tallapoosa R.: UAIC 1281 (3, 40-49), Elmore Co., 1.5 mi S of Thurlow Dam, Tal- lassee; UAIC 1514 (13, 40-47), Macon Co., Uphapee Cr. NW Tuskegee on Hwy 199; UAIC 1516 (29, 40-62), Macon Co., Up- hapee Cr., 0.2 mi N Franklin; UAIC 1368 (7, 41-46), Macon Co., Cubahutchee Cr. on Interstate 85, E of Montgomery; UAIC 1234 (38, 36-51), Montgomery-Macon cos., Line Cr. on Interstate 85, E of Montgomery; UAIC 1232 (14, 39-48), Montgomery Co., trib. on Wells Ferry Rd. near Mt. Meigs Station. Other tributaries: TU 40457 (15, 28-37), 9 Apr. 1966, RDS 3868, Dallas-Autauga cos., Big Mulberry Cr. 10.1 mi E of Selma, Hwy 14; TU 35066 (21, 32-46), Dallas Co., Oakmulgee Cr., 7.2 mi NW of Selma, Hwy 14; TU 35193 (6, 31-42); and TU 35187 (4, 32-39), Dallas Co., Pine Flat Cr. (Six-Mile Cr.) 6 mi S of Selma, Hwy Ane EUs 32455) (Cl. 30).) Wilcox-Co* Bear Cr. 3.1 mi NW of Lower Peach Tree, Hwy 1 WAIC 2403 %(245; “19=55), Dallas: Lowndes Cos., Old Town Cr., Y% mi NW Benton on US 80; UAIC 2366 (12, 24-36), Dallas Co., Cedar Cr., at Hwy 41, 4 mi S of Jct with Hwy 28; UAIC 2392 (11, 25- 39), Dallas Co., Bogue Chitto Cr., about 5 mi N of mouth. The following material was used for com- parison with the new species: Notropis blennius, USNM 67, lectotype (so desig- nated by Suttkus, 1958: 308), 55.9 mm in standard length, Arkansas River near Ft. Smith, collected by Dr. Shumard; USNM 171791 (syntypes, recataloged as paratypes on 27 June 1958) 5 specimens 50.5-64.0 mm in standard length (see Hubbs and Bonham, 1951: 103): TU 43167 (332: 13-60), Ark., Yell Co., Arkansas River at Dardanelle, Ark. Hwy 7 crossing, 1 February 1967, RDS 4080, Suttkus and Kenneth Relyea; TU 43174 (579: 27-65), same lo- cality as for RDS 4080, 2 February 1967, RDS 4081; TU 224221365: 17-50); Ark, Arkansas Co., Arkansas River at Pendleton’s Ferry, Ark. Hwy 1 crossing, 23 October 1959, RDS 2851, Suttkus, Myrna Andersson, Bangalore I. Sundararaj; UMMZ 180500 (16: 28-34), Manitoba, Seine River, T10, R3E, $34, 18 August 1955, J. J. Keleher; UMMZ ACC. 1947: XI: 28 (113: 31-58) Iowa, Muscatine Co., Mississippi River at Fairport, Miss. River Survey, 15 June 1946; No. 1 Uy 19296 (20, 17-57), lad, Posey ‘Co, Wabash River at old dam, about 4 mi SW of New Harmony, 28 August 1958, RDS 2744, Suttkus and Bruce B. Collette; TU 19356 (12, 22-61), Ind. Posey Co., Wa- bash River on west side of old dam, 2.5 mi S of New Harmony, RDS 2746, Suttkus, Collette, Dowell, Pugh; TU 43053 (15, 23-48), Ark., Crawford Co., Arkansas River 0.25 mi downstream from old Fort Smith bridge, 25 January 1967, KR-Ark 3, Relyea and Reimer; TU 48248 (856: 18-60) La., Madison Parish, Mississippi River across from Vicksburg, Miss., at U. S. Hwy 80 bridge, 11 October 1967, RDS 4236, Suttkus and) Glemmer, TU 47644 (= 53); La; Rapides Parish, Red River at River Mile 100 (U.S. Corps of Engineers Navigational Chart, 1958) 25 August 1967, RDS 4200, Suttkus and Clemmer; NLSC 5356 (105: 37-57), La., Concordia Parish, Mississippi River at Natchez, | to 4 miles north, 25-26 July 1966, Louisiana Wildlife and Fisheries Commission; Northeast Louisiana State Col- lege 5441 (19: 37-58), La., Concordia Par- ish, Mississippi River, 1 to 4 miles north of NIESG@ 5374 (47-30-55), La. East Carroll Natchez, Miss., 25-26 July 1966, LWFC; Parish, Mississippi River at Lake Providence, 27 \uly, 1966; LWEC; NUSC 5506 (41: 27-60), La., East Carroll Parish, Mississippi River at Lake Providence, 27 July 1966, EEG: INESG@ 53678 47— 22-63), Ea. East Carroll Parish, Mississippi River at Lake Providence, 27 July 1966, LWFC; UAIC 1830 (99, 24-59), Mo., Mississippi Co., T22N, RIZE, Sec. 6, Mississippi River 17 mi SE East Prairie, 14 August 1963, W. L. Pflieger and Robert Hentges; NLSC 5654 (43: 23-43), Ark., Jefferson Co., Arkansas River at Pine Bluff; NLSC 5574 (40: 20- 46), Ark., Jefferson Co., Arkansas River at Pine Bluff; TU 19022 (2, 16-22), Ky., Jefferson Co., Ohio R about opposite 38th St. in Louisville, just below K and I RR. bridge; TU 2242 (3, 43-46), Okla., Noble @o., Skeleton Cr; TU 13831 (6, 26-54), Okla., Noble Co., Salt Fork, 7 mi S of Ponca etn 155945— 639. 27-54), 2Okla., Wagoner Co., Verdigris R. at Okay bridge, TIGN, R19E, S19, TU 10250 (7, 29-43); MU 15637 01 20-52) 2.0) 14877- (143, 29-49); TU 15611 (90, 18-49); TU 15471 (9, 27-50); TU 39666 (3, 34-41); all from Ark., Pope and Yell cos. Arkansas R. at Notropis edwardraney1 25 Dardanelle, Hwy 7; TU 43164 (113, 17- 45), Ark., Arkansas Co., Arkansas R. at Pendleton Ferry crossing, 11.3 mi NE of Dumas, Hwy 1; TU 48267 (39, 16-61), La., West Baton Rouge Par., Mississippi R., 14, mi above US Hwy 190 bridge at Baton Rouge along west bank; TU 16786 (4, 24-43), La., St. Charles Par., Mississippi R. flood pools in front of Bonnet Carre Spill- way dam at Norco. Notropis pottert, TU 42251 (71: 20-58), Red River at River Mile 97, 6 mi down- stream from Alexandria, 9 November 1966, RDS 4039, TU 42304 (46: 16-51), Red River at River Mile 86, 17 mi downstream from Alexandria at Ryland Revetment, 9 November 1966, RDS 4043, TU 42395 (130: 17-58), Red River at River Mile 96.5, half mile downstream from Hudson Revetment, 23 November 1966, RDS 4049, TU 47536 (666: 13-52), Red River at River Mile 81.2, 25 August 1967, RDS 4194, TU 47649 (529, 11-48), Red River at River Mile 100, 25 August 1967, RDS 4200, Rapides Parish, La., Suttkus and Clem- mer, NLSC 5352 (483: 18-68), La., Bossier Parish, Red River 11 miles north of Bossier City, 8-12 August 1966, K. Burnside and J; Brantley, INIESC 5357 6050/96) eae Red River Parish, Red River at Coushatta, 8-12 August 1966, LWFC; NLSC 3700 (Gaik2 24551), an sRapides sParisha hed River north of Alexandria, 6 April 1966, EWEC? INESG 3839) (S51: 22-50). Lae Rapides Parish, Red River above Alexandria, 5 April 1966, LWFC; NLSC 7297 (46: 15-36), La., Natchitoches Parish, Red River 15 miles SE of Natchitoches; NLSC 7409 (54: 18-47), La., Red River Parish, Red River at Coushatta; NLSC 124 (36: 14- 32), La., Concordia Parish, Red River at Acme; NLSC 5442 (38: 26-46), La., Rapides Parish, Red River above Alexandria; NLSC 3393 (570: 18-45), La., Bossier Parish, Red River at Beene Place, Sec 7- TISNERIZW; NES© 5351) (6052 18-538)5 La., Red River Parish, Red River at Cou- shatta; NLSC 4966 (50: 23-60), La. Red River Parish, Red River at Coushatta, 1-2 miles north of bridge; NLSC 3291 (257: 18-58), La. Bossier Parish, Red River at Beene Place, 3-5 miles N of Bossier City; NESG AG07, (2622 12265). Tas Bossier Parish, Red River at Beene Place, Hwy 3, Sec 7-TI8N-R13W; NLSC 3840 (627: 23- Vol. 15 Tulane Studies in Zoology and Botany (¥9Z) (L9G) (Z9Z) 9LG-9SZ 18G-SSG 99G GLZ-CSS ILZ ysus, proH (ZIT) (901) (SOL) €SI-COl TII-101 Cll LOT-OOT IOI idop ysvof (GZS ) (913) (11d) 6GC-C1Z I€t-S6I LOS 6I1Z-86I LIG qysusz spuNnped yepney (9ST) (SST) (ZEST) 69I-9FT COI-LFT = P9OI-GFI igi api Apog (€€Z) (61Z) (¥GZ) p ISZ-GIZ SEC-10G PES GS-1 PCG yidop Apog (IGS) (€Te) (OE ) SZE-FOE PEE-LES POL OTE-06Z L6G aseq [Bpneo 0} UISLIO [vuy (60S ) (SOS) (OTS) GGG-G8P CIS-I6F 10¢ PCS-L6F 9I¢ Yasue otApedoid (608 ) (LOE) (662) 6IE-O0E 0ZE-G6S STE GIS-GLS 66G yNd1990 0} ULBIIO [es10C] (80 ) (¢0¢) (LOS) €CS-LOF OZS-Z8F QTc CGS-I6F CBr oseq [Bpnvo O} UISLIO [Vs..od (OTS) (LTS) (FIS) GGS-10S 8ZS-POS 60S CES-16F PGS Yysus] [PSIOpold (SSF) (CFS) (6'GS) « L'8S-€'SP #6 89-6 1S 6°SG #6 GS-0 TS OLS (Wu) YysUe] prepurys VMOT ‘IddIssissify sesueyly ‘sesuvy.ly vuueqgeypy “eueqeyy ayeYS puvw Woysdg AY sodAjodo J, 9d4}0}09'] sodajqvirgd adAjoloy] 7 ; Sa: IX'LP6T “OOV ZWD LOTEP AL L9 WNSO Sc0PF OL GSP6F OL JOQUINN, SOTRICS SnIUUd]q snuua}q Maups pion po sotoads “UOHRLIVA JO osuvI (sosoyyueied UL) MOfOG USAID st on[vA Uvosyy ‘YWSUeT prepurys JO syYypursnoyy ut s1doujoN JO syUoWoAMsvI]y [ aTav ardraneyt Notropis edu No. suoumoods ud} UO poseq . aqo] IOL19}sod JO pus 0} poANsvoUt - 9qO[ IOLIo} UB JO pus O} Ppoerusvout - Yysus] ULF OLApod Yysuo] ULF [LLOPoq -4asua] possaidop ‘uly [euy {yysues[ possoidep “uly [vs.0c] UPI [eyWqstoqus yysuey Mel teddy) Yeu] UIQ Yyysuey yous AYsof JSVOT “YAPUW [LPWGIO.1oJUT YPM pro]{ yydep prol{ (OST) (6FT) (9ST) PSI-SPI 9SI-cPrl OST 89I-SFI VST (68T) (961) (F6T) €OS-08 I 60G-08 I 161 LOG-68 I ssl (Z9T) (C91) (F9T) PLI-€ST 69I-9ST 191 SLI-9ST ect (91Z) (€16) (966) JEC-CVNG ScC—P0G PIG 6EC—-SIG OIG (66 (63) (0€ ) VEVG CE-VG LG 6E-LG 66 (¥8) (€8) (LL) I6~LL 06-92 6 C8—-L9 O08 (g9) (#9) (98) OL-09 L9—09 GL 16-62 06 (8L) (#8) (G8) C8sZ C6-8L 6L 68-91 L8 (86) (66) (68) LOT—06 POI-¥6 €6 96-E8 6 (OFT) (TST) (GST) ccl-rel 9SI-GPrI eri C9I-SFI PST (OLT) (SOT) (891) 9LI-G9T GLI-Sal S9T 9LI-c9T sol eMoOy ‘IddIssIssly sesuRy.ly “Sesuvyly vuregryy “euUeqelyy sodAjodo J, 9d4}0}09'T sodaAyeieg adAqopoy] 8G:IX‘L¥6I “DOV ZNWD. LOTEP OL L9 WNSf SCOP OL csr6r OL sniuua]q snuua]q thaupip.onpa (ponunuoy) ~ ATAVvy, AVIS puR WasAG IA IaG(UINNY BOpezv, soweds 28 Tulane Studies in Zoology and Botany Vol. 15 TABLE 2 Measurements of Notropis expressed in thousandths. Mean value is given below (in parentheses) range of variation. Species ical edwardraneyi- “blennius potteri . ee River System Alabama Arkansas Red Catalog Numbers TU 44028 TU 43167 NLSC 3700, 5352, 5357 Number of Specimens 50 50 50 Standard length (mm ) 44.0-55.5 45.9-60.0 41.8—-57.1 (50.5 ) (52.4) (47.8) Head length/Standard length 251-276 207-294 259-305 (263) (268 ) ( 285) Interorbital/Head length 312-372 347-413 324—400 (336) (374) (354) Snout/Head length 267-333 282-342 281-333 (307 ) (312) (307 ) Orbit/Head length 308-363 229-273 199-248 (333) (249) (223) Upper jaw/ Head length 267-323 285-348 320-382 (299) (320) (347 ) Postorbital/ Head length 369-436 44]-482 467-530 (402 ) (466 ) (498 ) Orbit /Interorbital 886-1102 589-755 558-714 (992 ) ( 667 ) (631) Orbit/Snout 978-1250 706-943 643-833 ( 1086 ) (799 ) (27) Orbit/Upper jaw 1023-1228 674-868 545-743 (1114) (781) (644 ) Orbit /Postorbital 727-907 478-579 381-508 (828 ) (535 ) (448 ) 51), La., Rapides Parish, Red River above Natchitoches Par., Red R. 5 mi N of Alexandria; Other material all from Red R., Rapides Parish, La. collected by the authors with Dr. Gerald E. Gunning and Jayson S. Suttkus: TU 42229 (61: 17-50), TU 47669 (100: 12-42), River Mile 105, 2 mi above Alexandria; TU 42244 (63: 17- 42), TU 45178 (22: 17-30), River Mile 100, 3 mi below Alexandria; TU 45172 (292023) peels 47628" 1225 15-40); River Miles 97) TU) 42267.) (51, 2053). TU 45163 (14: 19-30), River Mile 96.5, half mile below Hudson Revetment; TU 42277 (30: 20-49), TU 42374 (274: 15- AS), EO 4G. Clil=a20-32)) =. kU) 4/608 (86: 13-51), River Mile 94 at Grand Bend; EW 42287 ((202917-28)). EU 42365. 20: 15-54) TU 47584 9(220; 13-47). River Mile 90; TU 42345 (73: 16-59), TU 45140 (6: 21-35), TU 47571 (678: 13-36), River Mile 86, opposite Ryland Board Revetment; TU 42329 (14: 19-46), River Mile 82.7, 1 mi NE of Magda, 20.3 mi below Alex- andria; TU 42318 (32: 19-53), TU 45135 (28: 21-35), River Mile 81.5, at Roxana Revetment; TU 47556 (272: 12-40), River Mile 78; also TU 13403 (194: 20-57), La., Natchitoches; TU 13362 (124: 22-43), La., Rapides Par., Red R. at Alexandria, Hwy (Ae TO 3027 = S146) la Concorara Par., Red R. at Acme. Notropis potteri material from the Brazos R. system in Texas: TU 4936 (70, 28-86), Bosque Co., Brazos R. where trib. enters about 4 mi S Whitney Dam, 8 Apr. 1952, RDS 2278, Suttkus and Anderson; TU 20235 (5: 56-69), Stonewall Co., Double Mountain Fork, 11.3. mi S Aspermont, Hwy 83, 30 May 1959, RDS 2780, Suttkus, Negus, Shoop, Gould; TU 20257 (3: 54- 67), Wichita Co., Wichita R., 0.7 mi N Kadane Corner, Hwy 25, 29 May 1959, RDS 2778, Suttkus, Negus, Shoop, Gould: TU 4994 (11: 35-54), Bosque Co., Trib. 4.0 mi S Whitney Dam, 8 Apr. 1952, RDS 2277, Suttkus and Anderson; TU 35613 (6: 14-48), Brazos Co., Brazos R. at Mussel Shoals, 17 July 1964, RDS 3540, Env. Biol. Glass: TU 5013 (i 52), Williamson’ Gos San Gabriel R., 6.5 mi E Georgetown, 18 Apr. 1952, RDS 2282, Suttkus and Ander- son. Methods of counting and measuring de- No. 1 Notropis edwardraneyt 29 TABLE 3 Total number of vertebrae in three species of Notropis. Species eet Drainage 34 35 36 37 38 N X edwardraneyi Alabama R., Ala. 18 105 8 131 34.9 blennius Seine R., Manitoba 4 12 16 36.7 Mississippi R., lowa 6 9 15 36.6 Wabash R., Ohio R., Ind. 12 10 9} 36.4 Arkansas R., Ark. 1 4 47 26 78 36.2 Mississippi R., La. 11 6 1 18 36.4 Totals 1 4 80 63 ] 149 36.4 potteri Brazos R., Tex. 4 17 2) 23 35.9 Red R., La. al! 66 2, 89 36.0 Totals 15 83 14 112 36.0 edwardraneyi (TU 34614, 35269, 35323, 37698, 40320, 40497 ) blennius 43167; TU 48248 ) potteri scribed by Hubbs and Lagler (1958) were followed except for number of scale rows before the dorsal. The counts given repre- sent the number of rows crossing an imag- inary diagonal between the origin of the dorsal fin and the pectoral girdle, just above the anteriormost lateral line scale. Diagnosis. A moderately stocky shiner of medium length (largest specimen of 32,493 examined was 65 mm in standard length) ; usually 34 lateral line scales, often 33, occa- sionally 35; number of vertebrae usually 35, occasionally 34, rarely 36; eye very large, supralateral in position; diameter of orbit greater than length of upper jaw, about one- third length of head, nearly equal to fleshy interorbital distance (sometimes greater but more often slightly less than interorbital distance) and averages more than three- fourths of the postorbital distance. Description. Notropis edwardraneyi is similar to Notropis blennius in many proportions (Table 1) but differs pri- marily in the size of the orbit (Tables ieandie2eubios Al 2s 45) 7 eand 8). In profile the snout is blunter in edward- raneyi than in blennius and the eyes are more dorsal in position. The position and large size of the eye is reflected in the proportionally smaller interorbital distance. As shown in Tables 1 and 2, the relation- ship of the length of head to the standard (UMMZ 180500; UMMZ Acc. 1947:X1:28; TU 19296, 19356; TU 22422, 43053, (TU 4936; TU 42251, 42304, 47536, 47649 ) length is almost identical in edwardraneyi and blennius, but when the relationship of orbit to standard length is considered there is no overlap. Thus, in addition to the effect on size of interorbital distance, the large eye in edwardraneyi averages greater than three-fourths of the postorbital distance, whereas the diameter of orbit averages slightly more than half the postorbital dis- tance in blennius. The orbit is nearly always greater in diameter than length of snout in edwardraneyi and never as great (average about three-fourths snout length) in blen- nius. The mouth is smaller in edwardraneyt and the diameter of orbit is always greater than length of upper jaw. Notropis blennius has the opposite combination, 1e., larger mouth and smaller orbit that is never as great as length of upper jaw (about three- fourths ) . Notropis edwardraneyi typically has a pharyngeal tooth formula of 2, 4-4, 2. An examination of 50 sets revealed 47 spect- mens with 2, 4-4, 2; two with 1, 4-4, 1 and one with 1, 4-4, 2. All specimens used were taken from TU 33381. For comparison the same number of Notropis blennins (TU 22422, Arkansas River) and Notropis pot- teri (NLSC 3839, Red River) were exam- ined. The examination of pharyngeal teeth of blennius resulted in 46 specimens with 2, 4-4, 2; two with 1, 4-4, 2; one with 1, 30 Tulane Studies in Zoology and Botany Vol. 15 TABLE 4 Number of lateral line scales for three species of Notropis. Species and Drainage 32.) $3 784, 985 036, 37 (38. S39" 40 41 aN edwardraneyi Tombigbee R. 8 13 Ht ssh Black Warrior R. 5 9 14 33.6 Big Mulberry Cr. Be seal 1 see) Cahaba R. Olan wali 2 20 33.9 Alabama R.., Watts Bar 18 5 SO) Sss9) Hobbs Bar 2) 16 2 20 34.0 Yellow Jacket Bar Coli 7 30 634.0 Choctaw Bluff Bar Dy ily 12 Bi Ss} Totals 2, Sl alt 7 181 33.8 blennius Seine R., Manitoba 8 U 1 1G Be Mississippi R., Iowa ie) (ato) 5 0 0 0 1 Jal) 5.7 Mississippi R., Missouri wy XY) PAG) 2 50) S016 Arkansas R., Arkansas QD DY Sil 3 2 0) Sto. 7/ Mississippi R., Lake Providence 1426" 10 0 i 51 36.0 Vicksburg 2, 1S 323 6 1 50) 8}5357/ Natchez 4 17 19 10 50) B7/ Red R., Louisiana il 1 Totals 10 138 197 i 4 iL 0 Ih ehsts} Bis i/ potteri Red R., Bossier City, La. 4 14 2, 20 34.9 Coushatta, La. 9 4 2 ey B}5945) Alexandria, La. 3 7 5 sy Ball Totals 7 30 11 2 Oy Binge 4-4. 1; and one with 3, 4-4, 2. Of the 50 specimens of potteri, 48 had 2, 4-4, 2 pharyngeal teeth and two had 1, 4-4, 2 pharyngeal teeth. The total number of verte- brae, including Weberian apparatus (4) and urostylar vertebra (1), was as follows: 18 specimens with 34, 105 with 35, and eight with 36 vertebrae (Table 3). Except on the belly, the scales are rather large over most of the body, including the nape. The head and breast are naked. Although we found that most of the specimens of blen- nius had complete squamation of breast (Hubbs and Bonham, 1951: 105), we ob- served that some were incomplete either by having the extreme anterior portion naked or by having only a few scattered, partially imbedded, scales on the breast. The scales on the belly are progressively smaller an- teriorly and some of the anteriormost are imbedded. The lateral line is complete and slightly decurved anteriorly. There are 32 to 35 scales (some slightly elevated) along the lateral line (modally 34, frequently 33) (Table 4), 20 to 26 (typically 22 to 25) rows around body before dorsal and pelvic fins (Table 5), scales are not crowded be- fore dorsal, 13 to 15 rows before it on a diagonal to opercle (Table 6). The number of scale rows around the caudal peduncle was determined for 181 specimens. Nearly all (177) had five rows above and five rows below the lateral line (5-2-5). One specimen had five above and only four below (5-2-4), two specimens had 6-2-5, one had 7-2-5. The same 181 specimens had the following fin ray counts: 180 speci- mens had 8 principal dorsal fin rays, one had 7 rays; 178 had 7 principal anal fin rays, three had 8 rays; 175 had 19 principal caudal fin rays, two had 20 rays, three 18 rays and one had 16 principal caudal fin rays; 1 specimen had 12 left pectoral rays, five had 13 left pectoral rays, 57 had 14 rays, 93 had 15 rays, and 25 had 16 rays (Table 7); four specimens had 13 right pectoral fin rays, 62 had 14 rays, 92 had 15 rays, 22 had 16 rays, and one had 17 No. 1 Notropis edwardraneyt 1 TABLE 5 Body circumference scale row counts for three species of Notropis. Species and Drainage 0; SL Sh BES aE hy ONG} 7} 9) YO) BNL BO IN X edwardraneyi 4 Tombigbee R. 1 One oe BAL ppl Black Warrior R. too4& Mil L428 Big Mulberry Cr. I 8} iss DET Cahaba R. S Jul 4! 20 22:9 Alabama R., Watts Bar o> AL) 330) Ba Hobbs Bar Sia. © ai(s aapll 20m 2022: Yellow Jacket Bar iL oe Je 1@ & it 30 23.4 Choctaw Bluff Bar lL 2 » Is ig 8 Slee24.2 Totals lest D4 Come Om na Sie 3235 blennius Seine R., Manitoba 3 8 ®@ 2 iG} 213} Mississippi R., lowa 1 5) 489) IG) ISO)? yw) Ome 2a: Mississippi R., Missouri Wey ML ak GH HOMNZ5 3 Arkansas R., Arkansas il les 8) SI) 60 25.4 Mississippi R., Lake Providence 1@ I tk 8 i@ © © @ i Sil BE Vicksburg 1 OS al 4 3 1 bO Fy 25:0 Natchez Dt) of} AD Tk OL 50 26.0 Red R., Louisiana 1 ] Totals ORNS Oe Om OMNA Gl erOr, S 1 0 38S) 25:4 potteri Red R., Bossier City, Louisiana Je ee ONe SOT, ohne, 20 27.8 Coushatta, Louisiana 3 Bey Wes 7/355 Alexandria, Louisiana Ie eS Os ee IL Ii ATi A! Totals i § ihh 20) 1G 8 50 27.6 right pectoral rays (Table 8). The typical same rays. The double row of tubercles is number of pelvic fin rays was 8 on either side but two specimens had 9 on each side and five specimens had only 7 on each side (Table 9). The mouth is terminal to slightly sub- terminal, upper lip varies from slightly ex- ceeding snout to slightly subequal. The lower lip is subequal to or included by upper lip; maxilla is slightly curved. The mouth is oblique and does not quite reach to the anterior border of orbit. Anterior lobe of dorsal fin is longest in depressed po- sition, its posterior margin straight or slightly falcate. The posterior lobe of the moderately falcate anal fin is typically longer than anterior lobe in the depressed position of fin. Nuptial tubercles are well developed on pectoral fins only in adult males. There is a single row along the upper surface of the anterior ray. Proximally there is a single row on the second to the seventh or eighth ray but a double row more distally on these separated at the fork of each ray, one row continues out each of the two branches. Both males and females have fine tubercles on the head, more on the former. Coloration. As is typical with many large river forms Notropis edwardraneyi is a silvery shiner without any striking colors. The entire ventral surface of head and body is devoid of pigment except for a few deeply imbedded melanophores along the base of the anal fin and along the ventral portion of caudal peduncle. The dorsal and dorsolateral portion of body is pig- mented (Fig. 4) with small chromatophores. These chromatophores are concentrated on the central portion of most scales and the concentrations have a well defined posterior border which is submarginal to the posterior margin of the scales. Thus many scales in the dorsolateral area of the body have a pronounced light margin as illustrated in Figure 4. The lower margin of the above described area is bordered by a band of Uo ie) Tulane Studies in Zoology and Botany Vol. 15 TABLE 6 Predorsal scale row counts between dorsal fin and opercle in three species of Notropis. Species and Range 13 14 15 16 7 18 19 20 N Xx edwardraneyi Tombigbee R. ] 12 § Dil 14.3 Black Warrior R. 1 8 5 A ele Big Mulberry Cr. fe) 6 15 144 Cahaba R. 14 6 D0 ALS Alabama R., Watts Bar 18 12 30 14.4 Hobbs Bar 12 8 20 14.4 Yellow Jacket Bar 1 16 13 30 »=—- 14.4 Choctaw Bluff Bar 3 iil 7 31 14.1 Totals 6 110 65 181 14.3 blennius Seine R., Manitoba 5 10 0 1 16 15.8 Mississippi R., lowa 1 44 44 16 3 2 OR eal5:S Mississippi R., Missouri i 20 20 6 3 50) «15:8 Arkansas R., Arkansas 33 24 iy) ] 0) 15159 Mississippi R., Lake Providence 24 18 6 9) 0 1 Dill 15.8 Vicksburg il 20 22 4 2 1 50 15.8 Natchez ] 30 15 2, 2 50 55) Red R., Louisiana ] Totals AD aliGe alias 36 14 4 Th hats} 7 potteri Red R., Bossier City, Louisiana 1 10 6 iy) 1 20 ISIG Coushatta, Louisiana 1 3 9 2 15 15.8 Alexandria, Louisiana 3 9 2) 1 ey sy, IL Totals 5 29) 7/ 5 1 50 ~)=—s 15.5 larger chromatophores. This band extends posteriorly along the caudal peduncle to the base of the caudal fin. There is a faint dis- junct basicaudal, wedge-shaped spot in some specimens. Many juveniles and young have such a spot. The area just above and below the lateral line sensory pores is sparsely pigmented. Some specimens contained in the University of Alabama collection from Black Warrior River system are rather densely pigmented. The moderate-sized melanophores are scattered (Fig. 4) and typically are not concentrated immediately above and below the lateral line pores nor are they aligned with scale margins. There is a fine (sometimes only a single line of chromatophores) to moderately developed pre- and postdorsal, median stripe. The top of the head is rather uniformly pigmented, the pigmentation extending only to the upper part of the almost immaculate opercle and cheek. There is a narrow ring of pig- ment around the lower rim of the orbit and a concentration of pigment in front of orbit (preorbital bar hardly visible with unaided eye). The extent of pigmentation on lips is variable. In some specimens both lower and upper lips are immaculate, other specimens have just the anterior portion of upper lip pigmented and in others both lips are pig- mented but none has pigment on chin or isthmus. The pelvic and anal fins (a few specimens from Black Warrior River have pigmentation along rays of anal fin) are without pigment; the caudal and dorsal fins lack pigment on the membranes but have it along the rays. Pigmentation on the pec- toral fins is sparse, progressively diminish- ing from anterior to posterior rays. There is usually a single file of chromatophores along the leading and posterior edge of the first pectoral ray and only along posterior margins of succeeding rays. The chromato- phores are more widely spaced in the files, and the lengths of the files are shorter pro- gressively toward the posterior rays of the Notropis edwardraney1 5p) TABLE 7 Number of left pectoral fin rays in three species of Notropis. Species and Range Il 2, 13) 14 15 16 yi N x edwardraneyi Tombigbee R. 1 ll 10 3 a le 7 Black Warrior R. 1 5 8 14 14.5 Big Mulberry Cr. 6 1 2 ilI55 1M 7/ Cahaba R. 5 8) 6 XN) Alba S0) Alabama R.., Watts Bar 4 20 6 30 1554 Hobbs Bar i 9 8 2 D0 IAS Yellow Jacket Bar ll 16 3 Sy ME 7/ Choctaw Bluff Bar 1 2 10 15 3 31 14.5 Totals 1 5 Dil 3 25 18] Ai blennius Seine R., Manitoba 9 a Geli 4: Mississippi R., lowa 2 7 i) 2 30 =615.4 Mississippi R., Missouri 2 4+ 24 We 9) 49 15.3 Arkansas R., Arkansas 4 29 23 4 60 15.4 Mississippi R., Lake Providence 1 0 0 6 34 10 51 15.0 Vicksburg 1 9g 32 U i a0) 0) Natchez 1 at HI 16 1 50) ALBA Red R., Louisiana 1 1 Totals 1 0) 4 36 166 90 10 307 52 potteri Red River, Bossier City 5 Tf 8 2, 205 b:4 Coushatta I 8 5 1 [5 eS! Alexandria 2 6 5 2; | Saale Totals 6 21 18 5 50) lpr: fin. Usually the posterior half to two-thirds of the fin is immaculate. The peritoneum is silvery with scattered melanophores. Reproduction. Ripe females were collected from the Alabama River at the abandoned ferry landing across from the town of Ca- haba between 2300 and 0045 hours on 27-28 June 1964 (TU 35243). The spawning in- dividuals were taken in moderate current along the bank from over a sand bottom. The water varied from one to two-and-a- half feet deep and was 28 C at the time of capture. Ripe individuals were collected again at the same locality on the following night (June 28) between 2230-2330 hours (TU 35269). The next time that ripe in- dividuals were collected was 6 May 1965, from the Tombigbee River (TU 37585) northwest of Columbus, Mississippi, be- tween 2115-2330 hours. A number of speci- mens was checked at the time of collection and all proved to be ripe females. These were taken from over gravel or a mixture of gravel, sand and silt in one and a half to two feet of water. The water temperature was 21 C at time of collection. During the following spring the same locality on the Tombigbee was sampled. Distended females collected between 2300 and 2430 hours, 9 and 10 April 1966 (TU 40489) voided eggs only when firm pressure was applied with fingers (squeeze ripe). Water tem- perature was 15 C at time of collection. Females collected from Tombigbee River 3.9 miles west of Amory on 10 April 1966 between 1120 and 1200 hours (TU 40497 ) were squeeze ripe also. Water temperature was 15 C and depth of capture, type of as- sociated bottom and current was similar as described for other localities. On 31 May 1967, many ripe females were taken from the Alabama River at an un- named bar (River Mile 130.2) and Yellow Jacket Bar, the type locality (River Mile 129.8). The former collection (TU 46796) was obtained between 2123 and 2142 hours and the latter (TU 46802) 2150 to 2230 hours. Water temperature was 24 C at both 34 Tulane Studies in Zoology and Botany Vol as TABLE 8 Number of right pectoral fin rays in three species of Notropis. Species and Range 13 14 15 16 17 N Xx edwardraneyi Tombigbee R. Black Warrior R. Big Mulberry Cr. Cahaba R. Alabama R., Watts Bar Hobbs Bar Yellow Jacket Bar Choctaw Bluff Bar Totals blennius Seine R., Manitoba Mississippi R., lowa Mississippi R., Missouri Arkansas R., Arkansas Mississippi R., Lake Providence Vicksburg Natchez Red R., Louisiana Totals potteri Red R., Bossier City Coushatta Alexandria Totals 8 10 3 21) 4s 7 6 1 14 14.6 6 7 2 15° SIA? 6 10 4 20) e14s9 5 19 6 30. =-:15.0 2 8 8 1 ] 20 «6145 10 16 4 30,——«14.8 2 12 16 1 31 14.5 4 62 92 22, sil 14.7 11 a 1 16 $15.4 1 3 8 16 2 BO Mellor 3 6 27 12 | 49 15.0 2 29 23 6 60 15.5 ul 31 12 i OL lida 3) 34 8 50 ~=—-:15.0 11 23 15 i 30°) «Se 1 1 4 37 163 91 12 307") Na:2 1 8 if 4 20 dat 1 6 il IL 15 15.5 2 5 7 1 15 «15.5 4 19 21 6 50 §815.6 localities. In both areas ripe fish were taken from over gravel in a riffle area as well as from over sand bottom in moderate current. The areas of capture varied from one to two-and-a-half feet in depth. Thus in sum- mary, Notropis edwardraneyi was taken in spawning condition during May and June at water temperature ranging from 21 to Usyal Gy Range. All specimens were taken from the main channel of the Alabama and Tom- bigbee rivers and the lower reaches of their major tributaries (Fig. 9). The irregular distribution of Notropis edwardraneyi shown on the map is a reflection of the discon- tinuity of collecting and not discontinuities in distribution. Geographic Variation. No clines seem to be present in meristic characters, propor- tions, Or pigmentation. Specimens from the extremes of the range as well as from in- termediate areas and tributaries were used in making meristic counts. The only notable deviation was the number of lateral line scales in the sample from Choctaw Bluff (Table 4). However, the frequency dis- tribution for Choctaw Bluff specimens and the other distributions for the upstream (more northerly ) samples do not illustrate a north-south cline. Relationships. Notropis edwardraneyi ap- pears to be a close relative of Notropis blen- mus and may represent an off-shoot from blennius stock that gained access to the Ala- bama and Tombigbee systems through a former connection with the ‘Tennessee (Hayes and Campbell, 1900: 131-133; Simpson, 1900: 133-136; Suttkus and Ramsey, 1967: 138). The absence of ed- wardraneyi in the Pascagoula River, Pearl River, and other minor river systems be- tween the Alabama-Tombigbee systems and the Mississippi River would seem to negate an eastward movement of blennius stock from the Mississippi River basin to the Ala- bama-Tombigbee; we believe that the Ten- nessee River was the most likely route of access in the near geological past. A number of series of blennius from various localities along the Mississippi basin and from Mani- toba was examined and characteristics com- pared to determine presence of geographical No. 1 Notropis edwardraneyt 35 TABLE 9 Number of pelvic fin rays for three species of Notropis. 9-9 98 89 88 87 7-8 7-7 86 7-2 N edwardraneyi 2 3 l 163 3 4 a? ae 181 (TU 33381, 30; TU 35134, 20; TU 35323; 31; TU 37585, 21; TU 40457, 15; TU 43332, 14; TU 44028, 30; TU 47515, 20) blennius 5 3 3 232 9 0 2 2 ibe Anil (UMMZ 180500, 16; UMMZ Acc. 1947:XI:28, 30; TU 43167, 60; TU 48248, 50; NLSC 5356, 32; NLSC 5374, 25; NLSC 5441, 18; NLSC 5506, 20; NLSC 5367, 6) potteri 3 1 44 1 0 1 50 (NLSC 3700, 15; NLSC 5352, 20; NLSC 5357, 15) clines or populations which possessed inter- mediacies between other blennius popula- tions and edwardraneyt (Tables 1, 4, 5, 6, 7, 8, 10). Unfortunately, blennins material from the Tennessee River was not available for our study. The specimens (UAIC 1830) from the Mississippi River in southeastern Missouri seem to have a larger orbit than any other blennius material from more northern or southern localities. However, some specimens (see footnote, Table 10) were in a partially dehydrated condition and accuracy of measurements was not com- parable to that for other samples. Excluding the Missouri sample, the other three samples used for Table 10 demonstrate a weak north- south cline, with a slight increase in size of orbit toward the south. No samples examined indicated a population with close affinities (particularly with regard to size of orbit) to edwardraneyi. Thus we assume that dif- ferentiation occurred primarily or entirely after the stock became isolated in the Ala- bama-Tombigbee basins, if blennius and ed- wardraneyi are indeed close relatives and do not merely represent convergence because of their large river habitat. Geographical clines are hardly demon- strable for Notropis blennius (Tables 1, 4, 5, 6,7, 8 and 10) especially with regard to me- ristic characters. Perhaps the only propor- tions that could be considered as clinal are the diameter of orbit and length of upper jaw in relation to head length (Table 10). These data seem to be in contradiction to those presented by Hubbs and Bonham, 1951: 104 and comparison number 23, on page 107 in which they said Notropis blen- nius jejunus had a larger eye than N. J. blen- nius. However, they said also that in the far north the eye in jejwnus becomes reduced. Apparently they did not mention the source of their comparative material and so further discussion is unwarranted. The trend toward the south in an increase in length of jaw is in about the same magnitude as the increase in size of orbit. In view of the wide separation of the range in size of orbit of edwardraneyi and the most southern samples of blennius examined, there seems to be little or no reason to believe that a popula- tion of blennius with intermediate size orbit exists, particularly in the intermediate geo- graphical area. For the sake of completeness the following data are presented for 258 specimens of Notropis blennius (338 less 80 of the series; UMMZ ACC. 1947:XI: 28). One of the 258 specimens had 9 princt- pal dorsal rays, all the rest had 8; two of the 258 specimens had 8 principal anal rays, all the rest had 7; 243 speci- mens had 19 principal caudal rays, two had 20, ten had 18 and three had 17; 233 speci- mens had a scale count around the caudal peduncle of 5-2-5, one had 5-2-6, one had 5-2-7, ten had 6-2-5, one had 6-2-6, eight had 7-2-5, two had 7-2-6 and two had 7-2-7. Notropis potteri was mentioned as an ally of blennius by Metcalf (1966: 122) and as a Close relative of edwardraneyi in our introduction and perhaps is, but it 1s more distinct from edwardraneyi than blen- 36 Tulane Studies in Zoology and Botany TABLE 10 Measurements of Notropis blennius expressed in thousandths. Mean value is given below (in parentheses) range of variation. Vol. 15 Locality Miss. R., lowa Miss. R., Mo. Arkansas R., Ark. Miss. R., La. Catalog Number UMMZ Acc. UAIC 1830 TU 43167 NLSC 5356 1947: X1:28 NLSC 5441 Number of Specimens 30 50* 50 50 Standard Length (mm) 39.7-58.1 38.0-59.0 45.9-60.0 37.3-58.0 (45.1) (47.6) (52.4) (49.5) Head Length/S. L. 248-279 251-280 207-294 257-283 (265 ) (265 ) (268 ) (27Ab) Interorbital/ Head Length 342-391 351-405 347-413 328-393 (369 ) (379) (374) (358 ) Snout/Head Length 273-319 281-326 282-342 272-336 (295 ) (301 ) (312) (294) Orbit/ Head Length 228-258 243-287 229-273 234-279 (241) (262 ) (249) (253) Upper Jaw/Head Length 298-347 299-339 285-348 300-351 (316) (318) (320) (323) Postorbital/ Head Length 474-533 447-514 441-482 437-510 (494) (476) (466 ) (476) Orbit /Interorbital 600-731 620-778 589-755 618-828 (655 ) (693 ) ( 667 ) (708 ) Orbit /Snout 730-903 756-972 706-943 735-968 (819) (872) (799 ) (851) Orbit/Upper Jaw 700-854 750-897 674-868 711-906 (765 ) (826) (781) (783) Orbit /Postorbital 452-533 491-636 478-579 460-615 (489 ) (552) (535 ) (532) * Some specimens in partially dehydrated condition, orbit enlarged because of shrunken condi- tion of fleshy orbital rim. nius in a number of ways. Perhaps it too was an early offshoot of blennius stock that was isolated in the Brazos River and there dif- ferentiated. However, we believe a more plausible explanation of relationship is that potteri represents a descendent of a different but unknown stock, and that it has con- verged in a number of ways. Cross (1953: 258) concluded that the presence of No- tropis bawdi in the Brazos was not. at- tributable to recent introduction as a bait minnow, but that N. baird? is endemic there. In a similar way we differ from Hubbs and Bonham (1951: 107-109) about the pres- ence of Notropis potter: in the Red River as a result of bait introductions from the Brazos River. We believe that N. potteri gained access into the Red River system through natural means as the result of tribu- tary captures or stream connections. We say this on the basis of the thousands of specimens of N. potter: taken from the lower Red River that are available in the Tulane University and Northeast Louisi- ana State College collections. We do not maintain that these collections of potteri from the lower portion of Red River refute the idea of bait introductions especially be- cause most of the mentioned material was collected in recent years. The earliest collec- tions were obtained in 1955 and 1956 from the Red River at Acme, Natchitoches, and Alexandria, Louisiana. At the Natchitoches locality a total of 837 specimens represent- ing 18 species was seined, of which 194 specimens were poftert. The Alexandria col- lection had 15 species and 440 specimens, of which 124 were potter. Unfortunately we do not have collections available at this time from a large intermediate section of the Red River either before or after forma- tion of Lake Texoma. Our collections from the upper Red River system show Notropis bairdi to be the dominant ecological repre- sentative in that area. If potteri could not successfully compete with bairdz, this would Notropis edwardraneyt ~— Wo TAsERn) J Number of scales around the body below the lateral line. 9 10 in 12 13 14 15 N x edwardraneyi 33 63 76 fe) 18] 10.3 blennius l 13 198 69 54 3 2 338 MEAS) potteri 1 6 34 i 2, 50 U3} account for its absence in the upper Red River. Perhaps the reverse is true in the lower Red River where at least we have never taken bairdi. The occasional collec- tions of potter: from the Red River at Lake Texoma (Riggs and Bonn, 1959: 163) may be a reflection of more collecting ef- fort after construction of the dam, or it may be a result of reinvasion of the area because of the changes in the habitat which are more suitable for pottert and other fishes that have been maintaining populations in the continuously flowing lower Red River. Further discussion of natural dispersion versus introduction by man should include Notropis brazosensis (= shumardt), fol- lowing Gilbert and Bailey, 1962, and No- tropis oxyrhynchus (Miller, 1953: 33-34). Such a discussion was not planned for this paper. However, an additional remark with regard to man’s introduction versus natural occurrence of N. potteri in the Red River is here presented. If man introduced pofteri it has swamped out and completely, or nearly so, replaced blennius in the shallow mar- ginal areas of the lower Red River. Notropzs blennius may have been displaced to the deeper parts of the channel where it may be present as natural resident populations. The foregoing remarks are pertinent to the discussion of the relationship of potteri and blennius. Hubbs and Bonham (1951: 103) reported on a specimen of blennius taken from the Red River south of Ada, Oklahoma, by George A. Moore and J. M. Paden. Moore thought that it represented a bait introduction and perhaps this is true. A single specimen of blennins (TU 47644) was collected along with the hundreds of specimens of potteri from the lower Red River that are contained in the Tulane Uni- versity collection. Notropis potteri obvi- ously is the dominant ecological representa- tive in the areas that we are able to sample with seines. However, many of the series housed at Northeast Louisiana State Col- lege were collected by use of wire trap nets. The single specimen of blennius is typical of the species from the Arkansas River in counts, measurements, and pigmentation and was immediately distinguished from potters taken in the same collection. Northeast Louisiana State College collections from the Mississippi River at Natchez, Miss., repre- sent a mixture of blennius and potteri. Be- cause of the improper preservation (speci- mens probably died before they were pre- served) it is difficult to be certain that in- trogressive hybridization has or has not taken place. Some specimens (those in bet- ter condition ) are identifiable as blenniuws and others as pottert. Thus we record the pres- ence of Notropis potteri in the Mississipp1 River proper. The collection (RDS 4236) obtained by the authors from the Mississippi River at Vicksburg on 11 October 1967 con- tained no potters but 856 specimens of blennius (TU 48248), none of which re- sembled potteri. Of course this does not prove the absence of potteri in the Vicks- burg area other than perhaps at the particu- lar time and place of our collection. Collec- tions of blennius from the Mississippi River at Baton Rouge and New Orleans, La., ap- parently do not exhibit any potters traits. Further analysis of the relationship of the two species in the stretch of Mississippi River between Natchez and the mouth of the Red River is deemed impractical at this time. However, the relationship of pot- teri from the Red and Brazos rivers and blennius from the Arkansas and elsewhere in the Mississippi River drainage will be discussed. The ontogenetic changes in Notropis pot- teri are remarkable and are decidedly dif- ferent from those of either blennius or ed- wardraneyt. The size of orbit does not in- crease at the same rate as increase in length of head (Fig. 10). In specimens of 50 mm standard length the orbit is approxt- mately one-fourth the length of head (Table 38 Tulane Studies in Zoology and Botany 2) whereas the diameter of orbit is less than one-fifth the length of head in speci- mens over 80 mm standard length. For spect- mens which average about 50 mm standard length (Table 2) the orbit of potteri is considerably smaller than that of blennius and extremely so in comparison to that of edwardraneyt. Though the eye is smaller in potteri the interorbital distance is not pro- portionally so much greater than that in blennius because of the higher position of the eyes in pottert. The changes in the shape and length of snout deserve special mention. The young and small juveniles of potters have an ex- tremely sharp-pointed snout (sharper than either blennius or edwardraneyt) which gradually approximates the shape and rela- tive length of snout of blennius and ed- wardraneyz and then at its largest size has the bluntest snout of the three. Obviously larger individuals were under observation when Hubbs and Bonham (1951: 102) remarked upon the similarity of potteri to Semotilus atromaculatus. Moreover, there is a structural difference between potteri and blennius which apparently adapts potteri for a closer association with the bottom. The pectoral fins project laterally in a rather rigid fashion so that they are seldom seen on their side while flopping in the seine but are usually on their belly. Notropis blennius usually shows its side when alive in the net. Numerous field observations dis- close that potteri lives on the bottom and is usually closely associated with it; how- ever, blennius is often observed and taken from mid-depths, off the bottom. Some dif- ferences in proportions among the three species under consideration are illustrated in the outline drawings, Figures 6, 7, and 8. We are in agreement with Hubbs and Bonham (1951: Table VI, 109) on measure- ments of potteri and so did not duplicate or incorporate a set of measurements in our Table 1. Notropis potteri tends to be closer to blennius than to edwardraneyi in most of the scale and fin-ray counts pre- sented in Tables 4-8. Notropis potteri av- erages more scales around the body and usually has 13 or more below the lateral line (Tables 5 and 11). Many of the high counts in blennius relate to a high count above the lateral line and not below, the latter count (below) is modally 11. Vol 5 Hubbs and Bonham (1951: 107) gave a description and comparison of pigmen- tation of blennins and potter. Cognizant of some variability in pigmentation in the three species under consideration, we believe Figures 3, 4, and 5 represent typical pat- terns. Notropis potteri often has an inter- vening scarcely pigmented area between the lateral line pores with their associated pig- mentation and the more dorsal band of large chromatophores. In blennius the pig- mentation is continuous from the back, down the sides to the lateral line row of scales and often on the scale row below the lateral line. Again, in order to be more complete, the following are the data for the less variable structures. In 50 specimens of potteri from the Red River in Louisiana two had 7 principal dorsal rays, the rest had 8; two had 8 principal anal rays, one had 6, and the rest had 7; two had 18 principal caudal rays, the rest had 19; 20 specimens had 5-2-5 scale rows around the caudal peduncle, 12 had 6-2-5, one had 6-2-6, 11 had 7-2-5, two had 7-2-6, and four had 7-2-7. Other scale and fin-ray counts are given in Tables 4-9, 11. Etymology. We take pleasure in naming this new form in honor of Dr. Edward C. Raney in recognition of his many contribu- tions to North American ichthyology and his guidance and imparted enthusiasm toward a multitude of students. LITERATURE CITED Cross, FRANK B. 1953. A new minnow, No- tropis bairdi buccula, from the Brazos River, Texas. Texas J. Sci., 5(2):252-259. GILBERT, CARTER R. and REEVE M. BAILey. 1962. Synonymy, characters, and distribution of the American cyprinid fish Notropis shumardi Copeia, 1962 (4):807-819. Hayes, C. W. and M. R. CAMPBELL. 1900. The relation of biology to physiography. Science, 12(291 ): 131-133. Husss, Cart L. and KELsHAw BONHAM. 1951. New cyprinid fishes of the genus Notropis from Texas. Texas J. Sci., 3(1):91-110. Husss, Cart L. and Kari F. LAGLErR. 1958. Fishes of the Great Lakes region. Cranbrook Inst. Sci. Bull., 26:1-213. Metca.r, ArTIE L. 1966. Fishes of the Kansas River system in relation to zoogeography of the Great Plains. Univ. Kansas Publ., Mus. Nat. Hist., 17(3):23-189. MiLterR, Donatp R. 1953. Two additions to Oklahoma’s fish fauna from Red River in Bryan County. Proc. Okla. Acad. Sci., 34: 33-34. No. 1 Notropis edwardraneyi 49 Riccs, Cart D. and Epwarp W. Bonn. 1959. nal cyprinid species Moniana deliciosa Girard An annotated list of the fishes of Lake Tex- and Cyprinella texana Girard. Copeia, 1958 oma, Oklahoma and Texas. Southwest. Natur., (4):307-318. 4(4):157-168. SuttTkus, Roya D. and Joun S. Ramsey. 1967. Simpson, CuHas. T. 1900. On the evidence of Percina aurolineata, a new percid fish from the Unionidae regarding the former courses the Alabama River system and a discussion of the Tennessee and other southern rivers. of ecology, distribution, and hybridization of Science, 12( 291 ):133-136. darters of the subgenus Hadropterus. Tulane Surrxus, Royat D. 1958. Status of the nomi- Stud. Zool., 13(4):129-145. October 16, 1968 om ; — A\fi ( es TULANE STUDIES IN ZOOLOGY AND BOTANY ~~ Volume 15, Number 2 December 23, 1968 AGGREGATIVE BEHAVIOR AND HABITAT CONDITIONING BY THE PRAIRIE RINGNECK SNAKE, DIADOPHIS PUNCTATUS ARNYI HAROLD A. DUNDEE Department of Biology, Tulane University New Orleans, Louisiana 70118 an M. CLINTON MILLER II Department of Biostatistics, Tulane University School of Medicine, New Orleans, Louisiana 70112 : p. 41 A NEW TURTLE SPECIES OF THE GENUS MACROCLEMYS (CHELYDRIDAE) FROM THE FLORIDA PLIOCENE JAMES L. DOBIE Department of Zoology-Entomology Auburn University, Auburn, Alabama 36830 p. 59 STUDIES ON FROG TRYPANOSOMIASIS II. SEASONAL VARIATIONS IN THE PARASITEMIA LEVELS OF TRYPANOSOMA ROTATORIUM IN RANA CLAMITANS FROM LOUISIANA RALPH R. BOLLINGER, JOHN RICHARD SEED and ALBERT A. GAM Laboratory of Parasitology, Department of Biology Tulane University, New Orleans, Louisiana 70118 p. 64 STUDIES ON AMERICAN PARAGONIMIASIS VI. ANTIBODY RESPONSE IN THREE DOMESTIC CATS INFECTED WITH PARAGONIMUS KELLICOTTI JOHN RICHARD SEED, FRANKLIN SOGANDARES-BERNAL, and ALBERT A. GAM Laboratory of Parasitology, Department of Biology, Tulane University, New Orleans, Louisiana p. 70 TULANE UNIVERSITY NEW ORLEANS TULANE STUDIES IN ZOOLOGY AND BOTANY is devoted primarily to the biology of the waters and adjacent land areas of the Gulf of Mexico and the Caribbean Sea, but manuscripts on organisms outside this geographic area will be considered. Each number is issued separately and contains an individual monographic study, or several minor studies. As volumes are completed, title pages and tables of contents are distributed to institutions receiving the entire series. Manuscripts submitted for publication are evaluated by the editor or associate editor and by an editorial committee selected for each paper. Contributors need not be members of the Tulane University faculty. 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Authors may obtain separates of their articles at cost. Address all communications concerning manuscripts and editorial matters to the editor; communications concerning exchanges, and orders for individual numbers to the Meade Natural History Library. When citing this series authors are requested to use the following abbreviations: Tulane Stud. Zool. and Bot. Price per number (flat rate): $1.50 Price per volume (flat rate): $5.00 Gerald E. Gunning, Editor Arthur L. Welden Associate Editor Department of Biology Tulane University New Orleans, Louisiana 70118, U.S.A. Meade Natural History Library Tulane University New Orleans, Louisiana 70118, U.S.A. Tim M. Berra Assistant to the Editors £ & TULANE STUDIES IN ZOOLOGY AND BOTANY Volume 15, Number 2 December 23, 1968 AGGREGATIVE BEHAVIOR AND HABITAT CONDITIONING BY THE PRAIRIE RINGNECK SNAKE, DIADOPHIS PUNCTATUS ARNYI HAROLD A. DUNDEE og Department of Biology, Tulane University I New Orleans, Louisiana 70118 and M. CLINTON MILLER III Department of Biostatistics, Tulane University School of Medicine, New Orleans, Louisiana 70112 HARV ABSTRACT When ringneck snakes were released in laboratory arenas in which all cover ob- jects presented equalized physico-chemical conditions the snakes were found to follow a multinomial distribution with unequal probabilities of occupancy of each cover object. The resultant clumped distributions are the result of intrinsic aggregative drives which are not based on sexual attraction, as was evident from repeated aggregation formation among either males or females in experiments involving only one sex. Preliminary findings on effects of environ- mental variations in the experimental arenas indicate that aggregative behavior can be modified, at least by increasing or decreasing the dampness of the substrate. Persistent use of the same cover object by groups of ringneck snakes indicated that there might be a bias for particular locations in the arenas. Various experi- mental tests demonstrated that the bias was due to presence of an attractant sub- stance, possibly integumentary, cloacal gland, or excretory in origin, which is deposited by snakes that first frequent the particular cover site. Such sites are herein described as “conditioned habitat.” Snakes will seek and return to conditioned substrate even when it is translocated in the experimental arena. Snakes apparently use visual and olfactory and/or vomero- nasal sensory perception to locate the con- ditioned sites and to trail other snakes or locate trails left by the travels of other ringneck snakes. The advantages of aggregative behavior and habitat conditioning are discussed and compared to denning habits of large snake species. V AML UNIVERSITY INTRODUCTION Social behavior in snakes, at least in the form of epideictic display, is very limited. Passive social behavior such as aggregation is a common phenomenon, but studies on factors initiating and affecting aggregative behavior are few and are limited in scope. Reports of snake dens are common in litera- ture, the most exhaustive discussion of the topic being that by Klauber (1956). Den- ning, of course, is a seasonal phenomenon principally characteristic of large snake spe- cies. Aggregation is also common among small secretive species. It may occur in hi- bernacula during the winter, but is also observed at other times of the year. A com- prehensive study of aggregative phenomena in the northern brown snake (Storeria de- kayi) was reported by Noble and Clausen (1936). Otherwise the reports on aggrega- tion in small snakes have been casual state- ments about the number of snakes found under a single object, or descriptions of the responses of animals to extremes of tempera- ture, humidity, light, and so forth. The interpretation of aggregative phe- nomena must take into account extrinsic and intrinsic factors. The extrinsic factors relate to habitat selection in a varied en- vironment; the intrinsic factors involve in- stinctive social drive in the absence of al- ternative physical environments. A seasoned snake collector working in a familiar area EDITORIAL COMMITTEE FOR THIS PAPER: Dr. CHARLES C. CARPENTER, Professor of Zoology, University of Oklahoma, Norman Dr. FREDERICK B. TURNER, Laboratory of Nuclear Medicine and Radiation Biology, University of California, Los Angeles 4l 42 Tulane Studies in Zoology and Botany develops considerable ability to find his quarry. What he is recognizing is habitats and microhabitats and, crudely, the physio- logical needs of a snake. He also becomes aware that the environment is not uniform and that he is unable to satisfactorily evalu- ate the distinction between intrinsic and extrinsic factors bringing the snakes into the sites that he searches most exhaustively. The excellent study by Noble and Clausen (op. cit.) has provided many instructive clues for recognition of these conditions, but for reasons given below other approaches to the understanding of aggregative phe- nomena are required. In the south central United States there are several species of small secretive snakes, generally averaging under 12 inches long and usually found under sheltering objects in the mild, moist spring months. Some species are largely solitary (e.g., Tantilla gracilis), others tend to occur singly or in groups of two or three under individual sheltering objects (e.g., Tropidoclonion line- atum, Carphophis amoenus, Leptotyphlops dulcis, Sonora episcopa), and one species (Diadophis punctatus arnyi) is very gre- garious, often occurring in numbers of half- a-dozen or more under a relatively small piece of cover. Additionally, certain stones regularly have small snakes under them, sug- gesting that optimal conditions must prevail in these microhabitats. Published studies by various authors on the above mentioned species have largely confirmed these im- pressions. The distribution of small secretive snakes can, of course, be assumed to be non-random in the field. Since the natural environment is variable the snakes will ordinarily be found under naturally occurring suitable cover items; hence their distribution will not be random with respect to all cover. If all cover objects had equal probability of selection as a cover object by each snake the snakes would be unable, therefore, to select cover objects on the basis of physico-chemical attraction. Under such equalized conditions, three alternative distributions of the snakes appeared possible, given sufficient but not excessive time. The snakes could be dis- tributed as: 1) a multinomial distribution with equal probability of occupancy of each cover site or Vol. 15 2) a uniform distribution or 3) a multinomial distribution with un- equal probabilities of occupancy of each cover site. If the first distribution were observed no social relationships would be likely and dis- tribution could be termed as random. If the distribution were found to be uniform, terri- torial behavior would be suspected, par- ticularly if only one animal utilized an in- dividual cover item. Finally, if a multi- nomial distribution with unequal probabil- ities of occupancy for each disc were ob- served, there would be evidence of social relationships. The first and third distribu- tions would of course result in aggregations or clumps of the ringneck snakes, but the expectancy of clumps would be greatest in the third type of distribution. The ringneck snake, Diadophis punctatus arnyt, was selected for the present study be- cause of its abundance and its obvious gre- gariousness. Various reports call the species common, apparently assigning this category to finds of a dozen or so in an hour. Hurter (1911) reported finding 17 specimens under a rock about 24 X 18 inches. Brennan (1936) reported 14 under a flat rock. Rich- ard B. Loomis (personal communication ) knew of one case where 44 were found un- der a single stone. Loomis and Dundee col- lected 300 ringnecks, as well as many other reptiles, in one hour on April 30, 1950 in Douglas County, Kansas. On April 11, 1967 Dundee alone collected 238 ringnecks in 50 minutes in Douglas County, Kansas, from an area about 10 feet wide and 400 feet long. This latter collection included all sizes of ringnecks and counts of 15, 12, 11, 9, 9 were recorded from individual stones. The obvious questions arising from these observations on abundance are (1) were the sites optimum habitat; (2) were in- dividual stones particularly desirable, thus attracting more ringnecks during their wan- derings; (3) were the snakes gregarious because there were so few cover objects in relation to the population size; (4) were the aggregations a result of intrinsic drive? Not all of these questions are immediately answerable, but certain generalizations might be made providing partial answers to some of these questions. First, we can assume that the Douglas County physical setting is an optimum habitat because it regularly har- Prairie Ringneck Snake Behavior 43 Figure 1. bors large numbers of ringnecks. Second, snakes were found beneath various sizes of stones under which the soil ranged from dry to moist. And third, the stones were loose on the ground or partially embedded. These observations indicate that the micro- habitats selected were not markedly re- stricted. There were many flat limestone slabs that harbored various reptiles or none, but which could have served as cover for ringnecks. Hence we judge that population pressure was not severe. Only laboratory experiments can provide data for an ade- quate answer to the fourth question and hence the basis for the initial studies of our investigation. MATERIALS AND METHODS The basic design of our experiments derives from the hypothesis that ringneck Circular sand-filled arena used for experiments. Snakes seen are ringnecks to show relative size in comparison to the seven-foot diameter of tank and 12-inch diameter discs. snakes have intrinsic aggregative drives which may or may not be modified by en- vironmental factors. Artificial habitats were established in the laboratory by using circular livestock water- ing tanks seven feet in diameter. The tanks were mounted on platforms equipped with rollers so that they could be rotated at will. Thirty mm of mason’s sand was placed on the bottom to provide a non-packing sub- strate. The sand could be smoothed to unt- form depth. Circular discs, 12’ in diameter, made of 14” plywood and painted gray to simulate limestone were arranged radially concentric to the edge of the tank (Fig. 1). These discs could be rotated by hand, and pressed down to make them lie flush upon the sub- strate. Loose sand was brushed up around the perimeters of the discs to insure sealing dt Tulane Studies in Zoology and Botany of any irregularity that might offer easy access to the underside of the discs. The circular tank and circular discs eliminated angles that could provide biased thigmotactic satisfaction. Each disc offered equal condi- tions for selection as a cover site. Most experiments were conducted using dry sand. In certain experiments with moist sand as a substrate, Bouyoucos soil moisture blocks from Industrial Instruments, Inc. were buried in the sand at strategic locations. Conductivity in the blocks was measured by a Bouyoucos Moisture Meter, also from In- dustrial Instruments, Inc. The experimental arenas were kept in a room usually main- tained at 75 +2°F and humidity of 57+ 5%. Only a diffused outside light entered the room by day. Artificial light provided from the ceiling was regulated by an automatic timer to approximate normal day length, but was set to come on after dawn so that no sudden brightening would frighten the snakes. Experiments were run with adult animals only and almost exclusively with males. The use of males was almost mandatory because of the paucity of females in field collec- tions. Snakes used in experiments were sorted according to sex (double checked by determining presence of anal ridges and hemipenial sheaths in males). For certain experiments the animals were marked with either brightly colored lacquer or butyrate dope for convenient recognition, thus eliminating the need for handling the snakes. Markings used were yellow, orange, and white neck rings or short dorsal stripes. Animals were reused for experiments but were fed earthworms between experiments and were permitted to rest for about a week before being placed into arenas again. In certain experiments, particularly those 1n- volving dry sand, when snakes were gathered for re-release, water was provided ad libitum before release to offset dehydration. Snakes were released from the center from under an inverted can after they had become quiescent. All releases were made during daylight hours. Dispersal and distribution of the snakes were determined by daily daytime inspec- tions of the arenas. Procedures included, (1 ) initial release of snakes and the notation of their locations without handling or dis- Vols placing them, (2) daily pickup and _ re- release from the center, (3) daily pickup and re-release after the sand had been gathered, thoroughly mixed, redistributed, and discs washed in detergent and reset, (4) trans- location of specific discs and their under- lying substrate to other positions in the arena, (5) rotation of tanks to test for pos- sible compass or light orientations. RESULTS General Behavior of Snakes After Release When the inverted can was lifted to re- lease snakes there was little reaction and the animals usually lay quiescent for up to 10 minutes. Eventually one or more animals would begin to glide out from the center. When this movement began it was not un- usual for the moving animals to be joined by others so that they appeared to stream out in pairs or triplets. Movement away from the center seemingly was cautious and exploratory but within 15 minutes most snakes were active and gradually edging from the center, though some tended to remain for periods of an hour or more. The snakes often pressed their snouts to the substrate, suggesting the use of olfaction. Their actions suggested that vision, olfaction, and vomeronasal sensing were being em- ployed to enable the snakes to determine where they wished to move. Both vision and olfaction were stated by Noble and Clausen (1936) to be the primary senses used by Storeria dekayi to locate their own kind, but a later study by Wilde (1938) suggested that the vomeronasal organ (= Jacobson’s organ) rather than olfaction was the effective sensory device. We are con- ducting experiments to determine if these senses are the major ones used by individual Diadophis during their dispersal and search for cover and other Diadophis. Eventually the radial dispersal became a concentric one because of the limiting wall of the arena, and snakes ceased to stream in pairs and triplets. Generally most snakes were hidden by day, but when the substrate was smoothed at night numerous tracks could be found the following morning, showing that consider- able nocturnal movement had taken place. Snakes that might be abroad in the early morning frequently were under cover when disc checks were made in the early after- noon. No. 2 Prairie Ringneck Snake Behavior 45 TABLE | Dispersal of ringneck snakes in an arena containing dry sand, 10 twelve-inch discs, 40 male snakes. Ambient temperature 75—76° F. Relative humidity 54-64%. May. ; Disc Days after release 1 9) 3 4 5 6 7 8 9 10 open hidden 2 1 1 1 5 6 4 0 5 1 2 14 0 3 0 2 1 2, 4 rl 0) 6 10) 2, 16 0) 4 1 9) 0 1 4 14 0 6 0 4 8 0 5 I 2 ) 0 4 17 0 4 0 2 8 0) 6 1 1 1 2 4 12 0 6 1 2, 10 0 7 0 2 0 0 3 24 1 6 0) 0 4 0 Access to the underside of discs was ac- complished by pressing the snout repeatedly against the perimeter of the disc until the head was pushed under. Only rarely did any snake burrow into the open sand. Snakes and Substrate Left Undisturbed After Location of Snakes Noted 1. General statement of hypothesis. In our first experiments we assumed that if snakes were released from a central point they would distribute randomly, and that if no aggregative drive existed, snakes would show a random distribution from day to day. A series of experiments was designed to test this hypothesis of random distribu- tion following release. Noble and Clausen (1936) found that aggregations of Storeria dekayi could be forced by jostling the cages. They also found that snakes clustered in areas of presumed optimum temperature and humidity when left undisturbed, but if the clusters were disturbed the snakes would break into smaller groups. Thus it was de- cided that experimental animals should be left undisturbed after release. Initial experi- ments showed that if discs were lifted gently and the snakes counted quickly there was little evidence of excitement of the animals. After the discs were replaced the snakes were free to move at random. 2. Results, In the first experiment 40 male snakes were collected and released in an arena con- taining dry sand and 10 twelve-inch discs. The distribution of the snakes was recorded daily. The results of this experiment are given in Table 1. The chi-square test of the null hypothesis that the distribution was the same for each day was not significant [chi-square, 45 df. = 47.34 (p= .35) (Cramér, 1946, p. 443) ]. We concluded that there was no difference in the distributions for the various days. Therefore we decided to pool the snake counts across days to determine an overall distribution of the snakes under the discs. A chi-square of the null hypothesis of equal probabilities of occupancy for each disc by each snake was rejected by the chi-square test (chi-square 9 d.f.= 274.89) with prob- ability <.01. It was concluded that the male ringneck snakes did not follow a multi- nomial distribution based upon equal prob- abilities of disc occupancy in their disper- sion in the arena. The second experiment consisted of re- leasing twelve female ringneck snakes in an arena containing dry sand and 10 twelve- inch discs. Daily observations were again made. The results of that experiment are presented in Table 2. A chi-square test of the null hypothesis of homogeneous distri- butions for each of the 6 days was not sig- nificant [chi-square 45 d.f.=41.16 (p= 65), non-significant]. Therefore, it was concluded that there was no difference in the distributions among the days. Tests of the equality of the probabilities of occu- pancy for each disc by each snake were then undertaken. Chi-square tests for days 2, 3, 4, 5, 6 and 7 were 44.41, 35.45, 45.16, 90.00, 52.50, and 92.00 respectively. Each of these chi-squares is significant. Therefore we concluded that the probabilities of occu- pancy for the discs were not the same for each of the 10 discs tested. The next experiment in this series con- sisted of releasing 9 female ringneck snakes in an arena containing dry sand with 10 twelve-inch discs (Table 3). The initial 46 Tulane Studies in Zoology and Botany Vol.A5 TABLE 2 Dispersal of ringneck snakes in an arena containing dry sand, 10 twelve-inch discs, 12 female snakes. Ambient temperature 75-76° F. Relative humidity 54-64%. May. q Disc Days after release 1 9) 3 4 5 6 q 8 9 10. open hidden dead 2 0 0) 0) 0 0) 0 0) 6 0 5 Il 0) 0 3 0) 0) 0 0 0) 0) 1 ri 1 2 0 0 1 4 0 il 0 0) 0 0 1 0 5 0 2 0 2 5 0 0 0) 0) 0 0 0 5) 0 0 1 0 3 6 0) 0 0 0 0) 0 1 3 0 0 0 0 2, 7 0 0) 0 0 0 0 0) 2 0 0 iL 1 0 Many tracks under discs 3, 4, 7 on day 2; snakes appear to travel in groups. Never any tracks under 5 or 6 during entire run. phase of this experiment was a duplication of the previous experiment. The findings were similar. A chi-square test of the null hypothesis of similar multinomial distribu- tions for the various days was not significant (chi-square 30 d.f. = 11.74). Similarly, chi- square tests for days 2, 3, 4 and 5 were 24.00, 24.00, 33.13, and 60.75 respectively. Each of these chi-squares is highly significant (probability <.01). Therefore it was con- cluded as above that the probabilities of occupancy for each of the discs by each snake were not the same. Additional experiments, while not de- signed primarily for testing the above hypothesis, did however reveal similar find- ings. For example, one experiment consist- ing of the release of 34 male snakes, 20 female snakes, and 2 of undetermined sex in an arena containing moist sand and 14 twelve-inch discs, demonstrated the follow- ing results (Table 4). The distributions TABLE 3 Dispersal of ringneck snakes in an arena containing dry sand, 10 twelve-inch discs, 9 female snakes. Ambient temperature 76.5 + 1.5° F. Relative humidity 48-58%. June. Disc Days after release 1 2 3 4 5 6 u 8 9 10 open hidden dead Animals left where found. 2, 0 0 0 0 0) 0) 1 3 0 4 1 0 0 3 0) 0 0 0) 0 0) 0 3 i 4 1 0 0 4 0 0 0 0 0 0 1 5 0 2 0 I> 3) 0 0 0) 0 0 0 0 re (0) 1 0) 0 Day 5 animals picked up and released from center. 6 0) 0) 0 0) 0 0) 2 2. 1 3 0 0 0) a 2 0 0) 0) 0) 0) 0 4 ») 0 0 0 0 After animals picked up on day 7 arena rotated 180°; animals then released from center. fo) 1 0 0 0) 0 0 2 3 0) 1 1 0 0 9 0) 0 0 0) 0 0 0 lh 0 0) 1 0 0) Discs 3 and 8 and their underlying sand interchanged after pickup. 10 0) 1 0) 0) 0 0 0 0 5 1 1 0 14 Dises 9 and 5 and their underlying sand interchanged after pickup. 11 0 0 3 0) 0) 0 0) 1 0 2 1 0) 0 All snakes picked up and replaced with 18 males. 12 0 3 0 0) 0 0 0 0 i 0 14 0 0 13 0 4 1 0 0 0 0 0) il 1 11 0) 0 *from disc 7 # from open Z 9 TABLE 4 snakes in an arena containing moist sand, 14 twelve-inch discs, 34 male snakes, 20 female snakes, and 2 of undetermined sex. i) Dispersal of ringneck lative humidity 70-79%. June. Temperature 75 + 3° F. Re Disc Days after hidden dead 14 open 13 11 10 6 Snakes left undisturbed in locations where found. io | if release 0) 0) 19 0 io 0) 0 al 0 0 al 0) 0 17 11 0 Prairie Ringneck Snake Behavior 47 0 AN 0 io 0 11 10 16 13 it 0 13 10 0 N 0 14 15 0 0 0) 0) 0 0 At termination of experiment sand under discs remained slightly moist. 15 from one day to another were heterogeneous (chi-square = 131.61, significant at .05). An examination of the data revealed that only 5 of the discs were persistently occu- pied and the significance of the test is due to the mobility of the snakes among these discs. The chi-square test for the equality of the probability of occupancy for each of the 14 discs by each snake tested to be highly significant for each day. In another experiment, designed primarily to test the effects of crowding, the same 56 snakes referred to in Table 4 were released in an arena with dry sand with only 7 twelve-inch discs (Table 5). Again the dis- tributions observed (see days 3 and 4) were not significantly different, chi-square = 2.48. The probabilities of occupancy for each of the discs by each snake were not the same. Chi-square 6 d.f. = 63.07, significant at .O1. In a final experiment (See Table 6), 20 male ringneck snakes were released in an arena containing dry sand and 10 twelve- inch discs. Again strong aggregative tend- encies were observed when the snakes were left undisturbed. The distributions of day 2, 3, 4, and 5 did not test significantly dif- ferent. Chi-square 27 d.f. = 28.33 and again the probabilities of occupancy for the 10 discs tested to be heterogeneous [chi-square 9 df. = 286.94, significant at .01]. A random distribution of the snakes was not occurring. Snakes tended to occur in several exceptionally large groups as well as a number of small groups, with a con- tinuous range of cluster sizes being the ex- ception. Daily Re-Release From Center 1. General statement of hypothests. As earlier noted we assumed that in the absence of intrinsic aggregative drive snakes released from the center of the arena would disperse randomly. Hence, by gathering snakes each day and re-releasing them after resetting the discs, a new set of independent distributions should be obtained if the snakes were not excited by the daily capture and release. To test this hypothesis a series of experiments was conducted. 2. Results. In the first of these experiments, (See Table 7), 10 male ringneck snakes were released in an arena containing dry sand 48 Tulane Studies in Zoology and Botany Vol iS TABLE 5 Dispersal of ringneck snakes in arena with moist sand, 7 twelve-inch discs, 34 male snakes, 20 fe- males, and 2 of undetermined sex. Temperature 72.5 + 1° F. Relative humidity 60-72%. June. Disc Days after release 1 2 3 4 5 6 le open hidden dead Snakes left undisturbed in locations where found. 3 7 y M7 I, 4 2 2 10 0 0 4 4 1 Me it, 8 2 2, 10 0 0 End of day 4 all females and undetermined sex removed; continue with 34 males only, with these being released from center day 4 but left undisturbed where found on subsequent days. 5 2, 1 14 5 2; 5 2 0 0 0) 6 5 0 0 5 6 4 4 1 2 0 lh 4 1 9 5 3 3 5 3 0 0) 8 a 0) 9 4 3 3 4 4 0 0 Soil under discs still moist at termination of experiment. and 10 twelve-inch discs. The snakes were collected and re-released from the center each day. All of the snakes collected were under disc 3 on the first day and continued to return to this disc after release for 3 successive days. The results of another experiment, given in Table 8, in which 10 male ringneck snakes were re-released in an arena con- taining dry sand and 10 twelve-inch discs show that on days | through 4, disc 8 was re- peatedly sought as cover. A third experiment (see Table 6, days 6 and 7) in which 20 male ringneck snakes were released in an arena containing dry sand and 10 twelve-inch discs showed simi- TABLE 6 Dispersal of ringneck snakes in an arena containining dry sand, 10 twelve-inch discs, 20 male snakes. Temperature 76.5 + 1.5° F. Relative humidity 48-58%. June. Dise Days after release 2) (37 74° >" 6 7 8) 49" 10) open Shiddentdead Snakes left undisturbed in locations where found. 2 G @O © © 7 1 0) 1 8 0) 0 3 0) 0 0) 0 0 0 0 0 8) 15 3 0 0 4 0 0 0) 0 0 0 0 0 3} 12 5 0 0 5 0 0 @) 0 1 ] 0 0 2 10 6 0 2 (from open) After recording day 5 snakes picked up daily and released from center. 6 2, 0 0 0) 4 0) 1 1 5 4 1 0) 0) Of 0 0 0 0 5 0 0 0) 2 11 0) 0 0 After picking up snakes on day 7 tank rotated 180° before release. 8 2 0 0 0 2 2, 0 0 0 10 2; 0) 0 9 0 0) 0 0 0 0) @) 0) 3 ll 4 0 0) Discs 4 and 10 translocated, sand and all, to each others position. 10 0) 0 0 0 0 0) 0) 0 12 2 4 0) 0 Discs 3 and 9 translocated, sand and all, to each others position. ll 0 0 0 0 0 0 0) 0) 6 4 8 0 0 Snakes removed and replaced by 7 females. WP 2 0) 0 0) 0 9) 0) 1 0 I I 0) 0 13 2, 0 10) 0 2 2 0 1 0 0 O 0) 0 No. 2 Prairie Ringneck Snake Behavior 49 TABLE 7 Dispersal of ringneck snakes in an arena containing dry sand, 10 twelve-inch discs, 10 male snakes. Release of snakes from center each day. Ambient temperature 74 + 1° F. Relative hu- midity 49-55%. July. Disc : eee Days after ae release 1 2, 3 4 5 6 ra 8 9 10 open hidden il 0 0) 10 0) 0) 0 0) 0) 0) 0 0 0) 2 0 0) 10 0 0 0 0) 0) 0 0 0 0 3 0 0 10 0 0) 0 0) 0) 0) 0) (0) 0 4 0 0 10 0) 0) 0 0 0) 0 0) 0 0 Ten males from another experiment released here in place of the original ten. 5 0 0 9 0 0 0 0 0 0 0 1 0 After pickup of snakes sand from under discs 3 & 8 interchanged; disc 6 placed on old disc 3 site (now sand of 8); disc 3 placed on old disc 6 site (still disc 6 sand). 6 0 0 0) 3 0) 0) 0 3 0 0 4 0 After pickup sand scrambled and discs returned to original com- pass positions of 1-10. 7 0 0 0 ] 0 0) 0 0 0 0 9 0 After pickup sand scrambled again. 8 0 0 0 4 0 0 lar results. Discs 5 and 10 were preferred over any of the other discs. A test of the homogeneity of the occupancy distributions was not significant. Chi-square 9 d.f. = 9.66 (p=0.35). Obviously, the null hypothesis of day to day independence of the distributions was incorrect. The distributions obtained by re- leasing the snakes from a central point each day were not independent from one day to another. In fact, the snakes congregated under very few discs or all snakes congre- gated under a single disc. We also observed that on successive days the dispersion time from the center appeared to be more ab- breviated and wanderings fewer. A snake arriving near a disc that had been regularly used would tarry near that disc and attempt to burrow under it. Occasionally the snake might move off a short distance, but would soon return to the disc. In these cases olfac- tion or vomeronasal testing appeared to be the primary sense employed. There did not have to be another snake present under the disc, nor did another snake have to be seen to prompt the return to the disc. These observations along with the previous analy- sis prompted us to conclude that these data could be interpreted as the result of the trailing phenomenon that was noted by Noble and Clausen (1936). A trail used 2 0 0 0 4 0 heavily the first day seemed to be providing a strong sensory stimulus so that on sub- sequent days snakes followed this trail and came to a particular disc at the terminus of the trail. Once the disc had been used by a number of snakes the sand under it was presumably becoming saturated with some dermal or cloacal secretion or some other “conditioner.” We decided that a series of experiments should be designed to determine if the substrate was being “conditioned” so that snakes found that particular portion of substrate particularly attractive. Habitat Conditioning “Habitat conditioning” was first examined by introducing a different set of snakes into an arena containing “conditioned” sand. One would hypothesize that if the sand was be- ing “conditioned” the distribution of these snakes would follow the distribution of the snakes that previously occupied the arena. The results of such an experiment are given in Table 7. A chi-square test of the null hypothesis was not significant. Therefore, it was concluded that ringneck snakes are capable of “conditioning” sand. Next, a group of snakes that had previ- ously been allowed to disperse on an un- disturbed basis was tested by daily release from the center. These snakes had demon- 50 Tulane Studies in Zoology and Botany Vol mis TABLE 8 Dispersal of ringneck snakes in an arena containing dry sand, 10 twelve-inch discs, 10 male snakes. Ambient temperature 74+ 1° F. Relative humidity 50-56%. July. Disc Days after release L. 25235 4.5.6 i, 38. “8S. 10opens thidden: dead Released from center daily. 1 0) 0) 0) 0 0) 0 0 4 0) 0 6 0) 0 2, 0 0 0 0 0 0) 0) 5 0 0) 3 0 0 3 oY © © OO OO C€ @ FY WwW GCG Ib) © 1 (from open) 4 0 0 0 0) 0 0 0) 7 0 0 ” 0) 0) Scrambled sand after pickup; added a new 10th snake. 5 Oo © C IL © @ © © @ 7 BMuichy -© 1 (from open) Scrambled sand after pickup; added a new 10th snake. 6 0) 0) 4 0) 0 0) 0 0 4 0 2 0 0 Left sand intact, no scramble, but snakes released from center. th 0) 0) 0 0 0) 0 0) 0) il 0) 2 0 0) Accidentally omitted data on one snake. After recording left snakes where found and sand intact but sprinkled sand around disc 9 heavily with water. 8 0 0 0) 0 0) 0) 0) 0) 8 0 1 0 0) After recording left snakes where found and gave area around disc 9 a fresh sprinkling. 9 0 0 0 0) 0 0) 0) 0 8 0) Il 0) 0 strated strong aggregative tendencies (see days 1-5, Table 6). The results of this experiment are reported in Table 6 (see days 6 and 7). Although there was some disorientation and scattering by the snakes following the center releases, the distribu- tion did not test significantly different from that of the previous day [chi-square 9 df. = 9.66 (p= 0.35) ]. Substrate and disc translocations. Further experiments were conducted based upon the hypothesis that if habitat condi- tioning existed the conditioning substance was a material that could be sensed by ol- faction, taste, or vomeronasal receptors. Ac- cordingly we attempted to translocate con- ditioned substrate by scooping it up and placing it where a similar quantity of sub- strate had been removed, or by interchang- ing the substrate from beneath two discs. The results of the first of these experi- ments is reported in Table 3 (day 10). Discs 3 and 8 and their underlying sand had been interchanged the previous day. The snakes which had been showing up mostly at disc 8 were now at the immediately adjacent position: apparently they “trailed” to favored site 8, found no conditioned substrate there, and stayed in that vicinity where perhaps some attractant still remained. The next day (day 11) three snakes appeared at position 3. This position had not been occupied in the 10 previous days. On day 10, sand from discs 5 and 9 was interchanged. On day 11 disc 9 was heavily tracked as if the snakes were investigating it. Disc 9 had probably been located via trails leading to it, but in the absence of conditioned sand the snakes did not remain there. Snakes appeared un- der adjacent discs 8 and 10, probably be- cause some conditioning had developed in that area from previous days. Table 6 gives more compelling evidence that ringneck snakes respond to conditioned substrate. On day 9, discs 4 and 10 were translocated (sand and discs included). Af- ter the release many snakes appeared to start toward the old number 10 position, apparently following old trails. But by the next day no snakes were found under disc 4, even though it occupied the formerly popular site of disc 10. Disc 10, occupying the old disc 4 site, again had snakes, ap- parently having been found by some wanderers. Numerous snakes occupied po- sition 9, possibly because it had some con- No. 2 Prairie Ringneck Snake Behavior Di TABLE 9 Dispersal of ringneck snakes in an arena containing 10 twelve-inch discs, ten male snakes, dry sand. Ambient temperature 74 + 2° F. Relative humidity 50-57%. Snakes released daily from center and sand scrambled daily after pickup of snakes. August. : Disc Days after release 1 2 3 4 >) 6 a 8 9 10 open 1 0 5 0) 0 0) 0 0 0 0 2 3 2, 0 0 0) 0) 0) 0 0) 0) 2, 0 1 3 3 0 0 1 1 0) 0 ] So 0 1 4 0 Il 3 0 0 ] 0 0 4 1 0 5 2, 0 0 0) 0) 0) 0 4 1 3 0 6 0) 2 0) 0) 0) 1 0) 6 0 0) il ditioning and was next to the original disc 10 location. On day 10, discs 3 and 9 were interchanged (disc and sand included). On day 11, disc 9 was popular, even though it occupied the original site of 3 which had never been used. The results of these trans- locations present clear evidence that the substrate must retain attractant material. Therefore, an additional experiment was conducted to identify the respective roles of the sand and disc in the retention of the “conditioner.” The results are given in Table 7. On day 5, the sands from beneath discs 3 and 8 were interchanged. Disc 3, from the popular area, was placed on the original site of disc 6, disc 6 was placed on top of the sand from number 8, and disc 8 was placed atop the translocated disc 3 sand. On day 6, conditioned sand from 3 was occupied. Disc 3 (the conditioned disc) was unoccupied and disc 4 next to the old conditioned sand site, was now occupied. From the foregoing results we concluded that sand is a strong retainer of the con- ditioning material, whereas painted wooden discs are essentially non-absorbent. Also, snakes are not likely to go to a disc merely because it once covered conditioned sand. Scrambling of substrate. The previously described results showed that an undisturbed substrate would pro- duce aggregations of snakes and that snakes seemed to find sites via olfactory, vomero- nasal, and visual stimuli. Final evidence that an attractant was present in the pre- ferred sites was provided by picking up all snakes and discs and scrambling the sand thoroughly before smoothing the sand and replacing the discs. The attractant would no longer occur in concentrated form, and a random distribution of the snakes would be expected. Snakes would presumably aggre- gate due to the aggregative forces discussed previously, but the site of the aggregations would no longer be fixed. Table 9 shows the results of an experi- ment in which sand was scrambled daily from the first. No conspicuous aggregations showed up at any specific site but complete randomness did not occur. Aggregations of 3-7 snakes were frequent, reflecting the in- trinsic aggregative drive. Table 7 (days 7 and 8) shows the results of another experiment. After a conditioned substrate was scrambled disorientation of snakes was conspicuous, and numerous in- dividuals were in the open. Similarly in another experiment (see Table 8), an ag- gregation established under disc 8 for four days was disorganized on days 5 and 6 after the substrate was scrambled on days 4 and 5. Although these experiments were gen- erally consistent, contradictory results oc- curred when despite daily scramblings at the ends of each of the first three days, disc 10 remained popular (Table 10). The intro- duction of a new group of snakes at the end of day four, along with scrambling, did not break this pattern. Strangely enough the aggregation broke up on day 6 though no scrambling had occurred. The significance of this is unclear to the investigators, but it should be noted that these snakes were huddled in the open. Checks on Extraneous Bias in Arenas The snakes involved in different experi- ments did not necessarily select discs at the same compass position for their favored cover. However, there remained the pos- sibility that, during a given experiment, drafts, light variations, etc., could be lo- 52 Tulane Studies in Zoology and Botany Volais TABLE 10 Dispersal of ringneck snakes in an arena containing dry sand, 10 twelve-inch discs, 10 male snakes. Ambient temperature 74+ 1° F. Relative humidity 49-55%. Snakes released from cen- ter each day. July. Disc Days after release 1 2 3 4 5 6 it 8 9 10 open hidden it 0 0 it 0) 0 0) 0 2 1 5 1 0 After pickup sand scrambled. 2 0) 0 0 0 0 0) 0) 4 0 5 1 0) After pickup sand scrambled. 3 0) 0 0) 0 0 0 0 0 10) 8 2 0 After pickup sand scrambled. 4 0 0 0 0) 0 0) 2; 0 0 a ] 0 After pickup snakes replaced with 10 males from another experiment. Sand scrambled. 5 0 0 0 0) 0) 0 2, 0) 0 4 4 0 Sand left intact after pickup. 6 0 0) 0) 0 0) 0) 0 3 iL 0 6* 0) Sand left intact after pickup. Tank rotated 180° before snakes released. 1 0 0 0 0 0 0) 0 i 0) 0) 7 i 8 0) 0 8 0 0 0) 0 1 0) 0 1 0 * Five snakes huddled near disc 10, the “favored disc.” calized, or that snakes might have some in- trinsic tendency for compass orientation. Such effects were checked by simple 180° rotation of the arenas. Table 3 (days 8 and 9) shows that snakes continued to return to their usual discs, even though the discs were 180° from their old compass positions. Table 6 (day 8) shows a similar return to usual haunts. The day after a 180° rotation (Table 10, day 7), a possible compass orientation appeared. Some snakes were huddled near disc 3, the position of the previously popular disc 8. However, most snakes stayed near disc 10, a disc popu- lar for several days previous. On day 8, a large concentration of snakes at disc 3 represents a return to the compass area closer to the area popular through the days prior to ro- tation. Effects of Environmental Modifications Field observations have indicated that ringneck snakes are usually found at times when the upper strata of soil are moist, or if the immediate surface layers are dry the soil under stones is moist. Therefore, our early experiments were conducted with the arena substrate moist. The substrate was moistened with a sprinkler before setting the discs into position. After several days the exposed soil became dry, whereas soil under the discs retained the “pleasantly moist” feel for a week or more. The term “pleasantly moist’ conveys our impression that the soil felt cool to the touch and had a texture associated with dampness. Bouyou- cos moisture blocks used in later experi- ments provided quantitative indices of soil moisture without disturbance of the sub- strate. In experiments where the arena was pre- moistened (see Tables 4 and 5), relatively high percentages of animals were in the open for the first few days of observations. But with the passage of time and the dry- ing of the substrate more animals retreated beneath the discs. This behavior suggested that moist soil provided an optimal habitat, and that as exposed soil dried it became less attractive than the soil beneath the discs. Obviously, another interpretation for this behavior is that snakes find excessive mois- ture undesirable and thus tend to avoid the high moisture level under the discs. Field observations have consistently 1n- dicated that small snakes are not common under cover objects during very wet and rainy periods. This suggests that there may Prairie Ringneck Snake Behavior N No. AIp O[-9 Ap c-T 4sroul ¢-[ 4srour 0) 0) 10) 61 PI € 8 €1 € € 10) 0) € cI 01-9 Aip 01-9 Arp c-T stoul c-T 3sroulr 0 0 I 6 ST ¢ 6 91 V 0) G 0 I FI ‘sosIp Jopun apyurtds Jou prp ynq ¢-T apts pepyulids sayvus Jo staqumu pue SUOTRIOT SUIPAOIII IVY OI-9 Arp OI-9 Arp c-T 3sroul C-T youl ll 0) 0 cl S V G & al 9 GL V G Salt OI-9 Arp STAAL ayes }Stoul IL 0 0) OT 9 il hs 9 Jann 6 9 I c GI OI-9 Atp ¢-T jsroul }STOUL 0 0) 0) Pal I Il CI 8 Or ¢ 0) S 9 Lit OT-9 Arp c-T 4sroul }StOUL 0) 0) 0 VG Il I cl 8 tS G I ll G OL ‘ulay} topun apyuttds 0} sosip Sufi] “G-T sostp YA Fey papyutids soyeus Jo sioquinu pue SUONRIOT BUIPIODIL 1OIFY Ap }SLOUL 0 0 0} el 9 I ZI Cees ZI 8 Ca ans 8 AIp }STOUL 0) 0 0 OI 8 ll OT V b 9 8 I yb ) }StOuL }STOuL I G 0 LI I il 6 ¢ Mk 6 0 G eT P }StOuL }StOur O g ¢ GL Jb G P 0 P OL 0) 0 8 G [ros osIp peep uappry uado 9 6 g L 9 GC - 3 z T asvo]ot uodo Japun [fos Io}fe SALC] OSTC] ‘qunf{—AvJy ‘sayeus Suljpuey yoy WM sdnois osivy JO SuUNOD oyerNI9R Jo AyNoYFIp 0} enp syuNoo ur so~urdososiqT ‘skep yuanbasqns uo punoyz d1oYAr Zo] Woy} “1o}U9. WOIF ASvaTAI [RYIUT ‘pojou zou Ayprumy sayrpy “DO oT + 1G eNye1oduiay, ‘sexes YOq FO soyeus LQ ‘SoSIp Your-aajoay OT ‘purs pousystowoid Surureyuoo vuare uv ul soyeus Yoousurt Fo [esiodsiq] TT @1avL 54 Tulane Studies in Zoology and Botany 605 ON Ye N ha a > If 2 50-4 nee SR / = we” N / - 7 \ / E 40) \ = aiUa| 7 < jae N J Fea < TS \ 304. \ Yo a 1\ cam Ly We +S) = . 7 Va \ = 204 g = SS \ NN 10 Y A eno i alt 4 L 1 I 0 2 3) 4 5 6 7 day Figure 2. Response of snakes to excessive moisture conditions. On day 0 approximately same number of snakes on both halves of tank. At that time side with discs 1-5 (damp) was sprinkled with water, both around and under discs. On day 5 when both halves of tank had reasonably similar numbers of snakes the area around the discs on side with 1-5 was sprinkled. Side with discs 6-10, labeled dry, had slight moistness under discs until day 5. on occasion be too much moisture. In na- ture, of course, recourse to drier subterranean haunts is possible. In our arenas snakes may have selected the surface as an avoidance reaction to excess moisture despite the ex- posure to daylight because there were no dry deeper strata. In an experiment involving a large num- ber of snakes in a pre-moistened arena, the animals were well dispersed over the first few days. Only at the start were animals observed in the open or burrowed (Table 11). At the end of day 8 thirty-three snakes were on one side of the tank under discs 1-5 and 34 snakes were on the other side under discs 6-10. At this time the side con- taining discs 1-5 was re-sprinkled, both around and under the discs. This moistening was followed by a rapid exodus of snakes from the freshly moistened side to the drier side. After four days conditions became similar and both sides of the tank harbored approximately equal numbers of snakes. A second sprinkling of the side with discs 1-5 on day 5 again produced the exodus. Figure 2 graphically depicts the changes of numbers in relation to sprinkling time. Our data on actual soil moisture levels are limited at this time but they suggest that moisture content of soil is a habitat variable that can lead to modified aggregative be- havior. Volwts TABLE 12 Dispersal of ringneck snakes in an arena con- taining moist sand, 5 twelve-inch discs, 20 fe- male snakes. Ambient temperature 25 + 0.5° C. Room non-air conditioned, humidity high but not measured. June. Disc Days after release 1 2 3 4 5. open hidden 1 oO @ i ww @ 8 6 2 OP ORO 2 eal 2 15 3 Oro 10:50 5 12 4 OO AO 6 1? Animals removed from burrows at end of ex- periment. Six animals clustered in one group, three in another group, and three in a third group. Soil in burrows “pleasantly moist.” All groups between discs 2 and 3. In experiments conducted in a room with- out air-conditioning evaporation from ex- posed soil was almost nil. Table 12 shows results from one such experiment. Numerous animals remained in the open but many others burrowed into the soil. Burrowing might provide a desired level of darkness as well as moisture. Even while burrowing the snakes tended to form groups, showing that aggregative drive occurs even under these circumstances. Thus we see that under high moisture levels the discs are not essential for cover. No alternative situations to high moisture levels were provided so a plausible explanation for the occurrence of exposed snakes as well as burrowing snakes is not available. Studies on terrarium-confined Dzadophis by Clark (1968) showed that the snakes would burrow. They selected, in fact, a soil more moist than a supposedly more fos- sorial species, Carphophis amoenus. Clark noted, however, that the more moist soil tended to clump, and facilitated penetra- tion by Diadophis, Clark’s substrates were sand and clay loam mixtures; we had no soil clumping problem with our sand sub- strate. In another trial we tried “seeding” cer- tain discs with marked snakes to determine whether sexual attraction occurred or if snakes simply went to a disc that had been seeded. This trial was run in September, at which time sexual drive probably was mint- mal. On the first two days of the pickups, before the seeding began, we found that No. 2 Prairie Ringneck Snake Behavior 55 TABLE 13 Dispersal of ringneck snakes in an arena containing dry sand, 10 twelve-inch discs, 9 male and one female snakes. Maintained at 73 + 2° C. Relative humidity 73% first day, thereafter 55- 60%. Snakes released from center daily except where otherwise noted. Sand under discs smoothed at time of daily pickup. September. } Disc = Days after — release 1 2 3 4 5 6 fl 8 9 10 open 1 0 0 Bae 0 0 0 0 5 0 jis 2 0 0 562 0 0) 0 0 ( 0 1 3 Only males released from center; female placed under disc 6. 3 0 0 0) 0 0 3 0) 0 0 0 Bee Qe Only males released from center; female placed under disc 1. 4 662 0 0) 0) @) 0) 0) 0) 0 0 3 Only males released from center; female placed under disc 4. Males not released until day 6. 8 0 0 0 762 0 0 0) 0 0 0) yy 9 No records taken; only 8 males released frcm center, along with the female. One male placed under disc 5. 10 0) 0 0 3 2 0 0) 0 Q 2 2 Released 8 males from center, placed one male under disc 2 and the female under disc 7. 11 0 iL 0 0) 0 0 Lee 0 0 2, 5 * Only one male near 2 males aggregated with the female (Table 13). Subsequent observations after the seed- ing began showed no consistent patterns, regardless of whether the disc was seeded with a male or female, or both were em- ployed (though under separate discs). Ag- gregations seen on days 4 and 8, involving the female, could have been sexual or could have been reflections of the tendency to come to a regularly used shelter. However, no one disc ever became popular in this experiment. On day 9 the female was re- leased from the center with several males, and a single male was “seeded” under disc 5. By day 10 one male had joined the “seed” male under disc 5, but the female was 1so- lated. Day 3 results also show her essentially isolated. Whether the same results might be obtained during the spring breeding season remains for subsequent experimentation. DISCUSSION The formation of aggregations presum- ably confers some benefit to ringneck snakes. The advantages might include ease of find- ing sexual partners, reduction of exposed body surface, with reduced moisture loss and perhaps moderation of temperature change due to weather changes, and perhaps sensual satisfaction. Sensual satisfaction actually in- cludes a host of taxes which are more likely the mechanisms responsible for initiating aggregation rather than a result of the grouping. Clausen (1937) showed reduced oxygen consumption in aggregated Storeria dekayi but the advantages of this are not clear. Reduced Oz consumption could re- flect lowered metabolism, hence lower energy requirements. Certainly if the snakes were crowded under a stone or buried in the ground, O» would be limited. However, the animals seemingly are not forced to aggregate in mature to conserve Oz since numerous cover stones are available. From our experiments some factors can be discounted as the primary cause of ag- gregative behavior. For example, in our experiments no need for oxygen was evident because in most situations the snakes had ample cover of equal quality available but not utilized. In addition, extremely skewed sex ratios occurred in our field collections (six males to one female) suggesting that sex drive may play a fundamental role in aggregation. In our experiments sexual drive was ruled out as an advantage of aggregation 56 Tulane Studies in Zoology and Botany to ringneck snakes because it occurred among males in the absence of females. Henry Fitch (personal communication ) con- firms this sex ratio from his field studies in northeastern Kansas. He has observed that there may be a slight increase in the rela- tive abundance of females in the summer, however the number of female snakes found then is far too low to indicate an actual sex ratio of 1:1. In future collecting, a sum- mary of actual sex ratios for each aggrega- tion found in the field would be instructive. Tentative observations in the field (April 1968) indicate that many groups of five aggregated adult snakes are wholly male and a group of 10 was 9 male and one female. The lack of females in many of the aggre- gations tends to further confirm our in- terpretation that sexual attraction is not a factor in aggregation. It should be noted here that sex ratios in the populations of various races of Dziadophis punctatus re- ported by Blanchard (1942) range from 0.7:1 male to female to 3:1 (our interpola- tion). The extremes were based on small samples (under 17 animals); larger samples generally ran 1:1 to 2.4:1 (north-central Kansas) and 1.1:1 (northeastern Kansas). A possible basis for aggregative drive is moisture conservation. Noble and Clausen (1936), Carpenter (1952), and others have observed that snakes tend to gather when humidities are low, possibly congregating in those places offering adequate or partial shelter from dehydration. Noble and Clausen also found that Storeria dekayi selected rela- tively high humidities when offered a choice and formed aggregations more readily under drier conditions. Bogert and Cowles (1947) have shown that the resistance to desiccation of reptiles may be correlated with habitats occupied. We have limited evidence that ringnecks become more gregarious when pre-moistened arenas begin to dry, but also have noted that aggregations form even when snakes are provided with drinking water and when ample soil moisture and high humidities prevail in the arenas. Increased temperature can also produce higher metabolic rates and therefore in- creased water loss. Noble and Clausen (op. cit.) found that increased temperatures were accompanied by increased breathing rates and more moisture loss, and that under Vol wis the elevated temperature conditions aggre- gations were more likely to form. The possibility that reduced oxygen con- sumption might be advantageous should be mentioned again here. The use of more oxygen can be an indication of elevated metabolic rates, hence more loss of metabolic water. When Noble and Clausen did their studies it was believed that reptile integu- ments were essentially impermeable. The re- cent finding by Bentley and Schmidt-Nielsen (1966) that substantial loss of moisture may occur through the integument points up a need for careful analysis of moisture loss in relation to environmental conditions. Integumentary vs. respiratory loss notwith- standing, total moisture loss still remains the significant criterion. Fitch (1956) reported numerous body temperature records for ringneck snakes found in the field. The most favored temperature seemed to be around 27-29° C, several de- grees higher than our laboratory arenas. A laboratory temperature gradient response study by Clark (1968) closely agrees with Fitch’s values. Fitch’s records appear to be daytime readings (hour not stated) and body temperatures were generally 6-7° higher than optimum air temperatures. The values were not correlated. Since the other environmental variables were not quantified we cannot at this time know if temperature plays the more prominent role in stimulat- ing aggregative drive, but aggregation seems to characterize ringnecks in the field whether the day be warm or cool. Seibert and Hagen (1947) found that Thamnophis radix and Opheodrys vernalis selected cover on an availability basis, but in hot weather they would avoid metal and go under cover that offered more effective insulation. The observed aggregational phenomena in our arena were most pronounced during the daylight hours; tracks on the substrate and the different locations at which marked individ- uals were noted prove that wandering was prevalent at night. In mature nocturnal movements permit activity at times when the humidity is highest. However, in spring, when snakes are at the surface nocturnal temperatures tend to be rather low for efficient snake activity. In our laboratory neither temperature nor humidity changed noticeably during the night, only the absence No. 2 of negative phototaxis was a conferred ad- vantage. The aggregative behavior exhibited has been identified as an intrinsic drive since the animals not only selected each other's company but also tolerated considerable population pressure (up to 24 snakes under a single disc one foot in diameter). The strength of the drive is emphasized by the formation of groups during the first day after release. In addition, snakes consistently seemed to follow each other or the trails left on the substrate. A more dispersed distribution is illus- trated by the results given in Table 11. This experiment presented the greatest popula- tion density of any experiment (6.7 snakes/disc compared to a maximum of 4 snakes/disc in other experiments). We can- not therefore dismiss the possibility of crowding as a modifying factor in aggrega- tion. The aggregative denning behavior of large snakes obviously insures adequate winter shelter and facilitates mating after the spring emergence. Rising temperatures as the sea- son progresses are accompanied by de- parture from the den. This is adaptive be- cause increased temperatures mean higher metabolic needs, and necessitates more food than is ordinarily available in the vicinity of dens. Large snakes do not as a rule “clump” at the den site, usually they are separated while exposed to the sun. The aggregation of ringnecks occurs dur- ing the springtime at the same time that large snakes are still at the den. However, the ringnecks do not disperse appreciably to receive warmth. The paucity of surface activity records prevents us from knowing if ringnecks disperse during the warmer parts of the year. Thorough explanation of distributional phenomena will not be pos- sible until adequate data on responses to environmental variables and quantification of the physiological requirements of ring- necks are available. Regardless of what stimuli foster or hasten aggregation of ringneck snakes, and the ad- vantages of such aggregations, our experi- ments have demonstrated that these snakes show a definite preference for substrates that have been traversed and occupied by other ringnecks. Ringneck snakes have shown an ability to locate other ringneck Prairie Ringneck Snake Behavior 57 snakes, apparently by means of visual, ol- factory, or vomeronasal sensing. These snakes can apparently follow trails on the substrate and recognize soil that has regularly har- bored other ringnecks. The ability of ring- necks to detect each other probably is an inherited trait, not unlike the chemical cue perception for food organisms reported for newborn snakes by Burghardt (1967). Noble and Clausen (1936) found that Storeria dekayi can detect areas where skin alone from other Storeria snakes has been rubbed over clean glass. This is a truly re- markable chemical perception since the skin of snakes is said to be non-glandular and any secretions left as a snake crawls along must be miniscule. The habitat conditioning that results from occupancy of a substrate by ringneck snakes appears to serve purposes other than those previously assigned to various animals. In a discussion of environmental conditioning presented by Allee, et al. (1949), the major functions of conditioning were given as: detoxification, alteration of fecundity, pro- duction of growth promoting substances, and physical alteration rendering the sub- strate more porous or penetrable. In ring- necks it appears that conditioning may provide cues to other ringnecks that the conditioned site is, or was, a desirable site to occupy, at least in the field. Our experiments attempted to nullify habitat selection but the experiments may have inadvertently been conducted within the environmental range evoking aggrega- tive drive. Nevertheless, snakes re-released daily could have gone elsewhere to form their aggregations, and had every opportu- nity to use visual or other means to follow their kind. In natural settings ringnecks may use the same cover object year after year because it offers optimal microhabitat conditions. There is little reason to doubt, however, that the site may be conditioned habitat whose use is perpetuated on a con- tinuing basis. Little is known of the homing instinct in snakes but their ability to return year after year to a den site from great distances suggests that den sites may be conditioned habitat. Regular routes are evident (Klauber, 1956). Whether the use of such routes in- volves a homing instinct or the trailing phenomena as exhibited by Storeria dekayi 58 Tulane Studies in Zoology and Botany and ringneck snakes is not clear. The paral- lels between denning and aggregation are striking and are perhaps manifestations of comparable instinctive drives. ACKNOWLEDGMENTS Appreciation is extended to the American Philosophical Society for financial assistance with travel and equipment costs. We also thank Dr. Robert Clarke and Mr. Stanley Roth for providing some of the snakes used in our experiments, and Miss Gay Harmann who assisted with some of the experiments. Dr. Gerald E. Gunning kindly loaned us several of his watering tanks which were used for experimental arenas. LITERATURE CITED ALLEE, W. C., A. EMERSON, O. PARK, T. PARK, and K. P. Scumupr, 1949. Principles of animal ecology. W. B. Saunders Co., New York. xii + 837 pp. BENTLEY, P. J. and K. ScHMipt-NIELSEN. 1966. Cutaneous water loss in reptiles. Science 151: 1547-9. BiancHarp, F. N. 1942. The ringneck snakes, genus Diadophis. Bull. Chicago Acad. Sci. HOD) alee Bocert, C. M. and R. B. Cowes. 1947. Mois- ture loss in relation to habitat selection in some Floridian reptiles. Amer. Museum Novi- tates No. 1358:1-34. BRENNAN, L. A. 1936. A check list of the am- phibians and reptiles of Ellis County, Kan- sas. Trans. Kans. Acad. Sci. 37:189-191. BurcHarpt, G. M. 1967. Chemical cue _ pref- Vol. 15 erences of inexperienced snakes: compara- tive aspects. Science 157:718-20. CARPENTER, C. C. 1952. Comparative ecology of the common garter snake (Thamnophis s. sirtalis), the ribbon snake (Thamnophis s. sauritus ), and Butler’s garter snake (Tham- nophis butleri) in mixed populations. Ecol. Monogr. 22:235-58. CLARK, DONALD R., Jr. 1968. Experiments into selection of soil type, soil moisture level, and temperature by five species of small snakes. Trans. Kansas Acad. Sci. 70:490-6. CLAUuSEN, H. J. 1937. The effect of aggregation on the respiratory metabolism of the brown snake Storeria dekayi. J. Cell. and Comp. Physiol. 8:367-377. Cramer, H. 1946. Mathematical methods of statistics. 10th Printing 1963. Princeton Univ. Press, 574 pp. Fircu, H. S. 1956. Temperature responses in free-living amphibians and reptiles of north- eastern Kansas. Univ. Kansas Publ., Mus. of Nat. Hist. 8(7):417-76. Hurter, J. 1911. Herpetology of Missouri. Trans. Acad. Sci. St. Louis 20(5):59-274. KiauBer, L. M. 1956. Rattlesnakes. Their habits, life histories, and influence on man- kind. 2 vols. xxix + 1476 pp. Univ. Calif. Press, Berkeley and Los Angeles. Nose, G. K. and H. J. Cxiausen. 1936. The aggregation behavior of Storeria dekayi and other snakes, with especial reference to the sense organs involved. Ecol. Monogr. 6:269- 316. SErBERT, H. C. and C. W. HaAceEn, Jr. 1947. Studies on a population of snakes in Illinois. Copeia, 1947, no. 1:6-22. Wiwpe, W. S. 1938. The role of Jacobson’s organ in the feeding reaction of the common garter snake Thamnophis sirtalis — sirtalis (Linn.). J. Exptl. Zool. 77:445-465. December 23, 1968 A NEW TURTLE SPECIES OF THE GENUS MACROCLEMYS (CHELYDRIDAE) FROM THE FLORIDA PLIOCENE JAMES LL. DOBIE Department of Zoology-Entomology Auburn University, Auburn, Alabama 36830 ABSTRACT Remains of specimens which appear to represent a new species of Macroclemys were excavated recently from a mid- Pliocene deposit at McGehee’s Farm in Alachua County, Florida. The new species named Macroclemys auffenbergi was compared with M. schmidti, Miocene of Nebraska, and with the Pliocene-to-Recent species, M. temmincki. Macroclemys schmidti is presumably the progenitor of M. temmincki. This presump- tion is based on distributional data and on the ages of the known fossils. Presumably, M. temmincki gave rise to M. auffenbergi. The causative factor in the extinction of M. auffenbergi was perhaps the lowering of the water table during the mid-Pliocene. INTRODUCTION Pliocene remains of Macroclemys were excavated recently from a deposit at Mc- Gehee’s Farm near Newberry, Alachua County, Florida, by staff members of the Florida State Museum. This is the first rec- ord for the genus from a Pliocene deposit in the southeastern United States although Pliocene representatives have been recorded by Zangerl (1945) and Hibbard (1963) from South Dakota and Kansas, respectively. The assemblage of material from the McGehee site was compared with Pleisto- cene remains held by the University of Flor- ida (UF) and with recent specimens de- posited at Tulane University (TU), at the University of Michigan Museum of Zoology (UMMZ), and in the author's personal collection (JLD). Measurements by Zangerl (op: cit.) for the Miocene species, 1. schmidti, Chicago Natural History Museum, (CNHM, P26014) and for the South Da- kota M. temmincki (CNHM, P15823) were compared with the fossil remains from McGehee’s Farm. In addition, two skull measurements from each of these two turtles were kindly supplied by Dr. Zangerl (per- sonal communication). Hibbard’s data (op. cit.) on the shell of the Kansas Pliocene M. temminckt, University of Michigan Museum of Paleontology (UMMP, 47109) were compared with the McGehee material. The Pliocene remains from Florida appear to represent an undescribed fossil species of Macroclemys. | propose that it be named Macroclemys auffenbergi in honor of Dr. Walter Auffenberg for his contributions to our knowledge of fossil and recent turtles. Holotype: University of Florida (UF) 10992. A nearly complete specimen (Fig. 1A-C) collected by Robert Allen in Decem- ber, 1963. The carapace of the holotype lacks peripherals 8-11 on the right side, costals 3, 4, 5, and 8 on the left side, and parts of costals 6, 7, and 8 on the right side. The carapace is similar to the recent species in that three supramarginals are present on the left and right sides of the carapace and because scute sulci are the same. The cara- pace is approximately 455 mm _ long, straight line measurement. The skull lacks all bones except for the two parietals, the right quadratojugal, the left squamosal, a piece of the supraoccipital and a fragment which appears to be part of the quadrate. The right side of the mandibular ramus lacks all bones except the dentary, the left ramus is entire. The rest of the holotype is entire except for caudal vertebrae and vari- ous elements of the fore and hind feet (Garyey 1a) Paratypes: The paratypes consist of 16 humeri (UF 9224-9228, UF 13051-13061), a basal phalanx of the third finger from a fore foot (UF 13062), two basal phalanges of the third fingers from the hind feet (UF 13063, UF 13064), an incomplete carapace EDITORIAL COMMITTEE FOR THIS PAPER: Dr. RAINER ZANGERL, Chief Curator, Department of Geology, Field Museum of Natural History, Chicago, Illinois 60605 Dr. WILLIAM W. MILSTEAD, Professor of Biology, University of Missouri, Kansas City, Missouri 64110 60 Tulane Studies in Figure 1A. Holotype remains of M. auffen- bergi, UF10992. Zoology and Botany Vol. 15 \ b AF 0792 Figure 1C. View of the fifth through eighth costal bones of M. auffenbergi (holotype, UF10992). Figure 1B. A_ photograph showing that humeri ( holotype, UF 10992; paratype, UF9228 ) and phalanges (holotype, UF10992) of M. auffenbergi are wider than the same skeletal elements in M. temmincki (JLD 4 Florida). and plastron (UF 9242), a nearly complete skull, with an entire mandible (UF 11053), and two partial skulls (UF 9198, UF 9199; Bro elD)): Type locality and horizon: McGehee site; McGehee Farm, T9S, R17E, S14 of NW, Sec. 22, located approximately 120 yards east of State Hwy. 45 and 3 miles north of Newberry, Alachua County, Florida. The lo- cality is a mid-Pliocene deposit of Hemphil- lian age (Webb, 1964). Diagnosis and description: As the hu- merus of the new species increases in length (width measured at a point halfway down the length of the humerus), the width, length ratio also increases, i.e., width in- Figure 1D. Dorsal view of skulls of Para- types UF9198-9199 and UF11053 of M. auf- fenbergi. creases more rapidly than length. This ratio decreases for M. temmuinckti, i., length increases more rapidly than width. The former is positive allometry, the latter is negative. The largest ratios are found in the larger animals for M. auffenbergi and in the smaller animals for M. temmuincki (Fig. 2A). There is no overlapping of ranges in width/length ratios for phalanges of digits of the fore and hind feet (Fig. 2B). The difference among the three species in the relative length of the alveolar surface of the upper jaw is apparent in Figure 2C. However, Dr. Zangerl (personal communi- cation) stated the following for M. schmidti, “the estimate of the total length: posterior PLIOCENE = @ 200 pretstoceNe = ° RECENT = © HOLOTYPE = A 180 = j= oO Z. (es) — -160 \ PLIOCENE CORRELATION AT MORE THAN .0005 = e = e Fr 421 Q 140 = 7 -0917 + .0008x 17 HUMERI MEASURED PLEISTOCENE AND RECENT 120 CORRELATION AT MORE THAN .025 Fe 8.11 Y = .1846 + (-.0017)X 12 HUMERI MEASURED 0 20 40 60 80 100 120 140 LENGTH OF HUMERUS A Figure 2A. Regression analyses of ratios of width/length for humeri of M. auffenbergi and M. temmincki. M.temmincki(§) length of phalanges: 14— 3] M.ausrenbergi(3) length of phalanges: 25.5—31 380 400 420 440 460 480 RATIOOFWIDTHALENGTH OF 3rd PHALANGES OF HIND FEET M.temmincki(]) M.ausyenbergi(2) length of phalanges:11.5 28 4 0 RATIOOFWIDTHALENGTH OF 3rd PH ALANGES OF FORE FEET B Figure 2B. Ratios of width/length for the basal phalanges of the third fingers for the fore and hind feet of M. auffenbergi and M. temmincki. length og phalanges:10- -24.5 end of the parietal to front end of pre- frontal is, of course, a wild guess based on the position of the occipital condyle.” The two points for the M. schmidti were cal- culated using the two “estimated” lengths supplied by Dr. Zangerl. This datum is un- available for the Pliocene skull from South Pliocene Turtle 61 MIOCENE= A PLIOCENE= @ PLEISTOCENE = @ RECENT= * Alveolar Length Parietal To Prefrontal Length 1 90 110 130 180 170 190 PARIETAL TO PREFRONTAL LENGTH C Figure 2C. Ratio of alveolar length (posterior end of alveolar shelf to anterior attachment with the premaxilla) divided by length from the posterior end of the parietal to the anterior end of the prefrontal for skulls of M. schmidti, M. auffenbergi and M. temmincki. 70) PLIOCENE = @ PLEISTOCENE = @ RECENT = ®& °o HOLOTYPE = A 60 50 40 30 WIDTH OF 7thCOSTAL 20 30 40 50. = 60 WIDTH OF 6thCOSTAL D 70 Figure 2D. Widths of the sixth costal bones of the carapace for M. auffenbergi and M. temmincki. Dakota (Zangerl, personal communication ). The new species is separable from M. temmincki on the basis of one character of the carapace: in the former, the sixth costal is wider than the seventh (Fig. 2D). The eighth costal bones of the holotype and paratype UF 9242 are broken. Comparison 62 Tulane Studies in Zoology and Botany | =e =| = = Figure 3. Former and present distribution of the genus Macroclemys. The + is the M. schmidti record from the early middle Miocene of Nebraska; A and B are records of the South Dakota (early Pliocene) and Kansas (late Pliocene) M. temmincki; C is the Florida (mid- Pliocene) M. auffenbergi; I and II are Pleistocene records for M. circles are recent M. temmincki records. of M. auffenbergi with M. temmincki from the upper Pliocene of Kansas (Hibbard, op. cit.) is not possible. DISCUSSION Formerly the genus Macroclemys was more widely distributed than at present (Fig. 3). I suspect that M. schmidti, from the early middle Miocene of the northwest- ern corner of Nebraska, was the progenitor of the early Pliocene (Clarendonian ) -to- Recent species, M. temmuincki. 1 base this on the following facts: (1) the South Da- kota deposit is close geographically to the early middle Miocene deposit in Nebraska; (2) the South Dakota deposit is early Plio- temmincki; solid dark cene; the Florida site (Hemphillian age) is middle Pliocene. Macroclemys temmincki is thus older than the new species and _ pre- sumably gave rise to M. auffenbergi. The Pliocene river(s) inhabited by M. auffenbergi was probably a “typical” river with a fauna similar to those that now popu- late southeastern rivers. The two new fossil species of Chrysemys (= Psendemys) (Rose and Weaver, 1966) discovered at the Mc- Gehee deposit support this view. It seems probable, in view of the close structural similarity between these turtles, that habitat requirements for M. aaffen- bergi were similar to those of M. temminckzi, i.e., thoroughly aquatic, sedentary, omnivor- No. 2 ous, and necrophagous (Dobie, 1966; un- published dissertation ). The question arises as to why M. tem- mincki has persisted, while M. auffenbergi has become extinct. Perhaps the extinction of the latter resulted from a lowering of the water table during a period of arid Pliocene climate. Macroclemys temmincki survived because its range included the Mississippi River and certain major tributaries, that per- sisted through this arid period. Acknowledgments: I appreciate the re- view of this manuscript by Robert H. Mount of Auburn University and the loan of Plio- cene and Pleistocene specimens by Walter Auffenberg of the Florida State Museum at the University of Florida, Gainesville. Pliocene Turtle 63 LITERATURE CITED Doster, J. L. 1966. Reproduction and growth in the alligator snapping turtle, Macroclemys temmincki (Troost). Unpublished Ph.D. Dis- sertation, Tulane University. Hispparp, C. W. 1963. The presence of Mac- roclemys and Chelydra in the Rexroad Fauna from the upper Pliocene of Kansas. Copeia, (4):708-709. RosE, F. L. and W. G. WEAVER, JR. 1966. Two new species of Chrysemys (= Pseudemys ) from the Florida Pliocene. Tulane Stud. Geol., 5(1):41-48. Wess, S. D. 1964. The Alachua Formation. In Guidebook 1964 Field Trip. Soc. Vert. Paleontology in Central Florida. University of Florida, Gainesville. ZANGERL, R. 1945. Fossil specimens of Macro- chelys from the Tertiary of the Plains. Fieldi- ana: Geol., 10:5-12. December 23, 1968 STUDIES ON FROG TRYPANOSOMIASIS II. SEASONAL VARIATIONS IN THE PARASITEMIA LEVELS OF TRYPANOSOMA ROTATORIUM IN RANA CLAMITANS FROM LOUISIANA RALPH R. BOLLINGER, JOHN RICHARD SEED and ALBERT A. GAM Laboratory of Parasitology, Department of Biology Tulane University, New Orleans, Louisiana 70118 ABSTRACT Seasonal variations in the parasitemia level of Trypanosoma rotatorium infections of Rana clamitans are described. The sea- sonal cycle of parasitemia is not influenced by the sex of the frog. Preliminary data also suggest that in frogs over 5.1 cm long, the age of the frog had no apparent relation to the percentage of infected frogs or the parasitemia level. Various factors, i.e., water temperature, mating cycle, changes in photoperiod, which might in- fluence the seasonal cycle are discussed. The importance of these results in epi- demiological studies of parasitic infections is also discussed. INTRODUCTION Numerous species of trypanosomes have been described from various species of frogs in the United States. Much of the early taxo- nomic work has been reviewed by Diamond (1965). He also discussed the biology of trypanosomes infecting frogs, including methods of cultivation, vectors involved in transmission, laboratory maintenance of frogs, vectors, and so forth. Nigrelli (1945) described a morphologi- cal type of trypanosome infecting Rana clamitans. He identified this species as Tryp- anosoma rotatorium (Mayer, 1843). This species is the one reported in this paper. Diamond (1965) believes that T. rotatorium (Mayer, 1843) has not been described from frogs captured in America. He did not, how- ever, name the species which has been re- garded as T. rotatorium by Nigrelli (1945). Until more concrete evidence regarding the true identity of T. rotatorium (Mayer, 1843) of Nigrelli (1945) becomes available, we will continue to identify this trypanosome as T. rotatorium. There have been few or no reports on the incidence or intensity of T. rotatoriwm infections of R. clamitans. The work to be reported here describes the parasitemia levels of T. rotatorium infections of R. clamitans captured near Norco, Louisiana throughout a 11% year period. MATERIALS AND METHODS The frog, Rana clamitans, was collected throughout the year from swamps in the Goodhope oil field near Norco, Louisiana. It was maintained in glass aquaria and ex- posed to the normal daily fluctuations of light. Subsequent to its capture, each frog was weighed, measured from the tip of snout to vent, and if possible its sex determined from external characters. To determine para- sitemia levels, a vein in the upper corner of the jaw was punctured, blood extracted with a micro Pasteur pipette, and placed on a microslide. Slides were examined under low power (125 X) with the light microscope. Frogs were examined within 3 days of cap- ture, almost always between the hours of 12:00 noon and 7:00 p.m. This is the period of maximum parasitemia level (Southworth, et al, 1968). Trypanosomes contained in a drop of blood covered with a 22 mm square cover slip were counted and averaged for a minimum of 30 low power fields. Only those fields which contained a relatively con- stant number of red blood cells were exam- ined. In addition, a drop of blood under a 22-mm-square cover slip was systematically EDITORIAL COMMITTEE FOR THIS PAPER: Dr. FRANKLIN G. WALLACE, Professor of Zoology, University of Minnesota, Min- neapolis, Minnesota 55455 Dr. ROBERT G. YAEGER, Associate Professor of Parasitology, School of Public Health and Tropical Medicine, Tulane University, New Orleans, Louisiana 70112 64 No. 2 Studies on Frog Trypanosomiasis Il 65 Figure 1. Trypanosoma rotatorium in Rana clamitans from Louisiana (1800 ). ments made on trypanosomes from stained blood smears are as follows: The measure- Posterior end to kinetoplast, 4.92 » (9); Posterior end to center of nucleus, 16.5 « (38); Posterior end to anterior end, 46.2 » (49); with including undulating membrane, 14.5 » (49); No free flagellum. The numbers in parentheses are the number of measurements made. examined from those frogs which appeared negative for trypanosomes. In one collection (30 Jan. 1967), the frogs were examined for T. rotatorium every 4 hours over a 24- hour period. Trypanosoma rotatorium was differenti- ated from the other morphological types of trypanosomes found in the fresh blood preparations of R. clamitans by their charac- teristic size, shape, absence of a free flagel- lum, and type of movement. Trypanosoma rotatorium was the largest trypanosome ob- served, generally assumes a U-shape, and moves with closed end of the U leading. For the measurement of trypanosomes, blood smears were prepared and _ stained with Giesma’s stain. Measurements were made under oil immersion with the light microscope (1250 x ). RESULTS In examining the blood of 486 Rana clamitans collected over a period of 1% years 81 per cent were found to be infected with Trypanosoma rotatorium (Mayer, 1843). The measurement and morphology of this trypanosome corresponds exactly to the elongate forms of T. rotatorium de- scribed by Nigrelli (1945) from Rana clamitans collected in New York, Connecti- cut, New Jersey and Pennsylvania. The morphology of Trypanosoma rotatorium 1s shown in Figure 1. Rana clamitans was also found to be infected with at least 5 other different and discrete morphological types of trypanosomes. The incidence and intensity of infection of these other types were low, and they will not be discussed in this report. The parasitemia levels and incidence in frogs showing the presence of circulating blood T. rotatorium are indicated in Tables 1 and 2 and Figure 2. The difference in the parasitemia level of frogs captured in the late summer (17 August 1966) when com- pared to the parasitemia level of frogs cap- tured in the winter were highly significant (Table 2). Table 3 shows that the sex of the frog does not influence the parasitemia 66 Tulane Studies in Zoology and Botany Vols TABLE 1] Seasonal variation in the percentage of R. clamitans with T. rotatorium in the peripheral blood. Date Number of Percentage of Frogs Frogs Frogs with T. rotatorium Water Collected Collected! in Peripheral Blood Temperature °C 2, Auigs (65 3 100 ND? 27 Aug. 65 10 100 26.5 30 Sept. 65 15 93 ND 27 Ock165 23 78 15.0 3 Nov. 65 29 76 ND 17 Nov. 65 7 59 15.0 1 Dec. 65 6 66 10.0 6 Jan. 66 23 13 15.0 24 Feb. 66 if 0 9.5 10 Mar. 66 71 66 14.0 17 Mar. 66 28 93 19.0 24 Mar. 66 16 94 15.0 1 April 66 20 100 ND 14 April 66 14 93 20.0 12 May 66 21 90 93.5 9 June 66 2 100 ND 8 July 66 17 100 28.0 1 Aug. 66 13 100 28.0 10 Aug. 66 29 97 26.5 17 Aug. 66 30 100 27.0 27 Sept. 66 36 100 24.0 7 Nov. 66 16 100 24.0 30 Jan. 67 10 0 ND Totals 456 = pee Average _ $1 20.0 | All frogs greater than 5.0 cm in length. - not determined levels. Frogs of both sexes showed approxi- mately equal parasitemia levels throughout the year (probability < 10%). Preliminary data in Table 4 suggest that the age (size) of the frog does influence the parasitemia level. Frogs 4.0 cm or less in length were generally negative for trypanosomes, whereas a higher percentage of frogs above 4.1 cm captured at the same period showed detect- 130} _bo un a ee 2) cy aS y O Bye LW 420 O20 g ae ; fu ¢ 2A | \\ ‘q © © D Erol . @ ho 0.1 27 Oct. 65 23 0.43 + 0.58 0-2.0 SS Vall 3 Nov. 65 29 0.20 + 0.31 Os > 0.1 17 Nov. 65 17 0.06 + 0.05 0-0.1 SS (iil 1 Dec. 65 6 0.07 + 0.05 0-0.1 051 6 Jan. 66 23 0.01 + 0.04 0-0.1 S> Ws 24 Feb. 66 il 0 0 SSO, 10 Mar. 66 71 ONBEEOIS 0-2.0 SO 17 Mar. 66 28 0.12 + 0.08 0-0.5 SO. 24 Mar. 66 16 0.21 + 0.18 0-0.6 SS OI 1 April 66 20, 0.19 + 0.12 0.1-0.5 > Onl 14 April 66 14 0.16 + 0.16 0-0.6 > 0.1 12 May 66 oil 0.33 + 0.48 0-2.0 > 0.1 9 June 66 2 2.00 + 2.70 0.1-3.9 NS 8 July 66 17 1.24 + 1.94 0.1-7.8 NS 1 Aug. 66 13 1.10 + 0.62 OI 5 10 Aug. 66 29 0.96 + 0.71 0-2.3 SY 17 Aug. 66 30 ILS) S= ILO) 0.1-3.7 — 27 Sept. 66 36 est se il 0.2—4.5 NS 7 Nov. 66 16 0.24 + 0.16 0.1-0.6 Onl 30 Jan. 67 10 0 0 SSO. 1 All frogs greater than 5.0 cm in length. - Based on student’s “T” test. The parasitemia levels of frogs captured on 17 August 1966 were compared to the parasitemia levels of frogs captured at other times of the year. ° Not statistically significant (P = < 10%) results are preliminary and require further verification, however, they do indicate (1) that negative frogs probably are not truly negative but harbor low undetectable num- bers of trypanosomes; and (2) that tem- perature is not solely responsible for the variable parasitemia levels, since the labora- tory temperature was relatively constant and warmer than field conditions. The frog sea- sonal mating cycle and/or seasonal changes in photoperiod may play a role in controlling intensities of infection. Both of these pos- sibilities are currently under investigation. There have been few epidemiological studies on trypanosome infections of frogs. Diamond (1965) reported that 27% of Rana pipiens collected from Anoka and Ramsey County, Minnesota were infected with Trypanosome pipientis. Diamond also reported a 38.5% infection of R. pipiens from Anoka County, Minnesota collected on the 22 July 1949. These percentages are considerably lower than those reported here, and suggest that the incidence of infection and intensities of infection by frog trypano- somes may vary according to the geographi- cal area and time of the year in which the frogs are examined. The presence of a 24- hour cycle influencing varied parasitemia levels of T. rotatorium in R. clamitans has been described (Southworth, et a/., 1968), and suggests that the time of day must also be considered in any analysis. In other words, to compare any two host populations of one species in similar climatic areas for the incidence and intensity of infection it would be necessary to examine all hosts at comparable daily and seasonal times. ACKNOWLEDGMENTS The authors would like to thank Dr. Franklin Sogandares-Bernal who greatly en- couraged us in this work. This work was supported in part by a research contract (DA Contract 49-193-MD- 2817) from the U. S. Army Medical Re- search and Development Command, Office of the Surgeon General and by a research grant from the National Science Foundation (GB-5235). ‘(Ad'T/'ON ) [PAZ vIutoyseied asviaAy , ‘peye[[09 sso1z JO Ioquiny ; 2 k ‘poorg [vteydiieg Ul wn110j70j0.L * J, YYIM SBO1F JO aSvyUsdIOJ « ‘JUBA 0} NOUS JO dij WOLF poInsvaW YYSUs], Apog ; S ae HOSt at st Qo) oe =o — 99 — -3) = asRIOAY aaa el ee af ae SOL Sabauae ap eo s[etor COO cSVPI+98T OOT 2 LTT+cG8T OOT 8 COMES (00a OR ORO 99 3doeg 1g 0 0 0 SOT+ S861 O01 GI COLE COL S00l er O80 SiG | OOl G O00 99 ‘sty LT CUE 680s 001 4s SciOce Zc0) TG. 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Il OOT IG Ses coil OOT Sill 99 ‘deg Lz aS 9610 == S6iT OOT €1 VOT + GOT OOT LI 99 “snV LT S LL'0 + 86'0 P6 SI 69'0 + G60 OOT IT 99 ‘snV OT om G10 + 8a'T OOT L OF'0 + 06'0 OOT 9 99 ‘sny [| v 99'0 == 9Z'0 OOT 6 IL'G + €8'T OOT 8 99 4[nf{ 8 S 6F'0 + 8f°0 v6 L SLO seo CL P 99 API ZI ~— LT O\== ZT 0 68 6 Vi OOO OOT S| 99 [Ndy FI wR CLOEGEO OOT LI 610+ L10 OOT € 99 [Edy T CLO so0 OOT GI SiO = ei 0 GL V 99 “IRIN VG S0i0 + 110 r6 LI tL 0) ere SS] 0) 16 Il 99) FUING LT SEO) Se LEO) c9 Tg cr'0 + LTO OL OG 99 JBN OT F0'0 + GO'0 SI LI 0 0 9 99 “uel 9 (Ad'T/'ON ) Joao] poo [ersyduog ur pepe[[op (Ad'T/‘ON ) [240] pooTq [eteydueg ul = ;paye]JoD pepe]fop VIULOPSLIV J WULNLLOJDJOL * J, YALAN ssOly] PIULOPISVIL WMLLOJDIOL * JT YIM ssOLy SBOLy ODRIDAY SBOI JO 95v}UDIIN0g yo Toquinyy IDVIIAW SBOIy{ JO 95v}UBIIO J jo Iaquinyy avd ssOLy o[ePUWlay SBOLY O[PIY 68 “UNLLOPDJON “J, JO [OA] VIUMopISeIVd pooyq UI UOTLLIVA [BUOSvas PUL xag © aATaV No. 2 REFERENCES DiamMonp, L. S. 1965. A study of the mor- phology, biology and taxonomy of the trypano- somes of Anura. Wild. Dis. Microcard No. 44. 3 Microfiches. NicRELLI, R. F. 1945. Trypanosomes from North American amphibians with a description of Studies on Frog Trypanosomiasis II 69 Trypanosoma grylli Nigrelli (1944) from Acris gryllus (eConte). Zoologica 30:47-57. SouTHworTH, G. C., G. Mason, and J. R. SEED. 1968. Studies on frog trypanosomiasis. I. A 24 hour cycle in the parasitemia level of Trypanosoma rotatorium in Rana clamitans from Louisiana. J. Parasit. 54:255-258. December 23, 1968 STUDIES ON AMERICAN PARAGONIMIASIS VI. ANTIBODY RESPONSE IN THREE DOMESTIC CATS INFECTED WITH PARAGONIMUS KELLICOTTI.* JOHN RICHARD SEED, FRANKLIN SOGANDARES-BERNAL, and ALBERT A. GAM Laboratory of Parasitology, Department of Biology, Tulane University, New Orleans, Louisiana ABSTRACT Sera obtained approximately every 10 days from three domestic cats exposed to varied numbers of metacercariae of Para- gonimus kellicotti were examined for com- plement fixing and precipitating antibody for a period of 150 days after exposure. The responses obtained are shown and dis- cussed, INTRODUCTION Serological tests have been developed for the clinical diagnosis of human paragoni- miasis infections (Sadun et al., 1959), but there have been few attempts to determine the type of antibody response throughout the course of human or animal infections. Sadun et al. (1959) have studied the com- plement fixing antibody response in cats experimentally infected with Paragonimus westermant. These authors first detected complement fixing antibody 30 days follow- ing the inoculation of a single dose of meta- cercariae and remained positive for at least 130 days. In their work, however, there was no description of the antibody response between 30 and 130 days, or of possible fluctuations in antibody titers. Several recent attempts have been made to characterize the antigens from adult Para- gonimus westermani and P. kellicotti ob- tained from naturally and experimentally infected cats (Yogore, Jr. Lewert, and Madraso, 1965; and Seed, Sogandares, and Mills, 1966). These reports are in close agreement that adult worms have a mini- mum of 5 precipitating antigens as shown by agar-diffusion techniques. As in the pre- vious work on complement fixing antibody, however, no attempt was made to follow the type of antibody response throughout the entire infection. It is considered conceivable that failure to determine the antibody re- sponse throughout an infection might pos- sibly produce misleading results. It is also believed possible that the quantity and specificity of precipitating antibody mole- cules might vary throughout the infection. This work describes our observation of sera obtained from infected cats approxi- mately every 10 days throughout the course of 150-day infections. MATERIALS AND METHODS Three domestic cats (5 to 10 Ibs body wgt.) were exposed by pipetting, per os, varied numbers of metacercariae of P. ell1- cottt. The cats were partially sedated by an intraperitoneal injection (1 m. 4% solu- tion/S Ibs body wgt.) of thiamylal sodium (Surital®, Parke, Davis and Co.) and bled by heart puncture. The cats were bled prior to exposure to P. &ellicotti and at approxi- mately 10 day intervals until they were killed. Sera were stored at —30°C until needed. The adult worms were harvested from cats, their uteri removed, washed sev- eral times in normal saline, and then stored at —30°C until needed. The cats were ne- cropsied for helminthic infections of the intestine. These cats were maintained in “This work was initiated under sponsorship of a research grant (AI 03386-06 TMP) from the Na- tional Institutes of Health and was directly supported by research grants GB 3036 and GB 5235 from the National Science Foundation. EDITORIAL COMMITTEE FOR THIS PAPER: Dr. IsAO TADA, Associate Professor of Medical Zoology, Kagoshima University, Kagoshima, Japan Dr. RAYMOND T. DAMIAN, Immunologist, Southwest Foundation for Research and Education, San Antonio, Texas 78206 70 No. 2 1/256 > on * a =" 1/64 he te ~ | AG aN . Hf 4“ 1/16 fe x / / \ | / | lied cy eee / / \7 Complement Firing Antibody Titer 0 10 20 30 40 50 60 70 80 90 100 110 120 130 140 150 Days Figure 1. The time course of complement fixing antibody in cats infected with Para- gonimus kellicotti. 1. A chronic cough first ob- served in cats CFS-20 and 21. 2. Eggs of P. kellicotti first observed in fecal samples from cat CFS-21. 3. Eggs of P. kellicotti first ob- served in fecal samples from cat CFS-20. 4. CFS-18 fed a second metacercaria. X cat 18; © cat 20; A cat 21. relatively good health throughout the in- fection as shown by continuous weight gains. Stool samples were obtained weekly from the infected cats, and were examined by Ritchie’s (1948) ether-formalin concentra- tion techniques for eggs of P. kellicotti. Paragonimus extracts were prepared as previously described (Seed, Sogandares, and Mills, 1966). Approximately 20 adult worms were suspended in 8 ml of distilled water and homogenized with the aid of a hand glass homogenizer. The homogenate was then centrifuged at 12,000 x G for 10 min in a refrigerated International Centrifuge. The sediment was discarded and the super- natant was used as the antigen. All antigens were stored at —30°C until needed. Protein was assayed by the method of Lowry ef al, (1951). All antigen suspen- sions were made to contain 4.0 to 5.0 mgs protein per ml. In a previous publication (Seed, Sogandares, and Mills, 1966) we stated that our suspensions contained 0.4 to 0.45 mg protein per ml. This was an error in the placement of a decimal point in the manuscript and text and should have read 4.0 to 4.5 mg protein per ml. Serological tests: 1) Agar diffusion tests were performed as described by Ouchterlony (1958). The agar used consisted of a 0.5% distilled water Paragonimiasis in Domestic Cats 71 g “A tA kak + T T T T T T T Th T 1 2 0 40 50 60 70 80 90 100 110 120 130 140 150 Day Figure 2. The time course of precipitating antibody in cats infected with Paragonimus kellicotti. See Figure 1 for explanation of num- bers 1 to 4. X cat 18; A cat 21. solution of Oxoid Ionagar, No. 2 to which was added glycine (0.5%) and merthiolate (0.1%) (Thimerosal®, Eli Lilly and Co.). The agar was then adjusted to pH 7.0 with 0.5M NaOH. 20 ml of agar were poured into each petri plate and wells were made with a No. 5 cork borer. The distance be- tween the centers of adjacent wells was 14.0 mm. The wells were filled with ap- proximately 0.1 ml of antigen or antibody. The plates were kept at room temperature and the results recorded at 24, 48 and 72 hours. Precipitating antibody was titrated in agar diffusion plates using twofold dilutions. The end-point was taken as the last dilution at which a visible precipitin line was found. 2) The complement fixation test was performed by a modification of the Kolmer method for syphilis (Kolmer, Spaulding, and Robinson, 1951). A 4 dilution of anti- serum was the lowest dilution tested, due to the anti-complementary activity of some cat antisera. Glycerinated anti-sheep hemol- ysin, lyophilized guinea pig complement, and 2% sheep red blood cells, were all ob- tained from the Colorado Serum Company, Denver. RESULTS The presence of both complement fixing and precipitating antibody was detected in sera of infected cats, confirming previous results. A minimum of 6 precipitating bands was found in agar diffusion plates indicat- ing the presence of at least six antigens in extracts of adult P. &ellicott1z, This is one 4 iS) < C Ge | \ 4 d c o CFS-18 Va Figure 3. Gel diffusion patterns produced by infected cat serum and Paragonimus kellicotti extracts. The Paragonimus extract (PE) con- tained 4.0 mg protein per well; 0.1 ml antigen or antibody was used per well. Sera CFS-2, 3 and 4 have been described previously (Seed Sogandares and Mills, 1966). The plates were read at 24 and 48 hrs and a final reading was made at 72 hrs. Tulane Studies in Zoology and Botany Vol. 15 @ Figure 4. Gel diffusion patterns produced by sera, from an infected cat, taken at varied time intervals and tested against Paragonimus kel- licotti extracts. The methods and materials were identical to that described for Figure 4. The time in days at which serum was obtained: GFS-21A (0), CFS-21C (8), CFS-21F (27); CFS-21G (43), CFS-21I (65), CFS-21K (85), CFS-21M (115), and CFS-210 (150). \ No. 2 Paragonimiasis in Domestic Cats 1) TABLE |] Data on infected cats from which sera were obtained.! Maximum Number of Paragonimus Other Maximum complement Cat metacercariae recovered at helminth Patency of precipitation — fixation number fed autopsy infections* P. kellicotti titer titer CFS-18 2 oe unidentified none 1/64 1/32 nematode CFS-20 AQ 36° T. cati patent ~ 1/32 D. caninum CFS-21 21 Pile iecat patent 1/128 1/64 D. caninum 1 All cats were infected for 150 days. ° Fed one metacercaria 8 March 1966, and another 21 June 1966. ’ Unencysted migrating preadults recovered at autopsy. * Determined by fecal examinations and by autopsy © Adults only recovered. more antigen than reported previously (Seed, Sogandares, and Mills, 1966). These results are in agreement with those of Tada (1967) who observed a maximum of six precipitin bands in serum from a rat in- fected with Paragonimus miyazaku. The de- tection of an additional band is presumably due to the greater length of time allowed for pattern development prior to the final recording of our results (48 versus 72 hours). As previously shown, the precipi- tating antibody response appears to vary in infected cats. Serum CFS-3 contained at least one unique antibody specificity not found in CFS-18 and 21 (Figure 3). Serum CFS-20 contained no detectable precipitat- ing antibody, although the largest number of adult worms was recovered from this cat (Table 1). Figure 4 suggests that although the intensity and position of the precipitat- ing bands in agar diffusion plates changes during the early stages of the infection, the same bands are found throughout the in- fection. This suggests, assuming our meth- ods are sensitive enough, that the cat does not experience different antigenic stimuli from the adult worms during the course of the infection. Figures 1 and 2 show the precipitating and complement fixing antibody titers throughout the course of a Paragonimus infection of cats. Both the precipitating and complement fixing antibedy curves are simit- lar in that once antibody is detected there is a sharp rise in titer and then a plateau 1s attained. The antibody titers are then main- tained at a high level throughout the course of the infection. One obvious difference be- tween the precipitating and complement fixing antibody curves is that complement fixing antibody is detected considerably earlier in the infection. Cat (CFS-20) con- tained no detectable precipitating antibody by the agar diffusion test but significant complement fixing antibody titers were found. Figures 1 and 2 also show that there is no obvious correlation between the rise and titer of complement fixing and precipi- tating antibody, and egg production or clinical symptoms (chronic cough). Cat (CFS-18), which never showed the pres- ence of eggs in its stool, contained definite precipitating and complement fixing anti- body. The complement fixing antibody was detected considerably earlier than the pas- sage of eggs or a noticeable cough. CONCLUSIONS It is apparent from these data that both complement fixing and precipitating anti- bodies arise relatively early (17 and 37 days) during P. &ellicotti infections of the cat. The antibody titer then reaches a plateau and remains high throughout the infection. This is to be expected since the adult worms would presumably release antigen(s) throughout the entire infection. Our results are similar to those reported by Sadun e¢ al. (1959) for P. westermani infections in the cat using the complement fixation test, and also So (1959) who investigated the pre- cipitating antibody response in rabbits and dogs infected with P. ohirat. Tada (1967), however, reported a de- crease in the number of precipitating anti- 74 Tulane Studies in Zoology and Botany bodies in sera from rats with long-term P. miyazakw infections. He suggested that the cyst surrounding the adult worms might impede the passage of excretory substances and, therefore, account for the reduced anti- body response. It was of interest that cat (CFS-20) which had the highest worm burden failed to show detectable precipitating antibody. One ad- ditional cat, (CFS-2), which was reported upon previously and had the next highest worm burden (30 adults recovered), also showed an unexpectedly low precipitating antibody titer. Numerous suggestions can be made to explain the low level or absence of detectable precipitating antibody in heav- ily infected cats. However, the most interest- ing possibilities to us are the phenomena of immune paralysis or the absorption and removal of circulating antibody by the large number of adult worms and their antigens. Both explanations might readily account for the data obtained, and are amenable to further experimentation. Sogandares (1966) indicated that P. kel- licotti infections became patent only in cats whose lung cysts contained two or more worms. Cat CFS-18 had been fed a single metacercaria, followed by a second 106 days later; the infection never became patent even 44 days after feeding of the second meta- cercaria, and two unencysted migrating pre- adults were recovered after 150 days. Patency usually results 30 to 44 days, past exposure to moetacercariae, when two or more worms locate in a single lung cyst. In view of Sogandares’ (1966) findings and in our recovery of two unencysted immature worms from cat CFS-18, in whose sera both complement fixing and precipitating anti- bodies could be demonstrated, it is strongly suggested that some complement fixing and precipitating antibodies may be produced in the absence of eggs passing through the host lung tissues. Since the sera from cat CFS-18 had essentially the same precipitin patterns in agar diffusion plates as sera from other cats containing adult worms, our find- ings also suggest that preadults produce anti- gens in common with the adult worms. This is in agreement with the work of Tada (1967) who found precipitating antibody to adult worms in infected rats prior to the encystment of the adults in the lung. Vol. 15 The detection of complement fixing anti- body considerably earlier than the detection of precipitating antibody, as well as the detection of complement fixing antibody in cat (CFS-20) which failed to produce significant levels of precipitating antibody, might be due to the greater sensitivity of the complement fixation test. The possibil- ity should also be considered that the com- plement fixing antigen(s) and the precipi- tating antigens are not identical. The early rise in complement fixing antibody might prove the complement fixation test valuable as a possible diagnostic tool especially early in infection prior to patency. Sadun e¢ al. (1959) previously suggested a similar use of the complement fixation test based on their observations of P. westermani infec- - tions in cats. ACKNOWLEDGMENTS The authors are indebted to Sue Carlysle Dike for technical assistance, and to Mrs. E. Peter Volpe for execution of the figures. LITERATURE CITED KouMeEr, J. A., E. H. SPAULDING, and H. W. Rosinson. 1951. Approved Laboratory Tech- nic, 5th Ed. Appleton-Century-Crofts, Inc., New York. Lowry, O., N. J. Roseproucu, A. L. Farr, and R. RANDALL, 1951. Protein measurement with Folin phenol reagent. J. Biol. Chem. 193:265- 276. OucHTERLONY, O. 1958. Diffusion-in-gel meth- ods for immunological analysis. Progr. Al- lergy 5:1-78. Rircure, L. S. 1948. An Ether Sedimentation Technique for Routine Stool Examinations. Bull. U. S. Army Med. Dept. 8:326-329. Sapun, E. H., A. A. Buck, and B. C. WALTON. 1959. The diagnosis of Paragonimus wester- mani using purified antigens in intradermal and complement fixation tests. Mil. Med. 124:187-195. SEED, J. R., F. SOGANDARES-BERNAL, and R. R. Mitts. 1966. Studies on American Paragoni- miasis. II. Serological observations of infected cats. J. Parasit. 52:358-362. So, N. 1959. Immunological Studies of the Lung-Fluke, Paragonimus Ohirai Miyazaki, 1939. (Precipitin ring test and Sarles’ Phe- nomenon). Fukuoka Acta Medica 50:2594- 2623. Tapa, I. 1967. Physiological and_ Serological Studies of Paragonimus miyazakii Infection in Rats. J. Parasit. 53:292-297. Yocore, M. G., Jr., R. M. Lewert, and E. D. Mapraso. 1965. Immunodiffusion Studies on Paragonimiasis. Am. J. Trop. Med. Hyg. 14: 586-591. December 23, 1968 TULANE STUDIES IN ZOOLOGY AND BOTANY LIBRARY rT . Volume 15, Number 3 March 24, 1969 “APS VAR UNIVERSITY, PLANOPHILA TERRESTRIS, A NEW GREEN ALGA FROM TENNESSEE SOIL ROBERT D. GROOVER AND SISTER ADRIAN MARIE HOFSTETTER, O.P. Department of Botany The University of Texas, Austin p. 75 LIFE CYCLES OF CARNEOPHALLUS CHOANOPHALLUS N. SP. AND C. BASODACTYLOPHALLUS N. SP. (TREMATODA: MICROPHALLIDAE) JOHN F. BRIDGMAN Laboratory of Parasitology, Department of Biology, Tulane University, New Orleans, Louisiana p. 81 ETHEOSTOMA COLLETTEI, A NEW DARTER OF THE SUBGENUS OLIGOCEPHALUS FROM LOUISIANA AND ARKANSAS RAY S. BIRDSONG Institute of Marine Sciences, University of Miami Rickenbacker Causeway, Miami, Florida and LESLIE W. KNAPP Smithsonian Oceanographic Sorting Center Washington, D. C. p. 106 MYSIDOPSIS BAHIA, A NEW SPECIES OF MYSID (CRUSTACEA:MYSIDACEA) FROM GALVESTON BAY, TEXAS JOANE MOLENOCK Marine Laboratory Texas A&M University Galveston, Texas p. 113 TULANE UNIVERSITY NEW ORLEANS TULANE STUDIES IN ZOOLOGY AND BOTANY is devoted primarily to the biology of the waters and adjacent land areas of the Gulf of Mexico and the Caribbean Sea, but manuscripts on organisms outside this geographic area will be considered. Each number is issued separately and contains an individual monographic study, or several minor studies. As volumes are completed, title pages and tables of contents are distributed to institutions receiving the entire series. Manuscripts submitted for publication are evaluated by the editor or associate editor and by an editorial committee selected for each paper. 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Berra Assistant to the Editors le et ele TULANE STUDIES IN ZOOLOGY AND BOTANY Volume 15, Number 3 March 24, 1969 PLANOPHILA TERRESTRIS, A NEW GREEN ALGA FROM TENNESSEE SOIL ROBERT D. GROOVER! and SISTER ADRIAN MARIE HOFSTETTER, O.P.? Department of Botany The University of Texas, Austin ABSTRACT A new chlorosphaeracean alga, Plano- phila terrestris, is described from Tennes- see soil. Hofstetter (1968) has reported on the distribution and identification of previously described soil algae from twelve areas in Shelby County, Tennessee. During this in- vestigation an organism tentatively identi- fied as a new chlorosphaeracean alga was isolated into unialgal culture. Because of the first author's special interest in the Chloro- sphaeraceae (sensu Herndon, 1958), the or- ganism was studied jointly during the sum- mer of 1967. Unialgal cultures were maintained in Pringsheim’s soil-water medium (Starr, 1964). Axenic cultures were obtained using the techniques of Brown and _ Bischoff (1962). The axenic cultures were main- tained and studied in Bold’s Basal Medium (BBM) with and without 1.5 per cent agar (Brown and Bold, 1964). The cultures were grown in a culture room under standard conditions, namely 300 ft c light (cool-white fluorescent) on a 12-hr light, 12-hr dark cycle at 22 C. Morphological observations were made routinely on cultures varying in age from one day to three months. Acetocarmine, Haidenhain’s haematoxylin, aqueous iodine (IsKI), India Ink, and Sudan IV were used to elucidate certain aspects of the cellular morphology. The photomicrographs were made with a Bausch and Lomb microscope with a 35-mm Zeiss-Winkel camera attach- ment. OBSERVATIONS Planophila terrestris sp. nov. (Figs. 1-16) Cellulae vegetativae sphericae, 6-15.2 p diam ad paululum ellipsoideas, 14-18 p xX 18-26 »; membrana cellulae tenuis levisque, senescens paululum spissescens, chloroplastus parietalis, in cellulis maturis perforatus; una- tres pyrenoides omni in cellular; cellulae per crescentiam uninucleatae; vacuolae con- tractiles in cellulis vegetativis observatae; fasciculi cellularum duo-quattuor-cellulares, plerumque due-dimensionales, per divisiones cellularum vegetativarum successivas effecti; cellulae vetustae numerosas guttulas olei minutas habentes; cellulae brunneo-aurantiae; culturae senescentes colonias Hormotilopsis- formes effectae. Reproductio per dissociationem fascicu- lorum cellularum aut per zoosporas quadri- flagellatas aut per aplanosporas; zoosporae elongatae, per bipartitionem iteratam effec- tae, 3-6 p lat. X 7-16 p» long., sine mem- brana, nucleum anteriorem, stigma anterius lineare, duas vacuolas contractiles anteriores, 1 Present address: Department of Biology, Tulane University, New Orleans, Louisiana 70118. 2 Present address: Siena College, Memphis, Tennessee 38117. : The authors express their sincere appreciation to Dr. H. C. Bold for reading the manuscript. EDITORIAL COMMITTEE FOR THIS PAPER: Dr. TEMD R. DEASON, Associate Professor of Biology, University of Alabama, Unt- versity, Alabama Dr. H. WAYNE NICHOLS, Associate Professor of Botany, Washington University, St Louis, Missouri 76 Tulane Studies in Zoology and Botany Figures 1-8. Planophila terrestris. Figures 1-6. Cells from actively growing cultures on 3N BBM agar. Figure 1. Young cells; note parietal plastid, one pyrenoid per cell, and variation in cell size. < 600. No. 3 pyrenoidem posteriorem aut interdum me- diam, et plastidem parietalem habentes; zoosporogenesis per examinantem directam interdum effecta. Reproductio sexualis non observata. Massa plantalis in agaro 3N BBM dicto culta aetate 2 et 4 hebdomadum sine micro- scopio observata sicca, magnificatione 14 X observata aspera (minute glomerulata ), con- sistencia butyracea, in culturis veteribus in periodo immobili brunneo-aurantia facta. Origo: Plantae e solo horti culto in loco Memphis, Shelby County, Tennessee dicto. m. Nov., 1966 lectae; numerus culturae R-81. In actively growing cultures, young vege- tative cells, recently derived from zoospores, are spherical to slightly ellipsoidal (Fig. 1). Such cells, generally 6-11 » in diameter, at- tain a diameter of 15.2 » in cultures 4 days old. The chloroplast in young cells is typically parietal and covers most of the cell surface (Fig. 1). The plastid soon becomes perforate (Fig. 2) and contains from one to three pyrenoids (Figs. 1-5); the cells are unt- nucleate. Young cells contain numerous colorless lipid droplets (Fig. 2). One or two contractile vacuoles have been observed in vegetative cells. The walls of cells in ac- tively growing cultures are thin and smooth (Figs. 1-5) but become slightly thickened with age (Figs. 8 and 9). Solitary cells which fail to divide vegetatively or to un- dergo zoosporogenesis may be slightly ellip- soidal, 14-18 X 18-26 p. Vegetative cell division (sensw Herndon, 1958) is characteristic of log-phase cultures. During cytokinesis a partition is laid down across the center of the parent cell, dividing it into two daughter cells which remain in close association with the parent cell wall. The daughter cells, in turn, may undergo vegetative cell division. The direction of each division is usually perpendicular to the preceding division giving two-dimensional, four-cell packets (Figs. 2-6). Not infre- quently, three-dimensional, cuboidal, eight- Planophila terrestris ial celled packets are produced. The packets may dissociate either partially or entirely to form diads or solitary cells. In stationary-phase cultures, a unilateral deposition of wall materials results in the formation of branching Hormotilopsis-like colonies (Figs. 7 and 8). Cells from such cultures are brownish-orange due to the presence of colored oil droplets. Asexual reproduction is by zoospores and aplanospores (Figs. 10-15). Both solitary cells and those in packets may produce zoo- spores by successive bipartition of the ma- ternal protoplast. Usually four (Fig. 10), sometimes two or eight, zoospores are pro- duced by each parent cell. Zoospores are released through a pore formed by a gelatini- zation of the zoosporangial wall. Occasionally the protoplast of a single cell behaves as a single zoospore (Fig. 11). In such cases the remains of the parent wall are often vis- ible in the culture medium (Fig. 11). The zoospores, elongate in shape, are 3-6 » wide and 7-16 » long. Each has a parietal plastid, an anterior nucleus, an anterior linear stigma, two contractile vacuoles, and a pos- terior, sometimes medial, pyrenoid. The zoospores have four flagella of equal length (Fig. 12). They lack a wall (Protosiphon- type, Starr, 1955) and become spherical at quiescence. The period of motility is several hours. The eyespot is persistent and may be observed for several hours after the zoo- spore has become nonmotile. The zoospores may be rapidly motile or exhibit slow, amoeboid movements. Aplanospores (Figs. 14 and 15) are formed in the same manner as zoospores and are morphologically similar. They develop a wall prior to their release by a gelatinization and rupture of the aplanosporangial wall. Macroscopically the plant mass of Plano- phila terrestris on 3N BBM! agar is dry; at 14X it is rough and glomerulate (Fig. 1BBM with 3x the usual amount of NaNO:. < Figure 2. Cells showing perforate condition of chloroplast. The numerous droplets (arrow ) in the cells are Sudan-IV-positive and probably lipid. « 1500. Figures 3, 4. Typical packets. Note cells with more than one pyrenoid and dividing (arrow). Figure 3, x 1250; Figure 4, x 800. cell Figure 5. Cells illustrating contiguity of the parent wall around packet (arrow). x 1200. Figure 6. Cells arranged in typical packets. x 100. Figures 7, 8. Cells from stationary-phase cultures on 3N BBM agar; note branched Hormo- tilopsis-like organization. Figure 7, x 700; Figure 8, x 500. 78 Tulane Studies in Zoology and Botany Vol. 15 No. 3 16); its consistency on agar is butyrous. Actively growing cultures are green but change with age through various shades to a brownish-orange. DISCUSSION Only a small number of green, nonfila- mentous algae reproducing by quadriflagel- late zoospores have been described. In two genera, Fernandinella Chodat (1922) and Tetraciella Pascher and Petrova (1930), the cells are typically pyriform. Fott and Kalina (1965) consider the two genera to be synonymous. In the tetrasporalean alga Hor- motilopsis Trainor and Bold (1953), per- sistent packets are absent. The genus Plano- phila Gerneck (1907) is characterized by spherical cells which divide vegetatively in two directions to form flat (1.e., two- dimensional), two-eight-celled packets. In determining the taxonomic position of the organism herein described, the descriptions and illustrations of the four above-named genera and their species were studied in the original references and also in several other sources: Wille (1911), Arce and Bold (1958), Reisigl (1964), Fott and Kalina (1965), and Bourrelly (1966). The morphological attributes of this or- ganism, though not in complete agreement with the generic descriptions of any of the four genera, are sufficiently close to Plano- phila to warrant its classification in this genus. The isolate described here differs distinctly from the species of this genus treated by Gerneck (1907) and Reisigl (1964) and accordingly, it is described as a new taxon. The majority of species of Planophila have their cells in a flat, biseriate packet. The packets of P. terrestris are frequently Planophila terrestris 79 cuboidal but this characteristic is not of suf- ficient importance to remove it from the genus. The cuboidal packets of cells, the presence of a Hormotilopsis-like stage, and the much greater length of the zoospores distinguish P. terrestris from the other three species. The species of Planophila may be dis- tinguished as follows: 1. Stationary-phase cultures forming Hormo- tilopsis-like stages; zoospores elongate, up to 16 win length P. terrestris sp. nov. 1. Hormotilopsis-like stage not reported; zoo- sporesvless) than Oya) long eee 2 2. Zoospores without an eyespot —____ P. asymmetrica (Gerneck) Wille (1911) 2. Zoospores with an eyespot 3 3. Solitary cells 6-8 » in diameter; pyrenoids two; zoospores 6.5 w long, 4 u wide wine) Eas P. bipyrenoidosa Reisig] (1964) 3. Solitary cells 5-15 « (-22) uw in diameter; pyrenoid one; zoospores almost spherical, Gkwaneciame terme ae eee ee “2 ee tee P. laetevirens Gerneck (1907) A culture of Planophila terrestris has been deposited in the Culture Collection of Algae, Indiana University, Bloomington, Indiana and an herbarium specimen in the Field Museum of Natural History, Chicago, IIli- nois. REFERENCES CITED Arce, G. and H. C. Bop. 1958. Some Chloro- phyceae from Cuban soils. Amer. Jour. Bot. 45:492-503. BourRELLY, P. 1966. Les algues d’eau douce. Algues vertes. N. Boubée Cie. Paris. Brown, R. M., Jr. and H. W. Biscuorr. 1962. A new and useful method for obtaining axenic cultures of algae. Phycol. News Bull. 15:43-44. at EA , and H. C. Borp. 1964. Phycological Studies V. Comparative studies of the algal genera Chlorococcum and Tetra- cystis. Univ. Texas Publication No. 6417. Cuopat, R. 1922. Materiaux pour histoire des Figures 9-16. Planophila terrestris. Figure 9. Cells from stationary-phase cultures on 3N BBM agar; note thickened cell walls. x 725. Figures 10-15. Cells from actively growing cultures on 3N BBM agar. Figure 10. Zoospores prior to release, four per sporangium (arrow). < 1200. Figure 11. Direct swarming; note liberation of entire protoplast as a single zoospore and _per- sistence of parental walls. « 2000. Figure 12. Single zoospore. Note quadriflagellate condition and position of pyrenoid. Phase- contrast. « 1500. Figure 13. Zoospore. Note parietal plastid, position of pyrenoid, and position of eyespot (arrow). x 1800. Figures 14, 15. Aplanosporangia; eight aplanospores per parent cell in Figure 14 and four per cell in Figure 15. Figure 14, x 300; Figure 15, x 1000. Figure 16. Surface view of plant mass; 2-week-old culture on 3N BBM agar. x 16. 80 Tulane Studies in Zoology and Botany algues de la Suisse. Bull. Soc. Bot. Geneve 13:66-114. Fort, B. and T. Katina. 1965. Zur Klarung einiger tetrakonten Griinalgen. Preslia 37: 369-379. GeERNECK, R. 1907. Zur Kenntnis der niederen Chlorophyceen. Beih. Bot. Centralbl. 2(21): 221-290. Hernpon, W. R. 1958. Studies on chlorophaera- cean algae from soil. Amer. J. Bot. 45:298- 308. Horstetter, Sister A. M. 1968. A preliminary report on the algal flora of soils from selected areas of Shelby County. Jour. Tenn. Acad. Sci. 43(1):20-21. Perrova, J. 1930. Eine neue festsitzende Pro- tococcalengattung (Tetraciella nov. gen.). Arch. Protistenk. 71:550-566. Vol. 5 ReisicL, H. 1964. Zur Systematik und Oko- logie alpiner Bodenalgae. Osterr. Bot. Z. 4: 402-499. Starr, R. C. 1955. A comparative study of Chlorococcum Meneghini and other spherical zoospore producing genera of the Chloro- coccales. Indiana Univ. Publication Sci. Ser. No. 20. ee, Ta 1964. The culture collec- tion of algae at Indiana University. Amer. Jour. Bot. 51:1013-1044. Trainor, F. and H. C. Boup. 1953. Three uni- cellular Chlorophyceae from soil. Amer. Jour. Bot. 40:758-767. Witte, N. 1911. Chlorophyceen. In: Engler und Prantl, Die naturlichen Pflanzenfamilien. 1. Aufl., Nachtrag. LIFE CYCLES OF CARNEOPHALLUS CHOANOPHALLUS N. SP. AND C. BASODACTYLOPHALLUS N. SP. (TREMATODA: MICROPHALLIDAE)! JOHN F. BRIDGMAN2 Laboratory of Parasitology, Department of Biology, Tulane University, New Orleans, Louisiana ABSTRACT The life histories of two new microphal- lid trematodes Carneophallus choanophal- lus and Carneophallus basodactylophal- lus, from South Louisiana are reported. The natural hosts for C. choanophallus are: the raccoon, Procyon lotor (LL) and the black rat, Rattus rattus (L) [definitive hosts], the snail, Lyrodes parvula Guild- ing, 1928 (unspined form) [first intermedi- ate host], and the shrimp, Macrobrachium ohione Smith, 1874 and Palaemonetes pugio Holthuis, 1949 [second intermediate hosts]; for C. basodactylophallus: the raccoon [de- finitive host], the snail Lyrodes parvula Guilding, 1828 (spined form) [first interme- diate host] and the blue crab, Callinectes sapidus Rathbun, 1896 [second intermedi- ate host]. The cercariae of the two species differ primarily in stylet length and experi- mentally were not capable of cross-infect- ing the crustacean hosts. Cercarial penetration, development and growth of the metacercariae of C. choa- nophallus are described. A 15-month geo- graphical and ecological study of infected M. ohione in the Mississippi River is pre- sented. INTRODUCTION Freshwater shrimp, Macrobrachium ohione Smith, 1874, from the shallow water of the west bank of the Mississippi River at Ama, Louisiana were collected in July, 1966. The musculature of the abdomen and the cephalo- thorax of these shrimp was infected with metacercariae of an unidentified micro- phallid trematode. Further study showed that the metacercariae were larvae of a new species of Carneophallus Cable and Kuns, 1951. An intensive search for the other stages of the life-cycle and an ecological study of the infected shrimp led to the find- ing of high incidences of densely infected M. ohione in several areas near the mouth of the Mississippi River. It was subsequently found that black rats, Rattus rattus (L), from Fort Jackson, Louisiana and raccoons, Procyon lotor (L), from Pass a Loutre, Louisiana were naturally infected with adults of the new species. Further examination of the small animal runways in the marshes and ponds at Pass a Loutre, Louisiana led to the discovery of the snails, Lyrodes parvula Guilding, 1896 (unspined form) which harbor the cercariae. The grass shrimp, Palae- . monetes pugio Holthuis, 1949, was also found to harbor metacercariae in endemic areas. During studies on the life history of the microphallid from M. ohione, metacercariae of another new, very closely related, species of Carneophallus were discovered in the blue crab, Callinectes sapidus Rathbun, 1896. Adults of this form were also found in rac- coons, and the snail, Lyrodes parvula Guild- ing, 1896 (spined form) was found to har- bor cercariae which infect the blue crab. MATERIALS AND METHODS Mississippi River shrimp, Macrobrachiwm ohione Smith, 1874 and the brackish water grass shrimp, Palaemonetes pugio Holthuis, 1949, were used in studies and experiments with Carneophallus choanophallus, while the blue crab, Callinectes sapidus Rathbun, 1896, was used for those dealing with Carneophal- lus basodactylophallus. These crustaceans were collected from various natural habitats with baited traps, dip nets and by dredge- ‘This study was supported in part by NIH grant No. GM-669 and NSF Grants GB-3036 and GB-5235. 2 Present address: Shikoku Christian College, Zentsuji, Kagawa-Ken, Japan. EDITORIAL COMMITTEE FOR THIS PAPER: Dr. FRANK J. ETGES, Professor of Zoology, University of Cincinnati, Cincinnati, Ohio Dr. WALTER E. MARTIN, Professor of Biological Sciences, University of Southern California, Los Angeles, California 82 Tulane Studies in Zoology and Botany trawling from a small boat. M. ohione col- lected for studies of shrimp size, incidence, and intensity of infection, were collected with '%4-inch-mesh wire traps (baited) at- tached to the river bank with 10- to 25-foot- long lines at the ruins of the old Sellers Plantation House at Ama, Louisiana. The shrimp were transported to the laboratory for observation and study in aerated minnow buckets. Measurements of total length of the shrimp are in millimeters from the end of the rostrum to the tip of the telson. The total length measurements were grouped into classes of 5-mm increments. Metacer- cariae were counted by examination of the transparent shrimp body under low power of a dissecting microscope. Shrimp and crabs were maintained in the laboratory in aerated aquaria or in individual containers. They were fed dried dog food (Austin’s Baked Dog Food, Sunshine Biscuits, Inc., Long Island, New York) every other day followed by a change of water several hours after each feeding. Naturally and laboratory infected shrimp and crabs were used to infect suspected definitive hosts. Metacercariae for infection experiments were counted by placing small pieces of cyst-containing tissue under the dissecting microscope. Laboratory definitive hosts were exposed by feeding metacercariae in small pieces of shrimp or crab tissue. Excystment of metacercariae was done in 0.75 per cent saline with sharp needles; however, the worms usually excysted spon- taneously when allowed to stand in saline for several hours or overnight at room tem- perature. Natural definitive hosts were collected in the endemic areas by live-trapping along the bank of the river and by shooting by day and by night from a small boat equipped with a spotlight. In the laboratory, small mam- mals were killed by a sharp blow to the back of the head, larger mammals were killed by etherization. Within 30 minutes of killing or shooting, the mammal’s small in- testine was taken out and the adult trema- todes were removed by slitting and scraping the gut into containers of 0.75 per cent saline. For larger animals, the small intestine was usually cut into 4-inch lengths before slitting. Examination was done in_ petri dishes under the low power of a dissecting microscope. Vol. 15 Spined and unspined snails of the species Lyrodes parvula, which carried the two species of Carneophallus cercariae, were col- lected with the aid of a large tea strainer from fresh to brackish water marshes and ponds where small animal trails could be seen in the marsh grass (Spartina spp.), al- ligator weed (Alternanthera spp.) and the water weed (Cabomba spp.). Snails were taken to the laboratory and those observed to be shedding cercariae were isolated. Snails used for incidence studies were crushed on glass plates in a drop of 0.75 per cent saline and observed under a dis- secting microscope for cercariae. All snails were maintained in the laboratory in aquaria containing vegetation and water with the salinity adjusted to match that of the col- lecting area. Since snails were collected from areas near the mouth of the Mississippi and Pearl Rivers, the salinity was found to vary from time to time. The snails were induced to shed cercariae by placing them in small containers of water adjusted to the proper salinity and subjecting them to a slight in- crease in temperature under an ordinary reading light bulb for several hours. Shrimp or crabs were preadapted to water of the proper salinity and then exposed to cercariae. Injecting the cercariae into the gill chambers was also effective. If shrimp maintained in the laboratory for 15 days or more showed no metacercariae in their transparent bodies under the low power of the dissecting microscope, they were con- sidered uninfected and were used for in- fection experiments. Staged studies of meta- cercarial development in M. ohione were done in an incubator maintained at 27C. Crabs from certain localities (W4ullswood and Osgood Ponds 12 miles west of New Orleans on U.S. Highway 90) were found to be very lightly infected. These crabs were maintained in the laboratory for three weeks before being exposed to cercariae. Laboratory infections with thin cyst walls were then easily distinguished from natural infections. Eggs were teased from the uteri of adult worms and their development was followed in containers of water which was changed twice weekly. Both fixed and living material were used to study all stages of the life-cycles in the present study. Neutral red and _nile-blue sulfate vital stains were used. Whole mounts No. 3 Carneophallus Life Cycles 83 TABLE l. Geographical distribution, incidence and density of Macrobrachium ohione infected with Carneophallus choanophallus metacercariae from four Louisiana Rivers. Number Number Average of of Col- Incidence Density Location Mile! Shrimp lections (Per cent ) per Shrimp MISSISSIPPI RIVER Greenville, Miss. 540 47 1 93.3 0.32 Vicksburg, Miss. 435 10 1 30.0 0.60 New Roads, La. 260 93 2} 26.8 0.85 Plaquemine, La. 210 2.02 % aI 0.69 Wallace, La. 135 198 4 20.2 0.38 Bonnet Carre Spillway, La. 125 308 3 22.0 0.66 Ama, La. 115 9994 120 22.4 0.84 Empire, La. 30 13 1 46.1 2.69 Fort Jackson, La. 20 661 9 67.6 3.65 Baptiste Collette Pass, La. 10 246 4 64.4 5.28 Pass a Loutre, La. —5 484 8 71.0 4,23 ATCHAFALAYA RIVER Morgan City to Charenton, La. 125 2 0.8 0.008 RED RIVER Highway 107 at Vick, La. es 1 0.0 0.0 WEST PEARL RIVER Highway 90 to Pearl River, La. 0 3 0.0 0.0 1 Mile numbers indicate the distances above and below (—) Head of Passes, La. at the mouth of the Mississippi River (see Flood Control and Navigation Maps of the Mississippi River, Cairo, Illinois to the Gulf of Mexico, 34th Ed. (1966) prepared by the U. S. Army Corps of Engineers. 68 maps, 12 charts, 43 sheets ). were fixed in alcohol-formalin-acetic acid fixative and stained in Van Cleave’s (1953) combination hematoxylin, then mounted in Permount® (Fisher Scientific Company). Adult specimens used for growth studies were killed in hot water before being fixed to assure a standard state of body contrac- tion. Drawings were made free-hand and with the aid of a microprojector. Measure- ments are in millimeters unless otherwise specified. Carneophallus from Macrobrachinm ohione and Palaemonetes pugio The Life-Cycle In the fall of 1966, Mississippi River shrimp, Macrobrachium ohione Smith, 1874, were collected from the shallow water of the west bank of the Mississippi River at Ama, Louisiana. Twenty to 30 per cent of these shrimp had from 1 to 60 microphallid meta- cercariae in their transparent body muscula- ture (Figs. 1 and 2). Excysted metacercariae revealed a well differentiated preadult trema- tode which demonstrated all the character- istics of the genus Carneophallus Cable and Kuns, 1951. Metacercariae were fed to lab- oratory rats and mice and ovigerous adults obtained from the intestine 5 to 25 days post-exposure as well as excysted metacer- cariae were used to identify this trematode as a new species of Carneophallus. An intensive search for the other phases of the life-cycle was made in the Ama, Louisiana region. All snails and suspected definitive hosts taken from this study area were examined for microphallid cercariae and adults with negative results. A geo- graphical and ecological survey of the in- fected shrimp (Table 1) finally pointed the way to marsh and pond areas between the distributaries at the mouth of the Mis- sissippi River near the Louisiana State Fish and Wild-Life Camp at Pass a Loutre, Louisi- ana, where the incidence and density of shrimp infection was considerably elevated. Examination of numerous animals observed to be feeding on shrimp revealed that the raccoon, Procyon lotor (L), and the black rat, Rattus rattus (L), were naturally in- Vol. 15 Tulane Studies in Zoology and Botany 84 ‘aposnUL auolyo * WOLF PoyessIp vIBOIBDRJOUL snyDYydouvoYI °D JO (eInzeIoduu9} WOOL) UOTPaFUL A1OZLIOGeT P[O-Avp-AVfly “P [TBA SAO ay} Fo Sey au} Bune ASHO -Wlap aUOLYO “JY Ul BITROIBORJOU snjPDYdouDOYO *_D eINyeUI JO UOHeS UlfyeIe “gE ‘aUOrYyO “PY JO sINyeTMosnUT [eUTUTOpqe UI (MOIIe ) BLIBOIZORJOU SN7 -jpydouvoys snyvydoauiy “Z ‘(Mojeq) efeut pue (sAoqe) a[eUlof eord4y (dumzys toaty Iddississty) auoryo wnryooiqoiovy *[ “p-[ Semnsty No. 3 fected with species of Carneophallus, includ- the new species. Raccoons and rats were observed to make runways in the alligator weed (Alternan- thera spp.), marsh grass (Spartina spp.) and the water weed (Cabomba spp.) in the Pass a Loutre area. With the aid of a large tea strainer, amnicolid snails, the spined and unspined forms of Lyrodes parvula, were collected from runways and brought back to the laboratory where they were ob- served to be infected with microphallid cer- cariae. Although the cercariae shed from the two snail types appeared to be identical at first, only those from the unspined form of L. parvula were capable of infecting shrimp. Careful examination with vital stains later demonstrated morphological differences in the cercariae. Usually the unspined form of L. parvula was found on vegetation in the open ponds where Cabomba predomi- nates, whereas the spined form of L. parvula was found along the edges of the ponds and marshes in the dense vegetation. Young shrimp, M. ohione, and Palaemonetes pugio in these ponds were heavily infected with the new metacercariae. While field work was under way, labora- tory experimentation was conducted to de- termine which of several animal types could serve as possible definitive hosts. The re- sults of the survey were as follows: All mam. mals including white mice (ICR strain), white rats (Sprague-Dawley strain), cats, a raccoon, guinea pigs, meadow mice (Microtus montanus) and cotton rats (Szg- modon hispidus), which were fed meta- cercariae yielded good numbers of ovigerous adult worms 12 hr to 25 days post-exposure. Of eight chicks and four Pekin ducklings fed large numbers of metacercariae, only one chick upon necropsy yielded two nono- vigerous worms 24 hr post-exposure. The others sacrificed 24 to 96 hr post-exposure yielded no worms. Frogs, Rana pipiens and R. grylio, maintained at room temperature yielded nonovigerous worms upon necropsy 12 to 96 hr post-exposure. Turtles, Grap- temys spp., did not become infected. All mammals fed metacercariae from M. ohione and P. pugio were susceptible to infections producing ovigerous specimens. The frogs, turtles and birds were either not susceptible or unable to produce mature ovigerous in- fections under normal conditions. Carneophallus Life Cycles 85 These studies demonstrated that the rac- coon and the black rat are natural definitive hosts for Carneophallus metacercariae from M. ohione and P. pugio; however, any shrimp-eating mammal could probably serve as well. The first intermediate host is a snail, the unspined form of Lyrodes parvula, which shed cercariae demonstrated to pro- duce experimental infections in shrimp, M. ohione and P. pugio, identical to those found in nature. This life-cycle of the Carneophal- lus metacercariae from M. ohione and P. pugio is illustrated in Figure 5 A. The de- scription of this new species follows: Description of the Stages of the Life History of Carneophallus choanophallus n. sp. The Adult (Figures 6-8) SPECIFIC DIAGNOSIS: (based on 80 hot water-killed unflattened specimens from Procyon lotor, Rattus rattus, Rattus norvegt- cus albinus (Sprague-Dawley strain), Mus musculus albinus (ICR strain) and Felzs domesticus.) Microphallidae: Carneophal- lus. Body pyriform, sometimes with posterior notch, 0.355 to 0.520 long by 0.200 to 0.340 wide. Forebody 0.155 to 0.325 long. Integu- ment of anterior 1/2 of body spined. Oral sucker subterminal, 0.045 to 0.057 long by 0.050 to 0.062 wide. Prepharynx very short. Pharynx 0.025 long by 0.025 wide. Esopha- gus extending from pharynx to approxi- mately anterior 1/3 body, 0.065 to 0.137 long. Ceca two, extending posteriorly and obliquely from cecal bifurcation at posterior end of esophagus, 0.037 to 0.160 long. Ace- tabulum eguatorial, mesial, 0.042 to 0.062 long by 0.042 to 0.062 wide. Sucker ratio IOI O Ron HOS Genital pore sinistral to acetabulum, fol- lowed by genital atrium diameter approxi- mately equal to acetabulum. Testes two, side by side in posterior 1/3 body, edges smcoth, oval in outline: right testis 0.011 to 0.051 long by 0.027 to 0.079 wide; left testis 0.056 to 0.118 long by 0.037 to 0.071 wide. Vasa deferentia joining anterior to ace- tabulum, connecting with right posterior margin of seminal vesicle. Seminal vesicle club-shaped, preacetabular, transverse to longitudinal axis of body, tapered to form slender sperm duct surrounded by prostate gland cells at distal tip. Sperm duct con- necting with fleshy intra-atrial collared geni- 86 Tulane Studies in Zoology and Botany Vol als Figure 5, Diagrammatic representation of the life-cycles of (A) Carneophallus choanophallus and (B) Carneophallus basodactylophallus. tal papilla, 0.045 to 0.062 long by 0.040 to 0.062 wide. Ovary between seminal vesicle and dextral testis; oval to oblong in out- line, edges lightly lobed, 0.030 to 0.077 long by 0.037 to 0.087 wide. Uterus de- scending from mesial intertesticular Mehlis’ gland to fill posterior body, ascending along body wall to posterior tip of dextral cecum, descending to posterior body, ascending along body wall to posterior tip of sinistral cecum, connecting with slightly enlarged metraterm surrounded by gland cells which in turn connects with genital atrium. Vitel- laria composed of 6 to 11 coarse follicles on each side of body, extending from pos- terior ends of ceca to posterior 1/8 of body. Uterine eggs operculate, 16 to 17 microns long by 10 to 12 microns wide. Excretory vesicle V-shaped; main stem extending an- teriorly from mesial excretory pore at pos- terior end of body, forking at level of pos- terior border of vitellaria, extending to level of testes. Flame cell pattern 2 [(2 +2) + 22-2) | 6: No. 3 HOSTS: Procyon lotor (L) [type host] and Rattus rattus (L), [natural hosts]; Mas musculus albinus (ICR strain), Rattus nor- vegicus albinus (Sprague-Dawley strain), Felis domesticus, Cavia porcellus, Microtus montanus and Sigmodon hispidus, |labora- tory hosts]. LOCATION: Small intestine. TYPE LOCALITY: Mouth of the Missis- sippi River at Pass a Loutre, Louisiana. HOLOTYPE: U. S. Nat. Mus. Helmin- thological Coll. No. 70428. Carneophallus choanophallus is distin- guished from the other species of the genus by the collared male papilla and the promi- nent follicles of the vitellaria extending from the tip of two ceca, a position well anterior to the acetabulum, to past the testes well into the posterior region of the body. The very short prepharynx, even in extended living worms, distinguishes it from all types except C. turgidus Leigh, 1958. Growth in Different Hosts An attempt was made to determine if growth variations occurred within hosts of the same species. Four groups of 6 to 12 laboratory white mice (ICR strain) were fed approximate numbers of C. choanophal- lus metacercariae from natural infections of M. ohione, and the mice were sacrificed 48 hr (2 days), 5 days and 25 days post-in- fection. The worms were removed from the intestine, killed in hot water, fixed in alcohol- formalin-acetic acid, stained and mounted in Permount® for measurement. Computa- tions were based upon measurements of 20 worms for each determination. The compara- tive similarity of organ sizes is illustrated in Figure 13, in which the body length and width, oral and ventral sucker width and ovary length are compared. The ovary de- creased in size as the worm aged. For intraspecific infections, laboratory white rats (Sprague-Dawley strain) and mice, a laboratory domestic cat, a rac- coon (Procyon lotor) and a frog (Rana pipiens) were used. The raccoon was obtained from the Audubon Park Zoo, where it had been maintained for at least two months. However, it was kept in the laboratory 30 days before use in this experiment at which time it was assumed to be free of C. choanophallus infection as pre- liminary studies with other animal hosts had Carneophallus Life Cycles 87 shown that these short-term infections last no longer than 30 days in the laboratory. The other animals (laboratory reared) were assumed to be free of C. choanophallus, The hosts were fed approximate numbers of metacercariae from naturally infected M. ohtone, and were sacrificed 48 hr post-in- fection with the exception of the frog. Since C. choanophallus does not become ovigerous in frogs maintained at room temperature, a frog was placed in an incu- bator at 37 C for 15 hr post-exposure after which time it yielded ovigerous adults from the small intestine. Organs which showed the least amount of variation in dimension were the pharynx and the male copulatory organ. The signifi- cant variations in measurements are _illus- trated in Figure 13. It is seen that for these determinations the adult worms from the white rat and the raccoon were significantly and consistently smaller than those from the mouse and the cat. The causative factors for this difference are not known, but the rat and the raccoon have been found to be the natural definitive hosts. Since the differing dimensions noted are comparatively minor and the genital atrium and male copulatory organ showed no significant variation in di- mension or morphology as a result of host variation, the specific diagnosis of C. cho- anophallus is considered to be taxonomically valid. Longevity of Infection in the Laboratory Mouse To determine the longevity of C. cho- anophallus infections, 72 mature adult lab- oratory white mice of the ICR strain (sexes equally divided) were each fed 50 meta- cercariae from old natural infections of Palaemonetes pugio. The mice were divided into 6 groups of 12 mice each and sacri- ficed at 5-day intervals post-exposure. The small intestine was removed from each mouse, slit open and scraped into saline in a 10.3-cm petri dish for examination under a dissecting microscope. Searching for and counting the worms as they were removed from the petri dish was done 4 times over a 5- to 6-hr period in order to make sure that all worms freed from the intestinal mucosa had fallen to the bottom of the dish where they could be seen. Preceding each examination, the intestine was again scraped and agitated. 88 Tulane Studies in Zoology and Botany Vol. ———I —_—— 001 -500 Worm body length (mm) 350 200 Worm body width (mm) 050 Oral sucker width (mm) 060 -040 Ventral sucker width (mm) -080 040 Ovary length (mm) Figure 13. C. choanophallus standard length frequency distribution of body length, body width, oral and ventral sucker width, and ovary length (based on 20 hot-water-killed whole mounted specimens per sample). Figures on the left represent 25-, 15-, 5- and 2-day-old infec- tions taken from white mice. Figures on the right represent 48-hr-old infections from white mice (M), white rats (R), a cat (C) and a rac- coon (Rc). In each sample the horizontal line indicates the range of the measurements; the cross-bar, the mean; the hollow rectangle, 1 standard deviation on each side of the mean; the solid rectangle, 2 standard errors on each side of the mean. The results of this experiment are illus- trated in Figure 14. On days 5 and 10 post- exposure, all of the mice (24 of 24) were infected with an average of 25 to 28 worms. On the 15-, 20- and 25-day examinations, the number of infected mice was 6 to 7 out of 12, with the average number of worms recovered ranging from 3.5 to 5.8. On day 30, no worms were recovered. Although Carneophallus Life Cycles 89 ted mice fp inte Number of worms Number of 5 10 15 20 25 30 Days post exposure Figure 14. Mean number ( ) and range (vertical lines) of adult C. choanophallus from the small intestine, and the number of mice in- fected (____ ) of 6 groups of 12 mice each ex- amined 5 to 30 days post-exposure with 50 metacercariae per mouse. there were differences between the males and females, these were not considered sig- nificant. The general trend was to maintain the infection at a relatively high level for about 10 days, then the density fell to a low level which lasted in some cases up to 25 days but less than 30 days, thus demon- strating the short-term character of infection with C. choanophallus in this definitive host. Many of the 15-, 20- and 25-day-old worms were packed with uterine eggs and showed less activity than was observed in the 5- and 10-day infections. Reinfection of 6 mice 40 days post-ex- posure resulted in all of the mice becoming infected for a second time, suggesting that previous C. choanophallus infections did not make the host refractory to reinfection upon challenge. The Metacercarta (Figures 3, 4, and 15-24) C. choanophallus encysted metacercariae were without exception found embedded in < Figures 6-12. Carneophallus choanophallus. 6. Adult, dorsal view, drawn from flattened and stained whole mount. 7. Excretory system, drawn from observations on living excysted metacer- cariae and adults. 8. Frontal section of adult genital atrium showing the collared nature of the male papilla. 9. Egg, taken from the uterus of an adult. 10. Xiphidiocercaria after emergence from snail. Drawn from living specimen stained with neutral red. 11. and 12. Sporocysts from the un- spined form of Lyrodes parvula. 90 Tulane Studies in Zoology and Botany VolLw5 No. 3 the shrimp muscle tissue, usually in one or more of the abdominal segments (Fig. 2), occasionally in the cephalothorax and rarely in the appendages. Infection of the shrimp took place when cercariae from the unspined form of Lyrodes parvula entered the gill chamber and pene- trated the gill filaments by cutting a slit with the stylet. As the cercaria entered through the slit, its tail became detached and was lost in the current of water passing through the gill chamber. Fourteen shrimp (12 M. ohione and 2 P. pugio) were ob- served to be infected by many cercariae in this manner, after which they were placed in an incubator at 27 C. During the first 18 hr post penetration, the cercariae remained in the gills, and were seen to burrow into the hemocoel. They slowly made their way in the hemocoel to the abdominal musculature where by 24 hr some were found encysted in a very thin and fragile membrane (Fig. 16). The metacer- caria, although completely enclosed within this membrane, was seen to be migrating about the hemocoel as it made its way to the abdominal musculature. These newly formed cysts measuring 0.050 by 0.045 in size, grew to 0.090 by 0.040 by 72 hr, and increased in size to 0.095 by 0.050 by 96 hr. At this time, the stylet appeared motile and the oral sucker and excretory vesicle were clearly visible (Fig. 17). Excysted metacercariae of these early stages revealed no additional organs (Figs. 15 and 18). By day 9, the metacer- caria contained in its thin (less than 0.002 thick ) pliable membrane measured 0.160 to 0.220 long by 0.113 to 0.165 wide (Fig. 20). The cyst wall constantly changed shape as the metacercaria moved about. When the 9-day- old metacercariae were excysted, a remnant of the stylet, the oral and ventral suckers, the pharynx, a small bulbous male papilla sur- rounded by a thin-walled genital atrium and well-developed excretory vesicles as well as the anlagen of the digestive and reproductive organs appeared visible (Fig. 19). By day 18, the metacercariae measured Carneophallus Life Cycles 91 0.360 to 0.375 long by 0.240 to 0.245 wide. The cyst wall appeared less pliable and mea- sured 0.002 thick (Fig. 22). All the organs usually seen in metacercariae except the vitellaria and the uterus were observed in the excysted metacercaria although they were reduced in size. Cuticular spines were first observed at this stage. The phallus ap- peared as a small unlobed muscular bulb within the genital atrium (Fig. 21). On day 23, the encysted metacercariae measured 0.335 to 0.345 long by 0.290 to 0.300 wide. The cyst wall was composed of two mem- branes, the outer one measured 0.003 and the inner one less than 0.002 thick (Fig. 23). The excysted metacercariae (0.445 to 0.455 long by 0.265 to 0.275 wide) dem- onstrated clearly the well-developed organs of the preadult metacercaria except for the vitellaria. The collar lobe of the phallus was seen to be developing (Fig. 24). By day 30, the metacercariae were in- fective to definitive hosts. At this stage, the excysted metacercariae contained small vitel- line follicles and the seminal vesicles were filled with motile sperm. It is suggested that until the vitellaria appear, the meta- cercariae will not survive in the definitive host, as 4 of 4 attempts to infect mice with young cysts before this stage failed. The encysted metacercariae measured 0.325 to 0.375 long by 0.240 to 0.285 wide. The cyst wall was composed of an inner membrane 0.005 thick and an outer thicker granular wall 0.007 to 0.010 thick. The excysted metacercariae moved about in good condi- tion much as the adults, and measured 0.450 to 0.550 long by 0.305 to 0.320 wide, similar to the adults. In further development, no change was seen in the excysted metacer- cariae. The size of the encysted metacercariae within the inner membrane apparently reached maximum size about day 30 at 27 C. As time passed, the outer granular layer of the cyst developed into an outer fibrous coat 0.040 to 0.060 thick, external to an extremely tough inner hyaline layer 0.010 to 0.015 thick (Figs. 3 and 4). Encysted metacercariae in M. ohione have been main- < Figures 15-24. Carneophallus choanophallus staged development of metacercaria in Macro- brachium ohione. (All figures were drawn from living material. ) 17-18. 96-hr stage in muscle, 17 encysted, 18 excysted. 19- muscle, 15 excysted, 16 encysted. 15-16. 24-hr stage in gills and 20. 9-day stage, 19 excysted, 20 encysted. 21-22. 18-day stage, 21 excysted, 22 encysted. 23- 24. 23-day stage, 23 encysted, 24 excysted. 92 Tulane Studies in Zoology and Botany Vol. 15 TABLE 2. Incidence and density of infection of different size classes of Macrobrachium ohione by Carneophallus choanophallus from September, 1966 to November, 1967.1 Numbers of Shrimp Incidence , Number of Metacercariae . of Infection Frequency _ Total Shrimp Incidence = of Size =Number Size l—- 1l- 21—- 3l1- 41- 51- of Total Class (%) ot Class 0 10 20 30 40 50 60 Infection? Class Sample Total Samp. Cysts 20-24 2 —- =- = = = = 0/2 - - 0.02 - 25-29 23 ltl - = = = = 1/24 4.16 0.01 0.24 1 30-34 15 Q - - = = = 2M eG: “O02 0.17 5 35-39 60 ll - - - = = NON /Fall 15.49 0.11 0.71 18 40-44 560/138 54° 2° =| IF f= 145/705 = 20.56 1.45 7.05 437 45-49 ee) ZN PAN GS) 460/1790 25.69 4.60 17.91 1804 50-54 14857489) 323° 14 42 1 2 Hoo) 20lGm Os LoS 20.19 2376 55-59 1449392 2 so 2) = 421/1870 22.51 4.21 18.71 1504 60-64 OF 306° 18 —2 1 = = S27/1518 21.54 3.27 15.18 1071 65-69 Ul Wee NO Oe ey ley 201/988 20.34 2.01 9.88 859 70-74 AS See OG ee) Oe J Po ee 99/537 18.43 0.99 5.37 264 75-79 ee SOm eo ee 34/211 1611 0.34 NIL 92 80-84 107 5 - = = = = 5/112 4.46 0.05 iit 13 85-89 131 =- = = FF 0/131 — - Mor - TOTAL hiss 2080) 9% 34 13° 15) 36 2239/9994 8444 % of Total Mean No. Cysts Sample 77.9 20.8 97 .34 .13 .05 .06 22.4/100.0 per Shrimp 0.84 ‘ Collections made from the Mississippi River at Ama, Louisiana. * Number of infected shrimp/number of shrimp in specified class. tained in the laboratory in good condition for up to six months when the shrimp died. Excystment of the young metacercariae in the laboratory was easily done by simply pricking the thin pliable cyst membrane and teasing the worm out. By the time the fibrous and hyaline layers developed, considerable time and patience was needed to break the tough membranes mechanically. If the cysts were placed in saline at room temperature for 4 to 5 hr, bacterial and perhaps chemical activity would weaken the walls so that ex- cystment was easily done with a sharp needle. With longer time, the worms were usually able to excyst without mechanical assistance. Geographical Distribution of Infections im M. ohione During the summer of 1967, a geographi- cal survey of the M. ohione infected with C. choanophallus metacercariae was con- ducted in 4 Louisiana rivers. Shrimp collec- tions were made using traps and by dredge- trawling behind a small outboard motor boat. A summary appears in Table 1. The West Pearl River yielded no M. ohione, however, infected P. pugio were found in great abundance in the brackish water marshes near its entrance into Lake Borgne. M. ohtone from the Red River were unin- fected. From the Atchafalaya River, 2 col- lections were made upstream from Morgan City, Louisiana. One shrimp harbored a single metacercaria, therefore, the incidence is considered to be very light. In the Mis- sissippi River, the incidence and intensity of infection was found to be relatively con- stant from Ama, Louisiana (mile 115) to Greenville, Mississippi (mile 540). The in- cidence and intensity of infection increased in stations downstream from Empire, Louisi- ana (mile 30). The increase in the first intermediate host, the unspined form of Lyrodes parvula, and the definitive host populations near the mouth of the Missis- sippi River where great expanses of fresh to brackish water marshes and ponds contain- ing Cabomba spp., Alternanthera spp. (alli- gator weed ) and Spartina spp. (marsh grass ) remote from disturbances by humans offer the best explanation for the high incidence and intensity of infection of M. ohione with C. choanophallus metacercariae in these areas. No. 3 Incidence and Density of Infection in M. ohione Over a 15-Month Period at Ama, Loutstana From September, 1966 to November, 1967, systematic biweekly collections of M. ohione were made from the west bank of the Mississippi River near the ruins of the old Sellers Plantation House at Ama, Louisi- ana. During this 15-month period, a total of 9,994 shrimp were captured. Of these, 2,239 (22.4 per cent) were observed under a dissecting microscope to harbor from 1 to 60 metacercariae. Two thousand eighty (20.8 per cent) shrimp harbored 1 to 10 cysts, leaving only 155 (1.5 per cent) shrimp harboring 11 to 60 cysts. Cysts usually occurred in the transparent abdomi- nal musculature, but were also seen in the muscles of the cephalothorax. The trapped shrimp ranged in size from 20 to 85 from the tip of the rostrum to the tip of the tel- son. The infected shrimp ranged from 25 to 80 in length. The 50 to 54 size class con- tained the largest number of shrimp in the sample. This size class also showed the great- est incidence and density of infection. The mean density of infection for the total sample was 0.84 metacercariae per shrimp (Table 2). Figure 25 shows the per cent of infected shrimp compared with the per cent of the total sample by size classes. It is apparent that the incidence of infection with C. cho- anophallus metacercariae is proportional to the percentage of all shrimp in the size classes for the 15-month period. These curves are typical for any month during the collecting period. Monthly standard length frequency dis- tributions of the uninfected and infected shrimp (Fig. 26) demonstrate that there are variations in these categories during the 15-month sampling period. Graphic repre- sentation of the data (Fig. 27) shows that during the months of January to April, the mean sizes of the uninfected shrimp were less than the infected shrimp. In March and April, overlapping took place, and for the remainder of the study time, the mean size of the uninfected shrimp was greater than the infected shrimp. This disparity was more noticeable during the months of May to August. Figure 27 shows that during the 15-month collecting period there were two maxima Carneophallus Life Cycles 93 Percent of total sample 20 40 60 80 Size class (mm) Figure 25. Incidence of infected M. ohione (___ __) compared with the per cent of the total sample (______) by size classes for collections made September, 1966 to November, 1967 at Ama, Louisiana. (November and July) and two minima (January and September) in the mean sizes of the shrimp sample. This suggests two breeding periods. Female shrimp in berry were collected beginning in April and ending in September. As the young shrimp began to appear in the collec- tion samples, the mean size decreased. Growth of the shrimp is indicated by the increase in the mean shrimp size. The greatest growth occurred in the spring. Figure 28 shows the mean density per shrimp of the total sample compared with the incidence of infection for the collection period. Two maxima (spring and fall) and two minima (winter and summer) are noted with the summer to fall disparity being the greatest. The maximum density and incidence occurred in the early fall. Examination of Figures 27 and 28 together shows that when the sample was of maxti- mum shrimp size (composed of larger shrimp), the density and incidence were minimal. This suggests that recruitment in the shrimp population took place at these times (November to January and June to August), and that these young uninfected shrimp did not appear infected in the col- lection samples until approximately a month later. There seem to be two periods of shrimp growth, both of which were accompanied by a decrease in the incidence and density of infection by C. choanophallus metacercariae. 94 Tulane Studies in Zoology and Botany Vol gS al a ee ( 67) Oct a ae (129) Nov 20 40 60 80 Shrimp size in mm of standard length Figure 26. Standard length frequency distribution of uninfected (CH80) and infected (11) samples of M. ohione collected at Ama, Louisiana, September, 1966 to November, 1967. In each sample the horizontal line indicates the range of the measurements; the crossbar the mean; the hollow rectangle, 1 standard deviation on each side of the mean; the solid or dotted rectangle, 2 standard errors on each side of the mean. Figures in the parentheses equal the number of shrimp in the sample. The growth periods are followed by an in- crease indicates that most infections flux of young shrimp into the sample ac- probably occurred early in the life-cycle companied by an abrupt rise in the incidence of the shrimp, but it does not preclude and density of the infected shrimp approxi- the possibility that infection of the mately a month later. This biseasonal in- shrimp occurred continuously throughout No. 3 o o in mm wn 11) 50 Mean shrimp size Mean density per shrimp Month Figures 27-28. 27. Above. Monthly mean size of the uninfected ( M. ohione. 28. Below. Monthly incidence (___ ) and mean density (_ Carneophallus Life Cycles 5 incidence Percent ) and infected (_ ) ) of the sample of M. ohione. Collections were made at Ama, Louisiana September, 1966 to November, 1967. the year. Observations indicated that the conditions for infection of juvenile shrimp, which live in shallow relatively still water where the unspined form of Lyrodes parvula exists, appear to be more likely than those for the adult shrimp, which live in deeper swift moving currents. Laboratory experiments showed that large and small M. ohione are both 100 per cent susceptible to C. cho- anophallus cercariae and that superinfection takes place. The pathological effects of infection on the shrimp are not clear. The larger mean size classes of the uninfected shrimp versus those 96 Tulane Studies in Zoology and Botany of the infected shrimp for June and July suggest that infection interfered with growth, or that mortality was higher in 1n- fected shrimp. The Cercaria (Figure 10) Small “monostome” xiphidiocercaria with the characters of the Ubiquita group. The following dimensions were determined using living cercaria, stained with neutral red or Nile-blue sulfate. Body 0.075 to 0.120 long by 0.035 to 0.055 wide, ovoid to elongate. Tail 0.060 to 0.110 long by 0.010 to 0.015 wide at base, with fine cuticular annulations. Oral sucker 0.020 to 0.025 in diameter. Stylet 0.015 to 0.017 long by 0.004 wide, base squared, shaft cylindrical then tapered to slightly ventrally directed point. Diges- tive system not observed. Two thin pairs of penetration gland cell ducts 0.020 to 0.030 long by 0.003 to 0.005 wide, opening into the anterior portion of the oral sucker lateral to stylet; the lateral gland cell ducts opening posterior to the mesial gland duct apertures. Cystogenous glands not promi- nently visible. Excretory vesicle U-shaped. Excretory formulayZaliCle = i) ae all)a, Develop in oval or elongate sporocysts con- taining about 10 to 15 cercariae. HOST: Lyrodes parvula Guilding, 1928 (unspined form). TYPE LOCALITY: Pass a Loutre, Louisi- ana. The cercaria of C. choanophallus in gen- eral resembled other microphallid larvae but differed from the cercariae of the genus Microphallus in the possession of only two pairs of penetration glands and in the mol- luscan host. The stylet was symmetrical in dorsal aspect but asymmetrical when viewed from the side. The cystogenous gland cells were seen only in cercariae which had emerged from the snail several hours before examination. Emergence of the cercariae was accom- plished by placing the snail in water with the salinity matching that of the field col- lection site and then slightly elevating the temperature under an electric light bulb. Emergence into water of the incorrect salinity either did not occur, or occurred very lightly and resulted in weakened short-lived cer- cariae. Normally the cercariae lived up to 48 hr. Vol. 15 Upon emergence, the cercariae swam al- most continuously with the posterior part of the body flexed ventrally and the tail lashing vigorously in S-shaped movements. These swimming movements served the pur- pose of keeping the cercaria suspended in the water off the bottom of the container. If the water was jarred by movement of the container or by touching the surface with a probe, the cercariae stopped swimming for a moment during which time the body would extend and contract once or twice before swimming was resumed. This type of ac- tivity aided the cercariae as they were drawn into the gill chamber of the shrimp for penetration of the gills. Contact with the gill filaments stimulated the movements of the stylet as it slashed an opening through which the squirming cercaria penetrated the shrimp. The Daughter Sporocyst (Figures 11 and 12) There are probably two sporocyst genera- tions of C. choanophallus in the unspined form of Lyrodes parvula, but only the cer- caria producing one was observed in the naturally infected snails studied in the labora- tory. Out of 200 of the unspined form of L. parvula from a locality in which virtually 100 per cent of the P. pugio were densely infected, 13 snails (6.5 per cent) were ob- served to contain C. choanophallus cercariae and sporocysts when crushed. The sporocysts were mostly elongate, but some were saccate in shape. They contained 10 to 15 cercariae usually with two of the cercariae ready for release; the others being in various stages of development. The birth pore was termi- nal as under light cover glass pressure, cer- Cariae were seen to emerge from this po- sition. The opposite end of the sporocyst contained many small rounded structures considered to be developing cercariae and germ balls. The snails examined contained from 5 to more than 40 sporocysts. Counting be- yond 40 was difficult as the sporocysts broke up so that they could not be clearly identi- fied for counting. Since four of the crushed snails were seen to contain more than 200 cercariae, it might be assumed that these snails may have contained more than 40 sporocysts. No. 3 The sporocysts measured 0.045 to 0.350 long by 0.020 to 0.080 wide. The Eggs and Miracidium (Figure 9) The process of egg formation in the adult worm was observed to be similar to that described by Cable and Hunninen (1940). Three hours post-exposure the adult worms contained a few (1 to 10) eggs in labora- tory rats and mice. As seen earlier, it ap- pears that the longer the worm stays in the intestine of the definitive host, the larger the number of eggs found in the uterus. For egg studies the uteri of 72-hr worms from mice infections were used. Whole worms were cultured in stendor dishes of saline and dechlorinated tap water at room temperature for daily observation. Every other day the water was changed and any debris from the dead worms was removed. Development of all cultures was as fol- lows: On day 1, the eggs measured 17-18 microns long by 9-11 microns wide. The operculum was clearly visible. Living cells in the process of dividing were observed near either end of the egg. On day 5, the contents of the egg appeared to be divided into sections or clumps. By day 10, a miracid- ium with the anterior end directed against the operculum was observed. At this stage, it was estimated that 75 per cent of the eggs were viable. By day 15, there seemed to be no change in the appearance of the miracid- ium. No free swimming miracidia were observed. Infection of the unspined form of Lyrodes parvula by C. choanophallus was accom- plished by casting whole ovigerous worms, taken from 7 to 20-day mice and rat infec- tions, into aquaria containing the snails. The snails had been collected in a pond at Bonnet Carre Spillway, St. Charles Parish, Louisiana. Two hundred snails were crushed, examined, and judged to be uninfected. An- other 200 snails from this collection were exposed and kept in 2 large finger bowls containing dechlorinated water, Cabomba spp. and Alternanthera spp. Five months later 16 snails survived. Of the survivors, 8 (50 per cent) were infected with C. choanophallus sporocysts containing cer- cariae. It is believed that the high per- centage of infection in the surviving snails was the result of the laboratory exposures. Carneophallus Life Cycles Dy Carneophallus from Callinectes sapidus The Life-Cycle While searching the Pass a Loutre area for the snail harboring the cercaria of Car- neophallus choanophallus during the sum- mer of 1967, the spined form of Lyrodes parvula was found. These snails when brought into the laboratory produced large numbers of a small xiphidiocercaria of the Ubiquita type which fit the suspected de- scription of Carneophallus cercariae. The snails also came from a known endemic area. Since many unsuccessful attempts were made to infect Macrobrachium ohione with this cercaria, other possible second inter- mediate crustacean hosts from the Pass a Loutre area were brought into the labora- tory for examination. Of these, the blue crabs, Callinectes sapidus, for this collec- tion, were found to be 100 per cent in- fected in the hepatopancreas and muscles with microphallid type metacercariae. When these metacercariae were excysted or fed to rats and mice, the resultant trematodes were found to exhibit the characters of the genus Carneophallus Cable and Kuns, 1951. A detailed study of these worms revealed the discovery of a second new species of Carneo phallus. As the cercariae from the spined form of L. parvula were suspected of infecting the blue crabs from Pass a Loutre, a search was made to find a locality with a popu- lation of blue crabs uninfected or very lightly infected with Carneophallus spp. of meta- cercariae. Such a population was located 12 miles west of the Mississippi River bridge on U. S. Highway 90 at the Osgood and Willswood ponds. Ten of these crabs were exposed to cercariae from the spined form of L. parvula resulting in medium (50 or more metacercariae) to heavy (100 or more metacercariae) infections whereas the thirty unexposed crabs were either negative or harbored a maximum of three older meta- cercariae. In the Pass a Loutre area, all raccoons, Procyon lotor (L), examined harbored large numbers of C. choanophallus and the sec- ond new species. Several birds were also examined but none of these harbored Car- neophallus spp. No other suspected natural definitive hosts in this area were examined. The life-cycle is as illustrated in Figure 98 Tulane Studies in Zoology and Botany No. 3 5 B. The natural definitive host was the raccoon, Procyon lotor (L), but possibly other mammals may also serve. The first intermediate host was a snail Lyrodes par- vula Guilding, 1828 (spined form). The second intermediate host was the blue crab, Callinectes sapidus Rathbun, 1896. Description of the Stages of the Life History of Carneophallus basodactylophallus n. sp. The Adult (Figures 29-30) SPECIFIC DIAGNOSIS: [based on 10 hot water-killed, unflattened specimens from Rattus norvegicus albinus (Sprague-Dawley strain) and Mas musculus albinus (ICR strain) ]. Microphallidae; Carneophallus. Body pyriform, sometimes with posterior notch, 0.415 to 0.455 long by 0.285 to 0.330 wide. Forebody 0.187 to 0.212 long. Integument of anterior 1/2 of body spined. Oral sucker subterminal, 0.050 to 0.062 long by 0.050 to 0.062 wide. Prepharynx almost absent. Pharynx 0.022 to 0.025 long by 0.022 to 0.027 wide. Esophagus extending from pharynx to approximately anterior 1/3 body, 0.087 to 0.110 long. Ceca two, extending posteriorly and obliquely from cecal bi- furcation at posterior end of esophagus, 0.125 to 0.152 long. Acetabulum equatorial, mesial, 0.052 to 0.065 long by 0.052 to 0.062 wide. Sucker ratio 1:1.02 to 1.07. Genital pore sinistral to acetabulum, fol- lowed by genital atrium approximately 1.25 times diameter of acetabulum. Testes two, side by side in posterior 1/3 body, edges smooth, oval in outline: dextral testis 0.022 to 0.052 long by 0.065 to 0.095 wide; sinis- tral testis 0.027 to 0.050 long by 0.050 to 0.095 wide. Vas deferens joining anterior to acetabulum, connecting with dextral pos- terior margin of seminal vesicle. Seminal vesicle club-shaped, preacetabular, trans- verse to longitudinal axis of body, tapered to form slender sperm duct surrounded by prostate gland cells at distal tip. Sperm duct connecting with fleshy intra-atrial genital Carneophallus Life Cycles 99 papilla (0.060 to 0.062 long by 0.030 to 0.045 wide) with small posterior basal flap. Ovary between seminal vesicle and dextral testis; oval to oblong in outline, edges lightly lobed, 0.052 to 0.075 long by 0.050 to 0.075 wide. Uterus descending from mesial inter- testicular Mehlis’ gland to fill posterior body, ascending along body wall to posterior tip of dextral cecum, descending to posterior body, ascending along body wall to posterior tip of sinistral cecum, connecting with slightly enlarged metraterm surrounded by gland cells which in turn connects with genital atrium. Vitellaria composed of 6 to 11 coarse follicles on each side of body, extending from posterior ends of ceca to posterior 1/8 body. Uterine eggs operculate, 16-17.5 microns long by 10-12 microns wide. Excre- tory vesicle V-shaped; main stem extending anteriorly from mesial excretory pore at posterior end of body, forking at level of testes. Flame cell pattern 2 [(2+-2) + GE 2) | Ss16: HOSTS: Procyon lotor (L) (type host) [natural host]; Rattus norvegicus albinus (Sprague-Dawley strain) and Mus musculus albinus (ICR strain), [laboratory hosts]. LOCATION: Small intestine. TYPE LOCALITY: Mouth of Mississippi River at Pass a Loutre, Louisiana. HOLOTYPE: U. S. Nat. Mus. Helmin- thological Coll. No. 70430. Carneophallus basodactylophallus is dis- tinguished from the other species of the genus by the small posterior flap on the base of the male papilla. The very short pre- pharynx, even in extended living worms, distinguishes it from all types except C. tur- gidus Leigh, 1958 and C. choanophallus. The Metacercarta C. basodactylophallus metacercariae were found encysted in the hepatopancreas and muscle tissue of the cephalothorax and all the appendages of the blue crab Callinectes sapidus Rathbun, 1896. They are morpho- logically similar to C. choanophallus meta- cercariae. & Figures 29-33. Carneophallus basodactylophallus. 29. Adult, dorsal view, drawn from a flat- tened and stained whole mount. 30. Excretory system, drawn from observations on living excysted metacercariae and adults. 31. Frontal section of the adult genital atrium showing the posterior basal flap of the male papilla. 32. Xiphidiocercaria after emergence from snail. Drawn from living speci- men stained with neutral red. 33. Sporocyst from the spined form of Lyrodes parvula. 100 Laboratory infections of the blue crab took place when cercariae from the spined form of Lyrodes parvula were allowed to enter the gill chamber where penetration of the gills took place. Ten of ten (100 per cent) exposed crabs became moderately (50 or more metacercariae) to heavily (100 or more metacercariae) infected versus two of thirty (6.6 per cent) of the controls which were found to be very lightly (1 to 3 meta- cercariae) infected. Migration of the cer- cariae in the crab hemocoel was not ob- served. However, the cercariae wriggled their way to the hepatopancreas and the muscle where encystment took place in thin mem- branes. Development of C. basodactylophallus in the blue crab seemed to be slower than C. choanophallus in shrimp, since after 40 days post-exposure at room temperature the thick- ened cyst wall of the metacercaria had not yet developed. Newly formed encysted metacercariae measured 0.050 by 0.035 in diameter. Fully developed metacercariae measured 0.330 to 0.360 long by 0.325 to 0.355 wide. The cyst wall was composed of an inner mem- brane 0.005 thick and an outer hyaline layer 0.010 to 0.015 thick surrounded by a thick fibrous coat 0.040 to 0.060 thick. Of the geographical areas checked, the in- cidence of infection at Pass a Loutre was 85 per cent, and 9.2 per cent at the Wills- wood and Osgood ponds. Ten crabs taken from the canals of the Bonne Carre Spillway were all negative for C. basodactylophallus. The Cercarta (Figure 32) Small “monostome” xiphidiocercaria with the characters of the Ubiquita group. The following dimensions were determined using living cercariae stained with neutral red and Nile blue sulfate. Body 0.080 to 0.120 long by 0.040 to 0.066 wide, ovoid to elongate. Tail 0.060 to 0.110 long by 0.010 to 0.015 wide at base with fine cuticular annulations. Oral sucker 0.025 to 0.030 in diameter. Stylet 0.012 to 0.015 long by 0.004 wide, base rounded, shaft cylindrical then tapered to slightly ventrally directed point. Digestive system not observed. Two thick pairs of penetration gland cell ducts, lateral pair 0.035 to 0.040 long by 0.009 to 0.012 wide, mesial pair 0.030 to 0.035 long by Tulane Studies in Zoology and Botany Vol. 15 0.007 to 0.009 wide, opening into anterior portion of oral sucker lateral to stylet, lat- eral gland cell ducts opening anterior to mesial duct apertures. Cystogenous gland cells not prominently visible. Excretory vesicle U-shaped. Excretory formula 2 [| (1 + 1) + (1 +1) J. Develop in oval or elongate sporocysts containing about 40 to 60 cer- cariae, HOST: Lyrodes parvula Guilding, 1828 (spined form). TYPE LOCALITY : Pass. a) Loutte, Mouisi- ana. The cercariae of C. basodactylophallus closely resemble those of C. choanophallus. The notable differences were the shorter stylet with its rounded base, the enlarged penetration gland cell ducts, and the snail host; the spined form of L. parvula for C. basodactylophallus. The cercariae were in- capable of producing metacercariae in M. ohione. Conditions for emergence and sub- sequent activity of the cercariae appear to be similar to C. choanophallus. The Daughter Sporocyst (Figure 33) There are probably two sporocyst genera- tions of C. basodactylophallus in the spined form of Lyrodes parvula, but only the cer- cariae producing one was observed in natu- rally infected snails studied in the laboratory. Of 100 spined forms of L. parvula from Pass a Loutre, 2 were observed to contain C. basodactylophallus cercariae when crushed. The sporocysts were elongate to saccate in shape. They contained from 40 to 60 cer- cariae, many of which were ready for re- lease. The birth pore was terminal. The snails examined contained 5 and 30 sporocysts which measured 0.400 to 0.550 long by 0.150 to 0.300 wide. The sporocysts of C. basodactylophallus were longer and contained more cercariae than those of C. choano phallus. The Eggs and Miracidinm Morphologically, the eggs of C. basodacty- lophallus could not be distinguished from those of C. choanophallus. Measurements were 17-18 microns long by 9-11 microns wide. No attempts were made to culture the eggs and miracidia were not observed. Discussion The erection of the four sub-families within the Microphallidae (Deblock and Tran Van Ky, 1966) provides a realistic and convenient division of the numerous species of microphallids into related groups based on the morphology of the copulatory structures. Following Cable and Kuns (1951) the sub- family Maritreminae (Lal, 1939) Deblock and Tran Van Ky, 1966 would be con- sidered primitive. The sub-family Micro- phallinae Ward, 1901 could have arisen as one stem from Maritreminae. Evolution in this case is suggested to have proceeded from the primitive forms possessing a cir- rus and cirrus sac to forms in which these structures are lost. This loss is thought to be the result of evolutionary modifica- tion resulting in the numerous morphologi- cal variations of the genital atrium and male papilla presently demonstrated by these forms. The sub-families Gynaecotylinae Gus- chanskaia, 1952 and Sphairiotreminae De- block and Tran Van Ky, 1966 may be thought of as having arisen as another stem from a more primitive source or from the Maritreminae as a result of evolutionary modification to a vesiculo-prostatic sac which replaced the cirrus pouch. The discussion in this paper is concerned with the development of the genera and their species within the sub-family Micro- phallinae Ward, 1901. Biguet, Deblock and Capron (1958) have continued to follow the reasoning of Cable and Kuns (1951) in their listing of four genera of the Micro- phallinae (Leninseniella Stiles and Hassal, 1901; Spiculotrema Belopolskaia, 1949: Microphallus Ward, 1901; Endocotyle Belo- polskaia, 1952) based upon the morphologi- cal attributes of the genital atrium and its contained male copulatory papilla. Biguet, Deblock and Capron (1958) considered the genus Carneophallus Cable and Kuns, 1951 redundant. They argued that lobation of the male copulatory organ although a morpho- logical fact, is not significant enough to wafrant a separate genus. Cable, Connor and Balling (1960) objected to lumping of heterogeneous groups into the single genus Microphallus, contending that the adult morphology alone was not a_ satisfactory basis upon which to determine the taxo- nomic relationships of the Carneophallus group. This writer agrees with Cable, Con- Carneophallus Life Cycles 101 nor and Balling (1960) and considers the lumping of the many species listed under the genus Maicrophallus, based not upon shape and form of the phallus but upon its comparative dimensions (Biguet, Deblock and Capron, 1958) to be a deviation from the method used to this point. If the de- velopment of a thickened ornamented wall of the genital atrium is considered reason enough for the maintenance of the genus Levinsentella (Biguet, Deblock and Capron, 1958), then it also seems logical to sep- arate Out those species which have developed a large fleshy-lobed male papilla under the genus Carneophallus. Except for a few bor- derline cases (i.e. Carneophallus muellhaupti Coil, 1956 which possesses a lightly lobed male papilla), the lobation of the male papilla is strikingly obvious, illustrating further evolutionary modification of this structure as well as convenient division of a relatively large heterogeneous group of species. This reasoning has prompted this writer to place the two new species described in this paper into the genus Carneophallus Cable and Kuns, 1951. Cable and Kuns (1951) distinguished the genus Carneophal- lus as microphallids with a large unorna- mented thin-walled genital atrium which is almost filled with a lobed fleshy male papilla, one of the lobes being penetrated by the ejaculatory duct. Using this distinc- tion and the original descriptions, the fol- lowing twelve species are segregated: C. trilobatus Cable and Kuns, 1951. C. pseudogonotylus (Chen, 1944) Cable and Kuns, 1951. C. muellhaupti Coil, 1956. C. skryabini Caballero, 1958. C. turgidus Leigh, 1958. C. chabaudi (Capron, Deblock and Biguet, 1957) Cable, Connor and Balling, 1960. C. tringae (Capron, Deblock and Biguet, 1957) Cable, Connor and Balling, 1960. C. canchei (Biguet, Deblock and Capron, 1958) Cable, Connor and Balling, 1960. C. bilobatus Cable, Connor and Balling, 1960. C. lactophrysi Siddiqi and Cable, 1960. C. choanophallus n. sp. C. basodactylophallus n. sp. The specific name, choanophallus, indi- cates the collared nature of the phallus. The 102 term, basodactylophallus, describes a phallus possessing a finger or flap at its base. These two characters distinguish these two new species from all the others. Further support for the validation of the genus Carneophallus is obtained from growth studies in definitive hosts. Baer (1943) and Stunkard (1960) pointed out that growth in microphallids takes place in the second intermediate host, which is usually a crusta- cean. Staged development of C. choanophal- /us in shrimp demonstrated the remarkable increase in body and organ size that takes place at this stage in the microphallid life- cycle. It follows then that no significant growth takes place in the worm during its short-term infection in the intestine of the definitive host. Minor differences in worm dimensions were noted in worms recovered from different hosts. Such interspecific host growth variations have also been studied by Hunter (1952) for Gynaecotyla adunca. Hunter's (1952) observations agree with those of C. choanophallus in that in normal hosts the worms were smaller than in ab- normal hosts. In the case of C. choanophallus, these growth variations are minor, but they tend to cast doubt upon the classification system of Biguet, Deblock and Capron (1958) based upon comparative sizes of the male papilla and the ventral sucker rather than morphology for the species of the genus Microphallus. The shape and form of the male papilla, which was seen to be characteristically constant in C. choanophal- lus and C. basodactylophallus from all hosts observed, appears to be a character of enough stability to warrant the validation of the genus Carneophallus. Cable (1956) and Cable, Connor and Balling (1960) described differences in stylet and cephalic gland morphology of cercariae which they believed to be of tax- onomic significance. Cable, Connor and Ball- ing (1960) for this reason maintained that when the life-cycles of members of the genus Carneophallus became known, that addi- tional evidence to support the validity of the genus would appear. The cercariae of both C. choanophallus and C. basodacty- lophallus possess only two pairs of cephalic glands. Heard (personal communication ) also observed Carneophallus spp. cercariae with only two pairs of cephalic glands. Ex- cept for the cercaria of Maritrema caridinae Tulane Studies in Zoology and Botany Vol. 15 described by Shibue (1951), all microphal- lid cercariae are described as possessing four pairs of cephalic glands. It cannot be stated that all Carneophallus cercariae possess only two pairs of cephalic glands on the basis of the two cercariae seen out of the possible 12 species of these microphallids known. However, this peculiar difference observed in these two species lends further evidence to support the validity of the genus Car- neophallus Cable and Kuns, 1951. Experimental infection of Macrobrachium ohione and Palaemonetes pugio with cer- cariae of Carneophallus choanophallus dem- onstrated that this microphallid may utilize more than one species of shrimp as the sec- ond intermediate host. Microphallus minus Ouchi, 1928 in Japan and China is reported to encyst in three species of shrimp (Ouchi, 1928 and Yeh and Wu, 1951), Macro- brachium nipponensis, Palaemon asperulus and P. nipponensis. Repeated attempts to infect shrimp with C. basodactylophallus cercariae and crabs with C. choanophallus cercariae failed in this laboratory, thus con- firming the taxonomic relationship and sec- ond intermediate host specificity of these two very closely related species. The significance of the second intermedi- ate host specificity demonstrated by micro- phallids as it might reflect phylogenetic relationships is not clear. A case in point is Microphallus claviformis Brandes, 1888 (Syn. M. lamuli Stunkard, 1951). Deblock and Rosé (1965) discovered the metacer- cariae of M. claviformis in an isopod, Sphoe- roma serratum, following the invalidation of M. limuli by Biguet, Deblock and Capron (1958). Stunkard (1951, 1960) recognized the close resemblance of the excysted meta- cercariae of M. limuli and M. claviformis, but because the former were in the chelic- erate, horse-shoe crab, Limulus polyphemus, which is quite unrelated to the typical arthropod second intermediate host, he was reluctant to regard it a synonym. The manner in which microphallid cer- cariae enter the crustacean second intermedi- ate host was first described by Cable and Hunninen (1940). This description is not unlike that observed for C. choanophallus in shrimp with one interesting addition. Within 48 hr of penetration and passage into the hemocoel, the C. choanophallus cercariae have secreted thin protective mem- No. 3 branes which completely surround them, and within which they were seen to con- tinue to migrate toward the shrimp ab- dominal musculature. The early presence of this membrane suggests that it may serve to protect the cercariae from host anti- bodies. The presence of the cercariae and encysted metacercariae in the shrimp muscle apparently elicited no significant reaction as shrimp were not refractory to super- infection upon challenge to reinfection. The functions of the cyst wall and the methods by which nutrition and excretion across the wall are accomplished are not well under- stood at this time, but in view of the energy requirements of the developing larva would form the basis for an exciting study. Hunter and Vernberg (1953) and Ching (1962 a, b) demonstrated stages in the de- velopment and growth of microphallid metacercariae in the second intermediate host. This remarkable growth period in the life-cycle of C. choanophallus reached the stage of infectivity to mice when the vitel- line glands appeared on about day 30 at 27C in shrimp. The criteria for infectivity of the definitive host are not known. In this study, the muscular organs: the two oral suckers, the pharynx and the male papilla all appeared at the same time on about day 9. In general the staged develop- ment of C. choanophallus in shrimp is similar to that described previously for other micro- phallids. The necessity of approaching a trematode life history study from the ecological point of view is mentioned in Cable and Hun- ninen’s (1940) elucidation of the life history of Spelotrema nicolli as the key to finding the snail which shed cercariae. This was likewise true in the search for the cercaria of C. choanophallus, for it was not until a geographical survey led the way to marshes and ponds at the mouth of the Mississippi River where heavy infections of shrimp were found that snails bearing C. choanophallus cercariae were discovered. The question as to how shrimp taken from the Mississippi River at Greenville, Mississippi 540 miles from the mouth of the river be- came infected deserves some comment. It would not be possible for shrimp to migrate such a distance up-stream especially since young 20- to 25-mm shrimp were found in- fected. Infection must have occurred lo- Carneophallus Life Cycles 103 cally, but since infected snails were never found except at the mouth of the river, one may speculate that another yet-to-be- found snail is responsible, or that since shrimp infections are markedly lower in the regions up-river, the incidence in the snails is correspondingly so low as to make capturing an infected one unlikely. Since the Mississippi River rises and falls in water level several times a year, flooding into and communication with ponds and marshes along the banks, it is reasonable to assume that certain of these ponds not yet visited by this investigator could support popula- tions of snails infected with C. choanophal- lus cercariae. Rankin (1940) reported that Gynaeco- tyla nassicola, adults of which are harbored by migratory birds, showed a_ seasonal fluctuation in the snail Nasa obsoleta, the highest incidence being in the spring. No report was made on the incidence of in- fection for the second intermediate host. Studies on the life histories of Maritrema obstipum and Levinseniella amnicolae by Etges (1953) indicated that the highest incidence in the first intermediate host snail, Amnicola pilsbryi, and the second inter- mediate host, Asellus communis, infections were in the late summer and fall. Etges (1953) indicated that the periodicity of these infections was due to the migratory habits of birds which serve as the definitive hosts. In experiments with C. choanophallus, time has not permitted an extended survey of the incidence of infection in the first intermediate host, the unspined form of Lyrodes parvula, Present collections through- out the winter showed an incidence of 1 to 7 per cent suggesting that these snails may be continuously shedding cercariae when- ever conditions permit. Since this study demonstrated that infected shrimp are found throughout the year, the definitive hosts (raccoon and rat) may maintain continuous short-term infections seeding snails at all seasons of the year in the subtropical New Orleans, Louisiana area. It may be that the short-term infections of the definitive host in microphallids serves no other purpose than to disperse the eggs. Sogandares-Bernal and Lumsden (1964) suggested that in short-term infections of Ascocotyle leigh Burton, 1956, which encysts in the hearts of certain poeciliid and cyprinodont fishes, 104 and in microphallids, ovigerous worms may be passed in natural infections, much in the same way the gravid proglottids of certain cestodes are passed from the definitive host. They pointed out that the short-term infection of the definitive host is probably the result of an evolutionary sequence which has allowed for the worms to continue with their biological function of reproduction without harming the host. The usual long- term infection with these parasites could lead to the death of the definitive host if the minute eggs were to enter the circula- tory system with resultant complications (Africa, Garcia and de Leon, 1935a, b, 19365 19377). The 15-month study of M. ohione infected with C. choanophallus demonstrated two seasonal upsurges in incidence and intensity of infection with the heaviest infection oc- curring in the late summer and early fall. These two upsurges in M. oione infection were seen to follow two periods of recruit- ment of young shrimp into the samples sug- gesting that the infection in M. ohione oc- curred mainly in young shrimp twice a sea- son. The fact that juvenile shrimp in en- demic areas are usually found in shallow still water containing vegetation upon which the unspined form of Lyrodes parvula is normally found substantiates this point of view. This does not rule out the possibility that infection of older shrimp, which are found in deeper swift moving water may take place continuously at a low level throughout the year with activity declining during the cold winter months. Acknowledgments This study was done under the direction of Dr. F. Sogandares-Bernal. Acknowledg- ments are extended to Dr. Norman C. Negus, for advice on the ecological portions of this work, to Dr. John C. Hitt, for advice on computerization of the data and its analysis, to Dr. Harold W. Harry, for identi- fication of the snail hosts, to Dr. Emile A. Malek, for aid in the collection of snails, to Dr. W. A. Eggler, for identification of aquatic plants, and to Dr. A. E. Smalley, for confirming the shrimp identifications. I am especially indebted to Dr. T. B. Ford and Mr. Allan Ensminger, Louisiana Wildlife and Fisheries Commission, for use of facilities and other aid. Tulane Studies in Zoology and Botany Vol REFERENCES CITED Arrica, C. M., E. Y. Garcia, and W. DE LEON. 1935a. Intestinal heterophyidiasis with car- diac involvement. A contribution to the eti- ology of heart failure. Philipp. J. Publ. ae Poilory. ome = 1935b. Heterophyidiasis. II. Ova in sclerosed mitral valves with other chronic lesions in the myocardium. J. Philipp. Islands Med. 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Case, R. M. 1956. Marine Cercariae of Puerto Rico. Sci. Survey of Puerto Rico and the Vir- gin Islands. N. Y. Acad. Sci., 16:491-577. Case, R. M., R. S. CoNNor, and J. W. BALLING. 1960. Digenetic trematodes of Puerto Rican shore birds. Sci. Survey Porto Rico and Virgin Isl. N. Y. Acad. Sci., 17:187-254, 48 figs. CasBLe, R. M. and A. V. HUuNNINEN. 1940. Studies on the life history of Spelotrema nicolli (Trematoda: Microphallidae) with a description of a new microphallid cercaria. Biol. Bull., 78:136-167. CaB_e, R. M. and M. L. Kuns. 1951. The trem- atode family Microphallidae with the de- scription of Carneophallus trilobatus gen. et sp. nov., from Mexico. J. Parasit., 37:507-514. Cuinc, Hitpa Ler. 1962a. The description and life cycle of Maritrema laricola sp. n. (Trema- toda: Microphallidae). Canad. J. Zool., 41 881-888. ts 1962b. The life cycle and bionomics of Levinseniella charadri- formis Young, 1949 (Trematoda: Microphal- lidae ). Canad. J. Zool., 41:889-899. Deswock, S. and F. Rosé. 1965. Contribution a Tetude des Microphallidae Travassos, 1920 (Trematoda) des oiseaux de France. XI. Identification de Ja cercaire de Microphal- lus claviformis (Brandes, 1888). Bull. Soc. Zool. France, 90:299-314. Des.ock, S. et P. TRAN VAN Ky. 1966. Contri- bution a l’etude de Microphallidae Travassos, 1920 (Trematoda). XII. Especes d’Europe No. 3 occidentale. Creation de Sphairiotrema nov. gen.; considerations diverses de systematique. Annales de Parasitologie (Paris), 41:32-60. Erces, F. J. 1953. Studies on the life histories of Maritrema obstipum (Van Cleave and Muller, 1932) and Levinseniella amnicolae n. sp. (Trematoda: Microphallidae). J. Par- asit., 39:643-662. Hearp, R. W. 1968. ( Personal Communication ) University of Georgia, Athens, Ga. Hunter, W. S. 1952. Contributions to the mor- phology and life-history of Gynaecotyla adunca (Linton, 1905) (Trematoda: Micro- phallidae). J. Parasit., 38:308-314. Hunter, W. S. and W. B. VERNBERG. 1953. 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(Trematoda: Microphallidae ). Biol. Bull., 101:307-318. ‘ . 1960. Problems of the ge- neric and specific determination on digenetic trematodes with special reference to the genus Microphallus Ward, 1901. Lib. Ho- menaje Dr. Eduardo Caballero y C. Mexico, D. F., pp. 299-309. VAN CieEAvE, H. J. 1953. Acanthocephala of North American Mammals. III. Biol. Monogr., 23:1-179. YEH, G. and K. Wu. 1951. Progenesis of Micro- phallus minutus Ouchi (Trematoda: Micro- phallidae) in fresh water shrimps. Peking Nat. Hist. Bull., 19(203):194-209. March 24, 1969 ETHEOSTOMA COLLETTEI, A NEW DARTER OF THE SUBGENUS OLIGOCEPHALUS FROM LOUISIANA AND ARKANSAS! RAY S. BIRDSONG? Institute of Marine Sciences, University of Miami Rickenbacker Causeway, Miami, Florida and LESLIE W. KNAPP Smithsonian Oceanographic Sorting Center Washington, D. C. ABSTRACT Etheostoma (Oligocephalus) collettei is described from 308 specimens from the Ouachita River drainage in Louisiana and Arkansas and the Little, Red and Sabine river drainages of Louisiana. Etheostoma collettei appears to be most closely related to E. asprigene from which it is differenti- ated primarily by color pattern. A new species of the subgenus Olzgo- cephalus, herein described, is well repre- sented in collections from Louisiana and Arkansas, but has previously been confused with E. asprigene (Forbes). John D. Black and Carl L. Hubbs, in their studies of Arkan- sas fishes during the early 1940's, recognized this undescribed form and regarded it as a subspecies of E. asprigene. The new species is known from the Ouach- ita River in Louisiana and Arkansas and from the Little, Red and Sabine river drain- ages of Louisiana. It probably occurs in the Calcasieu River drainage, but no collections from this drainage have been examined. We gratefully acknowledge the assistance of the following: Neil H. Douglas, North- east Louisiana State College (NLSC), who provided specimens, Kodachrome transpar- encies and habitat notes on the type-locality; Bruce B. Collette, Bureau of Commercial Fisheries Systematics Laboratory, offered valuable suggestions and reviewed the manu- script; W. Ralph Taylor, U. S. National Museum (USNM), provided field data; Neil Hotchkiss, Patuxent Wildlife Research Center, identified aquatic plants; Ernest A. Lachner, United States National Museum (USNM), Edward C. Raney, Cornell Uni- versity (CU), Reeve M. Bailey, University of Michigan Museum of Zoology (UMMZ), Ralph W. Yerger, Florida State University (FSU), and Frank Cross, University of Kan- sas (UK) kindly loaned specimens; C. Rich- ard Robins, Institute of Marine Sciences, University of Miami, Royal D. Suttkus, Tu- lane University (TU) and William J. Rich- ards, Bureau of Commercial Fisheries Trop1- cal Atlantic Biological Laboratory, reviewed the manuscript. Measurements and counts were made fol- lowing Hubbs and Lagler (1958:8-15) ex- cept where otherwise indicated. It is our pleasure to name this new darter for Bruce B. Collette, Bureau of Commercial Fisheries Systematics Laboratory, in recogni- tion of his contributions to our knowledge of percid fishes. Etheostoma (Oligocephalus) collettei, new species Creole Darter (Fig. 1) Material. The description is based on 308 specimens from Louisiana and Arkansas. Holotype, USNM 201515 (¢ 56.3 milli- ! Contribution No. 1006 from the Institute of Marine Sciences, University of Miami. EDITORIAL COMMITTEE FOR THIS PAPER: 2 Present address: Old Dominion College, Department of Biology, Norfolk, Virginia 23508. Dr. REEVE M. BAILEY, Curator of Fishes, Museum of Zoology, University of Micht- gan, Ann Arbor, Michigan Dr. RoyAL D. SutTrKus, Professor of Biology, Tulane University, New Orleans, Louisiana 106 No. 3 meters in standard length), Dugdemona River (Little River drainage), TISN, R3W, Section 6, Jackson Parish, La., 12 December 1966, Neil H. Douglas and Jacob P. Yelver- ton. Paratopotypes, USNM 201516 (38, 299581) Lower Ouachita River paratypes: NLSC 6317 (20, 29.9-52.9), Choudrant Bayou at La. Hwy. 15 bridge, Ouachita Parish, La., 29 September 1966, Douglas, K. Burnside, and H. Stegall; Univ. Ala. 2575 (17, 28.8- 33.7), same data as NLSC 6317; CU 52108 (7, 40.8-61.5), Meridian Cr. on La. Hwy. 348, 1 mi. E. of Conway, Union Parish, La., T22N, R1W, Section 17, 12 November 1966, Douglas, Burnside and Yelverton; Tulane Univ. uncat. (10, 41.7-51.3), same data as CU 52108 (specimens apparently lost in transit); FSU 14678 (7, 40.5-60.0), Meridian Cr., T22N, R1E, Section 21, Union Parish, La., 15 June 1965, Douglas; USNM 172913 (14, 23.6-53.8), Meridian Cr., 1 mi. E of Conway, T22N, R1E, Section 17, Union Parish, La., 22 June 1965, Taylor; USNM 201517 (2, 51.9-52.1), Corney Lake Spill- way, Claiborne Parish, La., W. Harman and H. Hobbs; USNM 172538 (4, 40.1-54.6), Little Corney Bayou at La. Hwy. 15, T22N, R3W, Section 1, Union Parish, La., Taylor; TU 52876 (5, 44.9-54.8), Meridian Cr., 1 mi. E of Conway, T22N, R1E, Section 17, Union Parish, La., 21 May 1956, Taylor and Lowe. Upper Ouachita River paratypes: CU 42248 (71, 26.7-49.2), Middle Fork, Saline R. about 6 mi. W of Crows, Saline Co., Ark., 27 April 1962, Knapp and R. V. Miller; USNM 165915 (52, 25.5-48.5), South Fork, Saline R., 7.7 mi. SSW of Owensville, Gar- land Co., Ark., 22 April 1952, Lachner et al; UMMZ 123229 (2, 30.0-31.9), Gulpha Cr. at jet. with Lake Hamilton, 10 mi. E of Hot Springs, Garland Co., Ark. 19 June 1938, J. and R. Black; USNM 165953 (9, 36.6-42.9), Antoine Cr., Trib. of Little Mis- souri R., 2.5 mi. N of Kirby, Pike Co., Ark., 21 April 1952, Lachner et al. Little River paratypes: FSU 14676 (23, 35.5-45.6), Little Cr, T5N, RIE, Section 23, Rapides Parish, La, 21 June 1965, Douglas. Red River paratypes: UMMZ 113734 (0: 36:9-50:6), 72) mi. S ‘of ‘Dry Prong, Grant) Parish, Tas 29. May 1932; Creaser, Hedrick and Amell. Etheostoma collettei 107 Sabine River paratypes: FSU 14677 (8, 36.0-47.5), Sandy Cr. on La. Hwy. 11, T2N, R11W, Section 20, Vernon Parish, La., 24 June 1965, Douglas. Comparisons with Etheostoma asprigene were based primarily on the following ma- terial from the Ouachita River drainage: USNM 173058 (20, 28.2-35.6); USNM 172482 (4, 19:2-25.7); USNM 172759 (2, AS°3-49'5))/ WSINIMiily 2517 Ie Ol. 5522) USNM 172296 (16, 32.1-43.4); UK 3527 (1, 35.5); UMMZ 169795) (3, 343-419). Diagnosis. Distinguished from other mem- bers of the subgenus Oligocephalus by a combination of the following characters: dorsum with 3 or 4 prominent, dark blotches and 4 or 5 less conspicuous blotches; usually a prepectoral dark blotch; humeral spot pro- nounced; a dusky bar on body beneath pec- toral fin; darkly pigmented scale centers contribute to the formation of horizontal lines on the body that tend to disappear ventrally about 4-5 scale rows below the lateral line; anal spines 2; belly scale edges thickened as tubercles in a few male speci- mens; prepectoral area and breast naked; cheeks, opercles, nape and belly with scales; eye ovoid and breaks the dorsal outline in profile; branchiostegal membranes slightly joined to overlapping; lateral line nearly straight and incomplete (Table 2), 28-46 (usually 32-42) pored scales, 3-21 (usually 8-18) unpored scales, 44-60 (usually 47- 52) total scales; pectoral fin rays modally 13; coronal pore single; infraorbital canal complete, with 8 pores; preoperculoman- dibular canal with 10 pores; supratemporal canal usually complete in adults; snout blunt, slightly decurved; urogenital papilla mod- erately long in spawning females. Description. Etheostoma collettes is a large, robust species of Oligocephalus (Fig. 1). The largest specimen examined is a male 61.5 mm SL. Table 1 gives the proportional measurements for the holotype and nine paratopotypes. A moderately developed nuchal hump is present in both sexes, slightly more pro- nounced in males larger than 40 mm SL; the body is deepest at origin of spinous dorsal; the rostral frenum is well developed. The body, including nape and belly, is completely covered with scales in most speci- mens; however, in some (including holo- type) the scales of anterior 4 to % of belly 108 Tulane Studies in Zoology and Botany Vol. 15 10mm are partially to entirely embedded. The nape is nearly naked to fully covered with em- bedded scales; breast and prepectoral areas are naked (latter with a few em- bedded scales in large individuals); the opercle is fully invested with large, wholly to partially exposed scales; approximately 50 per cent of the cheek below and behind eye with small, partially exposed scales. Scales below lateral line 7-12 (usually 8-10); transverse scale rows 11-16 (usually 13-15); scales around caudal peduncle 18- 24 (usually 19-22). Vertebrae in 27 individuals from type- series: 36 (10 including the holotype), 37(15), 38(2). Palatine and vomerine teeth present. Dorsal spines number 8-12 (usually 10- 11); dorsal soft rays 11-14 (usually 12- 13); anal fin with two spines and 6-8 (usually 7) soft rays. Pectoral rays (left side only) 11-14 (usually 12-14). The shape of the spinous dorsal is variable but it is usually low and little rounded; the soft dorsal is somewhat larger than the anal fin, especially in large specimens; the pos- terior edge of the caudal fin is truncate. Coloration of males. Color descriptions are based on observations made by Knapp and on two Kodachrome transparencies pro- vided by Neil H. Douglas of a specimen approximately 55 mm SL, collected 29 June 1966, from Flem Branch (Little River drain- age), Grant Parish, La. and a specimen about 65 mm SL taken 15 October 1966, from Choudrant Cr. (Ouachita River drain- age), Ouachita Parish, La. The 8-9 bluish-brown blotches on the dorsum are a distinctive feature of the color pattern. The second, fifth, seventh, and eighth are often more pronounced than the others. The location and relative prominence of the blotches are as follows: 1—nape just behind the head, often faint or absent; 2— Nape just anterior to the spinous dorsal fin, pronounced; 3 and 4—below the spinous dorsal, faint; S—below the posterior end of the spinous dorsal, pronounced; 6—below < Figure 1. Etheostoma collettei, new species. Drawn from the holotype, USNM 201515, a male, 56.3 mm in standard length, collected in the Dugdemona River, Little River drainage, Jackson Parish, Louisiana. No. 3 Etheostoma colletter 109 TABLE | Proportional measurements of Etheostoma collettei, expressed in hundredths of standard length. USNM 201515 Character Holotype ( 2 ) Standard length (mm) 56.3 Predorsal length 35 Body depth 23 Caudal peduncle length 19 Caudal peduncle depth 10 Head length 30 Head depth at orbit 12 Orbit width (vertical ) 6 Orbit width (horizontal ) 7 Snout length 6 Upper jaw length 8 Width of gape ) Spinous dorsal base 30 Soft dorsal base 20 Anal base 14 Longest dorsal spine 12 Longest dorsal soft ray 16 Longest anal ray 15 First anal spine 9 Caudal fin length 18 Pectoral fin length 24 Pelvic fin length 20 Interpelvic distance 2 Transpelvic distance 8 USNM 201516 Paratypes 4¢4¢4 5229 Range Mean! Range Mean 46.8-53.1 ayllalt 41.5-58.1 49,1 35-36 35.4 34-36 Sow 18-20 20.0 18-21 19.4 17-19 18.4 16-18 17.4 9-10 9.4 9-10 9.2 29-30 29.8 28-30 29.2 12-13 | PA) WW=13 12.4 6 6.0 5-6 HIG 6-7 6.8 al 7.0 6-7 6.4 6=7 6.4 8-9 8.2 8-9 8.4 7-9 8.6 7-8 7.8 (sil 29.4 28-31 28.8 19-21 19.6 18-20 19.4 12-14 13.0 1O=13 11.6 11-13 12.0 IE} 11.6 13-16 14.8 13-15 14.4 14-16 15.0 14-15 14.8 9-12 9.6 7-9 8.0 18-20 18.6 18-20 18.5 22-94 23.0 22-—2,4 23.4 17-20 18.8 18-20 18.5 2 Z20 2 40) } 8.0 7-9 8.0 ' Holotype included in mean for males. the origin of the soft dorsal, faint; 7—below the mid soft dorsal, pronounced in some specimens but faint in others; 8—just be- hind the soft dorsal, pronounced; 9—mid- way the dorsal surface of the caudal pe- duncle, faint and sometimes absent. Most specimens have 5-8 vague, vertical bars on the posterior half of the body. These bars have a bluish cast in life but appear brown in preserved specimens. The area between the bars, the ventral surface of the caudal peduncle, and the belly are bright orange. The belly of most preserved male specimens is dusky. A poorly-defined, brown, basicaudal blotch is discontinuous with a small distinctive spot lying just below the dorsal procurrent rays of the caudal fin. A similar spot is present above the ventral procurrent rays of the caudal fin in small specimens but appar- ently is obscured in larger specimens by be- ing joined to the larger caudal blotch. Pos- terior to the caudal blotch at the base of the caudal rays, two roundish, orange spots lie just above and below the center of the caudal base. The axil of the pectoral fin is densely pig- mented with melanophores but character- istically has a bright orange bar. Posterior to the pectoral axis, a darkened bar of small melanophores begins immediately below the lateral line and extends ventrad to just be- hind the pelvic base. This bar is variable, but it is evident in most specimens. The breast of adult males is often covered with micromelanophores; however, some _ large adult males have few melanophores on the breast and in several specimens the breast is immaculate. The Choudrant Creek speci- men in the Kodachrome transparency has a tinge of orange on the breast. The prepectoral area, tinged with orange in life, often has a dark blotch. Pigmenta- tion of the urogenital papilla varies, but usually a few lateral melanophores are present. On the head there is a dark, well-defined subocular bar and a short, broad, postocular 110 Tulane Studies in Zoology and Botany Vols TABLE 2 Lateral-line scale counts in Etheostoma collettei and E. asprigene Total Lateral Line Scales Species and Drainage 44 45 46 47 48 49 50 51 52 53 54 55 56 57 58-60 N x E. collettei Upper Ouachita R. 3 2% GIS Sie 7 & a wt wl 106 oles Lower Ouachita R. L i @ IQ 4) gil 4 8 = & re, 49.0 Little R. lL © Ff Oilge es © 6 ah 4! 62 50.5 Red R. BB RR ses @ 8 B I = Il 20 olkG Sabine R. I. AL i & 8 50.5 E. asprigene Ouachita R. % HW MB Fig SS CG Ik Beal 44 49.2 Lateral-line scale counts in Etheostoma collettei and E. asprigene Pored Lateral Line Scales Species and Drainage 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46 N Xx E. collettei Wpper OucachitayRy onl lel OMlOMI2T0520 487 8 ba NOG BiB? Lower Ouachita R. Dy 8} i By we) Mey eG Ea! 2 ooul Little R. LW A eG Os) UO Gee IP Bh il 62 38.5 Red R. lia = 2) 32 16) 22 SS Se 20 = 39.0 Sabine R. 1 - So] = 84 =o = = =F il 8 36.6 E. asprigene Ouachita R. I Se DI ef SG Sah 2) 2) Si) ish) Lateral-line scale counts in Etheostoma collettei and E. asprigene ; Unpored Lateral Line Scales Species and Drainage mo teh TIO TEL US ee TUS TUG} Tlie TS}. IS) XO) AL N Xx E. collettei Upper Ouachita R.! 3) FL ss Psy ks} zt) GS) sl 103 16.0 Kower@uachita Ral, — 92) 5, 710 412910 6G 4 Ty 72 10.9 Little R. Be 8 IO il ako) & dil By @ G2MES Red R. De Aone AS 20° 12:6 Sabine R. eae —w i = 2 ee all 8 13.9 E. asprigene Ouachita R. I 4 8-6 1g) & 4 Sf Ss il = il 39 11.2 ‘Includes one specimen with 31 unpored scales. * Includes count for holotype. blotch which does not extend to the pre- opercular margin. A diffuse, dark stripe from the anterior edge of the orbit to the upper lip gives the effect, when viewed from above, of a pale stripe through the frenum. A small dark spot on the dorsal part of the orbit extends onto the eye. A thin line of con- centrated melanophores is present along the mandibular and maxillary grooves. The opercle is darkly pigmented as is the pos- terior edge of the preopercle. The branchios- tegal membranes are also heavily pigmented and tinged with red-orange where they over- lie the prepectoral area. The humeral spot is dark and well defined but is not enlarged. The spinous dorsal fin membrane has a narrow, dark, marginal band which appears to have a greenish-blue cast. Beneath the distal band is a broad, vivid, red-orange band followed by a narrower dark band. Along the base of the spinous dorsal fin is a very narrow, dusky-orange band. The pig- No. 34 Etheostoma collettei 111 TABLE 3 Comparison of Etheostoma collettei and E. asprigene from the Ouachita River drainage of Louisiana. Character E. collettei E. asprigene 8-9, 4 pronounced Ovoid Slightly decurved, blunt 5-7, indistinct above L. L. Well-developed Blotches on dorsum Eye shape Snout shape Body bars Postpectoral bar Spinous dorsal dial dark band Humeral spot Pronounced Postocular stripe Prepectoral area Pectoral rays Naked 11-14, modally 13 Wide orange-red band, narrow me- Short, broad; not reaching operculum 8-9, all equally developed Round Produced, acute 6—8, distinct above L. L. Absent or faint Narrow orange-red band; wide me- dial dark band Absent or faint Long, narrow; reaching operculum Partially exposed ctenoid scales 12-15, modally 14 mentation of all three bands is restricted to the membranous portion of the fin and does not go across the rays. The soft dorsal membrane has a dark, distal band similar to that of the spinous dorsal. Beneath the distal band is a broad, red-orange band which is dissected by the dusky rays. The base of the soft dorsal is marked by one, sometimes two, poorly- defined dark bands. The anal fin is greenish- blue along its base. Preserved specimens usually show an even gradation from a dark base to a light distal margin. In large pre- served specimens the entire anal fin is often dark. The caudal fin membrane has 5-7 poorly defined dusky bars and, in larger specimens, a dusky distal margin. The inter- radial membranes of the pelvic fins in large specimens are densely covered with micro- melanophores. The anterior interradial mem- brane is tinged with orange and the proximal portion of the other membranes is bluish- green. The pectoral fin displays several poorly-defined bars which are often obscure or absent. In life the proximal half of the pectoral fin is tinged with orange. Coloration of females. The following ac- count is based on preserved specimens. In general, the females are similar to the males, but, they differ in several subtle pigmentary features. The body above the lateral line is usually mottled. The vertical bars are less well defined than in the male. There are few, if any, micromelanophores on the ven- ter, making the pattern more highly con- trasted than in the male. The urogenital papilla is immaculate. The mandibular groove is more heavily pigmented and the branchiostegal membranes somewhat _ less pigmented in the female than in the male. The membranes of the soft dorsal are marked by 5 or 6 narrow dark bands. The anal and pelvic fins have little or no dark pigment. Habitat. Neil H. Douglas (pers. comm.) has kindly provided the following notes on the habitat at the type-locality in the Dug- demona River in Jackson Parish, La. “This area is near the headwaters of the river, where the depth is normally two inches to three feet, with a width of 10-20 feet. The shore line is heavily vegetated. The stream flows with moderate to fast current at this point, and many logs and cypress stumps create riffle areas. The bottom is of hard soil with no rocks or gravel.” Douglas fur- ther stated that the habitat at the type- locality is common in north central Louisi- ana, and that he has collected E. collettei throughout this area in streams with mod- erately strong current. In the Saline River of the upper Ouachita drainage, one of us (Knapp) has taken E. colletter in several habitats. Near Crows, Arkansas, it was taken from heavy growths of Podostemum ceratophyllum in rocky chutes along with E. blennioides Rafinesque and E. zonale (Cope). In other areas of the Saline River, it was taken from clear gravel riffles with E. whippli (Girard). Relationships. In some features Etheo- stoma collettei appears very similar to E. asprigene; however, a number of obvious differences exist (Table 3). Breeding tu- bercles, similar in type and arrangement to those described by Collette (1966:597) for 12 E. caeruleum Storer, have been found in a few males. The rather elongate urogenital papilla of the ripe female is quite unlike that of E. asprigene and may be an adapta- tion for deposition of eggs on aquatic vege- tation. In view of these dissimilarities, we do not feel that E. collette: can be placed in the E. asprigene species group as visualized by Ramsey and Suttkus (1965:74), but believe that it is more closely related to the asprigene group than to other groups in Oligocephalus. Tulane Studies in Zoology and Botany Vole LITERATURE CITED CoLLeTTE, BrucE B. 1966. Systematic signifi- cance of breeding tubercles in fishes of the family Percidae. Proc. U. S. Nat. Mus. 117 (3518) :567-614, 7 figs. Husss, Cart L. and Karu F. Lacier. 1958. Fishes of the Great Lakes Region. Cranbrook Inst. Sci. Bull. 26:213 pp., 44 pls., 251 figs. RaAMsEy, JOHN S. and Roya D. Surrxkus. 1965. Etheostoma ditrema, a new darter of the sub- genus Oligocephalus (Percidae) from springs of the Alabama River basin in Alabama and Georgia. Tulane Stud. Zool. 12(3):65-77, 3 figs. March 24, 1969 MYSIDOPSIS BAHIA, A NEW SPECIES OF MYSID (CRUSTACEA:MYSIDACEA ) FROM GALVESTON BAY, TEXAS! JOANE MOLENOCK2 Marine Laboratory Texas A&M University Galveston, Texas ABSTRACT A new species of mysid, Mysidopsis bahia, is described from West Bay, Galves- ton, Texas. This mysid was found in three areas within the Galveston Bay system. The salinity at the collection sites varied from 18%. to 29%, and the temperature, from 24°C to 32°C. M. bahia differs from M. almyra and M. bigelowi, which also have been reported from the Gulf of Mexico, by the armature of the telson, the number of spines on the uropod together with an un- segmented antennal scale. Specimens of a new species of mysid were collected during the week of November 22, 1965, near the northwest shore of Galveston Island in West Bay. A hand screen was used to collect them in water one-half meter deep, close to shore. The salinity was 28% and the temperature was 24°C. The follow- ing description is based on the laboratory- reared progeny of these specimens. Mysidopsis bahia,*? new species Figures 1-18 Description. Length of adults from an- terior margin of carapace to end of uropods, excluding setae, varies from 4.4 mm to 9.8 mm. Anterior margin of carapace roundly triangular, not produced into rostrum; pos- terior margin concave; somite 8 exposed in dorsal view. Eye rounded and slightly kid- ney shaped. Telson linguiform with bluntly rounded apex; lateral margins usually with 19-22 short spines on each side; apex with 4-5 pairs of closely set strong spines in adults; central pair longest, about “% as long as telson. Third segment of antenna 1 of male with lobe bearing thick fine setae. Scale of antenna 2 narrowly lanceolate com- posed of single segment which bears setae along entire length on each side. Labrum rounded anteriorly; posterior margin with small central indentation, middle portion with short setae. Molar of mandible reduced or absent; incisor of left mandible with 7 teeth, right with 6 teeth; left lacinia mo- bilis broad with 6 teeth; right lacinia smaller, constricted at base, with 7 teeth; medial row of closely set stout spines, 10-12; on left mandible this row ends with tuft of setae. Palp well developed. Outer plate of maxilla 1 with 8 spines at apex; inner plate with 2 setae at apex and | on outer margin. Proxi- mal lobe of maxilla 2 with 4 setae on trun- cate apex; exopod usually bears 6-10 setae. Thoracic leg 1 (maxilliped) short, stout. Endopod of leg 2 slender, ending in spine. Endopod of male pleopod 4 with single lobe bearing 4 stout spines, 1 short spine, exopod longer than endopod with long apical spine. Exopod of uropod about twice as long as telson, curved gently outward; endopod about *4 as long as exopod, armed on ventral sur- face with 2 to 3 spines near medial margin distal to statocyst. Color, Living specimens usually transpar- ent. However, color may be distributed over 1 Based on a thesis submitted in partial fulfillment of the requirements for an M.S. degree, Texas A&M University, 1966. 2 Present address, Scripps Institution of Oceanography, La Jolla, California. * From the Spanish bahia, bay. EDITORIAL COMMITTEE FOR THIS PAPER: Dr. WILLIAM D. CLARKE, Senior Scientist, Westinghouse Electric Corporation, San Diego, California Dr. ALFRED E. SMALLEY, Associate Professor of Biology, Tulane University, New Orleans, Louisiana 113 114 Tulane Studies in Zoology and Botany Vol. 15 Se 100 p» Figures 1-8. Mysidopsis bahia, new species. 1. Left mandible, gnathobasic process, oblique internal view. 2. Left mandible, gnathobasic process, internal view. 3. Right mandible, gnatho- basic process, oblique internal view. 4. Right mandible, gnathobasic process, internal view. 5. Left mandibular palp. 6. Maxilla 1. 7. Labrum. 8. Maxilla 2. > Figures 9-18. Mysidopsis bahia, new species. 9. Genital appendage, male. 10. Thoracic leg 8. 11. Antenna 1, ventral, male. 12. Antenna 2. 13. Thoracic leg 2. 14. Thoracic leg 1. 15. Pleopod 4, male. 16. Telson. 17. Right uropod, dorsal. 18. Male, dorsal. Mysidopsis bahia 115 No. 116 entire surface of animal in yellows, browns or blacks. Dorsally, preserved specimens have 2 contracted chromatophores at base of tel- son; ventrally, 1 pair at base of mouthparts, 2 pairs on thorax, 5 median chromatophores on abdomen. Types. The male holotype, USNM 113832, is a specimen reared in the laboratory from those originally collected in November 1965. Fourteen similarly reared specimens are designated as paratypes, USNM 113834. Twenty-two field specimens are also desig- nated as paratypes, USNM 113833. Occurrence. This mysid has been found at several locations within the Galveston Bay system. My original collection site was West Bay. Ruppia maritima occurred in patches on a sand bottom. Using a hand screen I collected this mysid and postlarval Penaeus, Tozeuma, Hippolyte and Palaemone- tes. Field paratypes were collected by Mr. Cor- nelius R. Mock in Lake Como, an inlet of West Bay. The specimens that I examined from this site were collected from July through September 1966. A net tow of 15 m along the bottom yielded hundreds of specimens of M. bahia. The bottom in Lake Como was sand, the salinity during this pe- riod varied between 22% and 29%, and the temperature ranged from 24°C to 32°C. Ad- ditional specimens given to me by Dr. David V. Aldrich were collected in September 1964 from Clear Lake, which is a secondary bay opening into Galveston Bay. The salinity in Clear Lake was 18% and the temperature, 29°C at the time of collection. Mock (1966) reported this site (A) with a bottom consisting of a uniform mixture of sand, silt and clay. Remarks, Two species of Mysidopsis have been reported from the Gulf of Mexico (Bowman, 1957, 1964; Clarke, 1956; Tat- Tulane Studies in Zoology and Botany Vol sis tersall, 1926, 1951), M. almyra Bowman and M. bigelowi Tattersall. An unsegmented antennal scale serves to distinguish M. bahia and M. bigelowi from M. almyra which has a segmented antennal scale. M. bahia and M. almyra have a slender second leg while in M. bigelowz this leg is robust. The number of strong spines on the apex of the telson also differentiates these species; M. bigelowi has 3 pairs, M. bahia, 4-5 pairs, and M. almyra, 6-7 pairs. An additional dis- tinguishing character is the number of spines on the endopod of the uropod; M. almyra has | spine, M. bahia, 2-3 spines and M. bigelowi, 5 spines. Acknowledgments I appreciate the advice and comments of Dr. Thomas E. Bowman upon my intro- duction to Mysidacea. Dr. William D. Clarke kindly discussed mysids with me and en- couraged my interest in them. LITERATURE CITED BowMAN, THoMas E. 1957. A new species of Mysidopsis (Crustacea: Mysidacea) from the southeastern coast of the United States. Proc. U.S. Nat. Mus., 107:1-7. 222 (aS ee 1964. Mysidopsis almyra, a new estuarine mysid crustacean from Louisi- ana and Florida. Tulane Stud. Zool., 12: 15-18. CLARKE, WILLIAM D. 1956. A further descrip- tion of Promysis atlantica Tattersall (Crusta- cea, Mysidacea). American Mus. Novitates 1755:1-5. Mock, Corne.tius R. 1966. Natural and altered estuarine habitats of penaeid shrimp. Proc. Gulf Caribb. Fish. Inst., 19:86-98. TATTERSALL, WALTER M. 1926. Crustaceans of the orders Euphausiacea and Mysidacea from the western Atlantic. Proc. U. S. Nat. Mus., 69: 10-12. —- __.............. 1951. A review of the My- sidacea of the United States National Mu- seum. U. S. Nat. Mus. Bull. 201:1-292. March 24, 1969 TULANE STUIDINES IN AZAOOLOGY AND BOTANY 5. VA-N Volume, 15, Number 4 June 26, 1969 MUS. COMP. zooL: LIBRARY, JUL 10 1969 HARVARD UNIVERSITY, DIGENETIC TREMATODES OF MARINE TELEOST FISHES FROM BISCAYNE BAY, FLORIDA ROBIN M. OVERSTREET Institute of Marine Sciences, University of Miami, Miami, Florida ip: VES oS ~ wl 2) in i, & \ ES TULANE UNIVERSITY NEW ORLEANS TULANE STUDIES IN ZOOLOGY AND BOTANY is devoted primarily to the biology of the waters and adjacent land areas of the Gulf of Mexico and the Caribbean Sea, but manuscripts on organisms outside this geographic area will be considered. Each number is issued separately and contains an individual monographic study, or several minor studies. As volumes are completed, title pages and tables of contents are distributed to institutions receiving the entire series. Manuscripts submitted for publication are evaluated by the editor or associate editor and by an editorial committee selected for each paper. Contributors need not be members of the Tulane University faculty. 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OVERSTREET2 Institute of Marine Sciences, University of Miami, Miami, Florida CONTENTS AA SSTIRYN GIP ca a eee, ng ee 2 te el Oe rc ee ee ee ed he ee 120 JAN CTS OM ETSI G] EINE RSs ae ol Se i a eels fe Ree See 120 NERDS OGIO Ngee ath eee ON es An A OLE ETE ge TO. RIL rays Nee ME 8 8 120 RIE OD Sweet ae. aN ee eel Ape. ON es Nee we PL Wb Oa 121 IDESCRIPAION CAND s OISCUSSION@ OF SPEGIES) 22-2222 t= ee 121 Sais rcOlicac wees ween samen sets, 2 Le NS i oe ee eee ee ee Al Brice pindlidacpameei Flees sense Re rRee eS es ae lat Relicdiscomatidace:=.2 mee cee =o a Pee eT ees ee eee 123 Param piiscomaticac poser seein ees etal: OG NTIE Se ey ee ee eee 128 Pronocephalidgeg=-aa20 Newel. f0eui Se are) ee eee 128 IM GrOScap niciicacye «seer tre Sy MD AO et ne 128 Pa ploponicac uae ive siti ser wew cB ews lew! reine JWEi yp SbeT 8 Si eevend 130 liaplosplanchnidacy tS Ware leediRA. . WeahE 2 tedden Bee 131 ie pocteacii acinar. ee TNA 8 ee eee 139 © isin eee cae a meme ere en Ee a a 143 OFSS SOS UG AYE ee rae ie ert en Cy REL elt okt wees oe CS pe 143 Gorpadenidacyi-2 = Asarh es: Wer ta ty 2) 4s Monto: i int et tee eee Sheol tne 149 EO SOU ACH ens ERE ee S02 me 2-4 et INS LOM, SNe ES el) LS ES 150 ROTOLCHIIG ACH: aa eee meee Bere) 2) PA ee OU et Ee © ee 152 Crp rOcominnidat eae waewest cake eee © eee Ee SANE Ee es 2 eee ee 158 PACAMENOCO PIC Ae ea wae tnwes Sul dea Se ee ee 159 IS FeoavitiercaVe Ae) Spee Ole UL eo eee ee Pee eRe elke ore eee 162 SGlerodistOmaticacte ee secs mew lene rN te ee A Ce a ee tes 166 | Se Kyraevo7 devi bzy= th Aus a a erat NP Breese a” Seger tere feet, Mer ese eee 167 PDESEW SSO Niger ee ee Se se ee Ye ed ee ee 167 EGS AIUASKEEIO Sire see oa ne OT) a ee eee 168 LEER AAR Pan D be to ake teal ret ete Se bis) Las te es a eee 172 BMIDEXO OF PARASITIC GENERA] 0 176 ! Contribution No. 1060 from the Institute of | ments for the degree of Doctor of Philosophy. Marine Sciences, University of Miami. This 2 Present address: Department of Parasitol- paper is part of a dissertation which was sub- ogy, Tulane University, School of Public Health mitted to the Graduate School of the University | and Tropical Medicine, New Orleans, Louisiana of Miami in partial fulfillment of the require- 70112. EDITORIAL COMMITTEE FOR THIS PAPER: Dr. HAROLD W. MANTER, Emeritus Professor of Zoology, University of Nebraska, Lincoln Dr. R. M. CABLE, Professor of Biology, Purdue University, Lafayette, Indiana 119 ABSTRACT This study includes 111 species of Di- genea from 69 of 113 species of teleost fishes examined from Biscayne Bay, Flor- ida. Biscayne Bay represents the reported northern limit along the Atlantic coast for all but 22 of the 111 species. It represents the southern limit for only one, Didymo- cystis scomberomori. From the known col- lections reported in the literature, the Digenea from Biscayne Bay appear to con- sist primarily of a tropical fauna. Species previously known from the Pacific Ocean only are Botulisaccus pisceus and Pseu- docreadium scaphosomum. There are 81 new host records. Two new genera are created: Claribulla (Fellodistomatidae) and Apertile (Opecoel- idae). Thirteen new species are described: Apocreadium cryptum, Cadenatella flori- dae, Claribulla longula, Diphtherostomum albulae, Hamacreadium confusum, Hurley- trema_ pyriforme, Hymenocotta manteri, Lasiotocus albulae, Lasiotocus haemuli, Lasiotocus mugilis, Neolepidapedon mac- rum, Nicolla halichoeri, and Opegaster pritchardae. Eleven new combinations are made: Apertile holocentri, formerly Neopecoelus holocentri Manter, 1947; Apocreadium foli- atum, formerly Homalometron foliatum Siddiqi and Cable, 1960; Bianium longi- pygum, formerly Diploproctodaeum longi- pygum Oshmarin, Mamaev, and Parukhin, 1961: Bianium macracetabulum, formerly Diploproctodaeum macracetabulum Osh- marin, Mamaev, and Parukhin, 1961; Diplomonorchis magnacetabulum, formerly Diplomonorcheides magnacetabulum Thomas, 1959; Helicometrina execta, for- merly Helicometra execta Linton, 1910; Hurleytrema malabonensis, formerly Pseu- dohurleytrema malabonensis (Velasquez, 1961); Hurleytrema shorti, formerly Pseu- dohurleytrema shorti Nahhas and Powell, 1965; Megasolena hysterospina, formerly Lepidauchen hysterospina Manter, 1931; Myosaccium opisthonemae, formerly Neo- genolinea opisthonemae Siddiqi and Cable, 1960; and Pseudopecoelus scorpaenae, for- merly Neopecoelus scorpaenae Manter, 1947. Five genera have been synonymized: Bilecithaster Siddiqi and Cable, 1960, synonym of Diplangus Linton, 1910; Diplo- monorcheides Thomas, 1959, synonym of Diplomonorchis Hopkins, 1941; Neogeno- linea Siddiqi and Cable, 1960, synonym of Myosaccium Montgomery, 1957; Neo- pecoelus Manter, 1947, synonym of Pseu- dopecoelus Von Wicklen, 1946; and Parahurleytrema Nahhas and Powell, 1965, synonym of Hurleytrema Srivastava, 1938. Twelve species have been synonymized: Antorchis holacanthi Siddiqi and Cable, 1960, synonym of Antorchis urna (Linton, Tulane Studies in Zoology and Botany Vol. 15 1910) Linton, 1911; Bilecithaster ovalis Siddiqi and Cable, 1960, synonym of Diplangus parvus Manter, 1947; Diplomo- norchis micropogoni Nahhas and Cable, 1964, synonym of Diplomonorchis leios- tomi Hopkins, 1941; Helicometra pretiosa Bravo-Hollis and Manter, 1957, a synonym of Helicometra torta Linton, 1910; Heli- cometrina parva Manter, 1933, and Heli- cometrina_ trachinoti Siddiqi and Cable, 1960, synonyms of Helicometrina execta (Linton, 1910); Hexangitrema breviceca Siddiqi and Cable, 1960, synonym of Hexangitrema pomacanthi Price, 1937; Manteria costalimai Freitas and Kohn, 1964, synonym of Manteria brachydera (Manter, 1940); Megasolena archosargi Sogandares-Bernal and Hutton, 1959, syn- onym of Megasolena hysterospina ( Manter, 1931); Opisthadena cortesi Bravo-Hollis, 1966, synonym of Opisthadena dimidia Linton, 1910; Paracryptogonimus neo- americanus Siddiqi and Cable, 1960, syn- onym of Paracryptogonimus americanus Manter, 1940; and Pseudohurleytrema ottoi Travassos, Freitas, and Buihrnheim, 1965, synonym of Hurleytrema _ shorti (Nahhas and Powell, 1965). ACKNOWLEDGMENTS I am especially indebted to Dr. Harold W. Manter and Mrs. Mary Hanson Pritchard of the University of Nebraska for verifying my identifications, answering many questions, and permitting me to use their library and collec- tions of trematodes for four days. I wish to thank Drs. Edwin S. Iversen, Frederick M. Bayer, W. Henry Leigh, C. Richard Robins, and Carl J. Sindermann, all of the University of Miami, for their suggestions and criticisms of the manuscript. Space and equipment were provided by Dr. Leonard J. Greenfield. I also wish to thank those who helped collect fishes for this study, especially Dr. Ronald Smith, Mr. Thomas Fraser, and Mr. Lucky McLeod. Iden- tifications or verifications of my identifications of the hosts were willingly made or obtained by Dr. C. Richard Robins, Mr. Thomas Fraser, Dr. Alan R. Emery, and Dr. Martin Roessler. I am grateful to those persons who lent or gave me specimens of Digenea; those people are acknowledged in the text under the appro- priate sections. INTRODUCTION The present study was undertaken to make the digenetic trematode fauna of fishes from the east coast of Florida better known. Emphasis has been placed on several of the numerous fishes that spend either all or a portion of their lives in Biscayne Bay. This report concerns 111 species of adult Digenea collected from 69 of 113 species No. 4 of teleosts represented by 333 individuals. A few of these trematodes have been re- ported from the Atlantic coast of Florida by Ward (1954), Schroeder (in press), Anderson (1965), Daigger and Lewis (1967), and Overstreet (1968). Descrip- tions of most of the previously named species are available in the literature based on collections from Beaufort, North Caro- lina (Linton, 1905; Manter, 1931; Pearse, 1949); Dry Tortugas, Florida (Linton, 1910; Manter, 1933a, 1933b, 1934, 1947); Woods Hole Region, Massachusetts (Lin- ton, 1900, 1901, 1940); Bermuda (Linton, 1907; Hanson, 1950); Bahama _ Islands (Sparks, 1957; Sogandares-Bernal, 1959); Curagao and Jamaica (Nahhas and Cable, 1964); Puerto Rico (Price, 1934; Siddiqi and Cable, 1960); Cuba (Pérez Vigueras, 1955a, 1955b, 1955c, 1957, 1958); and dif- ferent areas of the Gulf of Mexico, including the west coast of Florida (Sparks, 1958; Sogandares-Bernal and Hutton, 1959a, 1959b, 1959c; Nahhas and Short, 1965; Nahhas and Powell, 1965). Other known species encountered in this study have been reported from other localities and are dis- cussed in the text when relevant. METHODS Fishes for study were collected between January 1966 and May 1968 by hook and line, seine, trap, trawl, or spear and kept alive until they could be examined. Almost all the fishes were examined within two days after capture. Examination included the entire alimentary system and coelomic cavity of all fishes, and the gills, heart, swim blad- der, urinary bladder, and other tissues of many. Most trematodes were washed in 0.8 per cent saline; fixed in hot alcohol-formalin- acetic acid (AFA), using light coverslip pressure when necessary to prevent curling; stained with Van Cleave’s hematoxylin; and mounted in Permount. Lithium carbonate and butyl amine were added during dehy- dration in 80 per cent alcohol to prevent future fading. Sectioned material was stained with Mallory’s trichrome stain or Harris's hematoxylin with eosin as a counterstain. Measurements and computed ratios were taken using microns as the unit, but the values were rounded off for the text. All Digenetic Trematodes of Marine Teleosts 121 measurements are in millimeters unless otherwise indicated. Measurements were taken on fully-formed eggs unless the text indicates otherwise. The diameter of the oral sucker was compared with that of the acetabulum, with the oral sucker represent- ing one, in order to compute the sucker ratio. Asterisks indicate new host records. The common names, authors, and families of the fishes examined are included in the Host-Parasite List following the discussion. A camera lucida was used for all illustra- tions. Holotypes have been deposited in the Helminthological Collection of the United States National Museum, and the hosts in the Ichthyological Collection of the Insti- tute of Marine Sciences, University of Mi- ami, Miami, Florida. DESCRIPTION AND DISCUSSION OF SPECIES Except for new species, the discussion is limited to aspects that supplement the avail- able literature. A partial synonymy is pre- sented. FAMILY SANGUINICOLIDAE Graff, 1907 Deontacylix ovalis Linton, 1910 Host: Kyphosus sectatrix (6 of 6). Site: Body cavity. Discussion: Land (1967) considered the family Aporocotylidae Odhner, 1912, in which Deontacylix ovalis had previously been placed, as a synonym of Sanguini- colidae. My specimens agree with the orig- inal description by Linton (1910:83-84) and extended by Manter (1947:367-368). The testis is usually not as distinct as de- scribed by Manter, but is more reticulate with short lateral bulges extending from the longitudinal extracaecal extensions. Adjacent lateral bulges often overlap or touch each other. The lobed ovary occasionally has nu- merous perforations. The excretory system in- cludes a very short, terminal, sac-like vesicle with branched collecting ducts that extend anteriorly, accompanying the lateral nerves. The excretory pore is terminal. FAMILY BUCEPHALIDAE Poche, 1907 Bucephalus scorpaenae Manter, 1940 Figure 1 Figure 1. Bucephalus scorpaenae, ventral view. Hosts: Scorpaena grandicornis (2 of 2)*; Scorpaena plumieri (2 of 4). Site: Pyloric caeca and intestine. Discussion: Figure 1 shows at the base of rhynchus previously undescribed gland cells with large ducts which join the bases of the tentacles. Manter (1940c:5) and Win- ter (1960:183) described the dorsal wall of the anterior sucker as being vesicular and probably glandular. Other species of bu- cephalids, some without papillae or tentacles, have been reported to possess anterior gland cells. Hopkins (1954:356, figs. 1-2; 358, 6-7) described and illustrated such cells in Rhipidocotyle transversale Chandler, 1935, and Bucephaloides strongylurae Hopkins, 1954, as extending over and around the rhynchus, and subsequently opening an- teriorly. Bucephalus varicus Manter, 1940 Bucephalus polymorphus Baer, 1827, of Nagaty, 1937, and others. Bucephalus pseudovaricus Velasquez, 1959. Hosts: Caranx crysos (1 of 2); Caranx hip- pos (3 of 3). Site: Pyloric caeca, few in stomach. Discussion: Manter (1963c:228) dis- cussed the synonymy and identification of this variable species. The present specimens show further variability in that the testes are usually separated by the proximal por- tion of the uterus. Tulane Studies in Zoology and Botany Vol. 15 Prosorhynchus pacificus Manter, 1940 Gasterostomum sp. Linton, 1910 (from Mycteroperca bonaci and M. venenosa). Prosorhynchus atlanticus Manter, 1940. Hosts: Mycteroperca bonact (3 of 3); Myc- teroperca microlepis (2 of 2). Site: Intestine and pyloric caeca. Discussion: Hanson (1950:75) consid- ered Prosorhynchus atlanticus, apparently from a misidentified host, as a synonym of P. pactficus on the basis of the variability in the specimens she examined from Bermuda. She noted a wide range in the size of eggs. Nahhas and Cable (1964:174) reported nonintergrading egg lengths in specimens from Curacao, Jamaica, and Puerto Rico, as I find in my specimens, and they did not accept the synonymy. Winter (1960:187- 189), however, reported specimens from Mazatlan, Sinaloa, Mexico, with eggs 29 to 33 by 19 to 20 microns, about the same measurements as found in most Atlantic specimens. I am therefore accepting the synonymy. Rhipidocotyle adbaculum Manter, 1940 Hosts: Scomberomorus maculatus (1 of 2); Scomberomorus regalis (2 of 2). Site: Pyloric caeca and intestine. Discussion: My specimens differ slightly from the original description. The eggs measure 17 to 23 by 12 to 15 microns rather than 15 to 17 by 9 to 10, and the ovary does not always overlap the anterior testis. Bucephaloides bennetti Hopkins and Sparks, 1958 Bucephalopsis bennetti (nomen nudum). Melugin, 1940 Host: Paralichthys albigutta (1 of 1). Site: Pyloric caeca. Discussion: Sogandares-Bernal and Hut- ton (1959a:260) gave a review of B. ben- nettt, Bucephaloides arcuatus (Linton, 1900) Velasquez, 1959 Gasterostomum arcuatum Linton, 1900. Bucephalopis arcuatus (Linton, 1900) Eckmann, 1932. No. 4 Host: Scomberomorus regalis (2 of 2). Site: Pyloric caeca. Discussion: My specimens agree fairly well with the original description by Lin- ton. They have an excretory vesicle extend- ing to or near the rhynchus, and thus differ from specimens previously reported as Bz- cephaloides arcuatus from barracuda, as dis- cussed by Manter (1963c: 229-230) and confirmed by Nahhas and Cable (1964: 173). FAMILY FELLODISTOMATIDAE Nicoll, 1913 Tergestia pectinata (Linton, 1905) Manter, 1940 Distomum pectinatum Linton, 1905. Theledra pectinata (Linton, 1905) Lin- ton, 1910. Host: Caranx crysos (1 of 2). Site: Rectum. Discussion: Considerable confusion exists about the identification of some species of Tergestia. Manter (1947:323) believed that what Linton reported as T. pectinata from Auxis thazard and A. rochei is actually T. laticollis, and that T. laticollis of Yamaguti, 1951, is probably T. pectinata (see: Manter, 1963b:447). Two specimens 1.8 long from Caranx crysos have sucker ratios of 1:1.5 and 1:1.6, which would place them as T. laticollis in Manter’s key (1954:527). The specimens have a straight seminal vesicle, a character which Manter (1947:323) also used to differentiate T. Jaticollis from T. pectinata. However, Sogandares-Bernal and Hutton (1959b:64) noted a straight semi- nal vesicle in T. pectinata, and my specimens have a folded cirrus and eggs 19 to 23 by 12 to 14 microns, characters which Manter (1947:323) regarded as defining T. pecti- nata. 1 make the identification with reser- vation because of this confusion and because the specimens appear more like those from Selene vomer discussed below, except that the blunt-ended seminal vesicle in speci- mens from C. erysos extends posteriorly, rather than tapering off and curling back Digenetic Trematodes of Marine Teleosts 123 Figure 2. Tergestia sp., dorsal view. to approximately the middle of the posterior border of the acetabulum. Tergestia sp. Figure 2 Host: Selene vomer (1 of 2). Site: Intestine. Specimen deposited: U.S. N. M. Helm. Coll. No. 71296. Discussion: This could be a new species as suggested by Siddiqi and Cable (1960: 284) who found the flame cell pattern in a small specimen from Selene vomer to dif- fer from that recorded for Tergestia laticol- lis, It is similar to T. acuta, although the 124 vitellaria are not confluent in all my speci- mens. Tergestia pauca Freitas and Kohn, 1965, another similar species, also has nu- merous vitelline follicles which are dispersed both inter- and extracaecally, although it dif- fers from my specimens in having a sucker ratio of 1:1.97 to 2.34 and differently- shaped terminal genitalia. Nine mature individuals 1.2 to 2.4 long have 13 oral lobes, sucker ratios 1:1.5 to 1.7, eggs 16 to 23 by 11 to 16 microns, and width of pharynx 45 to 56% of the length. A prostatic vesicle is present in the cirrus sac. The uterus extends to a level posterior to the rear testis in the three larger speci- mens. Infundibulostomum spinatum Siddigi and Cable, 1960 Host: Haemulon sciurus (1 of 6)*. Site: Intestine. Discussion: A single specimen differs somewhat from the original description. It measures 1.4 long and has a sucker ratio of 1:0.30. The seminal vesicle is indistinctly bipartite, without an observable external seminal vesicle. Proctoeces lintont Siddiqi and Cable, 1960 Host: Lagodon rhomboides (2 of 5)*. Site: Rectum. Proctoeces maculatus (Looss, 1901) Odhner, 1911 Distomum maculatum Looss, 1901. Distomum subtenue Linton, 1907. Proctoeces erythraeus Odhner, 1911. Proctoeces subtenuts (Linton, 1907) Han- son, 1950. Host: Calamus bajonado (1 of 1). Site: Rectum. Discussion: In view of the variation in specimens from various parts of the world, it seems pertinent to give a few character- istics of my four specimens. They are 1.7 to 2.2 long with sucker ratios of 1:1.5 to 1.8 and eggs 45 to 62 by 19 to 24 microns. They have a long genital sinus and vitellaria extending to a level posterior to the testes. Manter and Pritchard (1962:115-116) gave a review of the species. Tulane Studies in Zoology and Botany Wolk 5) Antorchts urna (Linton, 1910) Linton, 1911 Mesorchts urna Linton, 1910. Antorchis holacanthi Siddiqi and Cable, 1960 (new synonym). Hosts: Holacanthus isabelita (1 of 1); Pomacanthus arcuatus (1 of 4); Poma- canthus paru (1 of 1). Site: Intestine and pyloric caeca. Discussion: The overlap in characteristics between Antorchis urna and A. holacanthi is such that the two appear to be synony- mous. Dr. R. M. Cable lent three slides with 25 specimens of A. wrna and four slides with 23 specimens of A. holacanthi from Puerto Rico, Curacao, and Jamaica. Siddiqi and Cable (1960:285) used the general size and shape of the body and pharynx, form of vitellaria, position of testes, extent of uterus, and shape of oral sucker to distinguish between the two species. I can separate Dr. Cable’s specimens into two groups by size of body, number and size of vitellaria, and shape of pharynx, although I do not con- sider them separate species. Of my speci- mens, eleven from or near the pyloric caeca of Holacanthus isabelita, the site and host for A. holacanthi, agree in all respects with others from the intestine of Pomacanthus spp., the site and hosts for A. urna. The lengths, are 1.1) to 15,and Al sto) di7eethe pharynx is spherical to elongate, the vitel- laria are relatively large, the posterior bor- der of the testes does not extend to a level anterior to the acetabulum, the uterus ex- tends to or to a level slightly anterior to the testes, and the oral sucker is usually funnel-shaped. Two differences between Dr. Cable’s specimens of A. holacanthi and mine of A. urna are (1) the length of body (up to 1.1 as compared to 1.1 to 1.7), and (2) number but not size of vitellaria (less than 10 as compared to about 20 to 30). The shorter length might be due to host-influ- ence. Also, Linton (1910:47-48) recorded a length of 0.84 for a specimen of A. urna. My smaller specimens have fewer vitellaria than the larger specimens, although never less than 10. In view of the variations, I consider A. holacanthi a synonym of A. urna. Both wholemounts and sagittal sections individual with few eggs, dorsal view. Figure 4. Botu- lisaccus pisceus, terminal reproductive organs of different individual, dorsal view. Figure 5. Botulisaccus pisceus, anterior end, ventral view. Figure 3. Botulisaccus pisceus, show an almost spherical, spined cirrus pro- jecting into a spinous atrium. Siddiqi and Cable (1960:286) incorrectly stated, in con- trast to Manter (1947:320), that “the muscular lining of the genital atrium is re- flected over the end of the cirrus sac, giving it an annulated appearance... .” Sterimgotrema corpulentum (Linton, 1905) Manter, 1931 Distomum corpulentum Linton, 1905. Hosts: Archosargus rhomboidalis (4 of 5) *; Lagodon rhomboides (1 of 5). Site: Pyloric caeca. Botulisaccus pisceus Caballero, Bravo-Hollis, and Grocott, 1955 Figures 3, 4, and 5 Host: Albula vulpes (7 of 7). Site: Intestine, few in pyloric caeca. Specimen deposited: U.S. N. M. Helm. Coll. Wo, 71297. Digenetic Trematodes of Marine Teleosts 125 Description (based on 18 wholemounts and 3 sectioned specimens): Body pyri- form, 1.0 to 1.8 long by 0.36 to 0.68 in maximum width, usually at or just posterior to level of acetabulum. Cuticle completely spinose. Oral sucker spherical, funnel-shaped, or cup-shaped; 0.07 to 0.18 long by 0.13 to 0.21 wide; sometimes retracted deeply into body (Figure 5). Acetabulum 0.21 to 0.34 long by 0.21 to 0.35 wide. Sucker ratio 1:1.2 to 1.9. Forebody 24 to 36% of body length; usually with a fold in the cuticle at a level just anterior to acetabulum. Pre- pharynx short. Pharynx large, muscular, oval to barrel-shaped, 0.13 to 0.19 long by 0.07 to 0.12 wide, anterior portion with diagonal rather than radial musculature; muscles ex- tending from a point at middle or posterior of pharynx to oral sucker. Esophagus 0.10 to 0.22 long; anterior region surrounded by glandular cells; with large epitheliated lateral outpouchings located midacetabular to a level immediately anterior to acetabulum. Primary intestinal bifurcation near level of posterior border of acetabulum. Caeca ex- tending to approximately middle of post- testicular region; prominently epitheliated except for some of internal portions. Testes symmetrical, elongate; right testis 0.16 to 0.29 long by 0.09 to 0.18 wide; left “testis 016 to 0.28: by 10:07 to 0X8: Posttesticular region 15 to 28% of body length. Cirrus sac muscular, straight to arcuate, 0.20 to 0.38 long, extending from a level immediately posterior to acetabulum to near ovarian level; constricted at level of prostatic vesicle. Cirrus sac containing bipartite seminal vesicle, portions oval, spherical, or irregular; prostatic vesicle wide, longer than seminal vesicle, surrounded by prostatic cells; cirrus short, wide, muscular, unarmed. Genital pore postacetabular, sinis- tral. Genital atrium wide but constricted at genital pore. Ovary spherical to ovoid, 0.10 to 0.18 long by 0.07 to 0.16 wide; between testes with 4 to “4 of its length above anterior margins of testes. Seminal receptacle lacking; sperm in proximal portion of uterus. Laurer’s canal opening dorsally posterior to ovary. Mehlis’s gland compact to diffuse. Uterus extensive, extending from cirrus sac to posterior end of body, leaving most of testes 126 visible. Metraterm short. Vitelline follicles numerous, extending from anterior portion of testes to or near acetabular level. Eggs operculated, thick-walled, 34 to 47 by 23 to 30 microns. Excretory vesicle V-shaped with arms terminating between midacetabular and pharyngeal levels. Excretory pore dorsal or terminal. Discussion: My specimens agree with the original description of Botulisaccus pisceus with two exceptions: first, I could not see the small spines on the cirrus mentioned by Caballero e¢ al. in their generic diagnosis, and, second, they misinterpreted the epi- theliated esophageal outpouching as the in- testinal bifurcation, which is located dorsal to the cirrus sac. Caballero et al. placed Botulisaccus in the Monorchiidae, and Yamaguti (1958:56) transferred it to the Zoogonidae, but Man- ter and Pritchard (1961:483) thought it belonged in the Monorchiidae. The presence of a V-shaped excretory vesicle, or Y-shaped with an extremely short stem, described here for the first time, and the absence of a spinose cirrus indicate that Botwlisaccus should be assigned to the Fellodistomatidae. The only unusual fellodistomatid charac- ters of this genus are the postacetabular cir- rus sac and genital pore and the esophageal outpouchings. This species was previously known only from Albula vulpes from the Pacific Ocean. Claribulla gen. n. Generic diagnosis: Body elongate, spinose. Oral sucker cup-shaped or funnel-shaped. Acetabulum preovarian, enclosed in genital atrium. Pharynx elongate. Caeca long or short. Testes diagonal to tandem. Seminal vesicle preacetabular. Pars prostatica present. Atrial papilla present. Ovary compact, pre- testicular. Seminal receptacle absent. Vitel- Jaria in acetabular-ovarian zone. Eggs with- out filaments. Parasites in marine fishes. Type and only species: Claribulla longula sp. n. Figures 6 and 7 Hosts: Albula vulpes (5 of 7), type hosts; Sphyraena barracuda (1 of 1). Tulane Studies in Zoology and Botany Vols Site: Pyloric caeca, few in upper intestine. Holotype: U. S. N. M. Helm. Coll. No. 71316, paratype: No. 71376. Description (based on 15 wholemounts and 1 sectioned specimen): Body bluntly rounded posteriorly, 0.9 to 2.7 long by 0.21 to 0.37 wide, the widest portion generally in hindbody; usually a slight constriction at acetabular level. Entire cuticle with minute spines. Oral sucker 0.10 to 0.24 long by 0.14 to 0.27 wide. Acetabulum enclosed in genital atrium, protrusible, 0.07 to 0.14 long by 0.09 to 0.13 wide. Sucker ratio 1:0.4 to 0.7. Forebody 29 to 44% of body length. Pharynx 0.07 to 0.19 long by 0.04 to 0.08 wide, in contact with oral sucker and con- nected to it by muscle fibers attached at or near equator of pharynx. Esophagus usually shorter than pharynx. Intestinal bifurcation much nearer oral sucker than acetabulum. Caeca sometimes swollen, extending to be- tween acetabulum and near posterior end of body. Testes diagonal to almost tandem, usually contiguous, spherical to slightly irregular; anterior testis 0.06 to 0.15 long by 0.08 to 0.18 wide, either sinistral or dextral to posterior testis; posterior testis 0.06 to 0.17 long by 0.10 to 0.16 wide. Posttesticular space 26 to 41% of body length. Cirrus sac absent. Seminal vesicle saccate, straight or bent, extending to a level anterior to or oc- casionally overlapping acetabulum. Pars prostatica sinuous or nearly straight, ex- tending posteriorly from anterior portion of seminal vesicle; surrounded by numerous free prostatic cells. Ejaculatory duct incon- spicuous, short and muscular. Atrial papilla large, muscular, located sinistrally, at a level anterior or lateral to acetabulum, protrud- ing into muscular canal which extends pos- teriorly to approximately midacetabular level, then joining large muscular genital atrium. Ovary rounded to slightly irregular, median or submedian, anterior to and al- most always in contact with anterior testis, 0.06 to 0.12 long by 0.08 to 0.12 wide. Laurer’s canal present. Proximal folds of uterus filled with sperm. Vitelline follicles clustered laterally, extending from ovarian level to or near acetabular level. Uterus ex- tensive, filling most of postovarian spaces and extending to anterior border of acetabu- No. 4 0.1 Figure 6. Claribulla longula, holotype, dorsal view. Figure 7. Claribulla longula, terminal re- productive organs, side view. Digenetic Trematodes of Marine Teleosts WAT lum before joining genital atrium. Eggs usually collapsed, noncollapsed specimens 16 to 30 by 10 to 13 microns; 25 to 29 by 11 to 13 in living specimens. Excretory vesicle Y-shaped; stem dividing at ovarian level with arms extending to pharyngeal level. Excretory pore terminal. Discussion: I believe the genus Claribulla belongs in the family Fellodistomatidae. If I were to accept the validity of Monod- helminthidae Dollfus, 1937, then C. longula, with its atrial papilla, or accessory organ as referred to by Yamaguti, could be placed in that family. However, I am treating this family as a subfamily in the Fellodistomat- idae. Yamaguti (1958:256-259) included the genera Monodhelmis WDollfus, 1937, Mehratrema Srivastava, 1939, Tandanicola Johnston, 1927, and Prosogonarium Yama- guti, 1952, in the Monodhelminthidae. Of these, C. /ongula resembles the species of Monodhelmis and Tandanicola by lack- ing a cirrus sac, although it aiffers from members of all four genera by having a preovarian acetabulum, an acetabulum en- closed in the genital atrium, and several other individual differences. The relation- ship of Tandanicola with the Fellodisto- matidae was previously reported by Cable (1953:417), when he amended Tandanico- linae Johnston, 1927, to include Megalomy- zon Manter, 1947, and Pseudosteringophorus Yamaguti, 1940, both recognized fellodisto- matids, and transferred the subfamily from the Brachycoeltidae to the Fellodistomatidae. Claribulla longula further links the monod- helminthids with the fellodistomatids by being similar to a group of the latter which includes Psewdobacciger Nahhas and Cable, 1964, Bacciger Nicoll, 1914, Pentagramma Chulkova, 1939, and Faustula Poche, 1926. Bacciger and Pentagramma were reviewed by Margolis and Ching (1965) without being assigned to a subfamily because of the unstable classification of the fellodisto- matids. Claribulla longula differs from species of the above four genera in the arrangement of the gonads, the absence of a seminal receptacle, and the presence of an atrial papilla. A cirrus sac is absent in members of Pseuwdobacciger and indis- tinct in those of Pentagramma. The absence of a cirrus sac in Psewdobacciger harengulae (Yamaguti, 1938) (= Bacciger h.) and in 128 the illustration of B. bacciger (Rudolphi, 1819) by Stossich (1889) caused Yamaguti (1938 and 1958) to place Baccrger in the Heterophyidae and later in the Cryptogoni- midae. The absence of a cirrus sac in conjunction with the atrial papilla in C. longula is sug- gestive of a cryptogonimid. In the super- family Opisthorchioidea, of which Crypto- gonimidae is a member, however, a con- sistent and conspicuous feature is a seminal receptacle. Also, all known cercariae in that superfamily which leave the snail have eye- spots which can be found, at least as scat- tered granular pigmented remnants, in the forebody of the adult (Cable, 1968: per- sonal communication). If the cercaria of C. longula is found to have eyespots, this spe- cies should be transferred to the family Cryptogonimidae. In Digenea lacking the usual protrusible cirrus, numerous modifications of terminal genitalia and adjacent portions are found in non-related groups, including the fello- distomatids. Convergent evolution of these modifications led parasitologists to place the presently recognized Microphallidae and Gymnophallinae as subfamilies of the Het- erophyidae. More recent studies on the life histories of these three groups indicate that they represent three distinct orders, with two (Microphallidae and Heterophyidae ) in the superorder Epitheliocystidia and one (Gymnophallinae) in the superorder An- epitheliocystidia. The single specimen from Sphyraena bar- racuda may represent an accidental in- fection. The name Claribulla is from clara (dis- tinct) and bulla (knot), and refers to the muscular atrial papilla. FAMILY PARAMPHISTOMATIDAE Fischoeder, 1901 Cleptodiscus reticulatus Linton, 1910 Host: Pomacanthus arcuatus (2 of 4). Site: Rectum. FAMILY PRONOCEPHALIDAE Looss, 1902 Barisomum erubescens Linton, 1910 Tulane Studies in Zoology and Botany Vol. a5 Pleurogonius erubescens (Linton, 1910) Prudhoe, 1944. Monostomum pomacanthi MacCallum, 1916. Pleurogonius pomacanthi (MacCallum, 1916) Price, 1931. Host: Pomacanthus arcuatus (1 of 4). Site: Rectum. Discussion: Linton (1910:70) implied, as I determine, that the dorsoventral muscu- lature of this worm was very unusual. The internal bundles are roughly situated in rows, separated from adjacent bundles by approxt- mately two bundle widths. They are most prominent in the intercaecal region. A few long crystals in the parenchyma apparently are sponge spicules that entered the worm mechanically. FAMILY MICROSCAPHIDIIDAE Tra- vassos, 1922 Hexangitrema pomacanthi Price, 1937 Figures 8 and 9 Hexangitrema breviceca Siddiqi and Cable, 1960 (new synonym). Host: Pomacanthus arcuatus (1 of 4). Site: Intestine and rectum. Specimen deposited: U. S. N. M. Helm. Coll. No. 71298. Description (based on 14 wholemounts and 3 sectioned specimens): Body mono- stomate, broadly fusiform, notched pos- teriorly, 1.5 to 2.4 long by 0.71 to 1.00 wide. Cuticle smooth. Eyespot pigment pres- ent, more conspicuous in smaller specimens. Oral sucker terminal, 0.09 to 0.16 long by 0.11 to 0.15 wide; retrodorsal pockets prom- inent or retracted and inconspicuous. Esopha- gus, including bulb at posterior end, 0.24 to 0.49 long, muscular; bulb spherical to elongated. Caeca wide, terminating blindly from 8 to 33% of length of body from posterior end. Testes tandem, lobed, intercaecal or slightly overlapping caeca, approximately 0.2 to 0.6 long by 0.3 to 0.5 wide. Genital pore near posterior border of oral sucker. Hermaphroditic sac thin-walled elongate; comprised of loosely woven fibrous strands; containing terminal portion of metraterm, prostatic cells, small or no internal seminal Figure 8. Hexangitrema pomacanthi, ventral view. Figure 9. Hexangitrema pomacanthi, an- terior end. Digenetic Trematodes of Marine Teleosts 129 vesicle, pars prostatica, and long (0.08 to 0.10), marrow, muscular hermaphroditic duct. External seminal vesicle long, sinuous, extending to or posterior to intestinal bi- furcation. Ovary smooth to irregular, 0.09 to 0.16 long by 0.06 to 0.13 wide, posttesticular by a variable distance. Mehlis’s gland posterior to ovary. Laurer’s canal present, inconspicu- ous. Vitellaria extracaecal and overlapping caeca ventrally, from between midlevel of either testis to postovarian region, some- times confluent posterior to ovary. Uterus intercaecal or slightly overlapping caeca; proximal coils filled with sperm. Metraterm long. Eggs thin-shelled, operculate, 79 to 92 by 49 to 59 microns. Excretory vesicle short, with two main collecting ducts, each dividing into promi- nent secondary ducts at a level between posterior testis and vesicle. Additional branching present, forming a network of ducts anteriorly. Excretory pore between caudal processes. Discussion: Hexangitrema breviceca Sid- digi and Cable, 1960, is distinguished from H. pomacanthi by the presence of lobed, somewhat separated testes and short caeca that do not reach the posttesticular region. My specimens show gradations in the sep- aration of testes from touching to being well separated, in the extent of the caeca which terminate between the midlevel of the posterior testis and beyond the ovary, and in the degree of lobation of the testes. The testes, however, are never completely spherical as illustrated by Price (1937, Figure 5). Many of my specimens were over 4 in length before fixation. Because of these intergradations and variations, I am placing H. breviceca as a synonym of H. pomacantht. The hermaphroditic sac is difficult to in- terpret in many wholemounts and has not previously been described for H. pomacantht. Sectioned material reveal its structure 1s similar to that which Looss (1902:676-682, figs. 121, 139) described and illustrated in species of Microscaphidium and Angiodic- tyum. These genera are related to Hexangy- trema and include species in marine turtles. Yamaguti (1958) placed these genera in Angiodictyidae Looss, 1902. I agree with 130 Stunkard (1943:143) in suppressing Angio- dictyidae and accepting Microscaphidiidae Travassos, 1922, as the only available name. FAMILY HAPLOPORIDAE Nicoll, 1914 Vitellibaculum spinosum (Siddiqi and Cable, 1960) Durio and Manter, 1968 Allomegasolena spinosa Siddiqi and Cable, 1960. Host: Chaetodipterus faber (1 of 2). Site: Posterior intestine. Discussion: Four specimens 1.8 to 2.4 long have cuticular spines extending to or almost to posterior end in addition to spines at terminal portion of body, pharynx 0.18 to 0.19 long by 0.12 to 0.14 wide and pyriform in shape, and eggs 62 to 75 by 39 to 43 microns. Sucker ratios are 1:1.3 to 1.9 and appear to be dependent on the state of con- traction of the oral sucker at the time of fixation. The forebody is attenuated in three of the specimens. In a 1.6 long curled speci- men with two eggs, lent by Dr. R. M. Cable, the pharynx is barrel-shaped and 0.12 long by 0.095 wide. The variations in shape of body, sucker ratio, and size of pharynx, in addition to the report of Vutellibaculum spinosum from both Chaetodipterus faber and Lutjanus apodus (see: Nahhas and Cable, 1964:179), all suggest that additional specimens of V. attenuatum (Siddiqi and Cable, 1960) from L. apodus, the type host, may well show V. attenuatum to be a synonym of V. spimosum. Dr. Manter has unreported specimens of V. spinosum from C. faber that he has identified from his collection from Beaufort, North Carolina. Megasolena hysterospina (Manter, 1931) comb. n. Figure 10 Lepidauchen hysterospina Manter, 1931. Megasolena archosargi Sogandares-Bernal and Hutton, 1959 (new synonym). Host: Archosargus rhombotdalis (4 of 5). Site: Intestine. Specimen deposited: U.S. N. M. Helm. Coll. INOW 71299: Tulane Studies in Zoology and Botany WoL IS O's QU IZ (eB SHO) Figure 10. Megasolena hysterospina, ventral view. Discussion: Examination of a paratype of Lepidauchen hysterospina, lent by Dr. H. W. Manter, revealed that species to have a hermaphroditic sac instead of a cirrus sac. This species is therefore in the genus Megasolena Linton, 1910, with M. /ystero- spina as a new combination. Prévot (1968) recently studied adult and larval L. steno- stoma Nicoll, 1913, and found that a seminal receptacle was present and a_hermaphro- ditic sac absent. He concluded that Lepz- dauchen Nicoll, 1913, belonged in the sub- family Lepocreadiinae Odhner, 1905. I consider M. archosargi Sogandares- Bernal and Hutton, 1959, a synonym of M. No. 4 hysterospina. Megasolena hysterospina is de- scribed as not having an esophagus or ex- ternal seminal vesicle. My specimens have both, but they are difficult or impossible to see in some individuals. A Laurer’s canal is present; it was not observed in M. archo- sargi. The hermaphroditic sac varies in shape from elongate to almost spherical, with its posterior extension lying between the an- terior and posterior borders of the acetabu- lum. A tubular prostatic vesicle is pres- ent. The vitelline follicles containing yolk granules agree with the descriptions; some gland cells not containing granules, how- ever, extend anteriorly to these. A ventral cuticular area, usually present as a groove and bordered by spines, extends between the suckers and includes their apertures. The bordering spines were observed on speci- mens lent by Dr. R. M. Cable from his Jamaican collection and were seen to extend only to the midacetabular level on Dr. H. W. Manter’s specimen. Additional measurements on 12 of my mature specimens are: Body 1.9 to 3.9 long by 0.99 to 1.56 wide. Oral sucker 0.33 to 0.68 long by 0.34 to 0.54 wide. Acetabulum 0.16 to 0.35 by 0.20 to 0.38. Sucker ratio 1:0.6 to 0.8. Forebody 26 to 34% of body length. Prepharynx contracted or up to 0.13 long. Pharynx 0.22 to 0.46 long by 0.19 to 0.34 wide. Anterior testis 0.20 to 0.55 long by 0.36 to 0.85 wide; posterior testis 0.38 to 0.73 by 0.37 to 0.74. Ovary 0.13 to 0.21 long by 0.32 to 0.65 wide; Dr. Cable’s specimens also have ovaries wider than long. Eggs 56 to 83 by 35 to 47 microns. FAMILY HAPLOSPLANCHNIDAE Poche 1925 Schtkhobalotrema sparisoma (Manter, 1937) Skrjabin and Guschanskaja, 1955 Haplosplanchnus sparisoma Manter, 1937. Host: Nicholsina usta (2 of 3)*. Site: Intestine. Schikhobalotrema kyphosi (Manter, 1947) Skrjabin and Guschanskaja, 1955 Haplosplanchnus kyphosi Manter, 1947. Host: Kyphosus sectatrix (3 of 6). Digenetic Trematodes of Marine Teleosts 131 Figure 11. Schikhobalotrema acutum, genital bulb and associated structures, ventral view. Site: Pyloric caeca, occasionally in upper in- testine. Additional measurements and discussion: Six specimens 1.9 to 2.8 long by 0.52 to 0.79 wide have sucker ratios of 1:0.6 to 1.1, depending on the expansion of the oral sucker. The oral sucker is 0.29 to 0.46 wide, and the acetabulum is 0.23 to 0.35 wide. Forebody is 26 to 36% of the body length. Pharynx is 0.11 to 0.17 long by 0.12 to 0.17 wide. Eggs are 80 to 92 by 51 to 64 microns. The anterior limit of the excretory vesicle may reach the level of the pharynx, and the extremely sinuous seminal vesicle may extend posterior to the testis. Sensory papillae are prominent on the oral sucker and anterior end, as Manter (1937b:385) predicted for all species of “Haplosplanch- nus. Schikhobalotrema acutum (Linton, 1910) Skrjabin and Guschanskaja, 1955 Figure 11 Deradena acuta Linton, 1910. Haplosplanchnus acutus (Linton, 1910) Manter, 1937. Host: Strongylura timucu (1 of 3). Site: Rectum. 132 Tulane Studies in Zoology and Botany Discussion: Sogandares-Bernal and Sogan- dares (1961:145-147) reviewed this species and considered Schikhobalotrema as a sub- genus of Haplosplanchnus. There is consid- erable range in length, an important charac- ter used to distinguish this species from S. adacutus. The previous authors reported a specimen of 0.6, without eggs, from Abu- defduf saxatilis (= A. marginatus as used by Manter, 1937), a host of the smaller S. adacutus, Siddiqi and Cable (1960: Figure 16) illustrated a specimen of 0.7 from a beloniform, the typical host for S. acutum. Manter (1937b:385) discussed specimens of 1.3 to 2.0 long and Caballero et al. (1953: 128) others 1.6 to 2.1 from the Pacific Ocean, all from beloniform fishes. My speci- mens are 2.1 to 2.5 with sucker ratios of 1:1.2 to 1.4 and have eggs 85 to 88 by 53 to 57 microns. Sensory papillae are present around the oral sucker. Manter (1937b: 386) described longitudinal striae dividing the muscular genital bulb. The dorsal por- tion of this bulb in my specimens ap- pears corrugated and the ventral portion is covered by a series of tubules which are filled with granules (Figure 11). The tubules extend posteriorly, dividing into two groups at the midlevel of the bulb. Both groups lead to large reservoirs of granules, which appear like yolk granules, located on each side of the acetabulum. Hymenocotta manteri sp. n. Figures 12 and 13 Host: Mugil cephalus (3 of 3), type host. Site: Intestine and pyloric caeca. Holotype: U. S. N. M. Helm. Coll. No. 71300, paratype: No. 71366. Description: (based on 16 wholemounts and 3 sectioned specimens): Body elongate, 0.9 to 1.6 long by 0.27 to 0.45 at maximum width, usually near acetabular level. Cuticle thick, with prominent rings, especially in tapering hindbody. Pigment granules scat- tered in forebody and occasionally through- out entire worm. Oral sucker replaced by > Figure 12. Hymenocotta manteri, holotype, dorsal view. Figure 13. Hymenocotta manteri, contracted oral disc, ventral view. No. 4 oral disc which may be flared out, or con- tracted (Figure 13) and superficially re- sembling spherical or cup-shaped oral sucker; 0.15 to 0.33 wide at greatest diame- ter, depending on individual variation and contraction. Numerous sensory papillae on disc. Acetabulum protrusible, without lobes, 0.16 to 0.25 long by 0.17 to 0.27 wide; aperture a circular to longitudinal slit-like opening. Sucker ratio 1:0.6 to 1.3. Forebody 26 to 36% of body length. Prepharynx about ' length of pharynx. Pharynx 0.07 to 0.10 long by 0.06 to 0.09 wide. Single caecum usually approaching or extending well into testicular level; highly cellular, often containing particulate matter. Testis elongate, spherical, or slightly ir- regular, not touching acetabulum; 0.13 to 0.19 long by 0.09 to 0.20 wide. Posttesticu- lar space 11 to 39% of body length, de- pending on contraction. Cirrus sac absent. Seminal vesicle tubular, long, extending near or well into testicular level; straight to highly sinuous; often with several loops directly anterior or posterior to acetabulum; connected to spherical prostatic vesicle by pars prostatica of greater length than vesi- cle. Genital atrium muscular. Genital pore median or submedian, at a level near pos- terior of pharynx. Ovary subglobular, 0.06 to 0.12 long by 0.05 to 0.12 wide, variable in position from anterior acetabular level to posterior border of testis, usually anterolateral to testis. Seminal receptacle dorsal, near ovarian level. Vitelline follicles usually in elongate groups, extending from level of acetabulum or slightly posteriorly to a level midway be- tween testis and posterior end of body. Uterus may extend to midtesticular level before turning anteriorly. Metraterm_pres- ent. Eggs usually partially collapsed, 66 to 89 by 35 to 48 microns; not containing fully-developed miracidia. Excretory pore terminal; vesicle bifurcat- ing at midtesticular level with arms extend- ing to near posterior border of pharynx. Discussion: Manter (1961:67-69) erected the genus Hymenocotta to accommodate a single species, H. mulli Manter, 1961, which differs from all other haplosplanchnids in having a cirrus sac and a six-lobed disc, which replaces the oral sucker. The present species is the second with a disc, but it is not six-lobed, and a cirrus sac is absent. Digenetic Trematodes of Marine Teleosts IN) What appears to be the cirrus sac is weakly developed and not evident in all of Dr. Manter’s specimens of H. mulli from New Caledonia (Manter, 1968:personal commu- nication). Pending further information, the present species is placed in Hymenocotta because of the oral disc which, with the absence of a well-developed bulb of pros- tatic ducts, distinguishes H. manteri from Schikhobalotrema elongatum Nahhas and Cable, 1964, an otherwise similar species from the same host. This species is named in honor of Dr. Harold Winfred Manter, in recognition of his contributions to the field of tremato- dology. FAMILY LEPOCREADIIDAE 1934 Nicoll, Thysanopharynx elongatus Manter, 1933 Host: Lactophrys quadricornis (1 of 3). Site: Intestine. Discussion: Manter (1963a:107) reduced the family Megaperidae Manter, 1934, which would include this and the following species, to a subfamily in the Lepocreadiidae. Nah- has and Cable (1964:179) tentatively ac- cepted Megaperidae and placed it in the superfamily Haploporoidea Mehra, 1961, along with the haploporids and_haplo- splanchnids. They also noted that Enenterum and Cadenatella, generally placed in the Lepocreadiidae, have features in common with members of the Haploporoidea and that they might later be transferred to that group. Pending life history studies, how- ever, they retained the two genera in the Lepocreadiidae. I also prefer to retain them, as well as Thysanopharynx and Megapera, with the lepocreadiids. Megapera sp. Host: Lactophrys quadricornis (1 of 3). Site: Intestine. Discussion: A single immature individual is included in this study because Lactophrys quadricornis is the final host for several species of Megapera., It is 0.51 long and has a constricted hindbody as in M. gyrina, al- though the vitellaria extend from the pos- terior end to the testes as in M. pseudogyrina and M. ovalis, The sucker ratio is 1:0.29. 134 Enenterum aureum Linton, 1910 Host: Kyphosus sectatrix (5 of 6). Site: Rectum and posterior intestine. Cadenatella americana Manter, 1949 Host: Kyphosus sectatrix (4 of 6). Site: Intestine. Discussion: The forebody, especially the the ventral region, contains individual gland cells opening to the exterior. The elongate or spherical prostatic vesicle generally ex- tends posteriorly, and dorsal to the acetabu- lum, rather than along its anterior border, as in the holotype. A sectioned specimen does not have a cirrus sac, confirming what Nahhas and Cable (1964:192) believed. A thin membrane, however, appears to sur- round the vesicle in some wholemounts. Cadenatella floridae sp. n. Figures 14, 15, and 16 Host: Kyphosus sectatrix (5 of 6), type host. Site: Pyloric caeca and occasionally anterior intestine. Holotype: U. S. N. M. Helm. No. 71301, paratype: No. 71367. Description (based on 10 mature whole- mounts and 2 sectioned specimens): Body elongate, 3.0 to 4.7 long by 0.31 to 0.44 wide; largest immature specimen 3.01. Cuti- cle spinose from anterior end to testicular level. Eyespot remnants present; brownish- yellow pigment scattered through paren- chyma. Oral sucker 0.10 to 0.16 in diameter with 8 muscular preoral lobes, 4 dorsal and 4 ventral. Acetabulum 0.15 to 0.19 long, 0.14 to 0.19 wide, and 0.12 to 0.17 deep; the surrounding cuticle with an inner ring of papillae adjacent to cuticular spines. Sucker ratio 1:1.1 to 1.5. Forebody 21 to 43% the length of body, with 8 to 11 midventral accessory suckers; occasion- ally 1 or 2 suckers ventral to posterior por- tion of pharynx, never anterior to pharynx; the second sucker anterior to acetabulum usually largest, 0.08 to 0.12 wide; smallest sucker 0.04 to 0.06 wide. Prepharynx 0.06 to 0.42, depending on contraction. Pharynx massive, pyriform, 0.22 to 0.32 long by 0.09 to 0.14 wide. Esophagus from 0.09 to 0.24 long; large muscular sphincter at an- terior end. Caeca extending to posterior end of body, joining excretory vesicle to form uroproct with terminal pore. Tulane Studies in Zoology and Botany Vol. 15 0.2 15 Wey Mi (sa Cs Figure 14. Cadenatella floridae, holotype, ventral view. Figure 15. Cadenatella floridae, acetabular region, ventral view. Figure 16. Cadenatella floridae, anterior end, dorsal view. Testis single, elongate, occasionally with indentation, 0.32 to 0.73 long by 0.10 to 0.21 wide. Posttesticular region 21 to 40% of body length. Cirrus sac absent. Seminal vesicle long, sinuous, extending to about midway between acetabulum and_ ovary. Prostatic vesicle spherical to ovoid, near an- terior border of acetabulum. Ejaculatory duct short. Genital atrium muscular, inconspicu- ous in wholemounts. Genital pore in in- vagination leading to posterior accessory sucker. Ovary globular to almost triangular; 0.08 to 0.14 long by 0.11 to 0.19 wide; slightly anterior to or overlapping forward portion of testis. Seminal receptacle of uterine type, at a level anterior to ovary. Mehlis’s gland near seminal receptacle. Laurer’s canal open- ing dorsal to ovary. Uterus preovarian, inter- caecal or occasionally overlapping caeca. No. 4 Metraterm present. Vitelline follicles dorsal, ventral, and lateral to caeca and testis; ex- tending from posterior ‘% of testicular level to end of body; filling most of posttesticular space. Gland cells not con- taining granules extending from near an- terior level of testis almost to pharyngeal level. Eggs thin-shelled, 41 to 72 by 30 to 40 microns for partially-collapsed eggs; 48 to 58 by 31 to 34 microns for noncollapsed eggs. Excretory vesicle Y-shaped, bifurcating at ovarian level; arms extending about mid- way between acetabulum and ovary; excre- tory canals extending from tips of arms to oral sucker then posteriorly through hind- body. Discussion: Five other species of Cadena- _ tella have been described, all from fishes of the genus Kyphosus. Only two, C. brumpti (Dollfus, 1946) and C. &yphosi Nahhas and Cable, 1964, have more than two ac- cessory suckers. The present species differs _ from both by having 8 to 11 rather than 14 to 17 accessory suckers, by the extension of spines on the ventral side of the body, and probably by possessing a muscular esophageal sphincter. It resembles C. kyphosi in size and oral lobes but differs in the lo- cation of the accessory suckers. Papillae near the acetabulum (Figure 15) probably are sensory in function; they have not been described for any other species of Cadena- tella. Nahhas and Cable (1964:191-192) dis- cussed the known species and considered _ Jeancadenatia a synonym of Cadenatella. - Sogandares-Bernal (1959:80) clarified some _ of the provisional description of C. brumpti by Dollfus (1946:124-126). Lepocreadinm trulla (Linton, 1907 ) Linton, 1910 Distomum trulla Linton, 1907. Host: Ocyurus chrysurus (3 of 5). Site: Intestine. Discussion: Six specimens 0.9 to 1.2 long have sucker ratios of 1:0.7 to 0.9 and eggs 49 to 56 by 29 to 35 microns. The pharynx is lobed anteriorly. The anterior extent of the excretory vesicle was observed in a liv- ing specimen to vary from the pharyngeal level to one a short distance posterior to the intestinal bifurcation. Sogandares-Bernal and Hutton (1960:282) discussed that fea- Digenetic Trematodes of Marine Teleosts i) [hes Figure 17. Lepocreadium pyriforme, ventral view. ture in fixed specimens and Hanson (1950: 78) gave measurements of small individuals, also from Ocyurus chrysurus. Lepocreadium pyriforme (Linton, 1900) Linton, 1940 Figure 17 Distomum pyriforme Linton, 1900. Host: Sardinella anchovia (1 of 3)*. Site: Pyloric caeca. Specimen deposited: U.S. N. M. Helm. Coll. Nos 71302: Discussion: Variations within Lepocrea- dium pyriforme make the status of that species uncertain, as discussed by Sogandares- Bernal and Hutton (1960:280-282) in re- gard to specimens from different hosts 136 which Linton identified as L. pyriforme. Nahhas and Cable (1964:188) and Nah- has and Short (1965:43) accepted as L. pyriforme only individuals resembling Figure 47 (Linton, 1940) or Figure 9 (Sogan- dares-Bernal and Hutton, 1960), both drawn from specimens from Palinurichthys perci- formis (Mitchill), the type host. Mine dif- fer from those figures illustrated in the extent of vitellaria, length of the prepharynx, shape and extent of the external seminal vesicle, and sucker ratio. In some of those respects, they are more like those illustrated by the above authors from other hosts. I am provisionally placing my specimens in L. pyriforme and describing them, so when adequate material is examined, they can be placed in the proper species. Description (based on 4 _ specimens): Body 0.96 to 1.06 long by 0.32 to 0.40 in maximum width. Oral sucker 0.10 to 0.12 long by 0.10 to 0.12 wide. Acetabulum 0.07 to 0.08 by 0.07 to 0.09. Sucker ratio 1:0.7 to 0.08. Forebody 42 to 45% of body length. Prepharynx approximately '2 as long as pharynx width. Pharynx 0.09 to 0.10 long by 0.07 wide. Esophagus as long as, or longer than, pharynx. Caeca ending at level of posterior border of rear testis to midlevel of posttesticular space. Testes tandem, slightly irregular; anterior testis 0.07 to 0.09 long by 0.10 to 0.12 wide; posterior testis 0.09 to 0.10 by 0.10 to 0.13. Posttesticular space 12 to 18% of body length. Genital pore anterior and sinistral to acetabulum. Cirrus sac 0.11 to 0.21 long by 0.05 wide, posterior extent from midway between acetabulum and ovary to ovarian level. External seminal vesicle elongate, large, approaching size of cirrus sac; termi- nating in zone of anterior testis. Ovary smooth, 0.05 to 0.08 long by 0.06 to 0.07 wide. Seminal receptacle sinistral to and usually touching anterior testis. Metraterm muscular, over *4 as long as cir- rus sac. Vitelline follicles from level of in- testinal bifurcation to posterior end of body. Eggs partially collapsed, 57 to 71 by 26 to 37 microns. Excretory pore terminal or subdorsal; vesicle extending intercaecally from pore, on dextral side, then crossing the left caecum near intestinal bifurcation and ending in the left side of forebody at anterior esopha- geal level. Tulane Studies in Zoology and Botany Vol. 15 Lepocreadium floridanum Sogandares- Bernal and Hutton, 1959 Host: Lagodon rhomboides (2 of 5). Site: Pyloric caeca. Discussion: Twenty-two specimens are 0.4 to 1.2 long and have sucker ratios of 1:0.9 to 1.2. The eggs are within the range given by Nahhas and Short (1965:43); those from living specimens were 62 to 66 by 32 to 40 microns. Extent of the excre- tory vesicle in a living individual was ob- served to vary between the bifurcal and an- terior esophageal level. Pseudocreadium scaphosomum Manter, 1940 Hypocreadinm scaphosomum 1940) Yamaguti, 1942. Host: Monacanthus hispidus (3 of 6)*. Site: Intestine. Discussion: Opinion differs as to synon- ymy between Hypocreadium Ozaki, 1936, and Pseudocreadium Layman, 1930, and also among the various species assigned to those genera. Sogandares-Bernal (1959:75) and Nahhas and Cable (1964:193) considered Hypocreadium a synonym of Pseudocrea- dium, and Manter (1946:414) questioned whether the same may be true of P. scapho- somum Manter, 1940, and P. lamelliforme (Linton, 1907). Also Bravo-Hollis and Man- ter (1957:38) were hesitant as to whether P. scaphosomum was a synonym of P. pa- tellare (Yamaguti, 1938). Pseudocreadinm lamelliforme (Linton, 1907) and P. myo- helicatum (Bravo-Hollis and Manter, 1957 ) are usually separated from P. patellare and P. scaphosomum by possessing some vitel- line follicles which overlap the caeca ven- trally. However, Sogandares-Bernal (1959: 76) found at least a few such follicles in a series of paratypes of P. scaphosomum and considered P. myohelicatum its synonym. My specimens can be placed in three groups: one of 5 individuals 1.0 to 1.3 long with eggs 51 to 80 by 35 to 47 microns are from 2 specimens of M. hispidus 9 cm long; another of 7 individuals 1.5 to 1.7 long with eggs 56 to 71 by 39 to 43 microns are from 1 specimen of M. hispidus 26 cm long; and a third of 17 individuals 0.9 to 1.6 long with eggs 56 to 75 by 32 to 43 microns are from 3 specimens of Balistes ( Manter, No. 4 capriscus. The three groups are alike in most respects: they have smooth to lobed gonads; long, sinuous, divided prostatic vesi- cle; sinuous cirrus when inverted; variable extent of caecal undulations; genital pore ventral or slightly anterior to caeca; and ex- tremely muscular distal portion of the metra- term. I place these specimens in two species, primarily on the basis of their vitelline ar- rangement; that is: P. scaphosomum from M. hispidus and P. lamelliforme from B. capriscus. Although vitellaria in a few speci- mens of P. scaphosomum are partially ven- tral to the caeca, the worms from M. hispi- dus are distinct from P. lamelliforme and five paratypes of P. myohelicatum on a slide lent by Dr. H. W. Manter. The vitellaria in P. myohelicatum are not intensive or in clusters and neither P. lamelliforme nor P. myohelicatum have vitellaria restricted from the caecal zone as in P. scaphosomum. For this reason, I consider P. myohelicatum a valid species, rather similar to P. balistis Nagaty, 1942 (originally P. balistes), which also has a short, reasonably straight prostatic vesicle, but a distinctly-partitioned external seminal vesicle. The internal seminal vesicle is extremely muscular, especially in P. scaphosomum, and may be spherical, bi- partite, or teardrop in shape, depending on contraction. Width of muscular constriction of the prostatic vesicle varies, and the ex- ternal seminal vesicle may be sinuous, tubu- lar, or saccate. Additional observations may reveal that P. patellare is conspecific with P. scaphoso- mum, and P. lamelliforme with P. myoheli- catum, or possibly that all four are the same species. Illustrations of P. scaphosomum from B. polylepis by Lamothe (1963:102- 108) and Caballero et al. (1953:117-121) indicate variability of that trematode. Pseudocreadium lamelliforme (Linton, 1907) Manter, 1946 Distomum lamelliforme Linton, 1907 (in part). Host: Balistes capriscus (2 of 4). Site: Intestine. Discussion: The excretory sphincter in my species of Pseuwdocreadium, especially P. lamelliforme, gives a false impression of 5 to 10 large, radial spines. Nahhas and Cable (1964:193-194) de- scribed P. lactophrysi and pointed out that Digenetic Trematodes of Marine Teleosts 7. Linton (1907:108-109) probably consid- ered that species, Dermadena lactophrysi, and P. lamelliforme as a single species. Dermadena lactophryst Manter, 1946 Distomum lamelliforme Linton, 1907 (in part). Host: Lactophrys quadricornis (2 of 3). Site: Intestine. Neoapocreadium coili (Sogandares- Bernal, 1959) Siddiqi and Cable, 1960 Apocreadium coili Sogandares-Bernal, LOD 2: Host: Balistes capriscus (1 of 4). Site: Intestine. Discussion: My single specimen is 3.3 long by 1.0 wide, with a forebody 37% the length of the body. The sucker ratio is 1:0.90, and the eggs measure 80 to 82 by 52 to 54 microns. The sucker ratio and amount of posttesticular space, characters used to separate N. coili from N. bravoi (Sogandares-Bernal, 1959), are intermediate between values for the two species. The confluent preacetabular vitellaria, however, are numerous, rather than sparse as_ illus- trated for N. bravoi. There appears to be a misprint in the description (Sogandares- Bernal, 1959:83): the body width of N. coi, as illustrated, should be about 0.69 rather than 0.225, as given in the text. When additional life history studies of trematodes belonging in the Lepocreadiidae and related families are collected, I believe the genera Apocreadium Manter, 1937, Neo- apocreadium Siddiqi and Cable, 1960, Post- porus Manter, 1949, and others will be transferred to a separate family. But un- til the limits of such a group are better defined, I prefer to leave the above three genera in the Lepocreadiidae. Apocreadium mexicanum Manter, 1937 Host: Balistes capriscus (1 of 4)*. Site: Intestine. Discussion: Three immature worms are 1.8 to 3.3 long, and even though Manter (1937a:11) reported a mature specimen of A. mexicanum 2.2 long, 1 believe only one species is involved. The lymphatic ves- sels are branched, and the vitellaria extend anteriorly to the posterior edge of the ace- tabulum. The largest specimen has a fore- 138 Tulane Studies in Zoology and Botany Vol. 15 body 23% the length of the body and a posttesticular space 43% that length; the sucker ratio is 1:1.6. The smaller specimens have longer forebodies, shorter posttesticular areas, and smaller sucker ratios. Siddiqi and Cable (1960:306) and Nahhas and Cable (1964:190) discussed some of the vari- ability of this species. Apocreadinm foliatum (Siddigi and Cable, 1960) comb. n. Figure 18 Homalometron foliatum Siddiqi and Cable, 1960. Hosts: Haemulon aurolineatum (1 of 7)*; Haemulon carbonarium (1 of 1)*; Haemulon parrai (3 of 7)*. Site: Intestine. Specimen deposited: U.S. N. M. Helm. Coll. No. 71303. Discussion: 1 place this species in the genus Apocreadium because it possesses a prominent lymphatic system. There are two pairs of longitudinal vessels which extend almost the entire length of the body and give off numerous branches, primarily in the hindbody. My 13 wholemounts and 2 sectioned specimens are 1.5 to 5.5 long by 0.44 to 0.85 wide, with an immature specimen 1.74 long. The cuticular spines cover the entire body of immature specimens but only near the testicular level in adults. The prepharynx may be as short as 1/3 that of the pharynx and the seminal vesicle overlaps or is slightly anterior to the ovary, predominantly on the left side. Eggs are 85 to 98 by 48 to 62 microns. With the exception of being larger worms with correspondingly larger organs, my specimens agree with those from Haemulon sciurus, lent by Dr. R. M. Cable from his Jamaican collection. Nahhas and Cable (1964:184) discussed specimens which had vitellaria extending to the ovarian level and had a larger body, wider range in sucker ratio, and larger eggs than in those of the original description. The excre- tory vesicle in my specimens and the bor- rowed ones extends only to or slightly be- yond the posterior border of the rear testis, rather than to the anterior testis. Apocreadinm foliatum differs from A. balistis Manter, 1947, A. caballerot Bravo- Hollis, 1953, A. longisinosum Manter, 1937, A. mexicanum Manter, 1937, and A. syna- Figure view. Rede ae ne GS 2 86 er as eCot ; FV eae AQ 5 wet ' a ae ene ek a i aN fe) g 6 8 HK) a 0.5 18. Apocreadium foliatum, ventral No. 4 gris Yamaguti, 1953, in usually having an acetabulum smaller than the oral sucker. In this respect, A. folliatum is more like the species of Apocreadium described below. Apocreadium cryptum sp. n. Figure 19 Hosts: Antsotremus virginicus (1 of 6); Haemulon parrai (1 of 7), type host; additional hosts from Florida Keys cited in discussion. Site: Pyloric caeca and intestine. Holotype: U. S. N. M. Helm. Coll. No. 71304, paratype: No. 71368. Description (based on 6 mature speci- mens from Biscayne Bay and Florida keys) : Body elongate, 4.5 to 9.3 long by 0.98 to 1.53 in maximum width, hindbody ex- tremely foliate usually with median inden- tation at posterior extremity. An immature individual 4.0 long. Cuticle thick, unspined. Little, if any, eyespot pigment in mature specimens; more conspicuous in immature individuals. Oral sucker funnel-shaped, 0.71 to 1.01 long by 0.68 to 1.37 wide. Acetabu- lum 0.34 to 0.54 long by 0.30 to 0.59 wide. Sucker ratio 1:0.4 to 0.5. Forebody 30 to 36% of body length. Prepharynx longer or shorter than pharynx. Pharynx 0.19 to 0.32 long by 0.20 to 0.36 wide, without promi- nent anterior circular muscle band. Length of esophagus variable, up to 80% as long as pharynx. Intestinal bifurcation midway between suckers or closer to oral sucker. Caeca marrow, terminating blindly near posterior end of body. Testes lobate, tandem, intercaecal, in con- tact or separated; separated to a greater ex- tent in larger specimens; anterior testis 0.15 to 0.31 long by 0.28 to 0.38 wide; posterior testis 0.19 to 0.41 by 0.26 to 0.35. Post- testicular region 31 to 40% of body length. Cirrus sac absent. Seminal vesicle saccate, extending to roughly midway between ace- tabulum and ovary. Entire length of pars prostatica surrounded by prostatic cells, more conspicuous and numerous anteriorly. Geni- tal atrium tubular. Genital pore median or slightly submedian, anterior to or ventral to acetabulum. Ovary smooth or slightly irregular, 0.11 to 0.30 long by 0.20 to 0.28 wide; anterior and slightly dextral to anterior testis, sep- arated by either more or less than length of testis. Seminal receptacle between seminal Digenetic Trematodes of Marine Teleosts 139 o 00. RASS 0! te ms LF Figure 19. Apocreadium cryptum, holotype, ventral view. vesicle and ovary. Mehlis’s gland between ovary and anterior testis. Vitelline follicles numerous, those containing yolk extending from level between seminal vesicle and ace- tabulum to posterior end of body; smaller 140 Tulane Studies in Zoology and Botany gland cells, without yolk granules, extend- ing forward to oral sucker. Uterus joining male duct near anterodorsal portion of ace- tabulum. Eggs 92 to 111 by 58 to 67 mi- crons. Excretory vesicle terminating anteriorly at posterior edge of rear testis. Excretory pore subdorsal. Lymphatic system conspicu- ous with longitudinal vessels, 3 pairs in 1 specimen, 2 in another, running the length of the body; numerous branches in hind- body. Discussion: This species differs from all others in Apocreadinm and Homalometron in having a funnel-shaped oral sucker and a more foliate hindbody. The only other un- spined Apocreadium is A. longisinosum Manter, 1937. Three larger specimens of A. cryptum were donated by Dr. Robert Schroeder, who collected them from the pyloric caeca of Haemulon sciurus and H. plumieri caught near Lower Matecumbe Key, Florida. They differ from smaller individuals by having a more contracted pharynx and a greater number of eggs with a larger mean size, but the largest individual from H. parrai also has large eggs. A large specimen has one anomalous caecum which is completely interrupted to form a short branch joining the normal caecum and another portion with two blind ends extending the length of the vitellaria and lacking well-developed epithe- lium of normal caeca. No evidence of injury or degeneration as the cause of this ab- normality was observed. The name cryptwm refers to the some- times secret habitat of this species in a pyloric caecum. Postporus epinepheli (Manter, 1947) Manter, 1949 Opisthoporus epinepheli Manter, 1947. Postporus mycteropercae (Manter, 1947) Manter, 1949. Host: Mycteroperca bonaci (1 of 3)*. Site: Intestine. Discussion: Siddiqi and Cable (1960: 305) reduced Postporus mycteropercae (Manter, 1947) to a synonym of P. epzi- > Figure 20. Neolepidapedon macrum, holo- type, ventral view. VolM> AQ \ \\ JN ) & a4 No. 4 nephelt. My three specimens are 2.8 to 3.1 long, have sucker ratios of 1:0.95 to 1.2 and have the prepharynx longer than the pharynx. They are like P. mycteropercae, except that the excretory vesicle reaches the pharyngeal level, thereby supporting the synonymy suggested by Drs. Siddiqi and Cable. Neolepidapedon macrum sp. n. Figure 20 Host: Mycteroperca microlepis (1 of 2), type host. Site: Intestine. Holotype: U. S. N. M. Helm. Coll. No. 71305. Description (based on 1 specimen): Body elongate, 4.7 long by 0.42 wide. En- tire cuticle spinose. Eyespot pigment pres- ent. Oral sucker slightly subterminal, 0.11 long by 0.12 wide. Acetabulum 0.11 long By e012) wide. ‘Sucker ratio a: 11. Pre- _ pharynx 0.10 long. Pharynx 0.06 long by 0.06 wide. Esophagus 0.30 long. Intestinal bifurcation nearer oral sucker than acetabu- lum; caeca extending almost to posterior end of body. Testes tandem, separated, smooth; an- terior testis 0.30 long by 0.20 wide; pos- terior testis 0.35 by 0.25. Posttesticular space 0.54 long. Cirrus sac extending well posterior to acetabulum; containing small, oval, internal seminal vesicle, large con- spicuous prostatic vesicle, and cirrus. Ex- ternal seminal vesicle tubular, sinuous, about twice length of cirrus sac. Prostatic cells few, free in parenchyma, mostly surround- ing anterior portion of external seminal vesicle. Genital atrium small. Genital pore sinistral near anterolateral border of ace- tabulum. Ovary entire, smooth, submedian, 0.19 _ long by 0.17 wide, anterior to and separated from anterior testis by about length of testis. Seminal receptacle postovarian. Uterus descending slightly posterior to ovary before extending anteriorly. Metraterm not as long as cirrus sac. Vitellaria extending from near posterior portion of cirrus sac to posterior end of body; confluent posterior to testes and between gonads, partially overlapping the gonads and excretory vesicle dorsally. Eggs 63 to 66 by 35 to 41 microns. Excretory vesicle tubular, ending nearer intestinal bifurcation than acetabulum; Digenetic Trematodes of Marine Teleosts 141 sphincter well developed; pore terminal. Discussion: Five other species of Neo- lepidapedon with an excretory vesicle ex- tending almost to the intestinal bifurcation have been described. This species resembles N. hypoplectri Nahhas and Cable, 1964, and N. mycteropercae Siddiqi and Cable, 1960. It differs from N. hypoplectri in body shape, sucker ratio, number of prostatic cells, posi- tion of genital pore, and, probably, length of prepharynx. Nahhas and Cable (1964: 186) examined 55 additional specimens of N. mycteropercae, some from Mycteroperca bonact, and discussed variability in the species. This specimen differs from N. mycteropercae in position of genital pore, anterior extent of vitellaria, and smaller sucker ratio. Caballero et al. (1955:131-134) de- scribed and illustrated what they believed was Lepidapedon elongatum (Lebour, 1908) Nicoll, 1915, from a serranid host. If their species actually belongs in Neolepidapedon Manter, 1954, a genus erected for species without a membrane enclosing the prostatic cells, then we are probably dealing with the same species. My specimen differs from their description slightly in measurements and by possessing cuticular spines over the entire body, rather than just the anterior portion. Caballero et al. discussed the possi- bility of loss of some spines from their specimens during preparation of the ma- terial. Bianium plicitum (Linton, 1928) Stunkard, 1931 Distomum sp. Linton, 1898 (from Lagocephalus laevigatus )—Linton 1905 (from Syngnathus fuscus, Cynoscion regalis, and Sphaeroides maculatus). Psilostomum plicitum Linton, 1928. Bianium concavum Stunkard, 1930. Bianium adplicitum Manter, 1940. Diploproctodaeum plicitum (Linton, 1928) Sogandares-Bernal and Hutton, L959: Hosts: Sphaeroides spengleri (1 of 2); Sphaeroides testudineus (4 of 5). Site: Intestine. Discussion: Manter (1947:279-281) dis- cussed the variability in small specimens from Tortugas, Florida, medium-sized ones from Beaufort, North Carolina, and large in- dividuals from Woods Hole and the Pa- 142 cific. My 10 mature specimens substantiate his data, as they also are intermediate as are those from Beaufort. They are 1.0 to 1.8 long with an immature individual reach- ing 1.2. The vitellaria never extend anterior to the acetabular level, and the acetabulum is larger than the oral sucker, except on the smallest and two immature specimens. This finding conflicts with Manter’s suggestion that an increase in the size of the oral sucker may be correlated with an increase in body length. Sogandares-Bernal and Hutton (1958) reviewed the status of Bzaniwm Stunkard, 1930, and considered it a synonym of Dzplo- proctodaeum LaRue, 1926. Travassos, Freitas, and Buhrnheim (1965) erected Amaroco- tyle as a new genus for the species A. simonet, which possesses a small unlobed ovary and lacks an external seminal vesicle and is similar to B. plicitum, also from Sphaeroides testudineus. The authors sep- arated three previously established species into Diploproctodaeum and Bianium and five others into Diploporetta Strand, 1942, and Diplocreadium Park, 1939. They did not include B. vitellosum (Sogandares- Bernal and Hutton, 1959) Gupta, 1968, and they accepted B. lecanocephalum Pérez Vigueras, 1955, which Sogandares-Bernal and Hutton (1958:566-567) considered a synonym of D. haustrum (MacCallum, 1918). Lamothe (1966:150-151) discussed the problem and recognized B. plicitum, B. holocentri (Yamaguti, 1942), D. haustrum, D. cryptosoma (Ozaki, 1928), D. hemz- stroma Ozaki, 1920, D. tetraodontis (Nagaty, 1956), and D. vitellosum Sogandares-Bernal and Hutton, 1959. He separated Brantwm from Diploproctodaeum on the basis of in- cluding species with a more or less longer esophagus and ventrolateral folds which do not unite beyond the acetabulum. Gupta (1968), unaware of the works by Travassos et al. or Lamothe, considered Diploprocto- daeum monospecific and separated D. haus- trum from all the species of Bianiwm by the shape of the body, length of esophagus, po- sition of genital pore, shape of ovary, and extent of vitellaria. He placed all the species discussed by Lamothe (except D. haustrum), Diplocreadium koreanum Park, 1939, and three new species all in the genus Braniwm. Oshmarin, Mamaev, and Parukhin (1961) described three new related species Tulane Studies in Zoology and Botany Vol. 15 from Abalistes stellaris in the Gulf of Tonkin, erecting a new genus for Sphincterts- tomum acollum. After considering their de- scriptions of the two new species of Dzplo- proctodaeum, 1 still accept that genus as monospecific; therefore, B. longipygum and B. macracetabulum become new combina- tions. All authors except Yamaguti (1958:134) and Gupta (1968:145) have apparently ac- cepted Diplocreadinm Park, 1939, until the presence of ani in D. koreanum is estab- lished. I think that in order to better understand the validity of the above genera, additional specimens of Diploproctodaeum haustrum, Diplocreadinm koreanum, Amarocotyle st- monet, and Sphincteristomum acollum should be examined. Bianium vitellosum (Sogandares-Bernal and Hutton, 1959) Gupta, 1968 Diploproctodaeum vitellosum Sogandares- Bernal and Hutton, 1959. Host: Chilomycterus schoepfi (3 of 6)*. Site: Intestine. Discussion: Four specimens show consid- erable variation among themselves and with the description. The ovary is not necessarily sinistral or in contact with the anterior testis. In fact, the testes are diagonal to al- most symmetrical, or overlapping medianly. As fixative was applied to one specimen, the testes changed from a tandem to almost sym- metrical position. The right testis is consistently more ventral than the left. Contrary to the description by Sogandares- Bernal and Hutton, the cirrus sac in my specimens of Bianiwm vitellosum is not al- most in contact with the ovary (as it is in my specimens of B. plicitum), and the external seminal vesicle does not overlap the ovary as it consistently does in my individuals of B. plicitum. The genital pore is located be- tween the level of the posterior and middle of the pharynx, rather than at the anterior border of the acetabulum. Vitellaria always converge anterior to the acetabulum and usually reach the oral sucker, but extend only to the midpharyngeal level in one specimen. The body length is 1.9 to 2.2, with sucker ratio of 1:1.0 to 1.2. There are a few delli- cate cuticular spines, primarily on the an- terior portion of the body, and the excre- tory vesicle apparently extends almost to No. 4 the ovary. The ejaculatory duct is not sinu- ous. In life, B. vitellosum is orangeish, with what is normally the posterior portion of the body usually located behind and between the oral sucker and acetabulum, which, along with the anterior body folds, act as the holdfast organ. The free end may take nu- merous shapes, including elongate and cau- date. In one living specimen the ani opened through two terminal protuberances. Multitestis inconstans (Linton, 1905) Manter, 1931 Distomum inconstans Linton, 1905. Host: Chaetodipterus faber (2 of 2). Site: Intestine and pyloric caeca. Crassicutis marina Manter, 1947 Host: Eucinostomus gula (1 of 3)*. Site: Midintestine. FAMILY OPISTHOLEBETIDAE Fukui, 1929 Pachycreadinm crassigulum (Linton, 1910) Manter, 1954 Lebouria crassigula Linton, 1910. Plagioporus crassigulus (Linton, 1910) Price, 1934. Host: Calamus bajonado (1 of 1). Site: Midintestine. FAMILY OPECOELIDAE Ozaki, 1925 Apopodocotyle oscitans (Linton, 1910) Pritchard, 1966 Hamacreadinm oscitans Linton, 1910. Podocotyle breviformis Manter, 1940. Pseudoplagioporus brevivitellus Siddiqi and Cable, 1960. Hosts: Antsotremus virginicus (2 of 6); Archosargus rhomboidalis (2 of 5). Site: Intestine and pyloric caeca. Discussion: Pritchard (1966:166-167 ) erected Apopodocotyle, with A. oscitans as the type species, to include Podocotyle-like species having a postbifurcal, median, or sub- median genital pore. The specimens from the intestine of Anisotremus virginicus are 0.9 and 1.3 long, whereas three specimens from the pyloric caeca of Archosargus rhombot- dalis are 2.1 to 2.8 long. One of the large specimens is unusually gravid, with the uterus extending to the posterior end of Digenetic Trematodes of Marine Teleosts | oy 0.5 PLEO = € 5508: f 3 2\ t CO QO® rs Wee ) a PSS a ODES B®, © - Oo Og Figure 21. Hamacreadium confusum, holo- type, ventral view. the body. Partially-collapsed eggs are 49 to 56 by 35 to 40 microns. Hamacreadium mutabile Linton, 1910 Hosts: Lutjanus griseus (1 of 3); Lutjanus synagris (1 of 7). Site: Pyloric caeca and stomach. Hamacreadium confusum sp. n. Figure 21 Hamacreadium mutabile Linton, 1910, of Siddigi and Cable, 1960 (in part). Host: Ocyurus chrysurus (1 of 5), type host. Oo 144 Site: Intestine. Holotype: U. S. N. M. Helm. Goll. No. 71306. Description (based on 2 mature and 2 immature specimens from Biscayne Bay and Puerto Rico): Body elongate, rounded at both ends, usually with a slight constric- tion at acetabular level, 1.2 to 2.2 long by 0.45 to 0.71 wide at level of anterior por- tion of acetabulum; hindbody slightly wider. Oral sucker subterminal, 0.14 to 0.17 long by 0.14 to 0.18 wide. Acetabulum 0.23 to 0.26 long by 0.22 to 0.29 wide, at or near equatorial level. Sucker ratio 1:1.4 to 1.7. Forebody 36 to 44% of body length. Pre- pharynx 0.02 to 0.05 long. Pharynx 0.06 to 0.12 long by 0.08 to 0.11 wide. Esophagus 0.06 to 0.16 long. Intestinal bifurcation usually nearer oral sucker than acetabulum. Caeca epithelial, terminating near posterior end. Testes slightly irregular, diagonal, con- tiguous or separated; left testis anterior, 0.20 to 0.28 long by 0.17 to 0.19 wide; right testis 0.19 to 0.33 by 0.17 to 0.28. Posttesticular space 9 to 13% of body length. Genital pore sinistral, close to or ventral to caecum, about midway between intestinal bifurcation and acetabulum. Cirrus sac arcu- ate, terminating posteriorly at anterior bor- der of acetabulum; containing large, folded seminal vesicle, short tubular prostatic vesi- cle, prostatic cells, and short muscular cirrus. Ovary dextral to median, lobed; usually 6 secondary lobes. Seminal receptacle in con- tact with ovary. Vitelline follicles extending from midesophageal or postpharyngeal level to posterior end, slightly confluent anteriorly. Uterus preovarian. Eggs thin-shelled, 72 to 85 by 43 to 49 microns. Excretory vesicle extending to ovarian zone; pore terminal, sphincter present. Discussion: One specimen with two eggs and one with none, both from Ocyurus chrysurus and slightly contracted, were lent by Dr. R. M. Cable and used in the above description. Of the four specimens, the smallest contains two eggs and all have well- developed organs, so the immature speci- mens are included in the description. All are similar to Hamacreadium mutabile and Cainocreadium gulella from related hosts. In fact, Siddiqi and Cable (1960:297) identified their specimens as H. mutabile, but they differ from that species in that Tulane Studies in Zoology and Botany Vol. 15 the excretory vesicle extends to the ovarian level only, rather than beyond the acetabu- lum. Yamaguti (1934:310-311) mentioned the taxonomic importance of the extent of the vesicle in H. mutabile. Because C. gulella has a median genital pore, Durio and Man- ter (1968:751) transferred that species from Hamacreadium to Cainocreadium. The present species is the only one in Hama- creadium reported to possess a short excre- tory vesicle. Of the several species for which the extent of the vesicle is unknown, H. con- fusum is distinguished from H. diacopae Nagaty and Abdel Aal, 1962, and H. lezperi Gupta, 1956, by the lobated ovary; from H. balistis Nagaty and Abdel Aal, 1962 (originally H. balistest), by the uninter- rupted vitellaria; and from H. morgani Baz, 1946, primarily by the size. The latter 1s 5.7 to 7.3 long and probably a synonym of H. mutabile. Helicometra torta Linton, 1910 Helicometra pretiosa Bravo-Hollis and Manter, 1957 (new synonym). Hosts: Epinephelus adscensionis (1 of 1)*; Epinephelus striatus (1 of 2). Site: Intestine and pyloric caeca. Discussion: Three specimens from the pyloric caeca of Epinephelus striatus have cirrus sacs that extend near or past the pos- terior border of the acetabulum, gonads close together, and lobed testes. Specimens from the midintestine of E. adscensionis have shorter extending cirrus sacs and rounded, separated testes. The excretory vesicle projects slightly past the ovary, de- pending on the contraction of the body. Because of the variation among my speci- mens, in addition to that found by Siddiqi and Cable (1960:300), I consider Helico- metra pretiosa Bravo-Hollis and Manter, 1957, a synonym of H. torta. Helicometrina execta (Linton, 1910) comb. n. Helicometra execta Linton, 1910. Helicometrina parva Manter, 1933 (new synonym ). Helicometrina trachinoti Siddiqi Cable, 1960 (new synonym). Hosts: Anisotremus virginicus (2 of 6)*; Bathygobius soporator (2 of 9); Blennius cristatus (1 of 6)*; Halichoeres bivittatus and No. 4 (10 of 11); Halichoeres pictus (2 of 2); Halichoeres radiatus (7 of 8); Labrisomus kalisherae (2 of 2)*; Trachinotus falcatus (nok 6): Site: Rectum and intestine. Discussion: 1 believe Helicometrina parva Manter, 1933, to be synonymous with Helz- cometra execta Linton, 1910, although H. execta is to be transferred to the genus Helicometrina as H. execta (Linton, 1910). When two testes are present, they are not tandem as they almost always are in species of Helicometra. There is intergradation of characters previously used to differentiate the two forms. Collections of 26, 15, and 14 mature specimens from Halichoeres radiatus, H. bivittatus and H. pictus respec- tively contain individuals with from none to five testes. Sucker ratios range from 1:1.6 to 2.1, and the acetabulum is situated any- where from the midlevel to just within the anterior 1/3 of the body. The body ranges from 0.86 to 2.07 in length, and the cirrus sac from pre- to midacetabular in posterior extent. The ovary has three to many lobes, with secondary lobing predominant. The characters originally defined as characteriz- ing H. parva are not constant, and Manter (1933a:179) apparently was correct in im- plying that H. parva may be a variation of H. execta. Eleven additional specimens from two H. bivittatus and one from H. pictus have nine testes, sucker ratios of 1:1.8 to 2.8, and further agree with my individuals of H. execta in having few ovarian lobes and the cirrus sac extending to midacetabulum. They also resemble H. trachinoti Siddiqi and Cable, 1960. Only worms with 9 testes were in the two H. bivittatus, whereas the speci- men from H. pictus was accompanied by several others with fewer testes. Five specimens from Trachinotus falcatus, four from Anisotremus virgimicus, and five from Labrisomus kalisherae were much like H. trachinoti with sucker ratios of 1:1.8 to 2.5. In specimens from all hosts, the loca- tion of the genital pore varies among posi- tions anterior, posterior, or directly ventral to the intestinal bifurcation. The cirrus sac may be on either the right or left of the acetabulum and varies in extent from the anterior margin to the midlevel of the ace- tabulum. There is at least one testis short of the predominant nine in some specimens Digenetic Trematodes of Marine Teleosts 145 from T. falcatus and A. virginicus, and L. kalisherae contained individuals with four, seven, eight, and nine testes. Two worms from Bathygobius soporator and one from Blennius cristatus have nine testes and com- pare well with others having that number. Thus, from the morphological variation and the low host specificity recorded for H. execta (Manter, 1933a:170), I also con- sider H. trachinoti synonymous with H. execta. A few specimens from Halichoeres radt- atus have poorly-formed eggs. Helicometrina mirzai Siddiqi and Cable, 1960 Hosts: Labrisomus nuchipinnis (1 of 1)*; Ogcocephalus cubifrons (1 of 2)*; Opsanus beta (3 of 6) *. Site: Intestine and rectum. Discussion: Twenty specimens from Labri- somus nuchipinnis 1.0 to 1.7 long have nine testes and sucker ratios of 1:1.9 to 2.2. The locations of the genital pore and the acetabu- lum vary as in Helicometrina execta, but most specimens differ from H. execta in having a long interruption of vitellaria at the level of the acetabulum. In a few speci- mens, the interruption is short or unilateral. Four individuals from Ogcocephalus cubt- frons compare well with those from L. nuchipinnis. I consider some specimens from Opsanus beta to be H. mirzai and designate others from that host as H. nimia because there is extensive secondary lobing of the ovary. Some confusion exists because both H. mirzai and H. nimia are present in one in- dividual of O. beta, and specimens with narrow interruptions of the vitellaria and weak secondary lobing of the ovary are present in another. Additional collections may show the two to be the same species. Helicometrina nimia Linton, 1910 Helicometrina orientalis Srivastava, 1936. Helicometrina elongata Noble and Park, SH Hosts: Lutjanus apodus (1 of 3); Lutjanus mahogont (1 of 2)*; Ocyurus chrysurus (1 of 5); Opsanus beta (2 of 6)*; Scor- paena grandicornis (1 of 2)*. Site: Pyloric caeca, intestine, and rectum. Discussion: Manter (1933a:176) reported 146 egg measurements for Helicometrina nimia from the Tortugas of 40 to 50 by 22 to 30 microns. In the present study, measurements of eggs from individuals from all hosts are 53 to 68 by 25 to 34 microns. An atypical specimen from Scorpaena grandicornis is 4.1 long and has only three testes. Deelman (1960:13) examined several hundred specimens from a single species of host from one locality, and reduced H. orientalis and H. elongata to synonymy with AH. nimaa. Manteriella crassa (Manter, 1947 ) Yamaguti, 1958 Horatrema crassum Manter, 1947. Host: Equetus acuminatus (3 of 3). Site: Intestine and bile duct. Discussion: | am following Mehra (1966: 23) and accepting the genus Manteriella. One host 9 cm long had six specimens in its bile duct in addition to 14 in its in- testine. The host for M. crassa and Pseudopecoe- loides equesi is the bold-striped island- species of Equetus common in the Florida keys. It is commonly referred to as E. pul- cher and is the species examined by Nahhas and Cable (1964). Pseudopecoeloides equesi Manter, 1947 Host: Equetus acuminatus (3 of 3). Site: Intestine. Discussion: The acetabulum may be larger than the oral sucker rather than smaller as described by Manter (1947:291). The cuti- cle, especially on and near the acetabular stalk and the terminal portion of the body, is ringed with minute striae bearing numer- ous pointed projections. Pseudopecoelus scorpaenae (Manter, 1947) comb. n. Neopecoelus scorpaenae Manter, 1947. Host: Scorpaena plumieri (2 of 4). Site: Posterior intestine and rectum. Discussion: Seven wholemounts agree fairly well with the description of Neopecoe- lus scorpaenae. They are 1.3 to 2.0 long by 0.33 to 0.51 wide, sometimes widest at the acetabular level. The forebody is 17 to 21% of body length. The oral sucker is 0.09 to 0.12 long by 0.09 to 0.10 wide and the acetabulum is 0.16 to 0.20 by 0.22 to 0.28, Tulane Studies in Zoology and Botany Vol. 15 with sucker ratios of 1:2.5 to 2.8. Eggs are partially collapsed and measure 49 to 54 by 30 to 34 microns. The genital pore is usually near the anterior level of the pharynx. The seminal vesicle passes to the left, right, or dorsal to the acetabulum before becom- ing strongly sinuous posterior to that sucker. Vitellaria extend from the middle of the acetabulum in one specimen, the posterior edge of the acetabulum in others. Temporary folds on the acetabulum of one specimen appear as but are not true papillae. The excretory vesicle extends a short distance anterior to the ovary. Serial sagittal sections of a specimen did not reveal ani. Manter (1947:294) stated that they “are inconspicuous in preserved specimens but were clearly observed in living specimens.” Nahhas and Cable (1964: 195) could not observe ani in either living or preserved material. Their absence places this species in the genus Pseudopecoelus where it differs from most species in lack- ing a lobed ovary. It is very similar, if not identical, to P. barkeri Hanson, 1950, from squirrel fishes. The posttesticular space in P. scorpaenae is slightly longer. There is apparently much variation in P. barkert. Siddiqi and Cable (1960, Figure 93) illus- trated a specimen with a sucker ratio of 1:1.6 and widely-interrupted vitellaria. Nah- has and Cable (1964:196) have specimens that differ from P. tortugae seemingly in size of eggs only. Psewdopecoelus scorpaenae differs from P. elongatus (Yamaguti, 1938) Von Wicklen, 1946, in anterior extent of vitellaria, shape and site of ovary, and size of pharynx; and from P. manteri Sogan- dares-Bernal and Hutton, 1959, in sucker ratio, anterior extent of vitellaria, and, prob- ably, position of testes. Because N. scorpaenae is the type species of Neopecoelus Manter, 1947, and I con- sider it a synonym of P. scorpaenae, Neope- coelus becomes a synonym of Pseudopecoe- lus Von Wicklen, 1946, and hence unavail- able, requiring the erection of a new genus for N. holocentri Manter, 1947. Apertile gen. n. Generic diagnosis: Body elongate, aspi- nose. Acetabulum without papillae. Pharynx large. Each caecum with anus. Testes lobed or not, tandem. Genital pore sinistral, at pharyngeal level. Cirrus sac absent, seminal No. 4 vesicle tubular, extending posterior to ace- tabulum. Ovary lobed or not. Seminal recep- tacle absent. Uterus preovarian. Metraterm present. Vitellaria from acetabular level to posterior end of body. Excretory vesicle short. Parasitic in intestines of marine fishes. Type species: Apertile holocentri (Manter, 1947) comb. n. Discussion: Apertile differs from Pseudo- pecoelus by having separate ani. Neohelico- metra Siddiqi and Cable, 1960, also has separate ani but differs from Apertile pri- marily in having a funnel-shaped oral sucker, a seminal receptacle, and filamented eggs. The significance of ani in some opecoelids is questionable. Sogandares-Bernal and Hut- ton (1959c:341) showed that in Opecoe- lides fimbriatus the ability to see ani or a uroproct depends on the contraction of the body. The name Apertile is from apertum (opened) and ze (intestine), and refers to the ani. Opegaster pritchardae sp. n. Figure 22 Host: Bathygobius soporator (1 of 9), type host. Site: Rectum. Holotype: U. S. N. M. Helm. Coll. No. 71307, paratype: No. 71369. Description (based on 11 wholemounts and 1 sectioned specimen from Biscayne Bay and Florida keys): Body elongate, 1.0 to 1.4 long by 0.32 to 0.47 wide, tapering slightly toward ends, decidedly pointed posteriorly in living specimens; a slight con- striction at acetabular level. Oral sucker 0.09 to 0.13 long by 0.09 to 0.15 wide. Acetabulum 0.18 to 0.25 by 0.19 to 0.26, with thickened anterior and posterior lips; papillae lacking. Sucker ratio 1:1.6 to 2.0. Forebody 30 to 38% of body length. Pre- pharynx short, pharynx 0.06 to 0.08 long by 0.05 to 0.08 wide. Esophagus shorter than, to over twice as long as, pharynx. Intestinal bifurcation usually closer to ace- tabulum than pharynx. Caeca uniting pos- teriorly and opening ventrally at an incon- spicuous common anus. Gonads _ postequatorial. Testes tandem, contiguous, slightly irregular; anterior testis 0.10 to 0.18 long by 0.20 to 0.26 wide; posterior testis 0.13 to 0.23 by 0.16 to 0.23. Posttesticular space 9 to 14% of body Digenetic Trematodes of Marine Teleosts 147 Figure 22. Opegaster pritchardae, holotype, ventral view. length. Genital pore inconspicuous, sinistral, slightly posterior to pharyngeal level. Cirrus sac short, slender; containing reduced pros- tatic vesicle and cirrus. Seminal vesicle ex- ternal, saccate, extending to or overlapping acetabulum anteriorly; surrounded by few prostatic cells. Ovary slightly irregular, 0.07 to 0.11 long by 0.11 to 0.20 wide, median or dextral, touching anterior testis. One ovary sinistral rather than dextral, with genital pore in that atypical specimen at midpharyngeal level. Seminal receptacle lacking. Laurer’s canal present. Vitellaria from anterior esoph- ageal level to posterior end of body, con- 148 fluent in forebody and posttesticular space. Metraterm weakly developed, extending to level of acetabulum. Eggs 54 to 64 by 31 to 39 microns. Excretory vesicle I-shaped, extending to ovarian level. Excretory pore terminal or subterminal. Discussion: Mr. Richard Heard provided eight mature specimens of Opegaster pritch- ardae which were collected from the intes- tine of Bathygobius soporator at Molasses Key, Florida. They agree in most respects with individuals from Biscayne Bay and are used in the description. The species differs from all others in the genus Opegaster Ozaki, 1928, except O. caulopsettae Manter, 1954, by lacking ace- tabular papillae. The anterior and posterior lips are too large and muscular to be papil- lae as in many opecoelids and not as muscu- lar as the lips in Labrifer Yamaguti, 1936. The caeca in O. caulopsettae are described as “seemingly uniting near posterior end of body and opening dorsally through an anus just anterior to the excretory pore near posterior end of body” (Manter, 1954:503). In my specimens the anus is ventral, the gonads are not lobed, the seminal vesicle does not extend past the acetabulum, and the posttesticular space is much_ shorter. They are most like O. gobii Manter, 1954, which has small acetabular papillae which sometimes are not evident (Manter, 1954: 503). Opegaster pritchardae is most easily distinguished from that species by the shorter posttesticular space. This species is named in honor of Mrs. Mary Hanson Pritchard, in recognition of her contributions to the field of tremato- dology. Genitocotyle cablei Nahhas and Short, 1965 Host: Hippocampus erectus (3 of 4)*. Site: Intestine. Discussion: Three specimens from H7p- pocampus erectus are identified as Genito- cotyle cablei and provide the following measurements which extend the ranges of those in the original description: length, 1.4 to 3.4: width, 0.36 to 0.66; diameter of oral sucker, 0.13 to 0.21; diameter of ace- tabulum, 0.20 to 0.29; sucker ratio, 1:1.4 to 1.5; forebody, 20 to 29% of body length; pharynx, 0.11 to 0.18 long by 0.10 to 0.16 wide; esophagus longer or shorter than Tulane Studies in Zoology and Botany 0.2 KE Figure 23. Nicolla halichoeri, holotype, dor- sal view. Figure 24. Nicolla halichoeri, dorsal view. pharynx. Diameters of anterior testis 0.21 to 0.37, posterior testis 0.19 to 0.45, ovary 0.17 to 0.25. Eggs, 46 to 48 by 19 to 37 mi- crons. The anterior extent of the excretory vesicle is not visible. Genitocotyle cablei was previously known from Ancylopsetta quadrocellata in Apala- chee Bay, Gulf of Mexico (Nahhas and Short, 1965:42). Nicolla halichoeri sp. n. Figures 23 and 24 Hosts: Halichoeres bivittatus (3 of 11); Halichoeres radiatus (3 of 8), type host. Site: Intestine. Holotype: U. S. N. M. Helm. Coll. 71308, paratype: No. 71370. Description (based on 14 wholemounts and 1 sectioned specimen): Body spindle- shaped, smooth, 0.7 to 1.2 long by 0.25 to 0.32 wide; widest at acetabular level. Oral sucker 0.07 to 0.10 long by 0.07 to 0.10 wide. Acetabulum slightly preequatorial, 0.11 to 0.17 by 0.12 to 0.19. Sucker ratio 1:1.8 to 2.1. Forebody 32 to 38% of body length. Prepharynx usually about % as long as pharynx. Pharynx 0.05 to 0.06 long by 0.04 to 0.06 wide. Esophagus 1 to 2 times as long as pharynx. Intestinal bifurcation ap- No. 4 proximately midway between acetabulum and pharynx. Caeca united slightly posterior to testes. Testes smooth to somewhat irregular, tandem, contiguous in all but one atypical specimen with ovary between them; anterior testis 0.05 to 0.10 long by 0.08 to 0.18 wide; posterior testis 0.07 to 0.12 by 0.09 to 0.15. Posttesticular space 18 to 23% of body length. Cirrus sac thin-walled, clavi- form; touching anterior border of acetabu- lum dextrally, sinistrally, or medially; con- taining large saccate seminal vesicle, short cirrus. Prostatic cells few, free in paren- chyma. Genital pore sinistral at esophageal level. Ovary subglobular, 0.03 to 0.07 long by 0.05 to 0.09 wide, either median or dextral, in contact with anterior testis. Seminal re- ceptacle usually indistinct, proximal coils of uterus full of sperm. Laurer’s canal pres- ent. Vitelline follicles well developed, from esophageal level to near posterior end of body, confluent anteriorly and _ posteriorly, one or more follicles usually between pos- terior testis and caecal loop. Uterus may extend posteriorly beyond caecal loop. Eggs partially collapsed, 49 to 62 by 26 to 34 microns. Excretory vesicle I-shaped, extending to ovarian level; pore terminal. Discussion: This species is similar to Nicolla gallica (Dollfus, 1941) Dollfus, 1959, in length of the body and in extent of the excretory vesicle. It differs from that species in having a uterus which may ex- tend posterior to the ovary. The uterus also extends posteriorly in N. wisniewskii (Slu- sarski, 1958), although the excretory vesicle in that species extends to near the posterior margin of the rear testis Nicolla halichoeri is probably the same as that Nahhas and Cable (1964:198) described and illustrated as Cottocaecum sp., in which the uterus ex- tended slightly posterior to the ovary and a seminal receptacle was lacking. Nicolla sp. Figure 25 Host: Halichoeres pictus (2 of 2). Site: Intestine. Specimen deposited: U. S. N. M. Helm. Coll. No. 71309. Discussion: Five specimens of Nicolla sp. 0.7 to 1.2 long by 0.25 to 0.32 wide Digenetic Trematodes of Marine Teleosts 149 Figure 25. Nicolla sp., dorsal view. with a sucker ratio 1:1.8 to 2.1, an esopha- gus shorter than the pharynx, the rear mar- gin of the posterior testis overlapping the united caeca, and a small prostatic vesicle appear to represent a new species. These specimens, however, may be N. halichoeri in an unsatisfactory host-parasite relation- ship since the vitellaria in three of the spect- mens are poorly formed and all the speci- mens are from a different species of wrasse than is N. halichoert. Nicolla sp. appears similar to N. extrema Travassos, Freitas, and Biihrnheim, 1965, in the appearance of the testes. It differs from N. extrema in length of the body and in the sucker ratio. FAMILY GORGODERIDAE Looss, 1901 Xystretrum solidum Linton, 1910 “Trematode allied with Phyllodistomum” of Linton, 1907. Xystretrum paptillosum Linton, 1910. Catoptroides aluterae MacCallum, 1917. Catoptroides magnum MacCallum, 1917. Macia pulchra Travassos, 1921. 150 Xystretrum pulchrum (Travassos, 1921) Manter, 1947. Hosts: Balistes capriscus (1 of 4); Mona- canthus hispidus (1 of 6)*; Sphaerotdes testudineus (5 of 6). Site: Urinary bladder. Discussion: Before Siddigi and Cable (1960:282-283) reduced Xystretrum pul- chrum to synonymy with X. solidum, the two were separated mainly by a sucker ratio of 1:1.5 or less in X. pulchrum, and a ratio of 1:1.5 or more with an abrupt widening of the hindbody in X. solidum (Manter, 1947:329-331). Ten specimens from Sphaeroides testudin- eus, with varying degrees of widening of the hindbody, have sucker ratios of 1:1.1 to 1.8; the lower ratios are from small worms and two of the three with larger ratios are poor preparations. The sucker ratio of seven specimens from Monacanthus hispidus is 1:1.3 to 1.6 and 1:1.3 in one worm from Balistes capriscus. Manter (1947) reported X. solidum from B. capriscus and X. pul- chrum from Sphaeroides splengert. FAMILY ZOOGONIDAE Odhner, 1911 Diplangus paxillus Linton, 1910 Hosts: Anisotremus virginicus (2 of 6); Haemulon carbonarium (1 of 1)*; Hae- mulon parrat (5 of 7); Haemulon sciurus Gon). Site: Rectum and posterior intestine. Discussion: There is much overlap of characteristics of Diplangus paxillus and D. parvus Manter, 1947. The length of D. parvis, 0.46 to 1.05, overlaps that of D. paxillus, 0.63 to 1.67. Also, testes may be tandem to diagonal in both species, and the size, shape, and number of coils in the seminal vesicle are not constant in either species. The posterior end of D. parvus is usually broadly rounded but may be pointed, whereas the generally pointed posterior end of D. paxillus may appear rounded. In both species, vitelline follicles vary in number and are situated between the anterior border of the acetabulum and that of the posterior testis, although they are usually restricted to the acetabular zone in D. parvus. The acetabulum in both species is protrusible and usually preequatorial. Extent of the caeca varies between the posterior edge of the acetabulum and the middle of the post- Tulane Studies in Zoology and Botany Vol. 15 testicular space, with the caeca of D. parvus usually shorter. In that species, the excre- tory vesicle reaches levels ranging from the middle of the posterior testis to the acetabulum. In D. paxillus, the vesicle ends near the ovarian level, although Manter (1947, Figure 71) illustrated it as being entirely posttesticular. The eggs of D. par- vus are 26 to 36 by 13 to 19 microns and those in D. paxillus 27 to 39 by 14 to 17 microns. Diplangus paxillus can be separated from D. parvus by having a sucker ratio of 1:1.2 to 1.7, as compared with 1:2.1 to 2.6. Also, D. paxillus occurs in the rectum or posterior intestine of the host, and D. parvus in or near the pyloric caeca. Diplangus parvus Manter, 1947 Bilecithaster ovalis Siddigi and Cable, 1960 (new synonym). Hosts: Anisotremus virginicus (3 of 6)*; Haemulon carbonarium (1 of 1); Haemu- lon parrai (2 of 7)*; Haemulon plumieri (1 of 5); Haemulon sciurus (5 of 6). Site: Pyloric caeca and anterior intestine. Discussion: 1 believe Bilecithaster ovalts Siddiqi and Cable, 1960, is a synonym of Diplangus parvus, and therefore Bilecithas- ter Siddiqi and Cable, 1960, a synonym of Diplangus Linton, 1910. Bilecithaster ovalis was apparently placed in the family Hemiurt- dae because the excretory arms seemed to unite at a level dorsal to the oral sucker. With the exception of that feature, B. ovalis fits the description of D. parvus, which 1s found in the same site, within the same host, from Tortugas, Florida, and Jamaica. Diphtherostomum anisotremt Nahhas and Cable, 1964 Hosts: Anisotremus virginicus (4 of 6); Haemulon plumieri (3 of 5)*; Haemulon scturus (5 of 6). Site: Rectum. Discussion: Nine specimens from the type host, Anisotremus virginicus, measure 0.53 to 1.08, have sucker ratios of 1:1.5 to 2.1, with the length of the esophagus three to eight times that of the pharynx, and the cirrus sac three to four times as long as wide. Specimens from Haemulon sciurus and H. plumieri have the same features except that some have an even larger esophagus-to- pharynx length ratio. No. 4 In the original description, Diphthero- stomum anisotremi was differentiated from D. americanum by having an esophagus at least three times as long as the pharynx rather than being about the same length, and a citrus sac about four times as long as wide instead of the width being '% to *4 the length. Diphtherostomum americanum Manter, 1947 Hosts: Archosargus rhomboidalis (2 of 5)*; Eupomacentrus leucostictus (1 of 6)*; Lagodon rhomboides (1 of 5). Site: Rectum and posterior intestine. Discussion: Sogandares-Bernal and Hut- ton (1959c:342-343) identified trematodes from Gobiosoma robustum, Lagodon rhom- boides, and Opsanus beta as D. americanum. Their illustration shows a short esophagus and a long, thin cirrus sac as do my speci- mens. In fixed ones, the length of the esoph- agus is 1.0 to 2.1 times the length of the pharynx; in a living specimen from Archo- sargus rhomboidalis it varied from slightly longer to 3.5 times longer than the pharynx. An egg released from that worm was 54 by 26 microns compared to 30 to 45 by 13 to 17 microns after the specimen was mounted. All my specimens identified as D. antisotremt or D. americanum have a completely spinose cuticle, an observable seminal receptacle, and variations in the positions of the gonads as reported by So- gandares-Bernal and Hutton (1959c:343). Apparently the only difference between the two species is the length of the variable esophagus. Another closely related, if not identical, species is D. macrosaccum Mont- gomery, 1957, which is completely covered with spines, has a short esophagus, and has a citrus sac about nine times longer than it is wide. This trematode is found in Neo- clinus uninotatus, a blenny from California. Diphtherostomum albulae sp. n. Figure 26 Host: Albula vulpes (4 of 7), type host. Site: Intestine and pyloric caeca. Holotype: U. S. N. M. Helm. Coll. No. 71310, paratype: No. 71371. Description (based on 13 specimens, 4 with newly-formed eggs): Body small, plump, tapered at ends, 0.43 to 0.73 long by 0.19 to 0.34 wide. Cuticle completely Digenetic Trematodes of Marine Teleosts x 7 y SSeS Figure 26. Diphtherostomum albulae, holo- type, dorsal view. spinose. Oral sucker terminal or slightly subterminal, 0.08 to 0.10 long by 0.10 to 0.13 wide. Acetabulum weakly developed, without lip-like thickening, 0.08 to 0.11 by O08" to- OMI Sucker ratio: 1707/5 ton 0:92: Forebody 38 to 57% of body length. Pre- pharynx very short. Pharynx 0.03 to 0.04 long by 0.03 to 0.04 wide. Esophagus 2 to 3 times longer than pharynx. Intestinal bi- furcation approximately midway between suckers. Caeca short, ending just anterior to or within acetabular zone. Testes roughly symmetrical, at or slightly anterior to level of acetabulum; left testis 0.08 to 0.12 long by 0.05 to 0.09 wide; right testis 0.09 to 0.13 by 0.05 to 0.10. 152 Genital pore at or near sinistral margin of body, at level of or slightly anterior to in- testinal bifurcation. Cirrus sac thick-walled, arcuate to straight, 0.14 to 0.21 long by 0.06 to 0.09 wide, usually ending at an- terior border of acetabulum; containing bi- partite seminal vesicle, wide prostatic vesicle, prostatic cells, and short, unarmed cirrus. Ovary smooth, 0.07 to 0.11 long by 0.07 to 0.11 wide, dorsal to acetabulum, at or slightly posterior to level of testes, usually close to right testis. Seminal receptacle not observed. Uterus filling most of body pos- terior to testes except extreme posterior re- gion. Metraterm weak. Vitellaria 2 small masses near posterior portion of ovary. Eggs extremely thin-shelled, 25 to 42 by 16 to 31 microns, when not compressed. Excretory vesicle short, saccate. Excretory pore at or near posterior end. Discussion: This species differs from others in the genus Diphtherostomum by lacking thickened acetabular margins and lip-like folds of the body overlapping the acetabulum, and by having an oral sucker larger than the acetabulum. In D. mzicroace- tabulum Schulman-Albowa, 1952, the suck- ers are about equal in size, the esophagus is shorter, the cirrus sac has a thinner wall than in D. albulae, and an obvious seminal receptacle is present. Steganoderma nitens (Linton, 1898) Manter, 1947 Distomum nitens Linton, 1898. Lecithostaphylus nitens (Linton, 1898) Linton, 1940. Steganoderma elongatum Manter, 1947. Host: Tylosurus crocodilus (1 of 7)*. Site: Intestine Discussion: Because of overlapping char- acters, Nahhas and Cable (1964:199) re- duced S. elongatum to synonymy with S. nitens. 1 find that the position of the ovary varies from midway between the testes and acetabulum to near that sucker, and the genital pore is sometimes almost marginal. In contrast to Nahhas and Cable's specimens, only three of my 48 have vitellaria that do not lie posterior to the testes. In those, the follicles, on one side only, end beyond the midlevel of the testes. The eggs measure 28 to 36 by 16 to 20 microns. The host, Tyloswrus crocodilus (Peron and Lesueur), was identified by Mr. F. H. Tulane Studies in Zoology and Botany Vol is Berry. This fish is separate from, but some- times identified as, T. acus. FAMILY MONORCHIIDAE Odhner, ae Genolopa ampullacea Linton, 1910 Genolopa longicaudata Siddiqi and Cable, 1960. Hosts: Anisotremus virginicus (4 of 6)*; Haemulon aurolineatum (5S of 7)*; Hae- mulon flavolineatum (2 of 2); Haemulon parrat (7 of 7); Haemulon plumieri (3 of 5); Haemulon sciurus (6 of 6). Site: Pyloric caeca and anterior intestine. Discussion: The considerable range in characteristics of this species has been noted by Manter (1942:351), Nahhas and Cable (1964:201), and others. The eggs are 13 to 20 by 9 to 13 microns and usually smaller than those described by Linton (1910:78) and Manter (1942:351). The atrial spines are up to 49 microns long, the ovary is subglobular to distinctly trilobed, and the oral sucker may be pyriform in shape. Lasiotocus longovatus (Hopkins, 1941) Thomas, 1959 Genolopa longovatum Hopkins, 1941. Proctotrema longovatum (Hopkins, 1941 ) Manter, 1942. Hosts: Anisotremus virginicus (2 of 6)*; Haemulon aurolineatum (1 of 7); Hae- mulon parrai (1 of 7)*; Haemulon sciurus (2 of 6); Orthopristis chrysop- terus (3 of 4). Site: Pyloric caeca and intestine. Discussion: There is considerable varia- tion in this species. The caeca may end at the testicular level rather than somewhat past it, the testis may be well removed from the acetabulum, and the cirrus sac of- ten extends beyond the acetabulum. The body is characteristically urn-shaped as orig- inally described but exceptions occur (Nah- has and Cable, 1964:201). Whether various species of monorchiids should be in the genus Proctotrema or Lasto- tocus has been discussed by Manter and Pritchard (1961:483-484) and Nahhas and Cable (1964:200). I am following Bartoli and Prévot (1966:406) who transferred to the genus Lasiotocus all species previously referred to as Proctotrema except P. bacullio- vatum. No. 4 Lasiotocus truncatus (Linton, 1910) Thomas, 1959 Genolopa truncatum Linton, 1910. Proctotrema truncatum (Linton, Manter, 1940. Hosts: Haemulon flavolineatum (2 of 2); Haemulon plumiert (4 of 5); Haemulon sciurus (5 of 6). Site: Pyloric caeca. 1910) Lastotocus longicaecum (Manter, 1940) Yamaguti, 1954 Proctotrema longicaecum Manter, 1940. Host: Anisotremus virginicus (3 of 6). Site: Rectum. Lasiotocus mugilis sp. n. Figures 27, 28, and 29 Host: Mugil cephalus (1 of 3), type host. Site: Intestine. Holotype: U. S. N. M. Helm. Coll. No. 71311, paratype: No. 71372. Description (based on 5 wholemounts ) : Body delicate, 0.8 to 1.3 long by 0.24 to 0.38 wide; forebody narrow; posterior end bluntly rounded. Cuticle spined. Eyespots lacking. Oral sucker weak, usually folded, 0.08 to 0.15 wide. Acetabulum weakly de- veloped, 0.10 to 0.13 long by 0.09 to 0.12 wide. Sucker ratio 1:0.8 to 1.1. Forebody 29 to 34% of body length. Prepharynx shorter or longer than pharynx. Pharynx 0.05 to 0.06 long by 0.05 to 0.06 wide. Esophagus 0.07 to 0.10 long. Intestinal bi- furcation usually nearer acetabulum than oral sucker. Caeca terminating near end of body. Testis slightly irregular, submedian, 0.10 to 0.25 long by 0.08 to 0.12 wide. Post- testicular space 20 to 34% of body length. Cirrus sac large, arcuate, 0.22 to 0.30 long by 0.07 to 0.10 wide, extending near or into gonadal zone; containing ovoid seminal vesi- cle, well-developed prostatic cells, large sac- cate cells opening into pars prostatica, long cirrus with thorn-shaped spines 8 to 19 microns long. Genital atrium unspined. Genital pore median or submedian, anterior to acetabulum. Ovary 0.10 to 0.20 long by 0.07 to 0.14 wide, lobed or unlobed; if lobed, with 3 or 4 lobes; slightly dextral to overlapping testis. Vitelline follicles in compact groups, Digenetic Trematodes of Marine Teleosts Figure 27. Lasiotocus mugilis, holotype, dor- sal view. Figure 28. Lasiotocus mugilis, ovary from different specimen. Figure 29. Lasioto- cus mugilis, anterior end. at gonadal level, intercaecal or overlapping caeca. Terminal organ partially collapsed, not over % as long as cirrus sac; anterior portion with spines 16 to 21 microns long, posterior portion unspined. Uterus not ex- tending past midposttesticular level, enter- ing terminal organ near junction of spiny and unspined portions. Eggs 11 to 17 by 9 to 10 microns. Excretory vesicle I-shaped, epithelial, ex- tending near or into testicular level; pore terminal. Discussion: Several species of Lasiotocus have caeca extending to near the end of the body and vitellaria at the gonadal level. This species can be differentiated from L. latus (Manter, 1942) by body shape and arrangement of gonads; from L. beawforti (Hopkins, 1941) by length of excretory vesicle, number of posttesticular uterine coils, and sucker ratio; from L. longicaecum Manter, 1940, by body shape and character of oral sucker; from L. chaetodipteri Thomas, 1959, by the body shape and amount of posttesticular space; from L. Jlongovatus (Hopkins, 1941) and L. pritchardae Nahhas and Cable, 1964, by sucker ratio and size of eggs; from L. malast (Nagaty, 1948) by 154 Tulane Studies in Zoology and Botany Vol. 15 the site of the vitellaria and length of esoph- agus and eggs; from L. himezi Yamaguti, 1951, L. odhneri (Srivastava, 1939), and L. cacuminata (Nicoll, 1915) by size of body and eggs and number of posttesticular uterine coils. Lasiotocus haemuli sp. n. Figure 30 Hosts: Haemulon plumieri (2 of 5), type host; Haemulon sciurus (2 of 6). Site: Intestine and pyloric caeca. Holotype: U. S. N. M. Helm. Coll. No. 71312, paratype: No. 71373. Description (based on 18 specimens) : Body elongate, 0.49 to 0.93 long by 0.16 to 0.22 wide, usually widest at midbody; posterior end pointed or rounded; usually a constriction immediately posterior to level of oral sucker. Cuticle completely spinose. Eyespots absent. Oral sucker terminal, fun- nel-shaped, 0.11 to 0.17 long by 0.10 to 0.15 wide. Acetabulum weakly developed, 0.05 to 0.06 long by 0.05 to 0.07 wide. Sucker ratio 1:0.4 to 0.6. Forebody 29 to 48% of body length. Prepharynx shorter than pharynx. Esophagus usually shorter than pharynx. Intestinal bifurcation a short distance anterior to acetabulum. Caeca ter- minating beyond middle of posttesticular space. Testis slightly irregular, 0.07 to 0.13 long by 0.05 to 0.10 wide. Posttesticular space 17 to 35% of body length. Cirrus sac arcuate, 0.11 to 0.18 long by 0.03 to 0.05 wide, extending around left or dorsal of acetabulum to or near ovarian level; con- taining large ovoid seminal vesicle, pros- tatic cells, saccate cells opening into pars prostatica, cirrus with thorn-shaped spines 4 to 5 microns long. Genital atrium short, unspined. Genital pore anterior to acetabu- lum, sometimes ventral to intestinal bifur- cation. Ovary varying from slightly irregular to triangular, never dextral, and occasionally sinistral to median line through testis, 0.05 to 0.10 long by 0.05 to 0.09 wide. Seminal receptacle absent. Vitelline follicles in lat- eral groups, 8 to 10 on each side, between levels of acetabulum and ovary. Terminal organ 0.05 to 0.10 long by 0.03 to 0.05 wide, 30 to 90% length of the cirrus sac; proximal vesicle unspined; anterior portion with spines 5 microns long. Uterus may Figure 30. Lasiotocus haemuli, holotype, ventral view. Or may not extend past caecal termination, entering terminal organ at junction of spiny and unspined portions. Eggs 17 to 24 by 11 to 13 microns. No. 4 Excretory vesicle short, usually extending to level of caecal termination; pore terminal. Discussion: Lastotocus haemuli is found with L. truncatus and resembles that species but differs from it by having a larger sucker ratio, a non-dextrally located ovary, and a larger pharynx compared with the width of the neck. Lastotocus haemuli differs from L. longovatus in the arrangement of the gonads and size and shape of eggs. Lasio- tocus beauforti and L. longicaecum differ from L. haemuli in that they are longer; also, L. beawforti has a longer genital atrium and L. longicaecum has longer caeca and a different position of the cirrus sac. Lastoto- cus pritchardae (Nahhas and Cable, 1964) is a larger species with a distinctly lobed, dextrally located ovary. Lasiotocus albulae sp. n. Figures 31 and 32 Host: Albula vulpes (6 of 7), type host. Site: Intestine and pyloric caeca. Holotype: U. S. N. M. Helm. Coll. No. 71313, paratype: No. 71374. Description (based on 15 wholemounts and 3 sectioned specimens): Body elongate, 0.7 to 1.2 long by 0.17 to 0.35 wide, slightly wider at acetabular level; neck region attenu- ated. An immature individual 1.1 long. Spines on entire cuticle, more numerous on neck region. Eyespots or dispersed pigment granules in pharyngeal to midesophageal region. Oral sucker sometimes _ slightly tapered posteriorly but not funnel-shaped, 0.06 to 0.09 long by 0.08 to 0.10 wide. Acetabulum weakly developed, 0.07 to 0.12 by 0.08 to 0.12. Sucker ratio 1:1.0 to 1.3. Forebody 32 to 52% of body length. Pre- pharynx contracted, or elongated up to 0.07 in length. Pharynx 0.05 to 0.06 long by 0.04 to 0.05 wide; widest just postequa- torial. Esophagus 2 to 5 times length of pharynx. Intestinal bifurcation anterior to acetabulum. Caeca terminating from level of posterior border of testis to near the end of the body. Testis elongate, 0.10 to 0.17 long by 0.06 to 0.13 wide, in posterior % to 4 of body. Posttesticular space 8 to 21% of body length. Cirrus sac slightly to strongly arcu- ate, 0.16 to 0.28 long; from midway be- tween acetabulum and ovary and ovary; pass- ing at a level dorsal to acetabulum; contain- ing unipartite seminal vesicle, large sac- Digenetic Trematodes of Marine Teleosts 55) aN *\ £4} Sey {jh | / \ 2 Figure 31. Lasiotocus albulae, holotype, ven- tral view. Figure 32. Lasiotocus albulae, with- drawn oral sucker. cate cells opening into pars prostatica, con- spicuous prostatic cells, muscular cirrus with wide-based spines 4 to 5 microns long. Gent- tal pore posterior to caecal bifurcation, median or slightly sinistral. Large, unspined genital atrium. Ovary subglobular to subtriangular, 0.04 to 0.09 long by 0.05 to 0.10 wide, median to slightly dextral, overlapping anterior edge of testis. Terminal organ well developed, about *4 as long as cirrus sac; anterior por- tion with slender spines up to 8 microns long; posterior portion unspined. Vitellaria 8 to 10 large or numerous small follicles on each side, from level of middle or posterior end of cirrus sac to near or beyond pos- terior border of testis. Many sperm in proxt- mal loops of uterus. Uterus filling body beyond middle or posterior end of cirrus sac; distal portion muscular, entering termt- 156 nal organ between middle of spiny portion and its junction with unspined portion. Eggs 13 to 17 by 8 to 11 microns. Excretory vesicle short, occasionally over- lapping testis posteriorly; pore terminal. Discussion: The body is delicate and may be extended greatly. The anterior portion of the worm may be retracted within the body (Figure 32). Lasiotocus albulae resembles L. delicatus Manter and Pritchard, 1961, in having a long esophagus and an oral sucker that is not funnel-shaped. It is, however, smaller than that species, has extensive vitellaria, and is most easily distinguished from L. delicatus by the presence of eyespot pig- ment. Postmonorchis orthopristis Hopkins, 1941 Pristisomum orthopristis (Hopkins, 1941) Yamaguti, 1958. Hosts: Anisotremus virginicus (1 of 6)*; Haemulon aurolineatum (4 of 7)*; Hae- mulon parrai (2 of 7)*; Haemulon plumieri (3 of 5); Haemulon sciurus (3 of 6). Site: Intestine. Discussion: Manter and Pritchard (1961: 483-484) discussed the invalidity of the generic name Pristisomum. Variability exists in some characters of my specimens. The vitellaria may be at the level of the testis in addition to being an- terior to this zone. The cirrus sac may be almost % instead of only % as long as the body. Hopkins (1941:396-397) did not observe the uterus posterior to the testis in any of 106 specimens, but in a few of mine it fills all the available posttesticular space. Diplomonorchis leiostomi Hopkins, 1941 Diplomonorchis micropogoni Nahhas and Cable, 1964 (new synonym). Hosts: Archosargus rhomboidalis (3 of 5); Lagodon rhomboides (2 of 5); Orthopris- tis chrysopterus (3 of 4). Site: Pyloric caeca and intestine. Discussion: Considerable variation exists in this species. The testes and vitellaria may be from equatorial in position to within the posterior half of the body. The uterus may have loops at the bifurcal level on one or both sides, the caeca may or may not ex- tend beyond the testes, the posterior portion Tulane Studies in Zoology and Botany Vol. 15 of the terminal organ may have a few spines or none, and the seminal vesicle may be spherical, oval, or tear-shaped. In most of my specimens, the testes are located toward the posterior end of the worm, and anterior loops of the uterus occur on both sides of the body. Nahhas and Cable (1964:206- 207) distinguished Diplomonorchis maicro- pogoni from D. letostomi by short caeca terminating near the posterior margin of the testes and by the extent of the uterus. Since my specimens show intergradation between the two species and the two are from the same or related hosts (also see Corkum, 1966:46-47; Nahhas and Powell, 1965:17; and Sogandares-Bernal and Hut- ton, 1959b:62), I am reducing D. micro- pogoni Nahhas and Cable, 1964, to synonymy with D., lezostomt. Diplomonorchis sphaerovarium Nahhas and Cable, 1964 Host: Ophichthus gomesi (1 of 1)*. Site: Intestine. Specimen deposited: U.S. N. M. Helm. Coll. No. 71314. Discussion: My specimens differ some- what from the published description. In some, the gonads are irregularly shaped and in others the ovary has three to four in- distinct lobes, suggesting that the species is not as atypical as Nahhas and Cable (1964:209) suspected. The large gland cells in the forebody may extend into the hindbody, and the esophagus is not neces- sarily shorter than the pharynx. Some ma- ture worms are 0.71 long, although imma- ture worms are often larger. Dr. R. M. Cable lent a slide with 12 immature speci- mens which revealed spines in the posterior portion of the prepharynx, as in my ma- terial. The seminal vesicle in some of my specimens appears indistinctly bipartite. In describing D. sphaerovarium, Nabhas and Cable (1964:209) broadened the concept of the genus Diplomonorchis so that it dif- fers from Diplomonorcheides Thomas, 1959, only in possessing species with a unipartite rather than bipartite seminal vesicle. Be- cause Thomas (1959, Figures 9 and 10) figured the seminal vesicle of Dzplomon- orcheides magnacetabulum as being clearly unipartite, I propose Diplomonorcheides as a synonym of Diplomonorchis, with Drplo- No. 4 monorchis magnacetabulum (Thomas, 1959 ) as a new combination. Hurleytrema eucinostomt Manter, 1942 Pseudohurleytrema eucinostomi (Manter, 1942) Yamaguti, 1954. Host: Eucinostomus gula (3 of 3). Site: Rectum. Discusston: Eight mature specimens dif- fer slightly from the original description. The eyespots are more anteriorly located than originally described and the caeca may extend almost to the midtesticular level. The ovary is usually distinct and dextral. The testis is elongate, with the width 40 to 80% the length. The specimens are 0.52 to 1.24 long, with an immature specimen 0.50 long. Sucker ratios are 1:0.8 to 1.0, usually 1:0.98. The eggs measure 18 to 27 by 14 to 17 microns. Nahhas and Cable (1964:204) gave measurements of specimens from Ja- maica, Curacao, and Puerto Rico. Hurleytrema shorti (Nahhas and Powell, 1965) comb. n. Pseudohurleytrema shorti Nahhas Powell, 1965. Pseudohurleytrema ottoi Travassos, Freitas, and Buhrnheim, 1965 (new synonym). and Host: Selene vomer (2 of 2). Site: Intestine, pyloric caeca, and stomach. Discussion: My specimens are extremely gravid. The smallest specimen is 0.46 long. Measurements of the organs are slightly less than those of the original description. The acetabulum is weakly developed and the sucker ratio is 1:1.0 to 1.5. Eggs measure 24 to 30 by 13 to 16 microns, with fila- ments up to at least 3.5 times the length of the eggshell. The caeca may extend posterior to the vitellaria. I agree with Manter and Pritchard (1961: 487) that Pseudohurleytrema Yamaguti, 1954, is a synonym of Hurleytrema Srivas- tava, 1939, and that Hurleytrema be tempo- rarily distinguished from Hwrleytrematoides Yamaguti, 1954, by including species with a unipartite rather than a bipartite seminal vesicle. The genus Hurleytrematoides accordingly includes H. chaetodoni (Manter, 1942), H. coronatum Manter and Pritchard, 1961, and H. curacaensis Nahhas and Cable, 1964; and the genus Huwrleytrema includes H. ovo- Digenetic Trematodes of Marine Teleosts y/ ATR, Prix a Lenn a, Figure 33. Hurleytrema pyriforme, holotype, dorsal view. caudatum Srivastava, 1939, H. eucinostomi Manter, 1942, H. /ongitestis Bravo-Hollis, 1956, H. trachinoti Thomas, 1959, H. mala- bonensis (Velasquez, 1961) comb. n., H. shorti (Nahhas and Powell, 1965) comb. n., and the new species described next. Pseudohurleytrema ottot Travassos, Freitas, and Buhrnheim, 1965, from Selene vomer in Brazil and H. shorts are the same species. The description of H. shorti appeared earlier in 1965 than did that of P. oftot. This synonymy is also recognized by Freitas, Tra- vassos, and Kohn (Freitas, 1968: personal communication ). Hurleytrema pyriforme sp. n. Figure 33 Host: Trachinotus falcatus (1 of 6), type host. Site: Pyloric caeca. 158 Holotype: U. S. N. M. Helm. Coll. No. 71315, paratype: No. 71375. Description (based on 5 apparently re- cently matured specimens): Body pyriform, 0.52 to 0.63 long by 0.28 to 0.33 wide; widest at gonadal level. Cuticle completely spinose. Eyespots at pharyngeal-esophageal level. Oral sucker 0.05 to 0.07 long by 0.07 to 0.08 wide. Acetabulum weakly developed, 0.04 to 0.05 by 0.05 to 0.06. Sucker ratio 1:0.8; 1:0.6 to 0.7 in 6 additional specimens without eggs. Forebody 40 to 48% of body length. Prepharynx 0.01 long. Pharynx 0.04 long by 0.03 to 0.04 wide. Esophagus 0.05 to 0.07 long. Intestinal bifurcation about midway between suckers. Caeca extending almost to or slightly past testis. Testis single, 0.07 to 0.10 long by 0.15 to 0.18 wide, sinistral, slightly irregular, 0.08 to 0.12 from acetabulum. Cirrus sac large, arcuate, over 0.20 long, extending to testicu- lar level; containing oval seminal vesicle, saccate cells opening into pars prostatica, numerous prostatic cells, cirrus with spines 6 to 11 microns long. Cirrus ’% or more the length of cirrus sac. Genital atrium unarmed. Genital pore immediately anterior to acetabulum, median to slightly sinistral. Ovary small, longer than wide, swollen at connection with oviduct, dextral and slightly anterior to testis. Seminal recep- tacle absent; proximal folds of uterus con- taining sperm. Uterus extending to a level posterior to testis, entering terminal organ near junction of its spiny and unarmed por- tions. Terminal organ over “4 as long as cirrus sac; distal portion tubular, heavily spined, anterior portion and sometimes al- most all of proximal chamber with spines 5 to 12 microns long with sharp points and wide bases, as spines of cirrus. Vitelline follicles in 2 lateral clusters, between or overlapping acetabulum and testis; may or may not overlap caeca. Vitelline ducts large. Newly-formed eggs 14 to 21 by 11 to 12 microns with filament 2 to 4 times the length of the eggshell. Excretory vesicle saccate, not reaching testis; collecting ducts sometimes swollen, extending, at least, to vitelline region. Ex- cretory pore terminal to slightly ventral. Discussion: Hurleytrema pyriforme tre- sembles H. trachinoti Thomas, 1959, which is found in the same genus of host. It dif- fers from H. trachinoti in that the oral Tulane Studies in Zoology and Botany Vol. 15 sucker is more spherical than funnel-shaped; the oral sucker, acetabulum, and pharynx are half as large; the excretory vesicle does not extend to the testis; and the distance between the acetabulum and testis is greater. Also, the cirrus sac in H. pyriforme is larger and contains a unipartite seminal vesicle, the spines in the terminal organ are situ- ated farther posteriorly, and the vitellaria are more compact. Nahhas and Powell (1965:19) erected the genus Parahurleytrema with P. trachinoti as type species, characterized by a bipartite seminal vesicle and the uterus entering the terminal organ near the junction of the spiny anterior and unarmed posterior por- tions. Since the present species, with a uni- partite seminal vesicle, shows such a close relationship to H. trachinoti and the illus- tration of the seminal vesicle of H. tracht- noti by Thomas (1959:106) reveals a rather indistinct bipartite condition, I place Para- hurleytrema as a synonym of Hurleytrema, P. coronatum (Manter and Pritchard, 1961) is therefore Hurleytrematoides coronatum, as originally described. FAMILY CRYPTOGONIMIDAE Ciurea, 1933 Siphodera vinaledwardsii (Linton, 1901) Linton, 1910 Monostomum vinaledwardstu Linton, 1901. Hosts: Lutjanus mahogoni (2 of 2)*; Lut- janus synagris (5S of 7); Opsanus beta (20tG)). Site: Intestine and pyloric caeca. Discussion: The size of eggs in this spe- cies is not well documented. It is 17 to 29 by 9 to 12 microns in my specimens. Metadena globosa (Linton, 1910) Manter, 1947 Stegopa globosa Linton, 1910. Hosts: Lutjanus griseus (1 of 3); Lutjanus mahogoni (1 of 2)*; Lutjanus synagris (3 of 7)*; Ocyurus chrysurus (4 of 5). Site: Intestine. Discussion: Manter (1947:333-335) and Hanson (1950:84) used several characters to distinguish Metadena globosa from M. adglobosa. After studying 27 specimens of M. globosa and 12 specimens of M. ad- globosa, 1 believe the two may be distin- guished best by uterine coils extending an- No. 4 terior to the acetabulum in M. globosa but not in M. adglobosa. The eggs are smaller in my specimens of M. globosa: 10 to 13 by 7 to 10 compared to 16 to 23 by 8 to 11 microns, although Manter (1947:334) re- ported 14 to 15 by 8 to 9 microns for M. globosa, and Hanson (1950:84) 10 to 27 by 7 to 12 microns for eggs of M. adglobosa. The body is usually more elongate in M. adglobosa. The diameter of the oral sucker in M. globosa is not always more than half the body width. The percentage is 34 to 55% compared to 35 to 45% in M. ad- globosa. The sucker ratios are 1:0.20 to 0.51 compared to 1:0.27 to 0.32 in M. ad- globosa. The seminal vesicle may also oc- casionally have coils anterior to the ace- tabulum in M. globosa; and even though the pharynx is relatively larger in M. glo- bosa, it is approximately the size of the acetabulum in both species. The caeca may extend to the posterior end in both species. Several specimens of M. globosa have small, narrow, nondescript, cuticular appendage- like structures on the lateral and posterior margins of the body which contain cyto- plasm. These processes were present on the trematodes when removed from the host and are not artifacts. Metadena adglobosa Manter, 1947 Hosts: Lutjanus apodus (2 of 3); Lutjanus griseus (2 of 3). Site: Pyloric caeca. Metadena sp. Host: Lutjanus griseus (1 of 3). Site: Pyloric caeca. Discussion: Specimens from a_ single _ Lutjanus griseus are similar to Metadena adglobosa, except they have a large, heavily- spined ventrogenital pouch. I consider these to be a new species which is described by Schroeder (in press) from the same host near Lower Matecumbe Key, Florida. Paracryptogonimus americanus Manter, 1940 Sid- Paracryptogonimus neoamericanus digi and Cable, 1960. Host: Ocyurus chrysurus (4 of 5). Site: Intestine and pyloric caeca. Discussion: Siddigi and Cable (1960: 280-281) distinguished Paracryptogonimus neoamericanus from P. americanus by its smaller size, 0.64 to 0.88, compared to 2.31 Digenetic Trematodes of Marine Teleosts 159 to 2.38, excluding a single specimen of 1.0; fewer oral spines, 46 to 51, compared to 52 to 57; terminal oral sucker; and smaller eggs, 16 to 18 by 10 to 15, compared to 20 to 25 by 9 to 10 microns. Dr. Robert Schroeder lent 19 specimens of P. amert- canus from Lutjanus griseus and two from Ocyurus chrysurus, collected near Lower Matecumbe Key, Florida, to compare with my nine from O. chrysurus. Mature speci- mens from L. griseus are from 0.60 to 2.32 long, have 43 to 49 oral spines, possess primarily terminal oral suckers, and have eggs 16 to 20 by 9 to 11 microns. My specimens are 0.84 to 1.17 long, have 46 to 54 oral spines, have oral suckers appear- ing either terminally or subterminally, and have eggs 12 to 23 by 9 to 11 microns, usually 16 to 19 by 9 to 10 microns. Dr. R. M. Cable lent three specimens of P. neoamertcanus from O. chrysurus which are 0.9 to 1.1 long. There are 45 to at least 56 oral spines, and the few noncollapsed well-developed eggs are 16 to 20 by 9 to 10 microns. This overlap between charac- ters of P. neoamericanus and P. americanus leads me to place the former as a synonym of the latter. Even though not indicated in the description of P. neoamericanus, the cuticle is very thick; in fact, over 0.06 in one specimen from L. griseus. The cuticle in Dr. Cable’s and my specimens is 12 to 25 microns at the thickest portion, in the forebody. The sucker ratios in the individ- uals from L. griseus are 1:0.5 to 0.8 and encompass the ratios of specimens from O. chrysurus. Pearse (1949:36) has previously re- ported P. americanus in Opsanus tau from Beaufort, North Carolina. FAMILY ACANTHOCOLPIDAE Luhe, 1909 Stephanostomum casum (Linton, 1910) McFarlane, 1934 Stephanochasmus casus Linton, 1910. Lechradena edentula Linton, 1910. Hosts: Lutjanus griseus (2 of 3); Lutjanus synagris (2 of 7); Ocyurus chrysurus (GikoE 5): Site: Rectum. Discussion: My specimens agree with the typical example of Stephanostomum casum with the exception of the sucker ratio. That 160 is, they have 36 oral spines; eggs are 61 to 78 microns long; and metraterm and cirrus are both long and spined, with the metraterm a little shorter than the cirrus sac. The sucker ratio is 1:1.3 to 1:2.2, rather than 1:2.4 to 2.7 (Manter, 1967:personal communication), depending on the amount of expansion of the oral sucker. Manter (1947:304-305) questioned the identifica- tions of S. caswm from the Pacific Ocean. Stephanostomum tenue (Linton, 1898) Martin, 1938 Distomum tenue Linton, 1898. Distomum tenue tenuissime Linton, 1898. Hosts: Lutjanus apodus (2 of 3)*; Lutjanus mahogoni (1 of 2)*; Trachinotus falca- tus (2 of 6) *. Site: Near or in rectum. Discussion: Manter and Van Cleave (1951:328) noted that Linton (1940) might have included more than one species in his description of Stephanostomum tenue. The specimens I designate as S. tenwe have a sucker ratio of 1:1.5, which is apparently typical for the species. The ratio in the specimen from Lutjanus mahogont is 1:1.2, but the oral sucker is expanded. The eggs, however, are not longer than about 72 mi- crons, whereas Linton (1940:57; 1889:536) reported 0.084 by 0.04 and 0.088 by 0.044 for eggs from specimens of Roccus saxa- tilus (=R. lineatus), the type host. The vitellaria in the two specimens from L. apodus extend to the acetabular level as in Figure 7, Plate LII (Linton, 1898) and vitellaria in the two specimens from Tracht- notus falcatus and one from L. mahogonti do not extend to the acetabulum, as Linton noted later (1940:57) for another worm from R. saxatilus. Three specimens have 42 oral spines and a fourth 40 or 42. Anderson (1965:71) reported S. tenue from the kidney of Pomatomus saltatrix from Sandy Hook, New Jersey, to Marathon, Florida but did not state whether it was a metacercarial or adult stage. Stephanostomum sentum (Linton, 1910) Manter, 1947 Stephanochasmus sentus Linton, 1910. Stephanostomum medtiovitellarum Pérez Vigueras, 1955. Stephanostomum lopezneyrai Pérez Vigue- tas, 1955: Tulane Studies in Zoology and Botany Vol. als Hosts: Calamus bajonado (1 of 1); Haemu- lon carbonarium (1 of 1)*; Ogcocephalus cubtfrons (1 of 2)*. Site: Rectum. Discussion: Three specimens agree with the diagnosis of Stephanostomum sentum by Manter (1947:306-307), but they differ from that given by Sogandares-Bernal (1959:89) in having sucker ratios of 1:1.6, 1ee7, and) led8* rathersthane lL: 1kOmtoml=5: The specimen from Ogcocephalus cubifrons is 2.4 long with eggs 74 to 78 by 39 to 41 microns, which is still larger than S. minu- tum (Looss, 1901) at 1.2 to 1.9 with eggs 47 by 36 microns. Vitellaria are poorly de- veloped anteriorly. In the specimen from Calamus bajonado, the anterior testis is not formed, and the posterior one is elongated. The excretory vesicle of this species ends immediately anterior to the posterior testis. Unlike Caballero (1952), Sogandares-Ber- nal (1959:89) did not consider S. sentum a synonym of S. minutum because he found immature specimens of S. sentum larger than mature S. minmutum occurring in the same host. Stephanostomum ditrematis (Yamaguti, 1939) Manter, 1947 Echinostephanus ditrematis Yamaguti, 1939. Stephanostomum 1940. Stephanostomum filiforme Linton, 1940. Stephanostomum manteri Pérez Vigueras, 195): Stephanostomum cubanum Pérez Vigueras, 19>): Hosts: Caranx crysos (1 of 2); Caranx hip- pos (2 of 3). Site: Rectum. Discussion: The number of oral spines varies in my specimens as in those of Man- ter (1947:308-309). One individual clearly shows the unspined metraterm joining the cirrus sac near the base of the acetabulum. The genital atrium is unspined. Sogandares- Bernal (1959:88-89) discussed possible synonyms of S. ditrematis. longisomum Manter, Stephanostomum megacephalum Manter, 1940 Host: Caranx hippos (2 of 3). Site: Rectum. ASX [Pr S re) @ EY: 0.2 34 Figure 34. Stephanostomum sp., dorsal view, vitelline follicles dorsal and ventral to gonads omitted. Stephanostomum sp. Figure 34 Host: Opsanus beta (1 of 6). Site: Intestine. Specimen deposited: U.S. N. M. Helm. Coll. Now 71317; Discussion: Linton (1901:468-469) re- ported Stephanostomum tenue from Opsanus tau and referred to the original description (Linton, 1898:535-536). His identification probably is incorrect because he considered more than one species as S. tenwe. The pres- Digenetic Trematodes of Marine Teleosts 161 ent specimen may well be a new species. It has the following characteristics: Body 1.0 long. Oral sucker 0.13 wide. Acetabulum 0.18 wide. Sucker ratio 1:1.33. Forebody 27% of body length. Apparently 38 oral spines. Prominent eyespots at level of an- terior of pharynx. Esophagus about as long as pharynx. Uroproct present. Ovary 0.11 from acetabulum, with cirrus sac extend- ing 7 that distance. Prostatic vesicle short. Posttesticular space 12% of body length. Gonads nearly contiguous. Vitellaria extend- ing from midacetabular level to posterior end of body; with follicles not shown in Figure 34 dorsal and ventral to gonads; meeting dorsally between testes, but not between ovary and anterior testis as in S. dentatum from flounders (Manter, 1947: 308). Longest collapsed eggs 68 by 31 mi- crons. Manteria brachydera (Manter, 1940) Caballero, 1950 Dihemistephanus 1940. Stephanostomum sp. Linton, 1940. Manteria costalimai Freitas and Kohn, 1964 (new synonym). Host: Oligoplites saurus (2 of 2). Site: Pyloric caeca. Discussion: The present material differs from Manteria brachydera as described by Manter (1940a:399-400) in the respects given by Siddiqi and Cable (1960:289). The pharynx is 0.05 to 0.07 wide instead of 0.08 to 0.09. Oral spines are smaller and number 38 to 45 rather than 50 to 60, and the vitelline follicles extend farther anteriorly, relative to the seminal vesicle. Bravo-Hollis (1954:234-238) described ad- ditional material from the Pacific coast of Mexico which compared well with Manter’s material from Ecuador (Manter, 1940a; Caballero, 1950). My specimens are more like the one illustrated by Siddiqi and Cable (1960, Figures 58 to 61) which reveals 32 oral spines and greater extent of vitellaria. Freitas and Kohn (1964) erected M. costalimat for specimens with a_ wide pharynx and variation in the extent of vi- tellaria anteriorly. They distinguished it from M. brachydera by having fewer oral spines, a slightly larger acetabulum, a smaller posttesticular region, and a longer genital brachyderus Manter, 162 atrium. Since my specimens show all but the number of oral spines to be integrated characteristics, I agree with Sogandares- Bernal and Hutton (1959a:268-269), who discussed the low number of oral spines in their material from the west coast of Florida and did not feel justified in erecting a new species on that basis. Manteria costalimai is therefore considered a synonym of M. brachy- dera with not more than subspecific status. My specimens do not show the dorsal interruption of the oral spines found in some of Manter’s material. Also, the tubular pros- tatic vesicle is very sinuous, a character not suggested in the illustrations by Siddiqi and Cable, Sogandares-Bernal and Hutton, and Freitas and Kohn. Partially-collapsed eggs measure 63 to 69 by 37 to 41 microns. FAMILY HEMIURIDAE Liihe, 1901 Aponurus elongatus Siddigi and Cable, 1960 Host: Chaetodipterus faber (1 of 2). Site: Stomach. Discussion: Six specimens 0.99 to 1.38 long have forebody 21 to 25% of body length, postovarian space 30 to 35% of body length, oral sucker 0.12 to 0.14 wide, ace- tabulum 0.26 to 0.28 wide, sucker ratio EON 23 wands egesi2 >). t01 52 by, e250 16 microns. Nahhas and Short (1965:45- 46) distinguished this species from Apo- nurus laguncula Looss, 1907, by the larger sucker ratio (1:2.5 compared to 1:1.7 to 2.1), more elongate body, more anterior ventral sucker, greater postovarian space, and longer-than-wide vitellaria. The mea- surements of eggs in my specimens lie be- tween those given by Siddiqi and Cable and those of Nahhas and Short. The hind- bodies are slightly contracted, giving a shorter postovarian space, and the sucker fatio is less than 1:2.5; it is 1:2.7 from the diameters of suckers illustrated in Figure 125 of Siddiqi and Cable (1960). The vitel- laria, anterior acetabulum, and postovarian space, however, are still useful in separating A. elongatus from A. laguncula. This species shows a marked similarity to A. callionymi Yamaguti, 1938, A. rhinoplagusiae Yama- guti, 1934, and A. acropomatis Yamaguti, 1938, all species from Japanese waters. A. elongatus differs from them in having slightly smaller eggs and convoluted caeca in un- extended specimens. Variation in the size and shape of the seminal receptacle and Tulane Studies in Zoology and Botany Vol. als vitellaria in my specimens suggests caution in appraising these characters. Parahemiurus merus (Linton, 1910) Woolcock, 1935 Hemiurus merus Linton, 1910. Parahemiurus parahemiurus Vaz and Pe- reira, 1930. Parahemiurus platichthyi Lloyd, 1938. Parahemiurus atherinae Yamaguti, 1936. Parahemiurus harengulae Yamaguti, 1938. Parahemiurus noblei King, 1962. Hosts: Caranx crysos (2 of 2); Caranx hip- pos (1 of 3); Lagodon rhomboides (2 of 5); Sardinella anchovia (3 of 3). Site: Stomach. Discussion: Travassos et al. (1967:31-33) gave an extensive synonymy for Parahemiu- rus merUus, Progenetic metacercariae of P. merus oc- cur in the coelom of Sagitta hispida from Biscayne Bay. I found a 3.2 per cent inci- dence of infection in 250 specimens of S. hispida collected by Mr. Gary Hendrix on 30 November 1967. Two of three infected chaetognaths maintained in separate beak- ers for 29 days had metacercariae with young eggs in the uterus. The body of the longest metacercaria of P. merus is 1.23. Parahemiurus anchoviae Pereira and Vaz, 1930 Figure 35 Host: Anchoa lyolepis (2 of 2)*. Site: Stomach. Specimen deposited: U.S. N. M. Helm. Coll. No. 71318. Description (based on 4 specimens): Body 0.51 to 0.63 long excluding ecsoma; 0.15 to 0.18 in maximum width, at vitelline level; ecsoma short, 0.06 to 0.12 long. Cuticular plications along entire body length. Oral sucker subterminal, 0.03 wide with small papillae about ventral mouth. Ventral sucker 0.07 wide. Sucker ratio 1:2.1 to 24. Forebody 16 to 17% of body length. Pre- pharynx absent. Pharynx spherical to slightly elongate, about 0.025 in diameter. Esopha- gus short. Caeca terminating mear or in ecsoma. Testes near the middle of the body, tan- dem to diagonal, in contact or slightly sep- arate. Genital pore ventral, at midlevel of 0.1 view. oral sucker, either sinistral or dextral. Semi- nal vesicle muscular, 0.06 to 0.09 long by 0.04 wide, ending posteriorly in zone of anterior testis. Prostatic vesicle sometimes sinuous, as long as or longer than seminal vesicle, extending anteriorly to near mid- acetabular level, surrounded by numerous prostatic cells. Sinus sac a slender, elongated muscular tube. Ovary spherical or slightly irregular, sep- Digenetic Trematodes of Marine Teleosts 163 arated from testes by more or less than length of a testis. Vitellaria 2 large, com- pact, slightly irregular masses, immediately postovarian. Postvitelline region 13 to 27% of body length. Uterus extending to or into ecsoma. Eggs 19 to 25 by 9 to 11 microns. Discussion: The present specimens differ in many respects from the original descrip- tion (Pereira and Vaz, 1930b:705-706), which lacks an illustration. Their specimens are from the intestine instead of the stom- ach, are 1.8 rather than 0.6 long, and ap- parently have a greater relative length of the “pars prostatica.” Also, those authors reported the cuticular plications as being more easily distinguished on the posterior 7 of the body, the genital pore median, the seminal vesicle not extending to the testes, the posterior extent of the uterus at the beginning of the posterior % of the body, and newly-formed eggs 28 by 8 microns. They could not determine the termination of the caeca. Because my specimens are from a related host, host specificity in this group is low, and considerable morphological variation exists, I am not inclined to ascribe specific magnitude to the differences between the present specimens and the original descrip- tion of P. anchoviae. This species was de- scribed from Anchovia olida, probably a synonym of Lycengraulis olidus (Gunther, 1874), also from Brazil and in the same family as Anchoa lyolepis. Parahemiurus anchoviae differs from other species with cuticular plications along the entire body. P. clupeae Yamaguti, 1953, is much larger and has a long ecsoma; P. equadori Manter, 1940, has the oral sucker larger than the acetabulum and an elongated pharynx; and P. australis Woolcock, 1935, has a more elongated seminal vesicle. The plications also extend near or to the end of the body in P. lovelliae Crowcroft, 1947, in which the seminal vesicle reaches the ovaty. Ectenurus americanus (Manter, 1947) Manter and Pritchard, 1960 Parectenurus americanus Manter, 1947. Magnacetabulum americanum (Manter, 1947) Yamaguti, 1954. Host: Synodus foetens (3 of 7). Site: Stomach. 164 Ectenurus virgulus Linton, 1910 Hosts: Caranx crysos (1 of 2)*; Caranx hippos (1 of 3). Site: Stomach. Sterrhurus musculus Looss, 1907 Sterrhurus laeve (Linton, 1898) of Man- cen 931 (Gngpart). Sterrhurus floridensis Manter, 1934 (in part). Brachyphallus musculus (Looss, 1907) Skrjabin and Guschanskaja, 1955. Hosts: Achiris lineatus (1 of 1); Aniso- trema virginicus (1 of 6)*; Caranx hip- pos (1 of 3)*; Diplectrum formosum (3 of 4); Epinephelus striatus (1 of 2)*; Eucinostomus gula (1 of 3)*; Haemulon aurolineatum (1 of 7)*; Haemulon par- rat (1 of 7)*; Lutyanus apodus (1 of 3); Lutjanus griseus (2 of 3); Ogcocephalus cubtifrons (2 of 2); Orthopristis chrysop- terus (2 of 4); Paralichthys albigutta (1 of 1); Scorpaena plumieri (1 of 4); Synodus foetens (5 of 7). Site: Stomach; also intestine of O. cubi- frons. Lecithochirinm parvum Manter, 1947 Sterrhurus floridanus Manter, 1934 (in part). Brachyphallus parvus (Manter, 1947) Skrjabin and Guschanskaja, 1955. Hosts: Archosargus rhomboidalis (2 of 5)*; Bathygobius soporator (3 of 9)*; Caranx crysos (1 of 2)*; Elops saurus (2 of 3) *; Eucinostomus gula (1 of 3)*; Haemulon flavolineatum (1 of 2)*; Lagodon rhom- boides (2 of 5)*; Lutjanus synagris (2 of 7)*; Mycteroperca bonaci (2 of 3)*; Mycteroperca microlepis (1 of 2)*; Scor- paena grandicornis (1 of 2)*; Synodus foetens (2 of 7). Site: Stomach. Discussion: Characteristics vary among specimens. The ecsoma may be extended, with the caeca terminating inside it, the testes may be separated, and the seminal vesicle is not always tripartite. Some speci- mens have slightly larger eggs and a slightly larger or smaller sucker ratio than described. I usually identify this species by the combi- nation of small size, preacetabular pit, weakly-developed sinus sac, and low num- ber of postovarian coils of the uterus. Leci- Tulane Studies in Zoology and Botany Vol. 15 thochirium parvum and Sterrhurus musculus should not be placed in the genus Brachy- phallus Odhner, 1905, because both lack cuticular plications, a diagnostic character of Brachyphallus. Lecithochirium microstomum Chandler, 1935 Sterrhurus monticelli (Linton, 1898) of Manter, 1931 (in part). Lecithochirium sinaloense ISS: Hosts: Centropomus undecimalis (2 of 4)*; Epinephelus striatus (1 of 2)*; Lutjanus synagris (1 of 7)*; Mycteroperca bonaci (2 of 3)*; Oligoplites saurus (2 of 2)*; Pomatomus saltatrix (1 of 1)*; Synodus foetens (2 of 7). Site: Stomach. Discussion: Several workers (Sogandares- Bernal and Hutton, 1959a:269; Manter and Pritchard, 1960a:94-95, 1960b:175-176; Reid, Coil, and Kuntz, 1965:203) have re- cently discussed variations and increased ranges of measurements in this species. Manter (1931:406) reported Sterrhurus monticelli from Pomatomus saltatrix at Beau- fort, North Carolina. In 1947 (:342) he changed the identification to a species of Lecithochirium, probably L. branchialis. He recently re-examined a specimen and found it to be L. microstomum (personal commu- nication, 1968). Examination of specimens of L. branchialis (Stunkard and Nigrelli, 1934) may show it to be the same as L. macrostomum. Bravo-Hollis, Lecithochirium synodi Manter, 1931 Hosts: Opsanus beta (2 of 6)*; Synodus foetens (3 of 7). Site: Stomach. Lecithochirinm sp. Host: Selene vomer (1 of 2). Site: Stomach. Discussion: Two specimens appear to be Lecithochirium texanus (Chandler, 1941) Manter, 1947, although they are 1.9 to 2.1 rather than 3.25 to 3.60 in length. The sucker ratio of both is 1:3.0. Leurodera decora Linton, 1910 Host: Haemulon aurolineatum (1 of 7)*. Site: Stomach. No. 4 Brachadena pyriformis Linton, 1910 Distomum bothryophoron (Olsson, 1868 ) of Linton, 1901 and 1905 (in part). Lecithaster antsotremi MacCallum, 1921. Lecithaster gibbosus (Rudolphi, 1802) of Linton, 1940 (in part). Aponurus symmetrorchis Siddiqi and Ca- ble, 1960. Hosts: Antsotremus virginicus (2 of 6); Calamus bajonado (1 of 1); Haemulon carbonarium (1 of 1)*; Haemulon parrai (Qlok 7)):. Site: Stomach. Myosaccium opisthonemae (Siddigi and Cable, 1960) comb. n. Neogenolinea opisthonemae Siddiqi and Cable, 1960. Host: Sardinella anchovia (3 of 3). Site: Stomach. Discussion: Twelve specimens ranging from 0.44 to 0.90 long with sucker ratios of 1:1.6 to 1.8 and eggs 21 to 26 by 9 to 11 microns compare well in all respects with what Siddigi and Cable (1960:313) called Neogenolinea opisthonemae except that eggs are smaller than 29 to 32 by 12 to 15 microns. Partially-collapsed eggs mea- - sure 21 to 26 by 9 to 11 microns. The cu- ticular plications extend to the posterior end of the body on small specimens and to about the level of the vitellaria on larger ones. I do not think at this time that enough difference exists to erect a new species. The genus Neogenolina is a synonym of Myosaccium Montgomery, 1957. I have ex- amined a paratype of M. ecaude, lent by Dr. H. W. Manter, and there is some confusion in the description of Myosaccium. The re- gion that Montgomery labels (1957, Figure 28) as the ejaculatory duct is the sinus or- gan. This is the same organ as in Erilepturus Woolcock, 1935, Dimurus Looss, 1907, and Ectenurus Looss, 1907, and is discussed by Manter (in press). The muscle fibers sur- rounding the prostatic vesicle are spirally arranged, rather than distinctly longitudi- nally, and the internal vesicular cells do not reveal nuclei. The specimens from Sar- dinella anchovia do not have filaments or spines on the eggs, although a look at col- lapsed specimens on a fixed plane strongly suggests their presence. Specimens from Opisthonema oglinum \ent by Dr. R. M. Digenetic Trematodes of Marine Teleosts 165 Cable also do not have filamented eggs. The paratype of M. ecaude has collapsed eggs crowded together, and I could not positively distinguish any filaments. Filaments are probably not present on the eggs of this species. These corrections in Montgomery’s description remove supposed differences of Myosaccium from Neogenolina. Kohn and Buhrnheim (1964), however, also report M. ecaude with filamented and spined eggs. Their specimens were collected from Sar- dinella aurita in Brazil and are 0.65 to 1.04 long with eggs 30 to 41 by 9 to 13 microns. M. opisthonemae could well be small or progenetic forms of M. ecaude. Study of ad- ditional specimens probably can settle this problem. Opisthadena dimidia Linton, 1910 Opisthadena cortesi Bravo-Hollis, 1966 (new synonym ). Host: Kyphosus sectatrix (6 of 6). Site: Stomach. Discussion: Bravo-Hollis (1966: 144) used six characters to separate Opisthadena cortest Bravo-Hollis, 1966, from O. dimidia. I do not find these differences to be valid because of the intergradation discussed be- low, and consider the two species synony- mous. My 18 mounted specimens have sucker ratios of 1:2.3 to 3.4 and diameter- of-pharynx to diameter-of-oral sucker ratios of 1:1.4 to 1.7. The genital pore is usually at a level just below the pharynx, and the sinus sac is along the anterior border of the acetabulum, but if specimens are fixed in certain positions, those features appear to be more anterior or posterior than usually observed. Also depending on fixation, the seminal receptacle may be anterior to the ovary, even with that gonad, or anywhere between. The eggs are 25 to 45 by 12 to 17 microns, some specimens having larger eggs than others. Dictysarca virens Linton, 1910 Host: Hippocampus erectus (2 of 4). Site: Swim bladder. Discussion: My two specimens have mea- surements and ratios which fall between those given by Linton (1910:58-59) and Manter (1947:364-365). A difference, how- ever, is that the ovary is not completely posterior to the vitellaria. One specimen 166 )) ty. \:+) J LIM LZ SE) MEX E BE LE Ne CORNEA A Tulane Studies in Zoology and Botany Vols Figure 36. Didymocystis scomberomori. Figure 37. Didymocystis scomberomori, anterior end. has one testis only. The eggs measure 24 to 35 by 12 to 19 microns. Daigger and Lewis (1967:8) reported Dictysarca virens in Biscayne Bay from the same host, under its synonym Hippocampus punctulatus. The trematode was previously known only from swim bladders of moray eels. Gonocercella trachinotit (MacCallum, 1913) Yamaguti, 1954 Distomum sp. Linton, 1905 (from Tracht- notus carolinus ). Distomum trachinoti MacCallum, 1913. Gonocercella atlantica Manter, 1940. Host: Albula vulpes (2 of 7)*. Site: Stomach and upper intestine. Discussion: Of five specimens with de- veloped gonads, the two largest are con- tracted specimens 2.4 and 2.5 long that have poorly-formed eggs, such as those re- ported for the type of Gonocercella pactfica (Manter, 1940a:437-439). MacCallum (1913:410-411) described Distomum tracht- noti from Trachinotus carolinus from which Linton (1905:367) described Dzistomum sp. Both authors were confused in interpret- ing the vitellaria. Apparently unaware of MacCallum’s paper, Manter (1940a:437- 439) erected G. atlantica and later (1947: 358) separated G. atlantica from G. pacifica by the presence of more profuse prostatic cells, longer and more coiled seminal vesicle, and shorter eggs (27 compared with 34 mi- crons). My specimens are like G. atlantica, except that eggs from the most mature specimens are 49 to 59 by 21 to 29 microns and the testes are symmetrical, as illustrated by Pearse (1949), rather than diagonal. MacCallum, however, described eggs as 0.09 long by 0.03 wide. Numerous gland cells surround the spherical excretory vesicle in my specimens. The study of fully-mature specimens will permit better understanding of the genus. Manter, Pearse, and Mac- Callum reported G. trachinoti from Mona- canthus hispidis, Fundulus majalis, and Roc- cus saxatilus (=R. lineatus). FAMILY SCLERODISTOMATIDAE Dollfus, 1932 Sclerodistomum sphoeroidis Manter, 1947 Hosts: Chilomycterus schoepfi (2 of 6); Sphaeroides testudineus (1 of 5)*. Site: Stomach. Discussion: Three specimens are 4.6 to 6.4 long and have eggs 24 to 31 by 19 to 24 microns. The genital pore is at the pharyngeal level in two specimens and % the distance from the oral sucker to the acetabulum in the other. Sogandares-Bernal and Hutton (1959b:66) discussed the prob- No. 4 able error in some records of sites for species in this genus. FAMILY DIDYMOZOIDAE Poche, 1907 Didymocystis scomberomori (MacCallum and MacCallum, 1916) Yamaguti, 1954 Figures 36 and 37 Distomum (Koellikeria) sp. Linton, 1901. Koellikeria scomberomori MacCallum and MacCallum, 1916. Hosts: Scomberomorus maculatus (2 of 2); Scomberomorus regalis (1 of 2)*. Site: Encysted in pairs in wall of stomach, intestine, and pyloric caeca. Specimen deposited: U.S. N. M. Helm. Coll. No. 71319. Description (based on 24 wholemounts and 1 sectioned cyst): Hermaphroditic, en- closed as pair in globular to subglobular cyst. Hindbody hemispherical to reniform, 0.43 to 1.32 long at median axis by 0.86 to 2.17 wide in wholemounts. Forebody attaching near anterior margin of hind- body, 0.23 to 0.92 long by 0.07 to 0.16 wide; widest at anterior level. Oral sucker re- placed by a spherical glandular organ, some- times overlapping pharynx. Pharynx muscu- lar, 0.021 to 0.035 long by 0.017 to 0.026 wide. Esophagus longer or shorter than % width of forebody. Caeca usually narrow in forebody, and vesicular in hindbody. Testes tubular, paired, near midlevel at side of hindbody; straight, arcuate, or sinu- ous. Genital pore subterminal or terminal, near pharynx. Vas deferens sinuous or not. Ovary tubular, either undivided or divid- ing near seminal receptacle into 2 stems, usually of unequal length, each extending in opposite directions, winding sinuously along lateral and posterior margins of hind- body, without secondary branching. Seminal receptacle saccate, site variable from almost median to midway to lateral margin, near base of hindbody or well interior. Mehlis’s gland lateral to receptacle. Vitellaria tubular, slender, sinuous, extending along posterior or dorsolateral margins of hindbody; branch- ing more in some specimens than in others; number of free ends variable; occasionally vitellaria occupying a little more than % of posterior margin of hindbody and a branch of ovary occupying the remainder of the margin. Uterus extensive, occupying almost all available space in hindbody. Reser- Digenetic Trematodes of Marine Teleosts 167 voir apparently present. Eggs slightly reni- form, 12 to 15 by 8 to 11 microns, usually 14 by 10 microns. Discussion: My specimens are apparently of the same species as that reported and illustrated by Linton (1901:447) as Dis- tomum (Koellikeria) sp. from the same host. Linton gave measurements on length, width, diameter of neck, and eggs from a single specimen. MacCallum and MacCallum (1916:153), not citing Linton’s report, gave almost identical measurements for K. scom- beromori but did not provide a detailed description. The anterior glandular organ might be the same as the “feebly muscular” oral sucker in D. submentalis Yamaguti, 1938. There is a thin membrane around the struc- ture, but the membrane is not muscular. DISCUSSION Three hundred and thirty-three individ- uals comprising 113 species of teleost fishes were examined for digenetic trematodes. Of these fishes, 214 (64.3 per cent), repre- senting 69 species, were parasitized by adult Digenea. This does not imply that the 44 species which were not parasitized are not definitive hosts for Digenea in Biscayne Bay, because they were represented by only 83 individuals, some of which were imma- ture and most of which were small. Imma- ture Digenea which might have been able tO attain maturity were present in some of those fishes. Of the 69 species which har- bored Digenea, only 36 of the 250 individ- uals examined did not have adult Digenea. Again, most of these were immature fishes or small species such as Blennius cristatus and Bathygobius soporator. Nineteen of the 69 species were infected by only one species of Digenea. Most of the previously-described Digenea from my collection have been reported from waters south of the mainland of Florida. Bucephaloides bennetti, Lepocreadinm flort- danum, Biantum vitellosum, and Genttoco- tyle cablei have been reported previously from the Gulf of Mexico only. I suspect that these trematodes will be discovered in areas between Miami and where they were te- ported, when more collections are made. Didymocystis scomberomori is known only from the Atlantic in areas north of Florida. Botulisaccus pisceus and Pseudocreadium 168 scaphosomum have been reported from hosts in the Pacific Ocean only. Not including Bucephaloides arcuatus from Pacific barra- cuda, which is a misidentification, there are 30 species of trematodes in my collection which are also found in hosts from the American Pacific coast. Manter, who first compared the Atlantic and Pacific faunas, listed (1940b and 1947:379) 24 common species. However, with the present knowl- edge, all but four species of trematodes from Biscayne Bay which are also found in the American Pacific are common to the Tortu- gas. Of the four, two are from hosts Dr. Manter did not examine. Only 22 species in my study, including Vitellibaculum spinosum and Lecithochwrium microstomum, have been reported from wa- ters adjacent to the American mainland north of Biscayne Bay. This does not in- clude species described from tropical fishes in the New York Aquarium. Fourteen ad- ditional species, however, have been re- ported from Bermuda. Another 18 species were reported from the Bahama _ Islands which were not reported from Bermuda. Sogandares-Bernal (1959:108-110) com- mented on the difference between Ba- hamian and Bermudian trematode faunas, noting that, from known collections, the Biminian fauna was more similar to the faunas of Tortugas and the American Pacific than to the Bermudian fauna. The trematode fauna reported from Bis- cayne Bay seems to be more closely affiliated with the tropical faunas than with the more temperate faunas. Even though some of these trematodes are specific to tropical fishes, several others are from fishes which are present, or whose related species are present, in more northern waters. Undoubt- edly, more trematode species will be re- ported from these and other hosts in the northern waters when parasitologists study additional collections. The significant findings of this study are the numerous undescribed species, inter- mediate forms, and new host records from Biscayne Bay. I do not believe that these species or populations are restricted to this area but rather that they indicate how little is known about the Digenea from the At- lantic and Caribbean, areas that are consid- ered to be some of the best studied in the world. Tulane Studies in Zoology and Botany Vol. 15 ALPHABETICAL LIST OF HOST-SPECIES WITH THE PARASITES FOUND IN EACH The first number following the name of the family indicates the number of individuals ex- amined, and the numbers in parentheses indi- cate the length or range of lengths of the hosts expressed as fork lengths in centimeters. Abudefduf saxatilis (Linnaeus), sergeant major (Pomacentridae )—2 (9) none Acanthurus chirurgus (Bloch), doctorfish ( Acanthuridae )—1 (11) none Achirus lineatus (Linnaeus), lined sole (Sole- idae )—1 (10) Sterrhurus musculus Acyrtops beryllinus (Hildebrand & Ginsburg), emerald clingfish (Gobiesocidae)—3 (2) none Albula vulpes (Linnaeus), bonefish (Albuli- dae )—7 (42-65) Botulisaccus pisceus Claribulla longula Diphtherostomum albulae Gonocercella trachinoti Lasiotocus albulae Alutera schoepfi (Walbaum), orange filefish (Balistidae )—1 (9) none Anchoa lyolepis (Evermann & Marsh), dusky anchovy (Engraulidae)—2 (7) Parahemiurus anchoviae Anisotremus virginicus (Linnaeus), (Pomadasyidae )—6 (15-20) Apocreadium cryptum Apopodocotyle oscitans Brachadena pyriformis Diplangus parvus Diplangus paxillus Diphtherostomum anisotremi Genolopa ampullacea Helicometrina execta Lasiotocus longicaecum Lasiotocus longovatus Postmonorchis orthopristis Sterrhurus musculus Apogon maculatus (Poey), gonidae)—1 (10) none Archosargus rhomboidalis (Linnaeus) (= A. unimaculatus), sea bream (Sparidae)—5 (19-28 ) Apopodocotyle oscitans Diphtherostomum americanum Diplomonorchis leiostomi Lecithochirium parvum Megasolena hysterospina Steringotrema corpulentum Ariosoma impressa (Poey), bandtooth conger (Congridae)—1 (24) none Arius felis (Linnaeus) (=Galeichthys f.), sea catfish ( Ariidae )—2 (28-39 ) none Atherinomorus stipes (Muller and Troschel), hardhead silverside (Atherinidae)—1 (6) none Balistes porkfish flamefish (Apo- capriscus Gmelin, triggerfish gray No. 4 (Balistidae )—4 (20-31) Apocreadium mexicanum Neoapocreadium coili Pseudocreadium lamelliforme Xystretrum solidum Bathygobius mystacium Ginsburg (?), island frillfin (Gobiidae)—1 (5) none Bathygobius soporator (Valenciennes ), fin goby (Gobiidae)—9 (5-11) Helicometrina execta Lecithochirium parvum Opegaster pritchardae Blennius cristatus Linnaeus, molly miller ( Blen- niidae)—6 (5-11) Helicometra execta Blennius marmoreus Poey, (Blenniidae )—6 (5-7) none Calamus bajonado (Bloch & Schneider ), head porgy (Sparidae)—1 (28) Host from ocean side of Sands Key Brachadena_ pyriformis Pachycreadium crassigulum Proctoeces maculatus Stephanostomum sentum Callionymus pauciradiatus Gill, dragonet (Cal- lionymidae )—3 (3) none Caranx crysos (Mitchill), blue runner (Caran- gidae)—2 (20-21) Bucephalus varicus Ectenurus virgulus Lecithochirium parvum Parahemiurus merus Stephanostomum ditrematis Tergestia pectinata Caranx hippos (Linnaeus), erevalle jack (Ca- rangidae )—3 (30-46) Bucephalus varicus Ectenurus virgulus Parahemiurus merus Stephanostomum ditrematis Stephanostomum megacephalum Sterrhurus musculus Carapus bermudensis (Jones), pearlfish (Ca- rapidae)—1 (14) none Centropomus undecimalis (Bloch), snook (Cen- tropomidae )—4 (50-86 ) Lecithochirium microstomum Chaetodipterus faber (Broussonet), spadefish (Ephippidae )—2 (7-22) Aponurus elongatus Multitestis inconstans Vitellibaculum spinosum Chilomycterus schoepfi (Walbaum), burrfish (Diodontidae)—6 (12-17) Diploproctodaeum vitellosum Sclerodistomum sphoeroidis Chriodorus atherinoides Goode & Bean, hard- head halfbeak (Exocoetidae)—1 (17) none Citharichthys spilopterus Gimnther, bay whiff (Bothidae)—3 (9-17) none Coryphopterus glaucofraenum Gill, bridled goby (Gobiidae )—4 (6) none frill- seaweed blenny jolt- Atlantic striped Digenetic Trematodes of Marine Teleosts 169 Coryphopterus thrix Bohlke & Robins, thread- fin goby (Gobiidae)—1 (5) none Corythoichthys albirostris Heckel, pipefish (Syngnathidae )—2 (10) none Dactyloscopus tridigitatus Gill, sand stargazer ( Dactyloscopidae )—2 (4) none Diplectrum formosum (Linnaeus), sand perch (Serranidae )—4 (12-22) Sterrhurus musculus whitenose Diplodus_holbrooki (Bean), spottail pinfish (Sparidae)—1 (19) none Elops saurus Linnaeus, ladyfish (Elopidae )—3 (36-45 ) Lecithochirium parvum Epinephelus adscensionis (Osbeck), rock hind (Serranidae)—1 (21) Helicometra torta Epinephelus striatus (Bloch), Nassau grouper (Serranidae)—2 (33-38) Helicometra torta Lecithochirium microstomum Sterrhurus musculus Equetus acuminatus (Bloch & Schneider) (= FE. pulcher), high hat (Sciaenidae )—3 (7-9) Manteriella crassa Pseudopecoeloides equesi Erotelis smaragdus (Valenciennes), emerald sleeper (Eleotridae )—1 (9) none Eucinostomus gula (Quoy & Gaimard), silver jenny (Gerridae)—3 (8-10) Crassicutis marina Hurleytrema eucinostomi Lecithochirium parvum Sterrhurus musculus Eupomacentrus fuscus (Cuvier) (= Poma- centrus f.), dusky damselfish (Pomacentri- dae )—1 none Eupomacentrus leucostictus (Miller & Tros- chel) (= Pomacentrus l.), beaugregory (Pomacentridae )—6 (5-9) Diphtherostomum americanum Eupomacentrus variabilis (Castelnau), cocoa damselfish (Pomacentridae )—1 (11) none Fistularia tabacaria Linnaeus, cornetfish ( Fis- tulariidae)—1 (19) none Floridichthys carpio (Gunther), goldspotted killifish (Cyprinodontidae)—8 (4-6) none Fundulus similis (Baird & Girard), longnose killifish (Cyprinodontidae )—4 (4-5) none Gobiesox strumosus Cope, skilletfish (Gobie- socidae )—1 (2) none Gobionellus smaragdus ( Valenciennes ), emerald goby (Gobiidae )—2 (4) none Gobionellus stigmaticus (Poey), marked goby (Gobiidae)—1 (5) none 170 Gymnothorax nigromarginatus (Girard), black- edge moray (Muraenidae)—2 (35) none Haemulon aurolineatum Cuvier (= Bathy- stoma a.), tomtate (Pomadasyidae)—7 (12- 79) Apocreadium foliatum Genolopa ampullacea Lasiotocus longovatus Leurodera decora Postmonorchis orthopristis Sterrhurus musculus Haemulon carbonarium Poey, (Pomadasyidae)—1 (21) Apocreadium foliatum Brachadena_ pyriformis Diplangus parvus Diplangus paxillus Stephanostomum sentum Haemulon flavolineatum (Desmarest), French grunt (Pomadasyidae )—2 (15-16) Lasiotocus truncatus Lecithochirium parvum Genolopa ampullacea Haemulon parrai (Desmarest), sailors choice (Pomadasyidae )—7 (15-24) Apocreadium cryptum Apocreadium foliatum Brachadena_ pyriformis Diplangus parvus Diplangus paxillus Genolopa ampullacea Lasiotocus longovatus Postmonorchis orthopristis Sterrhurus musculus Haemulon plumieri (Lacépéde), white grunt (Pomadasyidae )—5 (12-17) Diphtherostomum anisotremi Diplangus parvus Genolopa ampullacea Lasiotocus haemuli Lasiotocus truncatus Postmonorchis orthopristis Haemulon sciurus (Shaw), bluestriped grunt (Pomadasyidae)—6 (15-19) Diphtherostomum anisotremi Diplangus parvus Diplangus paxillus Genolopa ampullacea Infundibulostomum spinatum Lasiotocus haemuli Lasiotocus longovatus Lasiotocus truncatus Postmonorchis orthopristis Halichoeres bivittatus (Bloch), slippery dick (Labridae)—11 (14-19) Helicometrina execta Nicolla halichoeri Halichoeres pictus (Poey), painted doncella (Labridae )—2 (19-20) Helicometrina execta Nicolla sp. Halichoeres poeyi (Steindachner), wrasse (Labridae)—1 (12) none Halichoeres radiatus (Linnaeus), puddingwife (Labridae)—8 (12-24) Helicometrina execta Nicolla halichoeri Caesar grunt blackear Tulane Studies in Zoology and Botany Vol. 15 Hippocampus erectus Perry (= H. punctula- tus), spotted seahorse (Syngnathidae )—4 (13-14) Dictysarca virens Genitocotyle cablei Hippocampus zosterae Jordan & Gilbert, dwarf seahorse (Syngnathidae)—1 (6) none Histrio histrio (Linnaeus ), tennariidae)—1 (4) none Holacanthus isabelita (Jordan & Rutter), blue angelfish (Chaetodontidae )—1 (12) Antorchis urna Kyphosus sectatrix (Linnaeus), Bermuda chub (Kyphosidae )—6 (20-28 ) Cadenatella americana Cadenatella floridae Deontacylix ovalis Enenterum aureum Opisthadena dimidia Schikhobalotrema kyphosi Labrisomus kalisherae (Jordan), downy blenny (Clinidae)—2 (8-9) Helicometrina execta Labrisomus nuchipinnis (Quoy & Gaimard), hairy blenny (Clinidae)—1 (17) Helicometrina mirzai Lactophrys quadricornis (Linnaeus) (= L. tricornis ), cowfish (Ostraciidae )—3 (14-21) Dermadena lactophrysi Megapera sp. Thysanopharynx elongatus Lagodon rhomboides ( Linnaeus ), pinfish ( Spar- idae)—5 (14-20) Diphtherostomum americanum Diplomonorchis leiostomi Lecithochirium parvum Lepocreadium floridanum Parahemiurus merus Proctoeces lintoni Steringotrema corpulentum Lutjanus apodus (Walbaum), (Lutjanidae )—3 (14-22) Helicometrina nimia Metadena adglobosa Stephanostomum tenue Sterrhurus musculus Lutjanus griseus (Linnaeus), (Lutjanidae )—3 (18-25) Hamacreadium mutabile Metadena adglobosa Metadena globosa Metadena sp. Stephanostomum casum Sterrhurus musculus Lutjanus mahogoni (Cuvier), mahogany snap- per (Lutjanidae )—2 (18-23) Helicometrina nimia Metadena globosa Siphodera vinaledwardsii Stephanostomum tenue Lutjanus synagris (Linnaeus), (Lutjanidae)—7 (12-22) Hamacreadium mutabile Lecithochirium microstomum Lecithochirium parvum Metadena globosa sargassumfish (An- schoolmaster gray snapper lane snapper No. 4 Siphodera vinaledwardsii Stephanostomum casum Malacoctenus macropus (Poey), rosy blenny (Clinidae)—1 (5) none Monacanthus ciliatus (Mitchill), fringed file- fish (Balistidae)—1 (8) none Monacanthus hispidus (Linnaeus), planehead filefish ( Balistidae )—6 (9-26) Pseudocreadium scaphosomum Xystretrum solidum Mugil cephalus Linnaeus, striped-mullet (Mu- gilidae)—3 (23-36) Hymenocotta manteri Lasiotocus mugilis Mugil curema Valenciennes, white mullet (Mu- gilidae)—1 (16) none Mycteroperca bonaci (Poey), black grouper (Serranidae)—3 (22-41) Lecithochirium microstomum Lecithochirium parvum Postporus epinepheli Prosorhynchus pacificus Mycteroperca microlepis (Goode & Bean), gag (Serranidae )—2 (20-24) Lecithochirium parvum Neolepidapedon macrum Prosorhynchus pacificus Nicholsina usta (Valenciennes), emerald par- rotfish (Scaridae)—3 (18) Schikhobalotrema sparisoma Ocyurus chrysurus (Bloch), yellowtail snapper (Lutjanidae)—5 (15-20) Hamacreadium confusum Helicometrina nimia Lepocreadium trulla Metadena globosa Paracryptogonimus americanus Stephanostomum casum Ogcocephalus cubifrons (Richardson), batfish (Ogcocephalidae )—2 (25-28) Helicometrina mirzai Stephanostomum sentum Sterrhurus musculus Oligoplites saurus (Bloch & Schneider ), leather- jacket (Carangidae )—2 (26-27 ) Lecithochirium microstomum Manteria brachydera Ophichthus gomesi (Castelnau), shrimp eel (Ophichthidae )—1 (48) Diplomonorchis sphaerovarium Opsanus beta (Goode & Bean), gulf toadfish ( Batrachoididae )—6 (15-34) Helicometrina mirzai Helicometrina nimia Lecithochirium synodi Siphodera vinaledwardsii Stephanostomum sp. Orthopristis chrysopterus (Linnaeus), pigfish (Pomadasyidae )—4 (12-24) Diplomonorchis leiostomi Lasiotocus longovatus Sterrhurus musculus Paralichthys albigutta Jordan & Gilbert, gulf flounder (Bothidae)—1 (25) Bucephaloides bennetti Sterrhurus musculus Digenetic Trematodes of Marine Teleosts al Pomacanthus arcuatus (Linnaeus) (= P. au- reus), gray angelfish (Chaetodontidae )—4 (20-28 ) Antorchis urna Barisomum erubescens Cleptodiscus reticulatus Hexangitrema pomacanthi Pomacanthus paru (Bloch) (= P. arcuatus, in part), French angelfish (Chaetodontidae )—1 (18) Antorchis urna Pomatomus saltatrix (Linnaeus), bluefish (Pomatomidae )—1 (46) Lecithochirium microstomum Prionotus scitulus Jordan & Gilbert, leopard searobin (Triglidae)—2 (13-23) none Pristigenys alta (Gill), short bigeye (Priacan- thidae )—1 (6) none Sardinella anchovia Valenciennes, Spanish sar- dine (Clupeidae)—3 (12-13) Lepocreadium pyriforme Myosaccium opisthonemae Parahemiurus merus Scomberomorus maculatus (Mitchill), Spanish mackerel (Scombridae )—2 (30-35) Didymocystis scomberomori Rhipidocotyle adbaculum Scomberomorus regalis (Bloch), cero (Scom- bridae )—2 (30-35) Bucephaloides arcuatus Didymocystis scomberomori Rhipidocotyle adbaculum Scorpaena grandicornis Cuvier, lionfish (Scor- paenidae )—2 (18-23) Bucephalus scorpaenae Helicometrina nimia Lecithochirium parvum Scorpaena plumieri Bloch, spotted scorpion- fish (Scorpaenidae )—4 (21-27) Bucephalus scorpaenae Pseudopecoelus scorpaenae Sterrhurus musculus Selene vomer (Linnaeus), lookdown (Carangi- dae)—2 (19-21) Hurleytrema shorti Lecithochirium sp. Tergestia sp. Sparisoma aurofrenatum (Valenciennes), red- band parrotfish (Scaridae)—1 (18) none Sphaeroides nephelus (Goode & Bean), south- ern puffer (Tetraodontidae)—1 (26) none Sphaeroides spengleri (Bloch), bandtail puffer ( Tetraodontidae )—2 (11-15) Bianium plicitum Sphaeroides testudineus (Linnaeus), checkered puffer (Tetraodontidae)—5 (17-25) Bianium plicitum Sclerodistomum sphoeroidis Xystretrum solidum Sphyraena barracuda (Walbaum), great barra- cuda (Sphyraenidae)—1 (35) Claribulla longula Strongylura timucu (Walbaum), timucu (Be- lonidae )—3 (40-43) Schikhobalotrema acutum 172 Symphurus plagiusa (Linnaeus), blackcheek tonguefish (Cynoglossidae )—1 (15) none Syngnathus floridae (Jordan & Gilbert), dusky pipefish (Syngnathidae )—6 (14-19) none Syngnathus louisianae Ginther, chain pipefish (Syngnathidae )—3 (15-17) none Syngnathus scovelli (Evermann & Kendall), gulf pipefish (Syngnathidae)—2 (12) none Synodus foetens (Linnaeus), inshore lizardfish (Synodontidae )—7 (13-26) Ectenurus americanus Lecithochirium microstomum Lecithochirium parvum Lecithochirium synodi Sterrhurus musculus Trachinotus carolinus (Linnaeus), pompano (Carangidae)—1 (8) none Trachinotus falcatus (Linnaeus), permit (Ca- rangidae )—6 (7-10) Helicometrina execta Hurleytrema pyriforme Stephanostomum tenue Tylosurus crocodilus (Peron & Lesueur ), hound- fish (Belonidae)—1 (91) Steganoderma _nitens LITERATURE CITED ANDERSON, H. 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Prof. Mello-Leitao, Zool., (Sb) e542 Warp, H. L. 1954. Parasites of marine fishes of the Miami region. Bull. Mar. Sci. Gulf Caribb., 4(3):244-261. Winter, H. A. 1960. Algunos trematodos digeneos de peces marino de aguas del Oceano Pa- cifico del sur de California, U.S.A., y del litoral Mexicano. An. Inst. Biol. Univ. Méx. Year 1959, 30 (1-2) :183-208. YAMAGUTI, S. 1934. Studies on the helminth fauna of Japan. Part 2. Trematodes of fishes, I. Jap. J. Zool., 5(3):249-541. 1938. Studies on the helminth fauna of Japan. Part 21. Trematodes of fishes, IV. Publ. by author. 139 pp. 1958. Systema helminthum. Vol. I. Dige- netic trematodes of vertebrates, in 2 parts, Interscience Publishers, Inc., New York. 1575 pp. INDEX OF PARASITIC GENERA FROM BISCAYNE Bay The genera in parentheses indicate those used in junior synonyms only. ( Allomegasolena), 130 ( Lecithostaphylus ), 152 Antorchis, 124 ( Lepidauchen), 130 Apertile, 146 Lepocreadium, 135, 136 Apocreadium, 137, 138, 139 Leurodera, 164 Aponurus, 162, 165 (Macia), 149 Apopodocotyle, 143 (Magnacetabulum), 163 Barisomum, 128 Manteria, 161 Bianium, 141, 142 Manteriella, 146 ( Bilecithaster), 150 Megapera, 133 Botulisaccus, 125 Megasolena, 130 Brachadena, 165 (Mesorchis), 124 ( Brachyphallus), 164 Metadena, 158, 159 Bucephaloides, 122 (Monostomum ), 128, 158 (Bucephalopsis), 122 Multitestis, 143 Bucephalus, 121, 122 Myosaccium, 165 Cadenatella, 134 (Catoptroides), 149 Claribulla, 126 Neoapocreadium, 137 (Neogenolinea), 165 Neolepidapedon, 141 Cleptodiscus, 128 ( Neopecoelus), 146 Crassicutis, 143 Nicolla, 148, 149 Deontacylix, 121 Opegaster, 147 ( Deradena), 132 Opisthadena, 165 Dermadena, 137 (Opisthoporus ), 140 Dictysarca, 165 Pachycreadium, 143 Didymocystis, 167 Paracryptogonimus, 159 ( Dihemistephanus ), 161 Parahemiurus, 162 Diphtherostomum, 150, 151 ( Parectenurus ), 163 Diplangus, 150 (Plagioporus), 143 Diplomonorchis, 156 ( Pleurogonius ), 128 (Diploproctodaeum), 141 (Podocotyle), 143 (Distomum), 123, 124, 125, 135, 137, 141, Postmonorchis, 156 143, 152, 160, 165, 166, 167 Postporus, 140 (Echinostephanus ), 160 ( Pristisomum ), 156 Ectenurus, 163, 164 Proctoeces, 124 Enenterum, 134 (Proctotrema), 152, 153 ( Gasterostomum ), 122 Prosorhynchus, 122 Genitocotyle, 148 Pseudocreadium, 136, 137 Genolopa, 152, 153 ( Pseudohurleytrema ), 157 Gonocercella, 166 Pseudopecoeloides, 146 Hamacreadium, 143 Pseudopecoelus, 146 (Haplosplanchnus ), 131, 132 (Pseudoplagioporus), 143 Helicometra, 144 (Psilostomum), 141 Helicometrina, 144, 145 Rhipidocotyle, 122 (Hemiurus), 162 Schikhobalotrema, 131 Hexangitrema, 128 Sclerodistomum, 166 (Homalometron), 138 Siphodera, 158 (Horatrema), 146 Steganoderma, 152 Hurleytrema, 157 (Stegopa), 158 Hymenocotta, 132 (Stephanochasmus ), 159, 160 (Hypocreadium), 136 Stephanostomum, 159, 160, 161 Infundibulostomum, 124 Steringotrema, 125 ( Koellikeria), 167 Sterrhurus, 164 Lasiotocus, 152, 153, 154, 155 Tergestia, 123, 124 (Lebouria), 143 ( Theledra), 123 ( Lechradena), 159 Thysanopharynx, 133 ( Lecithaster ), 165 Vitellibaculum, 130 Lecithochirium, 164 Xystretrum, 149, 150 TULANE STUDIES IN ZOOLOGY AND BOTANY VOLUME 15 INDEX TO AUTHORS AND SCIENTIFIC NAMES (New taxonomic entities in boldface ) Abudefduf saxatilis, 168 Acanthurus chirurgus, 168 Acer negundo, 10 rubrum, 10 Achiris lineatus, 164, 168 Acyrtops beryllinus, 168 Albula vulpes, 125-6, 151, 155, 166, 168 Allomegasolena, 130 Alternanthera, 82, 85, 92, 97 Amunicola pilsbryi, 103 Anchoa lyolepis, 162, 168 Ancistrodon piscivorus, 16 Anisotremus virginicus, 139, 143-4, 150, 152-3, 165, 168 Antorchis, 124 Apertile gen. nov., 146 Apocreadium, 137-139 eryptum sp. nov., 139 Apogon maculatus, 168 Aponurus, 162, 165 Apopodocotyle, 143 Archosargus rhomboidalis, 125, 130, 143, 151, 164, 168 Ariosoma impressa, 168 Arius felis, 168 “scocotyle leighi, 103 Asellus communis, 103 stygius, 2 Balistes capriscus, 137, 150, 168 Barisomum, 128 Bathygobius mystacium, 169 soporator, 144, 147, 164, 169 Bianium, 141-2 Bilecithaster, 150 Birdsong, R. S., article, 106-112 Black, J. B., article, 5-9 Blennius cristatus, 144, 168 marmoreus, 168 Bollinger, R. P., article, 64-69 Botulisaccus, 125 Brachadena, 165 Brachyphallus, 164 Bridgman, J. F., article, 81-105 Bucephaloides, 122 Bucephalopsis, 122 Bucephalus, 121-2 Buteo lineatus, 16 Cadenatella floridae, sp. nov., 134 Calamus bajonado, 124, 143, 160, 164, 169 Callinectes sapidus, 81, 97, 99 Callionymus pauciradiatus, 169 Caranx crysos, 122-3, 160, 162, 164, 169 hippos, 122, 160, 162, 164, 169 Carapus bermudenis, 169 Carneophallus basodactylophallus, sp. nov., 81-2, 86, 98- 102 bilobatus, 101 canchei, 101 chabaudi, 101 choanophallus, sp. nov., 81-104 lactophrysi, 101 muellhaupti, 101 pseudogonotylus, 101 skryabini, 101 trilobatus, 101 tringae, 101 turgidus, 87, 99, 101 Carphophis amoenus, 42, 54 Catoptroides, 149 Cavia porcellus, 87 Celtis laevigata, 11 Centropomus undecimalis, 164, 169 Chaetodipterus faber, 130, 143, 162, 169 Chilomycterus schoepfi, 142, 166, 169 Chriodorus atherinoides, 169 Chrysemys, 62 Citharichthys spilopterus, 169 Claribulla longula, gen. nov. et sp. nov., 126 Clemmer, G. H., article, 18-39 Cleptodiscus, 128 Cobomba, 82, 85, 92, 97 Cole, G. A., article, 2-4 Corundus drummondi, 11 Coryphopterus glaucofraenum, 169 thrix, 169 Corythoichthys albirostris, 169 Crassicutis, 143 Dactyloscopus tridigitatus, 169 Deontacylix, 121 Deradena, 132 Dermadena, 137 Diadophis punctatus arnyi, 41-58 Dictysarca, 165 Didymocystis, 167 Dihemistephanus, 161 Diphtherostomum, 150-1 albulae, sp. nov., 151 Diplangus, 150 Diplectrum formosum, 164, 169 Diplodus holbrooki, 169 Diplomonorchis, 156 Diploproctodaeum, 141 Distomum, 123-5, 135, 137, 141, 143, 152, 160, 165-7 Dobie, J. L., article, 59-63 Dundee, H. A., article, 41-58 Durangonella, 2 Echinostephanus, 160 Ectenurus, 163-4 Elops saurus, 164, 169 Endocotyle, 101 Enenterum, 134 Epinephelus adscensionis, 144, 169 striatus, 144, 164, 169 Equetus acuminatus, 146, 169 Erotelis smaragdus, 169 Etheostoma asprigene, 106, 110-112 blennioides, 111 caeruleum, 112 INDEX TO AUTHORS AND SCIENTIFIC NAMES—Continued collettei, sp. noy., 106-112 whipplii, 111 zonale, 111 Eucinostomus gula, 143, 157, 164, 169 Eupomacentrus fuscus, 169 leucostictus, 151, 169 variabilis, 169 Felis domesticus, 85, 87 Fernandinella, 79 Fistularia tabacaria, 169 Floridicthys carpio, 169 Fundulus similis, 169 Galeichthys felis, 168 Gam, A. A., article, 64-69, 70-74 Gasterostomum, 122 Genitocotyle, 148 Genolopa, 152-3 Gobiesox strumosus, 169 Gobionellus smaragdus, 169 stigmaticus, 169 Gonocercella, 166 Graptemys, 85 Groover, R. D., article, 75-80 Gymnothorax nigromarginatus, 170 Gynarcotyla adunca, 102 nassicola, 103 Haemulon aurolineatum, 138, 152, 164, 170 carbonarium, 138, 150, 160, 165, 170 flavolineatum, 152, 164, 170 parrai, 138-9, 150, 152-3, 164-5, 170 plumieri, 152-4, 170 sciurus, 124, 150, 152-4, 170 Halichoeres bivittatus, 144, 148, 170 pictus, 145, 149, 170 poeyi, 170 radiatus, 145, 148, 170 Hamacreadium, 143 Haplosplanchnus, 131-2 Helicometra, 144 Helicometrina, 144-5 Hemiurus, 162 Hexangitrema, 128 Hippocampus erectus, 148, 165, 170 zosterae, 170 Hippolyte, 116 Histrio histrio, 170 Hofstetter, A. M., article, 75-80 Holacanthus isabelita, 124, 170 Homalometron, 138 Horatrema, 146 Hormotilopsis, 75, 77, 79 Hurleytrema, 157 pyriforme, sp. nov., 157 Hymenocotta manteri, sp. nov., 132 Hypocreadium, 136 Infundibulostomum, 124 Knapp, L. W., article, 106-112 Koellikeria, 167 Kyphosus sectatrix, 121, 131, 134, 165, 170 Labrisomus kalisherae, 145, 170 nuchipinnis, 145, 170 Lactophrys quadricornis, 133, 137, 170 Lagodon rhomboides, 124-5, 136, 151, 162, 164, 170 Lasiotocus, 152-155 albulae, sp. nov., 155 haemuli, sp. nov., 154 mugilis, sp. nov., 153 Lebouria, 143 Lechradena, 159 Lecithaster, 165 Lecithochirium, 164 Lecithostaphylus, 152 Lemna minor, 10-11 Lepidauchen, 130 Lepisosteus, 16 Lepocreadium, 135-6 Leptotyphlops dulcis, 42 Leurodera, 164 Levinseniella, 101 amnicolae, 103 Limulus polyphemus, 102 Lutjanus apodus, 145, 159-60, 164, 170 griseus, 143, 158-9, 164, 170 mahogoni, 145, 158-9, 160 synagris, 143, 158-9, 164, 170 Lynx rufus, 16 Lyrodes parvula, 81-2, 88, 91-2, 95-97, 99, 100, 1 Marcia, 149 Macrobrachium ohione, 81-85, 87, 91-95, 97, 100, 102, 104 nipponensis, 102 Macroclemys, 59-63 auffenbergi, sp. nov., 59-63 schmidti, 59-62 temmincki, 59-63 Magnacetabulum, 163 Malacoctenus macropus, 171 Manteria, 161 Manteriella, 146 Maritrema caridinae, 102 obstipum, 103 Megapera, 133 Megasolena, 130 Mesorchis, 124 Metadena, 158-9 Microphallus, 96, 101, 102 claviformis, 102 limuli, 102 minus, 102 Microtus montanus, 85, 87 Miller III, M.C., article, 41-58 Minckley, W. L., article, 2-4 Molenock, J., article, 113-116 Monacanthus ciliatus, 171 hispidus, 136, 150, 171 Monostomum, 128, 158 Mugil cephalus, 132, 152, 171 curema, 171 Multitestis, 143 Mus musculus albinus, 85, 87, 89 INDEX TO AUTHORS AND SCIENTIFIC NAMES—Continued Mycteroperca bonaci, 122, 140, 164, 171 microlepis, 122, 141, 164, 171 Myocastor bonariensis, 10 coypus, 10-17 Myosaccium, 165 Myrica cerifera, 10 Mysidopsis almyra, 116 bahia, sp. noy., 113-116 bigelowi, 113, 116 Nasa obsoleta, 103 Neoapocreadium, 137 Neogenolinea, 165 Neolepidapedon macrum, sp. nov., 141 Neopecoelus, 146 Nicholsina usta, 131, 171 Nicolla halichoeri sp. nov., 148-9 Notropis blennius, 18-22, 24-38 edwardraneyi, sp. nov., 18-39 potteri, 18-9, 21-2, 24-38 Nymphaea, 2 Ocyurus chrysurus, 135, 143, 145, 171 Ogocephalus cubifrons, 145, 160, 164, 171 Oligoplites saurus, 161, 164, 171 Opegaster pritchardae sp. nov., 147 Opheodrys vernalis, 56 Ophichthus gomesi, 156, 171 Opisthadena, 165 Opisthoporus, 140 Opsanus beta, 145, 158, 161, 171 Orconectes, 8 Orthopristis chrysopterus, 152, 164, 171 Overstreet, R. M., article, 119-176 Pachycreadium, 143 Palaemon asperulus, 102 nipponensis, 102 Palaemonetes, 116 pugio, 81, 83, 85, 87, 91-2, 96, 102 Paludiscala caramba, 2 Paracryptogonimus, 159 Paragonimus kellicotti, 70-74 miyazakii, 73-74 ohirai, 73 westermani, 70, 74 Parahemiurus, 162 Paralichthys albigutta, 122, 164, 171 Parectenurus, 163 Penaeus, 116 Plagioporus, 143 Planophila asymmetrica, 79 bipyrenoidosa, 79 laetevirens, 79 terrestris sp. nov., 75-80 Pleurogonius, 128 Podocotyle, 143 Podostemum ceratophyllum, 111 Pomacanthus arcuatus, 124, 128, 171 paru, 124, 171 Pomatomus saltatrix, 164, 171 Populus deltoides, 10 Postmonorchis, 155 Postporus, 140 Prionotus scitulus, 171 Pristigenys alta, 171 Pristisomum, 156 Procambarus ablusus, 5 lagniappe, sp. nov., 5-9 penni, 5, 8, 9 spiculifer, 5, 8, 9 versutus, 5 vioscai, 5 Proctoeces, 124 Proctotrema, 152-3 Procyon lotor, 81, 83, 85, 87, 97, 99 Prosorhynchus, 122 Protosiphon, 77 Pseudemys, 62 Pseudocreadium, 136-7 Pseudohurleytrema, 157 Pseudopecoeloides, 146 Pseudopecoelus, 146 Pseudoplagioporus, 143 Psilostomum, 141 Quercus virginiana, 11 Rana clamitans, 64-68 grylio, 85 pipiens, 67, 85, 87 Rattus norvegicus albinus, 85, 87, 99 rattus, 81, 83, 87 Rhipidocotyle, 122 Ruppia maritima, 116 Salix nigra, 10 Sambucus canadensis, 11 Sardinella anchovia, 135, 162, 165, 171 Schikhobalotrema, 131 Sclerodistomum, 166 Scomberomorus maculatus, 122, 167, 171 regalis, 122-3, 167, 171 Scorpaena grandicornis, 122, 145, 164, 171 pulmieri, 122, 146, 171 Seed, J. R., article, 64-69, 70-74 Selene vomer, 123, 157, 164, 171 Sigmodon hispidus, 85, 87 Siphodera, 158 Sogandares-Bernal, F., article, 70-74 Sonora episcopa, 42 Sparisoma aurofrenatum, 171 Spartina, 82, 85, 92 Spelotrema nicolli, 103 Speocirolana bolivari, 3-4 pelaezi, 3-4 thermydronis, 2-4 Sphaeroides nephelus, 171 spengleri, 141, 171 testudineus, 141, 150, 166, 171 Sphoeroma serratum, 102 Sphyraena barracuda, 126, 171 Spiculotrema, 101 Steganoderma, 152 Stegopa, 158 INDEX TO AUTHORS AND SCIENTIFIC NAMES—Continued Stephanochasmus, 159-60 Stephanostomum, 159-161 Steringotrema, 125 Sterrhurus, 164 Storeria dekayi, 41, 44-45, 55-57 Strongylura timucu, 131, 171 Suttkus, R. D., article, 18-39 Symphurus plagiusa, 172 Syngnathus floridae, 172 louisianae, 172 scovelli, 172 Synodus foetens, 163-164, 172 Tantilla gracilis, 42 Tergestia, 123-124 Tetraciella, 79 Thamnophis radix, 56 Theledra, 123 Thysanopharynx, 133 Trachinotus carolinus, 172 falcatus, 145, 157, 160, 172 Trifolium, 10 Tropidoclonion lineatum, 42 Trypanosoma rotatorium, 64-68 Trypanosome pipientis, 67 Tylosurus crocodilus, 152, 172 Ulmus americanus, 10 Vallisneria, 8 Vitellibaculum, 130 Warkentin, M. J., article, 10-17 Wolfia, 10-11 Xystretrum, 149-150 3 2044 093 361 053 Date Due ~ ~ . 7 ree : - ~ m ; ; : 0s ew ene ee epee ashes . oe rere thes hey tars salcye : PAA aan coe Aan a ie eae: a ed ches cereborwy AA Roeatartes