HARVARD UNIVERSITY m m LIBRARY OF THE Museum of Comparative Zoology VOLUME 7 1959 TULANE UNIVERSITY NEW ORLEANS TULANE STUDIES IN ZOOLOGY is devoted primarily to the zoology of the waters and adjacent land areas of the Gulf of Mexico and the Caribbean Sea. 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 exchanging the entire series. Manuscripts submitted for publication are evaluated by the editor and by an editorial committee selected for each paper. Contributors need not be members of the Tulane University faculty. MEMBERS OF THE EDITORIAL COMMITTEES FOR PAPERS PUBLISHED IN THIS VOLUME Joseph T. Bagnara, University of Arizona, Tuscon, Arizona Frank A. Brown, Jr., Northwestern University, Evanston, Illinois Raymond M. Cable, Purdue University, Lafayette, Indiana Fred R. Cagle, Tulane University, New Orleans, Louisiana Fenner A. Chace, Jr., United States National Museum, Washington, D. C. Robert Ph. Dollfus, Museum National d'Hstoire Naturelle. Paris, France William E. Fahy, University of North Carolina, Morehead City, North Carolina Kenneth L. Gosner, Newark Museum, Newark, New Jersey Horton H. Hobbs, Jr., University of Virgin-a, Charlottesville, Virginia Robert F. Hutton, Florida State Board of Conservation, St. Petersburg, Florida H. W. Manter, University of Nebraska, Lincoln, Nebraska M. Graham Netting, Carnegie Museum, Pittsburgh, Pennsylvania C. Richard Robins, University of Miami, Miami, Florida Robert B. Short, Florida State University, Tallahassee, Florida Franklin Sogandares-Bernal, Tulane University, New Orleans, Louisiana Royal D. Suttkus, Tulane University, New Orleans, Louisiana H. Marguerite Webb, Goucher C^ollege, Towson, Maryland Austin B. Williams, University of Nortii Carolina, Morehead City, North Carolina Mils. r?"\ ZOOL MAY 1 0 1960 UNIVERSiTY CONTENTS OF VOLUME 7 NUMBER PAGE 1. AN ILLUSTRATED KEY TO THE CRAWFISHES OF LOUISIANA WITH A SUMMARY OF THEIR DISTRIBUTION WITHIN THE STATE (DECA- PODA, ASTACIDAE) George Heni-y Penn 3 COMPARISON OF THE CHROMATOPHOROTROPINS OF TWO CRAY- FISHES WITH SPECIAL REFERENCES TO ELECTROPHORETIC BE- HAVIOR Milton Fingerman 21 2. A REVIEW OF THE SEABASSES OF THE GENUS CENTROPRISTES (SERRANIDAE) Rudolph J. Miller 33 3. DIGENETIC TREMATODES OF MARINE FISHES FROM THE GULF OF PANAMA AND BIMINI, BRITISH WEST INDIES Franklin Sogandares-Bernal 69 4. PARASITES OF THE COMMERCIAL SHRIMPS, PENAEUS AZTECUS IVES, P. DUORARUM BURKENROAD, AND P. SETIFERUS (LINNAEUS) Dwayne Nathaniel Kruse 123 THE LARVA OF THE OAK TOAD, BUFO QUERCICUS HOLBROOK E. Peter Volpe and James L. Dobie 145 Printed in the U.S.A. at New Orleans, by Hauser Printinc Co., Inc. a If Volume 7, Number 1 April 23, 1959 AN ILLUSTRATED KEY TO THE CRAWFISHES OF LOUISIANA WITH A SUMMARY OF THEIR DISTRIBUTION WITHIN THE STATE (Decapoda, Astacidae) GEORGE HENRY PENN, DEPARTMENT OF ZOOLOGY, TULANE UNIVERSITY, NEW ORLEANS, LOUISIANA COMPARISON OF THE CHROMATOPHOROTROPINS OF TWO CRAYFISHES WITH SPECIAL REFERENCE TO ELECTROPHORETIC BEHAVIOR MILTON FINGERMAN, DEPARTMENT OF ZOOLOGY, NEWCOMB COLLEGE, TULANE UNIVERSITY, NEW ORLEANS, LOUISIANA TULANE UNIVERSITY NEW ORLEANS TULANE STUDIES IN ZOOLOGY is devoted primarily to the zoology of the waters and adjacent land areas of the Gulf of Mexico and the Caribbean Sea. 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 ex- changing the entire series. Manuscripts submitted for publication are evaluated by the editor and by an editorial com- mittee selected for each paper. Contributors need not be members of the Tulane Univer- sity faculty. Manuscripts should be submitted on good paper, as original typewritten copy, double- spaced, and carefully corrected. Two carbon copies in addition to the original will help expedite editing and assure more rapid publication. An abstract not exceeding three percent of the length of the original article must accom- pany each manuscript submitted. This will be transmitted to Biological Abstracts and any other abstracting journal specified by the writer. Separate numbers or volumes may be purchased by individuals, but subscriptions are not accepted. Authors may obtain copies for personal use at cost. Address all communications concerning manuscripts and editorial matters to the editor; communications concerning exchanges, and orders for individual numbers to the Librarian, Meade Natural History Library. When citing this series authors are requested to use the following abbreviations: Tulane Stud. Zool. Price for this number: $0.60. George Henry Penn, Editor Meade Natural History Library, Tulane University, New Orleans, U. S. A. Assistants to the Editor: Robert K. Chipman Guy Marlow TULANE STUDIES IN ZOOLOGY KARvARD UNIVERSITY Volume 7, Number 1 April 23, 1959 CONTENTS An Illustrated Key to the Crawfishes of Louisiana with a Summary OF Their Distribution within the State George Henry Perm 3 EDITORIAL COMMITTEE: Fenner a. Chace, Jr., Curator, Division of Marine Invertebrates, United States National Museum, Washington, D. C. HORTON H. HOBBS, JR., Professor of Biology, University of Virginia, Charlottes- ville, Virginia Austin B. Williams, Associate Professor of Zoology, Institute of Fisheries Re- search, University of North Carolina, Morehead City, North Carolina Comparison of the Chromatophorotropins of Two Crayfishes with Special Reference to Electrophoretic Behavior Milton Fingerman . 21 EDITORIAL COMMITTEE: Joseph T. Bagnara, Assistant Professor of Zoology, University of Arizona, Tuc- son, Arizona Frank A. Brown, Jr., Professor of Biology, Northwestern University, Evans- ton, Illinois H. Marguerite Webb, Assistant Professor of Physiology, Goucher, College, Towson, Maryland AN ILLUSTRATED KEY TO THE CRAWFISHES OF LOUISIANA WITH A SUMMARY OF THEIR DISTRIBUTION WITHIN THE STATE (DeCAPODA, ASTACIDAE) ^ GEORGE HENRY PENN, Department of Zoology, Tulane University, New Orleans, Louisiana In 1940 the writer began a study of the crawfishes of Louisiana. Since that date he has collected in all parts of the State (Map 1 ) and examined over 20,000 specimens, about a third of which are currently in museum collections. A number of papers on this group of our fauna have been published. Four of these (Penn, 1950b, 1952a, 1956; Penn and Marlow, 1959) summarized in de- tail what was known at the time of the taxonomy, geographical and ecological dis- tribution, and life history of each of the species of the four genera represented in the State. In the present contribution my pur- pose is to gi\e a key that will be useful to any zoologist in identifying the 29 species and subspecies known to be present in Lou- isiana, and to bring the geographic data up- to-date. Taxonomic Characters In general, crawfishes of the subfamily Cambarinae (as defined by Hobbs, 1942a) are identified primarily on the basis of the structure of the first pleopods (or, swim- merets) of the adult males. Other super- ficial morphological characters of the males and females may be useful, but are of sec- ondary importance. Adult males crawfishes exhibit two distinctly different and alter- nating external appearances; these are re- ferred to as Form I and II ( or, first and second form) males. Form is related direct- ly to the seasonal reproductive cycle: males of Form I are capable of copulation, those of Form II ( and juveniles ) are not. A Form I male may be distinguished by the sharper appearance and corneous ( yellow- ish ) condition of one or more of the apical elements of the first pleopods (figs. 1, 2) and the presence of prominent hooks on the 1 Many of the specimens collected be- tween July 1954 and June 1957 were a "by-product" of researches supported by the National Science Foundation under g-rants G-947 and G-2330. Most of the illus- trations were completed from the writer's pencil sketches by Miss Sue Blackshear. Figures 1-4. First pleopod of male craw- fishes. 1. Hypothetical first pleopod: I^ mesial process, II = cephalic process, III = centrocaudal process, IV ^ caudal process, III-V =; central projection, C = caudomesial margin (after Hobbs, 1942). 2. Mesial view of first pleopod of form I male of Procam- barus blandvigi acutus. 3. Mesial view of first pleopod of form II male of P. b. acutus. 4. Mesial view of first pleopod of form I male of Orconectes palmeri longima)ius: A = total length, B ^ length of central pro- jection, C ^ length of mesial process, D ^ length of distal mesial shaft. ischiopodites of one or more pairs of the pereiopods (or, walking legs) (fig. 6 — /). Form II males have blunter apical elements on the first pleopods ( fig. 3 ) and have the hooks of the ischiopodites less well- developed. Although there are considerable differ- ences in the shapes and proportions of the apical parts of the first pleopods of the dif- ferent species, the pleopods have been ho- mologized with a four-parted structure (fig. 4 Tulane Studies in Zoology Vol. 7 "*""♦•" Figures 5-6. Crawfish terminology. 5. Whole crawfish, dorsal view: 1 = cephalothorax, 2 = abdomen, 3 = cephalic section of cephal- othorax, 4 = areola (or, thoracic section of cephalothorax) , 5 =^ rostrum, 6 = acumen, 7 ^ antennal scale, 8 = lateral spine of rostrum, 9 ^ lateral spine of cephalothor- ax, I-V ^ pereiopods (or, walking legs). 6. Third pereiopod of form I male of P. b. acHtns: D =: dactylopodite, P = propodite, C = carpopodite, M = meropodite, I = isch- iopodite (with hook), B ^ basipodite, Cx = coxopodite. 1) by Hobbs ( 1942b). All apical elements are present in most species of Procambarus (figs. 2, 11-22); the cephalic element is lacking in species of Cambarellus (figs. 23, 24); and, only the mesial process and cen- tral projection are present in species of Camban/s (figs. 25-28), and Orconectes (figs. 29-35). Other characters frequently used in craw- fish identification include the following ( fig. 5 ) : ( 1 ) rostrum ( length relative to cephalothorax length; and, with or without lateral spines); (2) lateral spines of cep- halothorax ( present or absent, number ) ; ( 3 ) areola ( length relative to length of cephalic section of cephalothorax; whether open or obliterated; and, if open, its length width ratio; (4) antennal scale (shape; length width ratio; length relative to cep- halothorax length; (5) chela (length width ratio; number and arrangement of tubercles on the fingers. The sculpture of the surface of the anmdus ventral is (figs. 36-55), or sperm receptacle, of the female is a useful taxonomic character particularly in the genus Procambarus. Key to Form I Males of Louisiana Crawfishes 1. First pleopod terminating in three or more distinct parts (figs. 2, 11-24) 2 First pleopod terminating in only two distinct parts (figs. 4, 25-35) 16 2. Hooks present on ischiopodites of the 3rd, or 3rd and 4th pereiopods (Genus Procam- barus) 3 Hooks present on ischiopodites of the 2nd and 3rd pereio- pods (Genus Cambarellus) 15 Figures 7-10. Examples of variation in structure of cephalothorax and chela. 7. Ce- phalothorax of Orc(i)i('ctcs hnicifer (ros- ti'um with lateral spines, areola obliter- ated). 8. Cephalothorax of O. cli/peains (rostrum without lateral spines, areola open). 9. Chela and carpus of Cambarus hedgpefhi (two prominent teeth on immova- ble finger). 10. Chela and carpus of Cam- barns diogciies (lioj/ciies (one prominent tooth on immovable finger). No. 1 Penn: Louisiana Crawfishes Figures 11-35. Apices of first pleopods of form I males; mesial view unless otherwise indicated. Pubescence removed from all structures; not drawn to scale. 11. P. tulanei. 12. P. s. simulcDis. 13. P. clarki. 14. P. bivittatns. 15. P. shermani. 16. P. viaeriridis. 17. P. planirostris. 18. P. hiuei. 19. P. vioscai (A — mesial, B — lateral). 20. P. penni (A — mesial, B — lateral). 21. P. natchitochae. 22. P. dnpratzi. 23. C. pner. 24. C. shnfeldti. 25. C. dissitiis. 26. C. oryktes. 27. C. hedgpethi. 28. C. d. Indovicianus. 29. O. beyeri (caudal view). 30. O. clypeatns (caudal view). 31. O. lancifer. 32. O. hobbsi. 33. 0. p. longimanus. 34. O. difficlis. 35. O. hathawayi. Tidane Studies in Zoology Vol. 7 3. Hooks present on ischiopodites of 3rd pereiopods only 4 Hooks present on ischiopodites of 3rd and 4th pereiopods 5 4. Central projection of first pleopod (fiji\ 1) extendinfr well distad of caudal knob (fig-. 11) ; basal structure of caudal knob (lateral view) stvaieht or nearl^' so I'rocaDtbanifi tnlanei Penn Central projection not extend- ing- distad beyond caudal knob (fig. 12); basal struc- ture of caudal knob (lateral view) more-or-less S-shaped Procambarus simidans simnlans (Faxon) 5. Areola obliterated (as in fig. 7), or ver-' narrow (length/ width ratio not less than 8) _. - 6 Areola open (as in fig. 8) and broad (length width ratio not greater than 5) 11 6. Areola obliterated in middle (fig. 58 t Procambams clarki (Girard) Areola open, but very narrow 7 7. Rostrum with lateral spines (as in fig. 7) 8 Rostrum without lateral spines (as in fig. 8) 9 8. Acumen (fig. 5) and lateral spines of rostrum long; are- ola about half length of ce- phalic section of cephalo- thorax (fig. 57) Procambarus bivittatns Hobbs Acumen short, lateral spines of rostrum inconspicuous; areola about one-third length of cephalic section of ce- phalothorax (fig. 56) ._ Procambarus blaudingi acutus (Girard) 9. Mesial process of first pleopod extending distally beyond apices of all other terminal elements (fig. 15) Procambarus sherm.ani Hobbs Mesial process not extending beyond all other terminal elements 10 10. Fiist pleopod with prominent "finger - like" knob on ce- phalic margin (fig. 16) Procrimbarus viacviridis (Faxon) First pleopod with shoulder, but not with prominent knob on cephalic margin (fig. 17) - Procambarus platiirostris Penn 11. Cephalothorax without lateral spine on each side; or, with a verv small one (fig. 60) ; all terminal elements of first pleopod small ; pleopod ap- pears twisted (fig. 18) Procambarus hinci (Ortmann) Cephalothorax with two later- al spines on each side; at least some terminal ele- ments of first pleopod con- spicuous; pleopod not twisted in appearance 12 12. Caudal knob of first pleopod rounded distallv in lateral aspect (fig. 19-B) Procambarus vioscai Penn Caudal knob of first pleopod subacute in lateral aspect (as in fig. 20-B) 13 13. Mesial process extending more- or - less straight distally (rever bent at more than 10° angle) (fig. 21) Procambarus uatchitochae Penn Mesial process extending cau- dad at or greater than 30° angle to shaft of pleopod 14 14. All terminal elements of first pleopod bent caudad at greater than 50° angle (fig. 22) P)-ocambarus dup)-atzi Penn Terminal elements bent cau- dad at about 30° angle (fig. 20-A) Procambarus penni Hobbs 15. All terminal elements of first pleonod straight or nearly so (fig. 24) Cambarellus sltufeldti (Faxon) All terminal elements of first pleopod bent caudad at about 40° angle to shaft (fig. 23) Cambarellus ))ucr Hobbs 16. Terminal elements of first ple- opod heavy, both bent at 90° angle to shaft (figs. 25-28) (Genus Cambarus) 17 Terminal elements straight or gently curved caudad, never both bent at more than 45° angle to shaft (figs. 29-35) (Genus Oiconecfes) 21 17. Hooks on ischiopodites of 3rd and 4th pereiopods Cambarus dissifus Penn Hooks on ischiopodites of 3rd pereiopods only __._ 18 18. Immovable finger of chela with two prominent teeth on op- posable margin (fig. 9) ; base of immovable finger bearded — 19 Figures 36-55. Annuli ventrales of mature females of some Louisiana crawfishes. Not drawn to scale. 36. P. b. acutus. 37. P. bivittatns. 38. P. clarki. 39. P. platiirostris. 40. P. hi)iei. 41. P. tulanei. 42. P. s. simulajis. 43. /'. dupratzi. 44. P. uatchitochae. 45. C. shufeldti. 46. C. hedgpethi. 47. C. orijktes. 48. C. dissifus. 49. (). clipeatus. 50. (). bcyeri. 51. (). hnurfer. 52. (). hohhsi. 53. O. p. lotigitiiauus. 54. O. hathau-aiji. 55. O. difj icilis. No. 1 Venn: Louisiana Crawfishes 8 Tulane Studies in Zoology Vol. 7 Immovable finoer of chela with one prominent tooth on op- posable margin (fig. 10) ; base of immovable finger not bearded 20 19. Antennal scale widest beyond middle; areola antennal scale ratio not greater than 3.5 Cautbarns hedgj)ctlii Hobbs Antennal scale widest at mid- dle; areola antennal scale ratio rot less than 4.0 Cainbariis oryktes Penn and Marlow 20. Ratio of cephalic section of cephalothorax to areola greater than 1.4; antennal scale length/ width ratio less than 1.45 ....Cambariis diogenvs diogenes Girard Ratio of cephalic section of cephalothorax to areola less than 1.3; antennal scale length width ratio greater than 1.45 Cambarus diogenes Indovicianus Faxon 21. The two rami of first pleopod unequal in length (central projection at least twice length of mesial process) (fig. 29, 30); areola wide (length width ratio less than 4.0) ; rostrum without later- al spines (as in fig. 8) 22 The two rami of first pleopod subequal in length (figs. 31- 35) ; areola narrow or oblit- erated; rostrum with lateral spines (as in fig. 7) ..-23 22. Central projection flattened, somewhat sickle-shaped, about twice length of mesial process (fig. 29) Orconectes beyeri Penn Central projection subcylindri- cal and more-or-less straight beyond basal bend, about three times length of mesi- al process (fig. 30) Orconectes clypeatns (Hay) 23. The two rami of pleopod short, stout, gently curved caudtid; central projection bladelike (fig. 31) ; rostrum and an- tennal scale exceptionally elongate (fig. 7) Orconectes Imicifer (Hagen) The two rami of pleopod long, more-or-less setiform, and at least mesial process strong- ly recurved caudad ( figs. 32-35) ; rostrum and anten- nal scale not exceptionally elongate 24 24. Central projection longer than distal mesial shaft (fig. 4) ...25 Central projection shorter than distal mesial shaft (fig. 4) 28 25. Areola narrowly open (fig. 68) ; ratio of cephalothorax length/pleopod length usu- ally less than 3.0; see also fig". 32 0)co)iectes hobbsi Penn Areola usually obliterated; ra- tio of cephalothorax length pleopod length usually great- er than 3.0 where ranges of O. p. creolainis and O. hobbsi overlap (Maps 11, 12) 26 26. West of the Mississippi River in tributaj'ies of Red River and rivers draining into Gulf of Mexico (Map 11) Orconectes palmeri longimanus (Faxon) East of Mississippi River 27 27. In streams tributary to Lake Pontchartrain and Peail Riv- er (Map 11) Orconectes palmeri creolanus (Creaser) In streams tributary to east side of Mississippi River (Map 11) Orconectes palmeri palmeri (Faxon) 28. Central projection greater than half length of distal mesial shaft (fig. 4) ; apex of mesial process bent cau- dad at about 90° angle to shaft of pleopod (fig. 34) Orconectes difticilis (Faxon) Central projection less than one third length of distal mesial shaft (fig. 4) ; apex of mesial process bent cau- dad at no greater than 30° angle to shaft of pleopod (fig. 35) Orconectes hathatvayi Penn Annotated List of Species and Subspecies In the following list, the paragraphs headed "recognition characters" supplement those "key" characters used above, and are intended to help students in associating females and form II males with form I males from the same collection; the para- graphs headed "distribution" include brief statements on ecological as well as geo- graphical range in Louisiana. A supple- mentary parish check list is included ( fig. 86) following this annotated list. Refer- ences to separately published studies of life cycles are cited under each species wiiere applicable, but no attempt has been made to cite references to studies of an experi- mental nature. Note should be made, how- ever, that three of our crawfishes ( P. clarki, C. sbujeldti. and O. clypeatns ) have proved No. 1 Pe7in: Louisiana Crawfishes Figures 56-70. Cephalothoraxes drawn to scale. 56. P. b. acutns. €0. P. hinei. 61. P. s. simidans. 65. C. oryktes. 66. C. dissitus. 67 hathawayi. of form I males of some Louisiana crawfishes. Not 57. P. hivittatus. 58. P. clarki. 59. P. planirostris. 62. P. dupratzi. 63. C. shiifeldti. 64. C. hedgpethi. . O. beyeri. 68. O. hobbsi. 69. O. ^J. longimayins. 70. O. 10 Tulane Studies in Zoology Vol. 7 useful in experimental studies of various aspects of physiology, behavior, and genetics. PROCAMBARUS BLANDINGI ACUTUS (Girard, 1852) (Figures 2, 3, 6, 36, 56, 71) Recognitio7i characters. — Rostrum with small lateral spines near apex ( absent in some specimens ) ; areola narrow, but open; one lateral spine on each side of cephalo- thorax; chela long and slender. Range of cephalothorax length of form I males: 25 to 57 mm. This is the so-called "white" or "river crawfish" of the markets; when alive, mature specimens have a pinkish over- all color with the abdomen bearing a wide, mid-dorsal, darker longitudinal stripe. Distribution. — The crawfish is ubiquitous in habitat, but has been found most abun- dantly in creeks, rivers, ponds, borrow pits, and roadside ditches. Statewide ( Map 2 ) and recorded from 53 of the 64 parishes, this crawfish eventually should be found in every parish when more intensive collecting is carried on. PROCAMBARUS BIVITTATUS Hobbs, 1942b (Figuresl4, 37, 57) Recognition characters. — Rostrum long, with conspicuous lateral spines and con- spicuous acumen; areola narrow, but open; one large lateral spine on each side of cep- halothorax; chelae long and slender. Super- ficially, preserved individuals resemble those of P. h. acutus, but when alive they are dis- tinct in having a pair of dark, longitudinal stripes along each side of abdomen. Range of cephalothorax length of form I males: 47 to 59 mm. Distribution. — Restricted in Louisiana to the Pearl River drainage (Map 5) where they have been taken only from the deeper, shaded pools of the smaller tributaries in St. Tammany and Washington parishes. PROCAMBARUS CLARKI (Girard, 1852) (Figuresl3, 38, 58,72) Recognition characters. — Rostrum with small lateral spines; areola obliterated in middle; one lateral spine on each side of cephalothorax; chela long but heavy; cep- halothorax conspicuously tuberculate. Range of cephalothorax length of form I males: 31 to 57 mm. Known as the "swamp" or "red" crawfish in the markets, this animal when mature has a dark red overall color; however, in recent years anomalous bright blue specimens have appeared in several parts of the state ( Penn, 1951 ). Distribution. — Statewide, and recorded from 41 of the 64 parishes (Map 3), this crawfish should be found in many more parishes when intensive collecting is done. It is most abundant in shallow, temporary bodies of water such as marshes, swamps, and roadside ditches, but it is taken com- mercially in the deep waters of the swamps near Pierre Part ( Assumption Parish ) , and a few individuals have been found even in slow-flowing streams and springs. The life history has been carefully studied by Viosca (1939, 1953) and Penn (1943). PROCAMBARUS PLANIROSTRIS Penn, 1953c (Figuresl7, 39, 59,73) Recognition characters. — Rostrum vv'ith- out lateral spines; areola narrow; lateral spine on cephalothorax reduced to the size of a small tubercle; chela narrow. Range of cephalothorax length of form I males: 22 to 28 mm. Not a conspicuously colored crawfish; it has a light tan to olive back- ground dorsally, and abdomen has a wide, mid-dorsal, longitudinal stripe of slightly darker hue. Distribution. — In Louisiana this crawfish is restricted to the Florida parishes ( Map 6) where it is usually found in shallow bur- rows and temporary surface waters. PROCAMBARUS VIAEVIRIDIS (Faxon, 1914) ( Figure 16) Recognition characters.. — Rostrum with- out lateral spines; areola narrow; without lateral spines on cephalothorax; chela long and narrow. Range of cephalotliorax length of form I males: 23 to 30 mm. Distribution. — Hitherto unrecorded from the State, this crawfish has been collected at one locality only, viz. Wham, Ouachita Parish (Map 6). No. 1 Venn: Louisiana Crawfishes 11 Fig'ures 71-76. Chelae of some form I male Louisiana crawfishes. Pubescence re- moved from all structures; not drawn to scale. 71. P. h. acutus. 72. P. clarki. 73. P. lAanirostris. 74. P. hinei. 75. P. s. simidans. 76. P. dupratzi. PROCAMBARUS HINEI (Ortmann, 1905) (Figures 18,40,60,74) Recognition characters. — Rostrum taper- ing sharply, with slight indication of lateral spines; areola broad; without lateral spines on cephalothorax (or, with only a small one on each side in some specimens ) ; chela subcylindrical, palm longer than fingers in males. Range of cephalothorax length of form I males: 11 to 20 mm. A redescrip- tion of the species was prepared by Penn (1953b). Dorsally it has a pair of con- spicuous dark, broad, parallel, longitudinal stripes that run on each side of the areola; on the abdomen the stripes converge slightly and become narrower. Distribution. — This crawfish is abundant in certain localities, but its distribution is spotty. It has been found primarily in semi- permanent ponds, borrow pits, and ditches in twelve parishes ( May 5 ) within the Quarternary lowlands of the state. PROCAMBARUS SHERMANI Hobbs, 1942b (Figure 15) Recognition characters. — Rostrum with- out lateral spines; areola relatively narrow; with one small lateral spine on each side of cephalothorax; palm of chela usually beard- ed in form I males. Range of cephalo- thorax length of form I males: 28 to 40 Figures 77-85. Chelae of some form I male Louisiana crawfishes. Pubescence re- moved from all structures; not drawn to scale. 77. C. shnfeldti. 78 C. onjktes. 79. C. dissitHS. 80. O. cly peat lis. 81. O. heijeri. 82. O. hobbsi. 83. O. lancifer. 84. O. p. longimanns. 85. O. difficilis. 12 Tulane Studies in Zoology Vol. 7 Maps 1-6. Distribution of Louisiana crawfishes. No. 1 Penn: Louisiana Crawfishes 13 mm. This is a drab-colored crawfish with- out conspicuous markings. Distribution. — Hitherto unrecorded from the State, this crawfish has been found in ditches in only several locahties; these are near Talisheek and near Hickory in St. Tammany Parish (Map 6). PROCAMBARUS TULANEI Penn, 1953a (Figures 11,41) Recognition characters. — Rostrum spatu- late and without lateral spines; areola rela- tively wide ( length width not greater than 10); without lateral spines on cephalo- thorax; chela long and narrow; palm of chela of form I males usually heavily bearded. Range of cephalothorax length of form I males: 43 to 55 mm. Distribution. — Mature individuals have been taken from burrows in the banks of streams tributary to the Ouachita and Red rivers in parishes lying between the Red River and the Arkansas-Louisiana line (Map 6). Juveniles have been found primarily in the streams of the same area. PROCAMBARUS SIMULANS SIMULANS (Faxon, 1884) (Figures 12,42,61,75) Recognition characters. — Rostrum spatu- late and without lateral spines; areola rela- tively narrow (length width not less than 15) ; one much reduced lateral spine on each side of cephalothorax; chela long, but palm wide. Range of cephalothorax length of form I males: 29 to 50 mm. Distribution. — This western crawfish has been found mainly in creeks and flowing ditches in the Sabine, Red, Calcasieu and Bayou Teche river systems, mostly west of the Red River (Map 6). PROCAMBARUS VIOSCAI Penn, 1946 (Figure 19) Recognition characters. — Rostrum with prominent lateral spines and usually with median carina; areola broad; two lateral spines on each side of cephalothorax; chela about three times longer than wide. Range of cephalothorax length of form I males: 18 to 42 mm. Basic color of this crawfish is light olive-green or tan; cephalic and thoracic sections of cephalothorax in dorsal view each with a pair of irregular, wide, dark olive-green, longitudinal stripes that curve toward the midline at the posterior margin of their respective sections (each is somewhat suggestive of a lyre); areola of same dark color; sripes of cephalothorax continue as narrower, lateral markings on abdomen. Distribution. — In Louisiana this crawfish has been found in streams west of the Mis- sissippi in the Ouachita and Red River drainages, and east of the Mississippi in tributaries of the Mississippi River and Lake Pontchartrain ( Map 4 ) . PROCAMBARUS PENNI Hobbs, 1951 ( Figure 20 ) Recognition charatcers. — Rostrum with prominent lateral spines and wtihout a me- dian carina; areola broad; two lateral spines on each side of cephalothorax; chela about 3.5 to 4 times longer than wide. Range of cephalothorax length of form I males: 23 to 43 mm. Basic color is light chocolate-tan; marking of cephalothorax essentially as in P. vioscai except that the color is blackish brown; in lateral view each abdominal seg- ment has a V-shaped marking of dark brown, and in the opening of each V there is an irregular spindle-shaped area of chimney red. Distribution. — In Louisiana this crawfish has been found only in streams of the Pearl River drainage (Map 4). PROCAMBARUS DUPRATZI Penn, 1953d (Figures 22, 43, 62, 76) Recognition characters.. — Rostrum with prominent lateral spines and some speci- mens with a weak median carina; areola broad; two lateral spines on each side of cephalothorax; chela about three times long- er than wide. Range of cephalothorax length of form I males: 23 to 42 mm. Basic color light olive-tan; pattern of cephalo- thorax essentially as in P. vioscai, but the dark color is olive; abdomen also similar to that of P. vioscai, but dark color is blue- black, and the stripes are wider and more irregular. Distribution. — In Louisiana this crawfish has been found only in streams of the Sa- bine and Calcasieu river systems (Map 4). 14 Tulane Studies in Zoology Vol. 7 PROCAMBARUS NATCHITOCHAE Penn, 1953cl (Figures 21,44) Recognition characters. — Rostrum with prominent lateral spines and usually with median carina; areola broad; two lateral spines on each side of cephalothorax; chela about 3-5 times longer than wide. Range of length of cephalothorax of form I males: 23 to 43 mm. Color and color pattern es- sentially as in P. dupratzi. Distribution. — In Louisiana this crawfish has been found in creeks tributary to the Red, Calcasieu, and Bayou Teche river sys- tems ( Map. 4 ) . CAMBARELLUS SHUFELDTI (Faxon, 1884) (Figures 24, 45, 63, 77) Recognition characters. — Rostrum with lateral spines; areola broad (length/width ratio 3.5 to 5 ) ; one lateral spine on each side of cephalothorax; chela relatively short, with palm inflated ( subcylindrical ) and longer than fingers. Range of cephalothorax length of form I males: 8 to 11 mm. Color pattern may be one of two types: two rows of dark spots on either side of the areola and down the abdomen, or two dark stripes in place of the spots. Distribution. — The dwarf crawfish has been found in all types of shallow, semi- permanent bodies of water in and east of the Mississippi valley and the Atchafalaya River, and in the Red River valley to Texas ( Penn and Hobbs, 1958) (Map 7). The life history has been studied by Penn (1942, 1950b). CAMBARELLUS PUER Hobbs, 1945 ( Figure 23 ) Recognition characters. — Rostrum with lateral spines; areola broad (length width ratio 5 to 6); lateral spines of cephalothorax small or absent; chela as in C. shufeldti. Range of cephalothorax length of form I males: 8 to 1 1 mm. Color pattern as in C. shufeldti. Distribution. — This dwarf crawfish has been found in the same types of habitats as C. shufeldti in and west of the Mississippi valley (Map 7) overlapping the range of C. shufeldti in the Mississippi valley and between the Mississippi and Atchafalaya rivers. CAMBARUS DIOGENES DIOGENES Girard, 1852 ( Figure 10) Recognition characters. — Rostrum with- out lateral spines; antennal scale length/ width ratio less than 1.45; sides of antennal scale nearly straight, and parallel; ratio of cephalic section of cephalothorax to areola greater than 1.40; chela not bearded, im- movable finger with one prominent tooth. Range of cephalothorax length of form I male: 32 to 54 mm. Ground color of cep- halothorax and abdomen shades dorsal to ventral from olive-green to brown; chela with red tips on fingers. Distribution. — Typically this crawfish is a chimney builder and adults may have an extensive system of underground burrows. The juveniles are found occasionally in shal- low burrows, but more frequently in a variety of aquatic habitats, particularly small creeks. The crawfish has been taken in twenty-two parishes throughout the state (Map 8) and should be anticipated in at least twenty more. CAMBARUS DIOGENES LUDOVICIANUS Faxon, 1884 (Figure 28) Recognition characters. — Rostrum with- out lateral spines; antennal scale length/ width ratio greater than 1.45, shape as in C. d. diogenes: ratio of cephalic section of cephalothorax to areola less than 1.30; chela as in C. d. diogenes. Range of cephalothorax length of form I males: 33 to 43 mm. Abdomen with three longitudinal red stripes; base of telson and uropods as well as margins of the uropods marked with red; basic color bluish; colors are most brilliant in smaller individuals. Distribution. — This burrowing, chimney- building crawfish is found only in the south- eastern part of the State (Map 8). CAMBARUS HEDGPETHI Hobbs, 1948 (Figures 27, 46, 64) Recognition characters. — Rostrum with- out lateral spines; areola obliterated; an- tennal scale length width ratio not over 2.5; lateral margin of antennal scale straight, No. 1 Penn: Louisiana Crawfishes 15 ^ C d'ogenes diogenes • C diogenes ludoncianus Maps 7-12. Distribution of Louisiana crawfishes. 16 Tulane Studies in Zoology Vol. 7 medial margin strongly curved; chela with prominent tuft of plumose hairs along base of immovable finger; immovable finger with two prominent teeth. Range of cep- halothorax length of form I males: 21 to 31 mm. This is a rather drab-colored craw- fish: cephalothorax and abdomen olive-tan dorsally, shading into a lighter hue of the same color laterally; abdomen with two faint, wide, bluish stripes extending its full length and onto the telson and uropods. Distribution. — About one-fourth of the specimens seen have been taken from bur- rows; the remainder were from a wide variety of shallow bodies of water, includ- ing roadside ditches, overflow puddles, sloughs, swamps, swamp pools and ponds. This crawfish has a statewide distribution; at present it is known from localities in twenty-seven parishes ( Map 9 ) • CAMBARUS ORYKTES Penn and Marlow, 1959 (Figures 26, 47, 65, 78) Recognition characters. — Rostrum with- out lateral spines; antennal scale not extend- ing beyond tip of rostrum ( length 'width ratio not over 2.5); areola obliterated; lat- eral spines on cephalothorax absent; chela relatively short (cephalothorax length/chela length ratio not less than 1.3); fingers of chela gently curved inward (i.e., ventrally); basal half of immovable finger heavily bearded in form I males. Range of cephalo- thorax length of form I males: 22 to 24 mm. This is an inconspicuously colored craw- fish in which the cephalothorax is blackish- brown dorsally, shading ventrally into tan; abdomen with a wide mid-dorsal stripe of blackish-brown that narrows posteriorly; lateral to the stripe the abdomen has a fine reticulate pattern of blackish-brown on a tan background. Distribution. — Most of the specimens ex- amined were dug from burrows in dry roadside ditches in the vicinities of Coving- ton and Alton, St. Tammany Parish (Map 9). CAMBARUS DISSITUS Penn, 1955 (Figures 25, 48, 66, 79) Recognition characters. — Rostrum with- out lateral spines; antenna! scale short, not reaching tip of rostrum ( length width ratio ranges from 1.9 to 2.8 j; areola obliterated in most specimens (if open, its length/ width ratio not less than 35); lateral spines of cephalothorax absent; chela relatively short ( cephalothorax length chela length ratio not less than 1.35 ) ; immovable finger without beard, and with only one prominent tooth; both fingers curved ventrally. Range of cephalothorax length of form I males: 24 to 32 mm. Distribution. — This crawfish is known only from specimens taken from shallow burrows in the upland shortleaf and longleaf pinehills of Caldwell and Lincoln parishes (Map 9). ORCONECTES CLYPEATUS (Hay, 1899) (Figures8, 30, 49,80) Recognition characters. — Rostrum spaai- late, without lateral spines; areola broad (length width ratio averages 4); lateral spines of cephalothorax absent (or, if pres- ent, greatly reduced and inconspicuous ) ; chela with palm inflated and subcylindrical, fingers and palm subequal in length. A re- description of the species was prepared by Creaser (1933). Range of cephalothorax length of form I males: 10.5 to 18 mm. Basic color olive-tan on cephalothorax and abdomen; two darker blue-black, longitu- dinal stripes extend from behind eyes, on either side of areola, to end of abdomen. Distribution. — This little crawfish is found in a variety of shallow aquatic habi- tats, including such diverse locations as roadside ditches, ponds, creeks, pineland sloughs, swamps, borrow pits, and burrows. With the exception of the recent alluvial deposits it has a statewide distribution ( Map 10). The life history has been studied by Smith (1953). ORCONECTES BEYERI Penn, 1950a (Figures 29, 50,67,81) Recognition characters. — Rostrum, lateral spines of cephalotliorax, and chela as in O. clypeatus; areola broad (length/'width ratio about 3.0). Range of length of cephalo- thorax of form I males: 11 to 18 mm. Distribution. — This crawfish is known only from several localities in DeSoto and Natchitoches parishes (Map 10). Appar- No. 1 Penn: Louisiana Crawfishes 17 <0 <0 k. 5 10 5 1 •** < 5 1 rn\T/7rTPC DiT:v!iriTi^ ^^^ genus Cambarus from Texas, with UKLUiytK^l c,c> UlttiHHd notes on the distribution of Cambarus (Faxon, 1898) fodiens (Cottle). Proc. U. S. Nat. Mus., (Figures 34, 55, 85) 98:223-231. ... r- • II 1951. A new crayfish of Kecogmtwn characters. — Superficially re- the genus Procambarus from Louisiana, sembling O.. hathaivayi in most respects ex- with a key to the species of the Spi- cept structure of first pleopod of male and culifer group. Jour. Wash. Acad. Sci., annulus ventralis of female. Range of cep- ^ .'t.'.^^,. ,,, , irr T 1 -^c Ortmann, a. E. 1905. A new species of halothorax length of form I males: .5 to Cambarus from Louisiana. Ohio Nat., 45 mm. 6: 401-403. Distrihution. — This crawfish is known Penn, George Henry 1942, Observations only from streams in the headwaters of «" ^J^^ biology of the dwarf crawfish, T3 -TL ji^i- T.- • Cam,oareihis shufeldtn (Faxon). Amer. Bayou Teche and the Calcasieu River in ^^^i ^„^_^ 28(3): 644-647. Louisiana (Map 12). 1943_ ^ study of the life cycle of the Louisiana red-crawfish, Cam- RefeRENCES Cited hams clarkii Girard. Ecology, 24(1): 1- Creaser, Edwin P. 1933. Descriptions of 1^. some new and poorly known species of 1946. A new crawfish of North American crayfishes. Occ. Pap. the genus Procambarus from Louisiana. Mus. Zool. Univ. Mich., No. 275: 1-21. Jour. Wash. Acad. Sci., 36(1) : 27-29. Faxon, Walter 1884. Descriptions of new 1950a. A new crawfish species of Cambarus to which is added a of the genus Orconectes from Louisiana, svnonymical list of the known species of Ibid., 40 (5) : 166-169. Cambarus and Astacus. Proc. Amer. Acad. 1950b. The genus Cam- Arts & Sci., 20: 107-158. ""'6are«Ms in Louisiana. Awer. Midi. Nat, ^— -^- - 1898. Observations on the 44(2) : 421-426. Astacidae m the United States National 1950c. A new Orconectes Museum and in the Museum of Compara- """from""the Pontchartrain watershed in tive Zoology, with descriptions of new Louisiana and Mississippi. Jour. Wash. species. Proc. U. S. Nat. Mus., 20: 643- Acad. Sci., 40(11): 381-384. 1951. A color anomaly of -....__.__ 1914. Notes on the cray- the Louisiana red crawfish, Procambarus fishes in the United States National Mu- darkii (Girard). Proc. La. Acad. Sci., seum and the Museum of Comparative 14; 66-67. Zoology with descriptions of new species * ' .,„_., mi. r\ and subspecies to which is appended a —---._- .1952a. The genus Oreo- catalog of known species and subspecies. "^^f^f ^"o^^T^'^"^- ''' ' Mem. Mus. Comp. Zool, 40(8): 347-427. 47(d): 743-/48. Girard, Charles 1852. A revision of the -... - 1952b. A new crawfish North American Astaci, with observa- of the Vtrtlis section of the genus Orco- tions on their habits and geographical nectes. Chicago Acad. Sci., Nat. Hist. distribution. Proc. Acad. Nat. Sci. Phila., Misc., No. 109: 1-7. 6: 87-91. 1953a. A new crawfish of Hagen, Hermann A. 1870. Monograph of the genus Procambarus from Louisiana the North American Astacidae. Illustr. and Arkansas. Jour. Wash. Acaid. Sci., Cat. Mus. Comp. Zool, 3: 1-109. 43(5): 163-166. Hay, William P. 1899. Description of two 1953b. A redescription of new species of crayfish. Proc. U. S. Nat. the crawfish Procambarus hinei (Oi-t- Mus., 22: 121-123. mann). Tulane Stud. Zool., 1(5): 61-68. HoBBS, HORTON H. Jr. 1942a. A generic re- 1953c. A new burrowing vision of the crayfishes of the subfamily crawfish of the genus Procambarus from 20 Tulane Studies in Zoology Vol. 7 Louisiana and Mississippi. Ibid., 1(6): 69-76. 1953d. Two new craw- fishes of the genus Procambarus from Texas, Louisiana and Arkansas. Amer. Mas. Novitates, No. 1636: 1-10. - 1955. A new Cambarus of the Diogenes section from North Lou- isiana. Tiilauc Stud. Zool., 3(4): 71-81. 1956. The genus Pro- camharns in Louisiana. Amer. Midi. Xot., 56(2) : 406-422. 1957. Variation and sub- species of the crawfish Orconectes palmeri (Faxon). Tulane Stud. Zool., 5(10): 229-262. Penn, George Henry snd Horton H. HoBBS, Jr. 1958. A contribution toward a knowledge of the crawfishes of Texas. Texas Jour. Sci., 10(4): 452-483. Penn, George Henry and Guy Marlow 1959. The genus Cambams in Louisiana. Amer. Midi. Nat., 61(1) : 191-203. Smith, Elsie Wayne 1953. The life his- tory of the crawfish Orconectes ( Faxon- ella) clypeatus (Hay). Tulane Stud. Zool., 1(7) : 77-96. VioscA, Percy Jr. 1939. Where to fish in Louisiana for crawfish. La. Cons. Rev., 8(1) : 17-18. 1953. All about crawfish: life history and habits. La. Conservation- ist, 5(6): 3-5. Abstract Recognition characters, a key, and maps of distribution records are given for the following 29 species and sub- species of crawfishes known to be pre- sent in Louisiana: Procambarus bland- ingi acutus, P. bivittatus, P. clai-ki, P. planirostris, P. viaeviridis, P. hinei, P. shermani, P. tnlanei, P. si)nulans simulans, P. vioscai, P. penni, P. du- pratzi, P. natchitochae, Cambarellns shnfeldti, C. piier, Cambarus diogenes diogenes, C. d. Indovicianus, C. hedg- pethi, C. oryktes, C. dissitus, Orconectes clypeatus, O. beyeri, O. lancifer. O. hobbsi, O. palmeri palmeri, O. p. cre- olanus, O. p. lo^igimanus, O. hatlunvayi, and O. difficilis. Eighty-five figures illustrating taxcnomic characters of these crawfishes, and twelve maps showing distribution are included. COMPARISON OF THE CHROMATOPHOROTROPINS OF TWO CRAYFISHES WITH SPECIAL REFERENCE TO ELECTROPHORETIC BEHAVIOR ' MILTON FINGERMAN, Department of Zoology, Newcomb College, Tulane University, New Orleans, Louisiana The chromatophore systems of two species of crayfishes, Caynharellus shi/feldti and Or- conectes clypeatus, commonly found in the New Orleans area have been the subject of a series of investigations (Fingerman, 1957a, b, 1958; Fingerman and Aoto, 1958a, b; Fingerman and Lowe, 1957a, b, 1958). Sev- eral differences exist between the behavior of the dark red chromatophores of these two crayfishes under a variety of experimental conditions. The ability of the dark red chromatophores to respond to black and to white backgrounds may be used to illustrate one of these differences. The dark red pig- ment of Cambarellus disperses maximally in specimens on a black background and con- centrates maximally in specimens in a white container (Fingerman, 1957a). The dark red pigment of Orconectes will also disperse maximally in specimens on a black back- ground, but this pigment is intermediate be- tween the fully concentrated and fully dis- persed conditions in specimens on a white background (Fingerman, 1958). The red pigment in both species is con- trolled by pigment dispersing and concen- trating hormones found in the eyestalks and supraesophageal ganglia plus the circum- esophageal connectives. The current con- cept of the origin of chromatophorotropins is that they are neurosecretory products. The sinus gland appears to be merely a storage and release center for neurosecretory ma- terial produced elsewhere, e.g. in the optic ganglia (Knowles and Carlisle, 1956). How- ever, there is no reason why at least some of the chromatophorotropic material found in central nervous tissues may not be re- leased directly into the cirailation rather than first being transported to the sinus gland. Evidence in support of this state- ment has been obtained by Fingerman and Lowe (1957b). Stimulation of the eye- stalk stubs of eyestalkless Cambarellus re- 1 This investigation was supported by Grant No. B-838 from the National Insti- tutes of Health. suits in transitory concentration of the dark red pigment, presumably due to release of a pigment concentrating hormone from the remaining central nervous organs. Fingerman and Aoto ( 1958b) studied the electrophoretic behavior of chromatophoro- pins in Cambarellus. These investigators found that the dark red pigment concentrat- ing and dispersing substances from the su- praesophageal ganglia plus the circum- esophageal connectives were different from those in the eyestalk. The primary object of the experiments described herein was to study the electrophoretic behavior of the chromatophorotropins in Orconectes and to compare them with the chromatophorotro- pins in Cambarellus. Since the dark red pig- ment of Orconectes is in an intermediate stage of dispersion in specimens on a white background, we have also an ideal situation in which to study antagonism between pig- ment concentrating and dispersing sub- stances. Materials and Methods Specimens of the crayfishes Cambarellus sbufeldti and Orconectes clypeatus were col- lected at least once a week at Hickory, Lou- isiana, for use in the experiments described below. The crayfishes were kept in aquaria that contained aerated tap water approxi- mately 2.5 cm. deep. The dark red chromatophores in the por- tion of the carapace dorsal to the heart were staged according to the system of Hogben and Slome (1931). Stage 1 represented maximal pigment concentration, stage 5 maximal dispersion, and stages 2, 3, and 4 the intermediate conditions. The exoskele- tons were sufficiently transparent to allow direct, accurate observation of the under- lying chromatophores. Crayfishes into which chromatophorotro- pins were injected had had one eyestalk re- moved at least 12 hours prior to use in an experiment. Brown, Webb, and Sandeen (1952) and Fingerman (1957a) found that 22 Tulane Studies in Zoology Vol. 7 the responses of one-eyed individuals to chromatophorotropins were greater than the responses of intact specimens, presumably because both eyestalks made the organisms more capable of antagonizing injected hor- mones. In every experiment the dark red chromatophores of crayfish that received chromatophorotropins as well as of those injected with saline as a control were staged at the time of injection and 15, .^0, 60, 90, and 120 minutes thereafter. Filter paper electrophoresis was performed in essentially the same manner described earlier by Fingerman and Aoto ( 1958b). A model E-800-2 Filter Paper Electrophoresis Apparatus manufactured by the Research Equipment Corporation was used. The volt- age was held constant at 500 volts and the current was 0.1-0.2 milliampere. One-tenth molar sodium hydroxide — boric acid buffer of pH 7.2 was used. When extracts were prepared, the desired number of organs (40 eyestalks or 20 supraesophageal ganglia with the circumesophageal connectives at- tached ) were dissected out, triturated, and suspended in 0.1 ml. distilled water. All extracts of eyestalks used in the experiments described below were centrifuged to remove the bits of exoskeleton and retinal pigments. Each extract was gradually applied to a 0.5 inch wide Whatman No. 1 filter paper strip. A cool-air blower was used to evaporate the water as the extract was applied thereby preventing spread of the extract over a band more than one-quarter inch wide along the strip. The entire strip was then moistened with buffer and placed into the electro- phoresis migration chamber. To minimize inactivation of the hormones the chamber was kept in a refrigerator maintained at an average temperature of 7°C. After electro- phoresis had proceeded for two hours the filter paper strips were removed from the chamber. The strips were cut into predeter- mined lengths. Each section was immediate- ly placed into 0.3 ml. Van Harreveld's solu- tion (Van Harreveld, 1936) in a covered container to minimize evaporation and then kept in the refrigerator for 30 minutes to wash the chromatophorotropin from the paper. The fluid was then collected in a syringe and injected into one-eyed crayfish. The dose was always 0.02 ml. per crayfish. Four-inch sections of the strip on the posi- tive side of the zone of application of ex- tracts of the supraesophageal ganglia plus the circumesophageal connectives of Cam- barelha were used. All other sections of the strips used in the experiments were three inches long. These lengths were chosen be- cause Fingerman and Aoto (1958b) found that the chromatophorotropins of Camba- rellus would not migrate further than four inches in two hours. As a control in the elctrophoresis experiments a strip of paper as long as the longest section used in the experiment was moistened with buffer and then placed into the same volume of Van Harreveld's solution as used on the strips with chromatophorotropins. Tissue extracts injected immediately after preparation rather than after electrophoresis were prepared as follows. The desired num- ber of organs were extirpated, placed with a minimum of fluid into a glass mortar, triturated, and suspended in sufficient Van Harreveld's solution to yield the desired concentration of one-third of an organ per 0.02 ml. In every experiment that Orcon- ectes served as a recipient of extract, the extract was assayed on 10 specimens. Pure Van Harreveld's solution was used as a con- trol for these experiments. Student's t test was used to determine the level of significance. The 95 "^r level was taken as the minimal value for a significant difference between two means. Experiments and Results Comparison of the distribution of dark red pigment concentrating hormone in the circumesophageal connectives of Cambarel- lus and Orconectes. — Fingerman and Lowe (1958) found that the chromatophorotro- pins in the tissues of Orconectes were effec- tive on the dark red pigment of Cambarel- lus. These investigators also noticed that the red pigment concentrating hormone in the extracts of the supraesophageal ganglia plus the circumesophageal connectives of Orconectes was effective over a longer pe- riod of time than was the hormone with the same function from the tissues of Cam- barellus. The aim of the first group of ex- periments was to analyze this difference with extracts of the circumesophageal con- nectives. Extracts of the circumesophageal connec- tives of Cambarellus and Orconectes were prepared as described above. These extracts No. 1 Fingerman: Chromatophorotropins of crayfishes 23 and Van Harreveld's solution as a control were injected into one-eyed Cambarellus whose dark red pigment was maximally dispersed as a result of having been kept in black containers. The experiment was performed twice. The results are presented in the top portion of Figure 1 where each point represents the average of 10 individ- uals. No response was elicited by physio- logical saline. The response produced by the circumesophageal connectives of Camba- the circumesophageal connective of Camba- rellus was qualitatively and quantitatively similar to that seen so many times previ- ously ( Fingerman, 1957a; Fingerman and Lowe, 1957a, 1958). The maximal response to red pigment concentrating hormone from the circumesophageal connectives of Camba- rellus when assayed on Cambarellus always occurs 15 to 30 minutes after injection. The response then disappears rapidly. In contrast, the hormone from the circum- esophageal connectives of Orconectes has a prolonged effect upon the red chromato- phores of Cambarellus, just as was reported for the supraesophageal ganglia plus the circumesophageal connectives ( Fingerman and Lowe, 1958). The average chromato- phore stages of the two groups of crayfish injected with chromatophorotropin deter- mined 60 minutes after injection of the extracts were significantly different from each other. The circumesophageal connectives of both species were then separated into four por- tions and extracted for use in the remaining experiments of this series. The portions were: (1) the tritocerebral commissure Figure 1. Responses of the dark red chro- matophores of Caynbarellus shufeldti to ex- tracts of the circumesophageal connectives and components of this organ taken from Cambarellus shufeldti and Orconectes cly- peatus. Top; open symbols, whole circum- esophageal connectives of Cambarellus; sol- id symbols, whole circumesophageal con- nectives of Orconectes; half-filled symbols, control. Middle; components of the circum- esophageal connectives of Cambarellus. Dots, connective ganglion; circles, portion of connectives anterior to ganglia; circles half-filled on left, tritocerebral commissure; circles half-filled on right, portion of con- nectives posterior to ganglia; circles half- filled on bottom, control. Bottom, compo- nents of the circumesophageal connectives of Orconectes. The symbols are the same as those used in the middle portion of the figure. that runs posterior to the esophagus from one connective to the other, (2) the con- nective ganglia, (3) the portions of the connectives that lie anterior to the connec- tive ganglia and (4) posterior to these ganglia. These extracts were likewise in- jected into Cambarellus with maximally dis- persed red pigment. The results are pre- sented in the middle ( Cambarellus ) and bottom {Orconectes) portions of Figure 1 HOURS 24 Tulane Studies in Zoology Vol. 7 where each point represents the average of of Figure 2 represents, therefore, the mean 10 individuals. of 35 specimens. The experiments with As is evident from inspection of the fig- tissues of Orconectes were performed twice, ure, the connective ganglia of both species The average of 10 specimens is represented contained more red pigment concentrating by the points in parts C and D of Figure 2. hormone than the other portions of the Inspection of the figure reveals that the circumesophageal connectives. Furthermore, sjnus glands and optic ganglia of Cambarel- the pattern of response to the extracts of the /^^j contain large quantities of dark red pig- connective ganglia was the same that was ,-nent dispersing hormone and small amounts observed when intact circumesophageal con- „£ concentrating substance. The correspond- nectives were used (top portion of Figure jng curves in parts A and B of Figure 2 1 ) . The two curves depicting the response ^ere not significantly different from each to extracts of the connective ganglia were other. On the basis of the volume of tissue, significantly different from each other. The ^hg concentrations of chromatophorotropins posterior portion of the circumesophageal j^ ^he sinus gland are much higher than in connectives of both species ranked second ^he optic ganglia since the sinus gland repre- in titer of this hormone. The tritocerebral sents about one percent of the tissue in commissures contained much less of this xhe eyestalk. chromatophorotropin than did the ganglia. The results with tissues of Orconectes a striking contrast to the situation in ^ere qualitatively similar to those shown the shrimp Crago septemspinosus wherein [„ parts A and B of Figure 2 (Cambarel- the activity resides almost exclusively in the i^j). The sinus glands and optic ganglia tritocerebral commissure together with the had more dispersing than concentrating sub- immediately adjacent medial aspect of the stance just as has been reported for extracts connective lying between the origins of of whole eyestalks ( Fingerman and Lowe, the commissure and the connective ganglia 1958). Interestingly, the peak of dispersing (Brown, 1946). activity of the optic ganglia from Orcon- Comparison of the effects of extracts of ectes was displaced towards the right. Pre- the sinus glands and optic ganglia of Cam- sumably, this was due to the presence of barellus and Orconectes. — Whole eyestalks the statistically significant quantity of con- of both species of crayfish were assayed on centrating substance ( part D of Figure 2 ) Ca7Jibarellus by Fingerman and Lowe in the optic ganglia that must have had to (1958). They found that the eyestalks of run its course before the dispersing sub- both species contained red pigment dis- stance could express itself maximally, an- persing and concentrating hormones. Much other case of hormonal antagonism. The more of the former hormone than of the curves in part C of Figure 2, in contrast to latter was present in the eyestalks of both those in part A of this figure, were sig- crayfishes. Since the sinus gland in deca- nificantly different from each other as pods is a storage and release center for revealed by the t test. The red pigment neurosecretory products and is not thought of crayfish injected with extracts of tissues to produce anything directly ( Bliss, Du- of Orconectes did not return to the fully rand, and Welsh, 1954), a comparison of concentrated condition as rapidly as the red the effects of extracts of the sinus glands pigment of crayfish injected with extracts and optic ganglia of both crayfishes would of tissues of Catnbarellus. The same effect be worthwhile. was apparent with whole eyestalks. (Finger- Extracts of sinus glands and of optic ^'^^^ ^'^'^ Lowe, 1958). ganglia were prepared in the manner and Electrophoretic analysis of the chroniato- concentration described above. The extracts phorotropins of Cambarellus assayed on Or- were assayed on one-eyed Cambarellus on conectes. — This group of experiments was black and on white backgrounds in order to designed to determine the effects on Orcon- determine the relative amounts of dark red ectes of ( 1 ) red pigment dispersing and pigment dispersing and concentrating hor- concentrating hormones of Cambarellus that mones in those tissues. The experiments had been separated by filter paper electro- with tissues of Cambarellus were performed phoresis and ( 2 ) mixtures of these sub- seven times. Each point in parts A and B stances in order to shed some information No. 1 Fingerman: Chromatophorotropins of crayfishes 25 on the nature of the antagonism between cumesophageal connectives was negatively these hormones. Fingerman and Aoto ( 1958b) found that at pH 7.5 the red pig- ment dispersing substance in the eyestalks of Cambarellus was positively charged but that the substance with the same function in the supraesophageal ganglia plus the cir- charged. The red pigment concentrating material in the latter tissues was positively charged. For the first experiment extracts of eye- stalks and of supraesophageal ganglia plus the circumesophageal connectives were pre- U O < u q: O I Q- I O < ^ O cc I O 4 B - D I 2 0 I HOURS Figure 2. Responses of the dark red chromatopnores of Cambarellus on white (A and C) and on black (B and D) backgrounds to extracts of the sinus glands (dots) and op- tic ganglia (circles) of Cambarellus (A and B) and Orconectes (C and D). Half-filled circles, control. 26 Tulane Studies in Zoology Vol. 7 pared and applied to filter paper strips in the manner described above. After electro- phoresis had proceeded for two hours the first three inches on the negative side of both strips were each washed for 30 minutes in 0.3 ml. Van Harreveld's solution. These extracts were given an arbitrary concentra- tion of 100*"^. Twelve hundredths of a milliliter of each solution was placed into a watch glass, mixed, and taken up in a sy- ringe. The remaining pure materials were diluted by half with saline. Each of these solutions plus a control was injected into one-eyed Orconectes that had been kept on a white background so that the red pig- of the strip to which the extract of the supraesophageal ganglia with the circum- esophageal connectives attached was applied significantly concentrated this pigment. The mixture produced a statistically significant concentration of the red pigment. However, the degree of concentration produced by the mixture was significantly less than that caused by the pure material, presumably because of the presence of the added dis- persing substance. The remaining experiments of this group were similar to those described immediately above with the one exception that red pig- ment dispersing substance from the supra- HOURS Figure 3. Responses of the dark red chromatophores of Orconectes to extracts of eye- stalks and of supraesophageal ganglia with the circumesophageal connectives attached of Camharelbis after filter paper electrophoresis. Dots, negative portion of strip con- taining .supraesophageal ganglia plus the circumesophageal connectives; circles half- filled on bottom, negative portion of the strip containing eyestalk tissues; circles half-filled on left, control; circles, positive portion of strip containing supraesophageal ganglia plus the circumesophageal connectives; circles half-filled on right, mixture of equal volumes of extracts containing pigment dispersing and concentrating hormones. ment was in an intermediate stage ( chro- matophore stage 3 ) . The experiment was performed twice. The results are presented in Figure 3A, where each point represents the average of 20 specimens. The electric charge on each hormone was the same as reported by Fingerman and Aoto ( 1958b). The material in the eyestalk extract that migrated toward the cathode dispersed the red pigment of Orconectes significantly just as it did in Cambarellus and the substance from the negative side esophageal ganglia with circumesophageal connectives attached of Camharell//s was used instead of dispersing hormone from the eyestalks. The results are shown in Fig- ure 3B where each point represents the average of 20 specimens. The pure extracts behaved as expected. The curves depicting pigment dispersion and concentration pro- duced by the pure extracts were significant- ly different from the control curve. The curve showing concentration produced by the mixture was significantly different from No. 1 Fingerman: Chromatophorotropins of Crayfishes 27 HOURS Figure 4. Responses of the dark red chro- matophores of Orconectes to extracts of eyestalks and of supraesophageal ganglia plus the circumesophageal connectives of Orconectes after filter paper electrophore- sis. A, ejestalks; B, supraesophageal gan- glia plus the circumesophageal connectives applied to the filter paper strip immediate- ly after extraction; C, supraesophageal gan- glia plus the circumesophageal connectives applied to filter paper strip after the ex- tract had been kept at room temperature for two hours. Circles, fraction that mi- grated toward the positive pole; dots, frac- tion that migrated toward the negative pole; half-filled circles, control. both the control curve and the curve de- picting the response to concentrating hor- mone alone, indicative of mutual antagonism between the two hormones. Electrotrophoretic analysis of chromato- phorotropins of Orconectes assayed on Or- conectes.— The object of this series of ex- periments was to analyze by filter paper electrophoresis the red pigment concentrat- ing and dispersing substances in Orconectes in order to compare them with the chroma- tophorotropins of Cambarellus. Fingerman (1958) showed that the eye- stalks of Orconectes contain a red pigment dispersing hormone and an extremely small quantity of red pigment concentrating hor- mone. Assay of eyestalk extracts after elec- trophoresis revealed that the pigment dis- persing substance was electronegative ( Fig- ure 4A). The amount of pigment disper- sion was statistically significant. Since this experiment was done twice each point in Figure 4 represents the mean of 20 speci- mens. No concentrating substance was ap- parent. Several explanations are possible for the failure to find the concentrating sub- stance, e.g., irreversible adsorption to the filter paper and inactivation of the chroma- tophorotropin during the process of elec- trophoresis. In the next experiment extracts of supra- esophageal ganglia with the circumesopha- geal connectives attached of Orconectes were subjected to electrophoresis. Three inch portions on both sides of the region of application were removed from the strips for analysis in the manner described above. The principal substance present was a sta- tistically significant electropositive red pig- ment concentrating one (Figure 4B). A slight but not statistically significant dis- persion was produced by the material that had migrated toward the positive pole. The CHROMATOPHORE STAGE — ro oj -t^ Figure b. i.espo.ises ox tne uark red chro- matophores of Orconectes to extracts of eyestalks and supraesophageal ganglia plus the circumesophageal connectives of Orco- nectes after filter paper electrophoresis. Circles, positive portion of strip containing eyestalk tissues; dots, negative portion of the filter paper strip containing supra- esophageal ganglia plus the circumesopha- geal connectives; circles half-filled on right, mixture of equal volumes of the extracts signified by the circles and dots; circles half-filled on left, control. 28 Tulane Studies in Zoology Vol. 7 lack of an appreciable degree of pigment dispersion was not surprising since Finger- man and Aoto ( 1958a) and Fingerman and Lowe (1958) had found that freshly pre- pared extracts of the supraesophageal gang- lia and circumesophageal connectives of Orconectes never caused pigment dispersion, only pigment concentration, when injected into Orconectes. Fingerman and Aoto (1958a), however, showed that extracts of supraesophageal ganglia plus the circumesophageal connec- tives of Orconectes that had been kept at room temperature for 120 minutes would disperse the red pigment of Orconectes. Presumably while the extracts remained at room temperature the concentrating sub- stance deteriorated thereby unmasking the dispersing material. In view of this infor- mation, the next experiment was designed as follows. An extract of supraesophageal ganglia plus the circumesophageal connec- tives was prepared for electrophoresis in the concentration and manner described above. However, instead of applying the extracts to the filter paper strips immedi- ately after preparation, as is routine, the extract was kept at room temperature for 120 minutes and then applied. The results of this experiment are shown in Figure 4C where each point represents the mean of 20 specimens. A statistically significant dis- persing effect was now apparent and the concentrating material had disappeared, as has been predicted. For the final experiment extracts of pig- ment dispersing and concentrating sub- stances were obtained by electrophoresis and mixed in the same manner as described above for the chromatophorotropins of Camharellus in order to learn something of the antagonism between these substances found in Orconectes. The dispersing sub- stance was obtained from the first three inches of the positive side of a strip of filter paper to which an extract of eyestalks had been applied and the concentrating material from the same length on the nega- tive side of a strip to which an extract of supraesophageal ganglia plus the circum- esophageal connectives had been applied. The experiment was performed three times. The results are presented in Figure 5 where each point represents the average of 30 specimens. The results obtained with the pure ex- tracts confirm the findings shown in parts A and B of Figure 4 and are statistically significant. The dispersing substance in the eyestalks of Orconectes is electronegative and the concentrating material in the supra- esophageal ganglia and circumesophageal connectives is electropositive. The mixture produced no concentration but a slight dis- persion that was not statistically significant. Discussion The results presented above provide fur- ther information concerning the chromato- phore systems of the crayfishes Camharellus shu]eldti and Orconectes clypeati/s. The dis- tribution of red pigment concentrating hor- mone in the circumesophageal connectives of Caniharellus and Orconectes ( Figure 1 ) was strikingly similar. The connective gang- lia in both species have by far more of this hormone than any of the other portions of the circumesophageal connectives. The sta- tistically significant difference in the time course of the hormones from the two cray- fishes suggests that these substances may differ from one another in some respect. This difference is not due to a difference in titer of red pigment dispersing hormone in the circumesophageal connectives of the two crayfishes. Such an hypothesis is not tenable because Fingerman and Lowe (1958) found the same difference in time course when they used supraesophageal ganglia plus the circumesophageal connec- tives of both species and, furthermore, the tissues from Orconectes contained much more red pigment dispersing material when assayed on Ca^nbarellus than did the tissues of Camharellus. Fingerman and Aoto ( un- published observations ) studied the cyto- logical features of the circumesophageal con- nectives of Camharellus and found neuro- secretory cell bodies in the connective gang- lia alone. Presumably much hormone is pro- duced and stored in this region. Another interesting comparison can be drawn from the electrophoretic studies. The red pigment dispersing substance in the eye- stalk of Camharellus has a positive charge whereas the substance with the same func- tion in the supraesophageal ganglia plus the circumesopliageal connectives of this species is negatively charged at pH 7.2 and 7.5. The pigment concentrator in the supra- No. 1 Fingerman: Chromatophorotropins of Crayfishes 29 esophageal ganglia plus the circumesopha- geal connectives of Cambarellus has the same charge as the disperser in the eye- stalk. Ccvnbarellus, therefore, must contain two different pigment dispersing substances. In contrast, the dark red pigment dispersing substances in the eyestalk and supraesopha- geal ganglia plus the circumesophageal con- nectives of Orconectes are both negatively charged and are presumably one substance. The pigment disperser in the eyestalk of Cambarellus must differ from that in Or- conectes. The pigment concentrating sub- stance in the supraesophageai ganglia plus the circumesophageal connectives of Or- conectes is positively charged just as is that in Cambarellus. The results obtained with electrophoresis of tissues of Cambarellus confirmed the findings of Fingerman and Aoto (1958b). In Figure 5 are shown the results of mix- ing pigment dispersing and concentrating substances from the eyestalks and supra- esophageai ganglia plus the circumesopha- geal connectives respectively of Orconectes that had been separated by electrophoresis. Although the potencies of the pure extracts were approximately equal as judged by the areas subtended between the experimental and control curves, no pigment concentra- tion was apparent. The dispersing hormone apparently is able to inhibit fully the con- centrator. This experiment points out the advantage of using Orconectes kept on a white background for studies of antagonism between chromatophorotropins. In one speci- men pigment concentration or dispersion could occur. In Cambarellus the pigment can be kept fully dispersed or fully con- centrated. If no pigment concentration oc- curs, for example, then we must distinguish between two possibilities, absence of chro- matophorotropins or the presence of a pre- ponderance of dispersing substance. To dis- tinguish between these alternatives requires a second assay with animals whose red pig- ment is fully concentrated as a result of having been kept on a white background. Summary and Conclusions 1. The chromatophore systems of the crayfishes Orconectes clypeatus and Cam- barellus shufeldti were investigated further. 2. The circumesophageal connectives of both species contain large quantities of a hormone that concentrates the dark red pig- ment. The connective ganglia contain more of this hormone than other portions of the circumesophageal connectives. The time course of the dark red pigment concentrat- ing hormone in the circumesophageal con- nectives of Cambarellus was statistically dif- ferent from that produced by this hormone in extracts of tissues from Orconectes when assayed on Cambarellus.. These hormones may, therefore, be different entities. 3. The sinus glands and optic ganglia of both species contain large quantities of red pigment dispersing hormone. Very little red pigment concentrating hormone is pres- ent in the eyestalks. 4. Electrophoretic analysis of chromato- phorotropins in the two crayfishes was per- formed at pH 7.2. The red pigment dis- persing substance in the eyestalks of Or- conectes and in the supraesophageai ganglia plus the circumesophageal connectives of Orconectes and Cambarellus was electro- negative. The substance with the same func- tion in the eyestalks of Cambarellus was oppositely charged. Cambarellus, therefore, contains two dark red pigment dispersing substances whereas Orconectes appears to have one only. The red pigment concentrat- ing substance in the supraesophageai ganglia plus the circumesophageal connectives of both crayfishes was electropositive. 5. Mixtures of dark red pigment dis- persing and concentrating hormones ob- tained from both crayfishes after filter paper electrophoresis revealed a mutual antagonism. References Cited Bliss, Dorothy E., James B. Durand, and John H. Welsh 1954. Neurosecretory systems in decapod Crustacea. Zeitschr. Zellforsch., 39: 520-5.36. Brown, Frank A., Jr. 1946. The source and activity of Cro^ro-darkening- hormone. Physiol. ZooL, 19: 215-223. , H. Marguerite Webb, and Muriel I. Sandeen 1952. The action of two hormones regulating the red chro- matophores of Palaemonetes. .Jour. Ex- per. ZooL, 120: 319-420. Fingerman, Milton 1957a. Endocrine con- trol of the red and white chromatophores of the dwarf crawfish, Cambarellus shu- feldti. Tulane Stud. ZooL, 5: 137-148. 1957b. Physiology of the red and white chromatophores of the dwarf crayfish, Cambarellus shufeldtii. Physiol. ZooL, 30: 142-154. 30 Tulane Studies in Zoology Vol. 7 1958. The chromatophore system of the crawfish Orconectes clype- atns. Amer. Midi. Nat., 60: 71-83. FiNGEKMAN, MiLTON and TOMOJI AOTO 1958a. Chromatophorotropins in the cray- fish Orconectes cUjpcatiis and their rela- tionship to long-term background adapta- tion. Physiol. Zool, 31: 193-208. 1958b. Electrophoretic analysis of chromatophorotropins in the dwarf crayfish, Camhurellus shnfeldti. Jour. Exper. Zool., 138: 25-50. FiNGERMAN, MiLTON and Mildred E. Lowe 1957a. Influence of time on background upon the chromatophore systems of two crustaceans. Physiol. Zool., 30: 216-231. 1957b. Hormones con- trolling the chromatophores of the dwarf crawfish, Cantbarellns shitfeldti: their secretion, stability, and separation by fil- ter paper electrophoresis. Tulane Stud. Zool., 5: 149-171. „_.„ _... 1958. Stability of the chromatophorotropins of the dwarf cray- fish, Cambarelliis shnfeldti, and their ef- fects on another crayfish, Biol. Bull., 114: 317-333. HoGBEN, Lancelot T. and David Slome 1931. The pigmentary effector system. VL The dual character of endocrine co- ordination in amphibian colour change. Proc. Roy. Soc. Lond., Ser. B, 108 : 10-53. Knowles, Francis G. W. and David B. Carlisle 1956. Endocrine control in the Crustacea. Biol. Reviews, 31: 386-473. Van Harreveld, A. 1936. A physiological solution for fresh water crustaceans. Proc. Soc. Exper. Biol. & Med., 34: 428- 432. Abstract Basic differences exist between the chro- matophore systems of two crayfishes found in the New Orleans area, Cam- barelliis shnfeldti and Orconectes clype- atns. The object of this investigation was to examine further the chromato- phore systems of these crayfishes to determine the extent of these differ- ences. The circumesophageal connec- tives of both species contain large quantities of a hormone that concen- trates dark red pigment. The connec- tive ganglia contained a greater amount of this hormone than did other portions of the circum.esophageal connectives. The time course of the dark red pig- ment concentrating hormone in the cir- cumesophageal connectives of Cambar- ellus was statistically different from that produced by this hormone in tis- sues of Orconectes when assayed on Camba)-ellus. These hormones may, therefore, be different entities. The sinus gland and optic ganglia of both species contain large quantities of red pigment dispersing hormone. Very little red pigment concentrating hor- mone is present in the eyestalks. Elec- trophoretic analysis of chromatophoro- tropins in the two crayfishes was per- formed at pH 7.2. The red pigment dispersing substance in the eyestalks of Orconectes and in the supraesopha- geal ganglia plus the circumesopha- geal connectives of Orconectes and Cambarelliis was electronegative. The substance with the same function in the eyestalks of Cambarellus was op- positely charged. Cambarellus, there- fore, contains two dark red pigment dispersing substances whereas Orco- nectes appears to have one only. The red pigment concentrating substance in the supraesophageal ganglia plus the circumesophageal connectives of both crayfishes was electropositive. TULANE STUDIES IN ZOOLOGY VOLUME 1, 1953-54 Number Price 1 On a new genua and species of mysid from Louisiana (Crustacea, Malacostraca), by Aibert H. Banner, pp. 1-8 (June 1, 1953) . .I$0.25 2 A contribution on the life history of the lizard Scincella laterale (Say), by Richard M. Johnson, pp. 9-27 (July 3, 1953) 75 3 An outline for the study of a reptile life history, by Fred R. Cagle, pp. 29-52 (July 28, 1953) , 75 4 A population of Holbrook's salamander, Eurycea lonyicauda guttolineata (tlol- brook), by Robert E. Gordon, pp. 53-60 (August 15, 1953) 25 5 A redescrlptlon of the crawfish Procam- barus hinei (Ortmann) (Decapoda, Astaci- dae), by George Henry Penn, pp. 61-68 (September 1, 1953) 25 6 A new burrowing crawfish of the genus Procamiarus from Louisiana and Missis- sippi (Lecapoda, Astacidae), by George Henry Penn, pp. 69-76 (September 15, 1953) 25 7 The life history of the crawfish Orconectes (Faxonella) clypeatus (Hay) (Decapoda, Astacidae), by Elsie Wayne Smith, pp. 77- 96 (October 23, 1953) 50 8 Ostrincola gracilis C. B. Wilson, a parasite of marine pelecypods in Louisiana (Cope- poda, Cyclopoida), by Arthur G. Humes, pp. 97-107 (December 21, 1953) 25 9 Hybrid inviability between Rana pipiens from Wisconsin and Mexico, by E. Peter Volpe, pp. 109 123 (February 8, 1954) 35 10 The butterflies and skippers of Louisiana, by Edward Nelson Lambremont, pp. 125- 164 (April 30, 1954) GO 11 Two new species of the genus Oraptemys, by Fred R. Cagle, pp. 165-186 (August 26, 1954) 75 12 The taxonomic status of the mid-Gulf Coast Atnphiuma, by Imogene R. Hill, pp. 189-215 (August 26, 1954) 35 Complete volume, including title page and table of contents, (unbound) $4.25 VOLUME 2, 1954-55 1 A new Eulimnadia from the rice fields of Arkansas with a key to the American species of the genus (Conchostraca, Lim- nadildae), by N. T. Mattox, pp. 1-10 (Sep- tember 14, 1954) 25 2 Studies in the ecology of the narrow- mouthed toad, Microhyla carolinensis caro- linensis, by Paul K. Anderson, pp. 13-46 (November 8, 1954) 50 3 A new species of Diaptomus from Louisi- ana and Texas with notes on the subgenus Leptodiaptomus (Copepoda, Calanoida), by Mildred Stratton Wilson, pp. 47-60 (No- vember 29, 1954) 25 4 Three new species and new records of southern mllUpeds, by Nell B. Causey, pp. 61-68 (December 28, 1954) .25 5 Notropis haileyi, a new cyprlnid fish from Pascagoula and Mobile Bay drainages of Mississippi and Alabama, by Royal D. Suttkus and Edward C. Raney, pp. 69-86 (January 12, 1955) 30 VOLUME 2, 1954-55-Continued Number 6 A reconsideration of the racer. Coluber constrictor, in eastern United States, by Walter Auffenberg, pp. 87-155 (February Price 25, 1955) 1.00 Notropis hypsilepis, a new cyprinid fish from the Apalachicola River system of Georgia and Alabama, by Royal D. Suttkus and Edward C. Raney, pp. 157-170 (Feb- ruary 25, 1955) 30 The fishes of two Gulf Coastal marsh areas of Florida, by John D. Kilby, pp. 171-247 (May 4, 1955) 1.00 Complete volume, including title page, table of contents and index (unbound) .•• -$3.75 VOLUME 3, 1955-56 1 Notropis asperifroiiSj a new cyprinid fish from the Mobile Bay drainage of Alabama and Georgia, with studies of related spe- cies, by Royal D. Suttkus and Edward C. Raney, pp. 1-33 (July 8, 1955) 50 2 A new Louisiana copepod related to Diap- tomus (Aglaodiaptomus) clavipes Schacht (Copepoda, Calanoida), by Mildred Strat- ton Wilson, pp. 35-47 (August 1, 1955) ... .30 3 A new species of Sternotherus with a dis- cussion of the Sternotherus cariiiatus com- plex (Chelonia, ICinosternidae), by Don- ald W. Tinkle and Robert G. Webb, pp. 51-67 (August 30, 1955) 50 4 A new Cambarus of the Diocjenes section from North Louisiana (Decapoda, Astaci- dae), by George Henry Penn, pp. 71-81 (September 30, 1955) 25 5 Notropis euryzonus, a new cyprinid fish from the Chattahoochee River system of Georgia and Alabama, by Royal D. Suttkus, pp. 83-100 (December 28, 1955) 50 G Factors influencing the rate of oxygen con- sumption of the dwarf crawfish, Camba- rellus shufeldtii (Decapoda Astacidae), by Milton Fingerman, pp. 101-116 (December 28,1955) 35 7 Identification and geographical variation of the cyprinodont fishes Fundulus oliva- ceus (Storer) and Fundulus notatus (Rafi- nesque), by Jerram L. Brown, pp. 117-134, (February 3, 1956) 50 8 The physiology of the melanophores of the isopod Idothea exotica, by Milton Finger- man, pp. 137-148 (April 12, 1956) .30 9 Osmotic behavior and bleeding of the oyster Crassostrea virginicOj by Milton Fin- german and Laurence D. Fairbanks, pp. 149-168 (April 12, 195G) 50 10 Anatomy of the eyestalk of the white shrimp, Penaeus setiferus (Linn. 1758), by Joseph H. 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AYilson, pp. 113-170 (September 5, 1958) $1.00 4 The copepod genus JIalici/clops in North America, with description of a new species from Lake Pontchartrain, Louisiana, and the Texas coast, by Mildred Stratton Wil- son, pp. 170-189 (December 31, 1958). Ontogenv of the first and second pleopods of the liiale crawfish Orconectes cliipeutiix (Hay) (Dedapoda, Astacidae), by Joe B. Black, pp. 190-203 (December 31, 1958) ... .60 Complete volume, includini;- title page, table of contents and index (unbound) .$3.50 Orders should be addressed to Meade Natural History Library, c/o Department of Zoology, Tulane University, New Orleans, 18, La., USA Please make remittance payable to "Tulane University" 3 JUL 4::^ 1959 HARVARD UNIVERSITY Volume 7, Number 2 July 9, 1959 A REVIEW OF THE SEABASSES OF THE GENUS CENTROPRISTES (SERRANIDAE) RUDOLPH J. MILLER, DEPARTMENT OF CONSERVATION, CORNELL UNIVERSITY^ ITHACA, NEW YORK TULANE UNIVERSITY NEW ORLEANS TULANE STUDIES IN ZOOLOGY is devoted primarily to the zoology of the waters and adjacent land areas of the Gulf of Mexico and the Caribbean Sea. Each number is issued separately and contains an individual study. As volumes are completed, title pages and tables of contents are distributed to institutions exchanging the entire series. Manuscripts submitted for publication are evaluated by the editor and by an editorial com- mittee selected for each paper. Contributors need not be members of the Tulane Univer- sity faculty. Manuscripts should be submitted on good paper, as original typewritten copy, double- spaced, and carefully corrected. Two carbon copies in addition to the original will help expedite editing and assure more rapid publication. An abstract not exceeding three percent of the length of the original article must accom- pany each manuscript submitted. This will be transmitted to Biological Abstracts and any other abstracting journal specified by the writer. Separate numbers or volumes may be purchased by individuals, but subscriptions are not accepted. Authors may obtain copies for personal use at cost. Address all communications concerning manuscripts and editorial matters to the editor; communications concerning exchanges, and orders for individual numbers to the Librarian, Meade Natural History Library. When citing this series authors are requested to use the following abbreviations: Tulane Stud. Zool. Price for this number: $0.75. George Henry Penn, Editor Meade Natural History Library, Tulane University, New Orleans, U. S. A. Assistant to the Editor: Robert K. Chipman TULANE STUDIES IN ZOOLOGY Volume 7, Number 2 July 9, 1959 A REVIEW OF THE SEABASSES OF THE GENUS CENTROPRISTES (SERRANIDAE) RUDOLPH J. MILLER, Department of Conservation, Cornell University, Ithaca, New York CONTENTS I. Introduction 35 II. Methods 35 III. Historical Review of the Species of Centropristes 36 IV. Generic Description 38 V. Key to the Species of Centropristes 39 VI. Species Accounts 42 A. Centropristes striatus 43 1. Material Examined 41 2. Description 44 3. Distribution 47 4. Population Analysis 48 5. Biology 50 B. Centropristes ocyurus 50 1. Material Examined 50 2. Description 52 3. Distribution . 57 4. Population Analysis 58 5 . Biology 58 C. Centropristes pbiladelphicus 59 1. Material Examined ____ 59 2 . Description 60 3. Distribution 61 4. Population Analysis 62 5. Biology .. .... 62 VII. Internal Anatomy 63 VIII. Summary 66 IX. Acknowledgements 67 X. References Cited 67 Abstract 68 MUS. COMP. ZOOL LIBRARY JUL 2 2 1959 HARVARD UNIVERSITY EDITORIAL COMMITTEE FOR THIS NUMBER William E. Fahy, Associate Professor of Zoology, Institute of Fisheries Research, University of North CaroHna, Morehead City, North CaroHna C. Richard Robins, Research Assistant Professor, The Marine Laboratory, Uni- versity of Miami, Miami, Florida Royal D. Suttkus, Associate Professor of Zoology, Tulane University, New Or- leans, Louisiana No. 2 Miller: Review of Centropristes 35 I. Introduction Prior to 1952, three species were recog- nized in the serranid genus Centropristes. The type species, Centropristes striatus, was the largest and best known form, ranging from Cape Cod, Massachusetts, to Pensacola, Florida. The rock sea bass, Centropristes philadelphicus. originally was thought to be limited to deep waters off Charleston, South Carolina, but later was found commonly elsewhere, especially off the Texas and Lou- isiana coasts. The gulf sea bass, Centro- pristes ocyurus, was described from the snapper banks off Pensacola in 1886, but it has been found since from Cape Hatteras, North Carolina, to the Yucatan shelf. Gins- burg (1952) described a new species, Cen- tropristes melanus, based on 13 specimens from the west coast of Florida. Before 1952, melanus had been considered an extreme western population of striatus. He also found differences between Gulf and Atlantic populations of both ocyurus and philadel- phicus but considered them to be "of low degrees, below the subspecies level". The present study attempts to analyze more fully differences between Gulf and Atlantic populations of Centropristes. De- scriptions of body and fin proportions, color patterns, and meristic characteristics are pre- sented. Much confusion in the literature on Centropristes resulted from a lack of knowl- edge of variation within species. II. Methods Twenty-one measurements and nine counts were made on each specimen examined, ex- cept in a few samples obtained at a late date in the study, where only selected counts and measurements were taken. Methods em- ployed in measuring and counting were those outlined by Hubbs and Lagler (1947) except as listed below: 1. Dorsal- fin origin to caudal base. — Measured from the dorsal origin to the point where the middle caudal ray disappears into the fleshy caudal base. 2. Dorsal-fin origin to occiput. — Meas- ured from the origin of the dorsal fin to the most anterior point of the scaled portion of the dorsal surface. In adults of ocyurus there is a noticeable scaleless gap between scales of the head and those of the body (fig. 16). There is also a gap between the .scales on either side of the head. The meas- urement in this case includes the most an- terior head scale on the left side. 3. Pelvic-fin insertion to snout. — Meas- ured from the insertion of the most anterior pelvic fin ray to the anterior tip of the pre- maxillary. 4. Anal- fin origin to caudal base. — Meas- ured from the origin of the first anal spine to the point where the middle caudal ray meets the fleshy covering of the anterior part of the caudal fin. 5. Body depth. — Measured from a point immediately anterior to the origin of the dorsal fin to a point immediately anterior to the pelvic-fin insertion. This measure- ment of body depth is more reliable than that used by Weed (1937) because the shoulder girdle prevents distortion. 6. Dorsal-fin origin to lateral line. — Least distance from the origin of the dorsal fin to the center of the nearest lateral-line scale. 7. Pelvic-fin insertion to lateral line. — Least distance between the insertion of the pelvic fin and the center of the nearest lateral-line scale. 8. Caudal-peduncle length. — Measured from the point where the last dorsal ray meets the body to the origin of the first upper rudimentary caudal ray. In philadel- phicus and ocyurus a small black spot ex- tends slightly forward from the latter point. 9- Interorbital ividth. — Least fleshy width. Weed measured the width of the bony bridge between the eyes. In larger speci- mens it is often very difficult to determine the exact limits of this bony bridge due to thickness of the covering flesh. In most specimens the fleshy width is well defined because the shelf of flesh that extends be- yond the bony bridge is tough and almost cartilage-like in texture. 10. Snout length. — Measured from the anterior tip of the snout to the point where a line extended backward from the tip of the snout along the axis of the body meets a line dropped from the anterior rim of the orbit perpendicular to the axis of the body. 1 1 . Length of caudal-fin rays. — Measured from the posterior tip to the point where they disappear into the fleshy caudal base. In forms with a tri-lobed caudal fin four measurements were often taken. The long- est ray in each of the three lobes and the 36 Tulane Studies in Zoology Vol. 7 shortest ray in the upper interlobe were measured. In forms with rounded caudals the longest ray was measured. 12. Predorsal scale rows. — Head scales are counted as one row in ocyurus because only one row approaches the mid-dorsal line (fig. 16). III. Historical Review of the Species of Centropristes Linnaeus (1758) described two species of perciform fishes that have since been placed in the genus Centropristes ( Labrus striatus and Perca philadelphica) . Both descriptions were brief and that of philadelphica was in- adequate. Linnaeus (1766) used Perca atraria for striatus and redescribed Perca philadelphica as Perca trifurca. The new names were used through most of the nine- teenth century, and it was not until late in the century that the descriptions in the tenth edition were recognized. In the sixty years following the original descriptions, the two common species of Centropristes were redescribed under va- rious names and by several authors. Walbaum (1792: 3.36) listed striatus as Perca furva from New York (after Sch(')- epff, 1788). Bloch and Schneider (1801: 297 ) described striatus from New York and named it Coryphaena nigrescens. Lacepede (1803: vol. 4, 196, 246) described phila- delphicus and striatus as Lutjanus tridens and Lutjanus trilobatus, respectively. Mit- chill (1815: 415, 416) presented a satis- factory color plate of striatus and described the form as Perca varia. Centropristes was first proposed by Cu- vier and Valenciennes (1829: vol. 3, 27- 40). According to Bailey (1951) this sec- tion was written by Cuvier who is accorded authorship in the following account. Cuvier (1829b) listed three species from the New World: nigricans (=:striatus) from New York, trifurcus ( =:philadelphicus ) , and rufus (probably striatus) from Martinique. Some question exists about the spelling of the generic name, which appeared as Cen- tropristis in Cuvier's Le Regne Animal, ap- parently published the same year ( 1829) as the Histoire. It is generally accepted that the latter preceded Le Regne Animal and that the correct spelling is Centropristes. DeKay (1842: 24-25) employed Centro- pristes nigricans for striatus and Centro- pristes trifurca for philadelphicus. obviously following Cuvier (1829b). Storer (1846: 9,35) used Cuvier's (1829a: 145) spelling of Centropristis and included therein nig- ricans {=striatus) and trifurca {=phila- delphicus ) . Holbrook (I860: 45-52) accepted Cen- tropristes but used atrarius for striatus. His figures of Centropristes atrarius and Centro- pristes trifurca i^^philailelphicus) are good. Here Holbrook first stated his belief that the northern and southern forms of the black sea bass (striatus) were distinct. Sup- posedly the northern form had the pectoral fin and the pelvic fin contemn inus and a non-sacculated air bladder, while the south- ern form had a sacculated air bladder and the pectoral longer than the pelvic. The line of demarcation between the two was said to be near Cape Hatteras. Various authors accepted Holbrook's ideas and used separate names for populations of striatus north and south of Hatteras until late in the nine- teenth century when Jordan and Eigenmann ( 1890) proved him wrong. Gill ( 1861 ) placed Cuvier's Centropristis in the family "Percoidae". He recognized two forms of the black sea bass ( striatus ) and called the northern form nigrescens and the southern form atrarius. He established the genus Triloburus to include Storer's Centropristis trifurca which he listed as Triloburus trifurcus without a description. The name apparently is based on the tri- lobed caudal fin of adults. Gill (1873) listed Centropristis atrarius ( by implication combining northern and southern populations of striatus) and Tri- loburus trifurctis. Jordan and Gilbert (1882a) placed the two known species of Centropristes in the larger genus, Serranus. and called them atra- rius and trifurcus. They ( 1882b) also gave a good description of a series of fishes taken from the mouths and stomachs of red snappers at Pensacola, Florida, under the name Serranus trifurcus. They noted some differences but could not separate specific- ally the Pensacola bass from Serranus tri- furcus from Charleston, South Carolina. Jordan and Gilbert (1882c) resurrected philadelphicus, listed Serranus atrarius and Serranus philadelphicus as common, and noted that the description of Perca philadel- phica "could not have been identified with No. 2 Miller: Revieiv of Centropristes 37 Figure 1. Sedimentai-y provinces of the Gulf of Mexico (from: U. S. Fish and Wildlife Sei-vice, Fishery Bulletin 89.) this species had not Linnaeus himself in his specimens of Centropristes n/fus Cuvier twelfth edition revised his description . . ." from Martinique. Though he listed the Jordan (1885a) listed S err anus philadel- form as Serranus rufus, he expressed some phicus and gave separate names to northern doubt as to its validity. and southern forms of striatus, calling them Jordan and Evermann (1886) described Serranus furi'us and Serranus atrnrius, re- a third form, Serranus ocyurus. They iden- spectively. In 1886 he examined the two tified fishes from Pensacola (1882b) as 38 Tulane Studies in Zoology Vol. 7 Table 1. yrequency distributions of total gill rakers and pectoral rays in populations Centropristes of Species Number of gill rakers 17 18 19 20 21 22 23 24 25 26 27 28 29 N s. striatus 5 7 15 46 29 16 4 122 26.2 s. melamis 1 7 15 25 12 4 3 67 22.9 ocyurus Gulf Atlantic 1 8 2 32 20 27 17 14 5 1 82 45 19.6 19.5 philadclphicHs Gulf Atlantic 3 3 11 7 33 13 36 25 20 14 9 5 112 67 19.8 19.8 Number of pectoral rays 15 16 17 18 19 20 N s. striatus s. melamis ocyurus Gulf Atlantic philadelphicus Gulf Atlantic 58 188 3 249 18.8 2 79 51 132 17.4 14 137 15 166 17.0 5 78 7 90 17.0 2 18 185 16 1 223 17.9 3 108 23 134 18.1 ocyurus and referred to the previous publi- cation as part of the description. They dis- tinguished ocyurus and atrarius (=:stria(us) which they regarded as allied forms but ignored philadelphicus. Jordan and Eigenmann (1890) used the spelling Centropristis after Cuvier (1829a) and first employed the name striatus ( Linn. Ed. X) for both northern and southern forms of the black sea bass, stating that they could not distinguish them. They listed Centropristis rufus but noted that it could as easily have come from New York as from Martinique. Centropristis philadelphi- cus was described from the south Atlantic coast of the United States, and Centropris- tis ocyurus was listed as a possible deep- water variety of striatus. They applied all references from the Gulf to Centropristis ocyurus. Boulenger (1895), apparently ignoring the earlier Linnaean names, listed Centro- pristes atrarius. ocyurus, and trijurca. Jor- dan and Evermann (1896) gave keys and descriptions for Centropristes striatus, rufus. ocyurus, and philadelphicus and limited ocyurus to the Gulf of Mexico and phila- delphicus to the Atlantic. In Jordan, Ever- mann, and Clark ( 1930) the same species were given but philadelphicus was placed in Triloburus Gill. Since Trilohurus was never described, few people used the name after the turn of the century, and its appearance in the check-list did little to revive it. Weed (1937) described Centropristes springeri from the Gulf of Mexico. Appar- ently he was confused by the large amount of color-pattern variation in Centropristes ocyurus and was convinced that the type of ocyurus was a Gulf specimen of philadel- phicus. He based the name springeri on a specimen with a slightly different color pat- tern. His springeri is a synonym of ocyurus. Ginsburg (1952) described Centropristes melanus from the Gulf and compared it with its Atlantic counterpart, striatus. His species was based primarily on number of gill rakers and pectoral rays. He believed that the Florida peninsula had isolated the Gulf and Atlantic populations and that speciation had occurred subsequently. Currently, four species of Centropristes are recognized from the coastal waters of the United States. A fifth from Martinique is doubtful. IV. Generic Description Centropristes Cuvier Body robust, somewhat compressed, cov- ered with rather large ctenoid scales. Mouth large, teeth in bands, no canines. Tongue smooth. Preopercle serrate, the lower teeth No. 2 Miller: Review of Centropristes 39 Table 2. Frequency distrihutions of scales in populations of Centropristes. Number of scak ! rows from I dorsal-fin origin to lateral' line Species 5 6 7 8 N X s. striatus 36 75 9 120 6.8 s. melanus 4 28 25 1 58 6.4 ocyurus Gulf Atlantic 15 3 57 32 7 7 79 42 6.9 7.1 philadelphicus Gulf Atlantic 84 11 24 52 3 108 66 6.2 6.9 Number of sea le rows from anal-fin origin to lateral line 12 13 14 15 16 17 18 19 20 21 N X s. striatus 7 26 42 29 8 5 117 15.2 s. melanus 1 28 24 9 4 66 13.8 ocyurus Gulf Atlantic 1 2 1 19 4 30 17 21 12 3 6 1 76 41 17.0 17.5 philadelphicus Gulf Atlantic 1 12 34 38 21 15 32 6 18 1 1 1 107 73 16.5 18.0 Number of predorsal scale rows 9 10 11 12 13 14 15 16 17 18 N X s. striatus 1 1 8 20 16 20 26 10 7 109 13.8 s. melanus 9 11 16 7 2 2 47 11.7 ocyurus Gulf Atlantic 2 1 17 7 31 20 20 12 1 1 71 41 12.0 12.1 philadelphicus Gulf Atlantic 2 6 2 22 12 35 18 15 11 20 13 8 8 1 108 64 14.4 14.8 somewhat antrorse. Gill rakers rather long and slender. Supraoccipital and parietals with strong crests extending forward to a point between postfrontal processes; frontais posteriorly with an angular transverse ridge in front of supraoccipital connecting the parietal crests; posterior processes of pre- maxillaries not reaching frontais. Smooth area on top or cranium short and small. Dorsal fin X, 11, the spines with fleshy filaments at tips; anal fin III, 7; caudal fin typically 17, rounded in young, trilobed or double concave in adults; pelvics I, 5, close together, inserted under or slightly in ad- vance of pectorals. The species of Cetitro- pristes are closely related. I agree with Weed (1937) in synony- mizing Trilobun/s Gill (based on philadel- phicus) with Centropristes. Further, the evidence refutes the retention of Triloburus as a subgenus. C. philadelphicus differs from ocyurus in about the degree ocyurus differs from striatus and ocyurus is reason- ably intermediate in general structure be- tween the two. The data in Tables 1-10 show that ocyurus is approximately inter- mediate in most of the characteristics ex- plored. The species of Centropristes have been found in coastal waters from the Gulf of Maine to the Yucatan banks. Though es- sentially a temperate and subtropical group, certain forms have apparently been able to acclimatize themselves to the colder waters off Maine and the tropical climes of the southern Gulf of Mexico. In the following discussion Centropristes rufus. the doubtful species based on two specimens supposedly from Martinique, will not be dealt with because of the absence of additional records from the West Indies. V. Key to the Species of Centropristes 1. Ground color dark, usually dark brown in alcohol, smoky gray to dusky brown or blue black 40 Tulane Studies in Zoology Vol. 7 Figure 2. Distribution of C. s. striatus. in life. Exposed parts of scales paler than margins, giving appearance of being barred with longitudinal ser- ies of pale dots. Seldom with traces of seven vertical bars on sides. Belly only slightly paler than sides, never white. Fins dark, dorsal marked with several series of whitish spots and bands. Deimal flaps ex- tend only slightly past tips of dorsal spines ( Black Sea Basses, C. striatus) sides, usually white, occasion- ally with traces of dusky pig- mentation. Fins pale, some- times fairly dusky but never black; soft dorsal and caudal with dark to inky spots — Total gill rakers and tubercles on first arch usuallv 22 to 24 (20-26) ; pectoral rays 17 or 18; jaw usually 18-19 (per cent of standard length) ; pectoral usually 26-28 (2.3- 29) ; young with dark stripe from opercular flap to cau- dal, but with black spot at base of last dorsal spines rather indistinct to almost absent; four dai'k blotches on upper and lower jaws promi- nent in young (fig. 23). West coast of Florida from Pensacola to Placida Gulf Black Sea Bass, C. striatus melanus Ginsburg Gill rakers and tubercles usu- ally 25 to 26 (23-29) ; pec- toral rays 18 or 19; jaw usu- ally 16 or 17 (14-18) ; pec- toral usually 28-29 (26-30); young with dark lateral stripe and distinct black spot at base of last dorsal spines; dark blotches on jaws usually absent in young. Atlantic coast, common from Cape Cod south to Cape Canaveral; oc- casionally found north to Gulf of Maine and south to Florida Kejs Atlantic Black Sea Bass, C. striatus striatus (Linnaeus) — iW^""^^ Ground color very pale to white, with .seven diffuse to distinct dark vertical bars on sides. Belly considerably paler than Figure 3. Distribution of C. s. melanus. No. 2 Miller: Review of Centropristes 41 3. Dorsal spines with short dermal flaps rarely extending beyond tips of spines. No distinct black spot at base of last three dorsal spines. Distinct black blotch at center of fourth vertical bar just below lateral line (the lateral spot). Lateral bars usually well marked, often broken up by two pale stripes into seven series of three black blotches each. Lowest third of bars often represented by groups of small, inky spots. Pectoral rays usually 17 (16-18) ; bodv depth usually 31-34 (28-38); interorbital usually 6-8 (2- 9) ; pelvic fin usually 24-26 (20-29). Atlantic arid Gulf of Mexico, from Cape Hat- teras to Yucatan, usually over hard bottom Gulf Sea Bass, C. ocyurus (Jordan and Evermann) Dorsal spines with long fleshy filaments, often as long or longer than spines, pale in al- cohol, yellow, orange or red life. Distinct black spot at base of last three dorsal spines (the dorsal spot). No distinct black blotch at cen- ter of fourth vertical bar. Lateral bars usually poorly marked, diffuse; seldom bro- ken up into three series. Low- est third of bars never as inky spots. Pectoral rays usu- ally 18 (14-19); body depth usually 27-30 (25-33) ; inter- orbital usually 4-5 (4-6) ; pel- vic fin usually 20-22 (19-24). Atlantic and Gulf of Mexico, from Cape Hatteras to Texas, straggling north to Chesa- peake Bay. Often over hard bottom Rock Sea Bass, C. philadelphicus (Linnaeus) Figure 4. Distribution of C. ocyurus. 42 Tulane Studies in Zoology Vol. 7 VI. Species Accounts Abbreviations used for collections: ANSP, Academy of Natural Sciences of Philadel- phia; AMNH, American Museum of Nat- ural History; CHML, Cape Haze Marine Laboratory; CNHM, Chicago Natural His- tory Museum; CU Cornell University; FU, University of Florida; MCZ, Museum of Compartive Zoology, Harvard University; UNC, Institute of Fisheries Research, Uni- versity of North Carolina; TU, Tulane Uni- versity; Ore. sta. no.. United States Fish and Wildlife Service M. V. Oregon stations; and USNM, United States National Museum. Collections listed were examined and are represented in the data, at least in part. Numbers in parentheses refer to the number of specimens and their ranges in standard length in mm. Species descriptions are modified after Jordan and Evermann (1896) and Gins- burg ( 1952). Characters used are presented in identical sequences for the several species. Counts and measurements are based on the samples presented in Tables 1-10 except where otherwise noted by enclosure with brackets. Extreme ranges for the several meristic characters are given in parentheses. In order to assess clearly differences be- tween Gulf and Atlantic populations of each form, discussions of data will deal primarily with three kinds of measurable character- istics. Meristic data are the easiest and most exact to procure. They are presented in the form of frequency distributions in Tables 1 and 2. The proportional measurements are not so exact because they are influenced by the condition and shape of each speci- men examined. All measurements were con- verted to percentages of standard length, and the resulting data were plotted against standard length on graphs (figs. 25-37) and tabulated ( to the nearest whole per cent of standard length) in Tables 3-11. Color pattern cannot be analyzed in the same manner as other characteristics and valid comparisons can be made only after exact descriptions have been made of the existing range of variation. Consequently, pigmentation comparisons are based on vir- tually the entire available sample of Cen- tropristes. During the course of this study certain information on the biology and natural his- tory of fishes in the genus was revealed. When possible my observations have been correlated with those described in the liter- ature. A scattergram plotting the per cent of standard length against standard length was made for each measurement of Gulf and Atlantic populations of the three species. Figure 5. Distribution of C. philadelphicus. No. 2 Miller: Review of Centropristes 43 Since total scattergrams number over a hun- dred, only those showing some variation with size will be discussed. Scattergrams for Gulf and Atlantic populations are usu- ally very similar and have, as a rule, been combined. In certain instances, however, both will be presented to illustrate the dif- ferences present. A. Centropristes striatus (Linnaeus) Black Sea Bass (Figures 8-12) For pertinent synonymy for this and other species see Jordan, Evermann, and Clark ( 1930) and previous section on history of the genus. Pigures 6-8. 6. (upper) Female C. philadelphicus, UNC, 157 mm SL. 7. (middle) Male C. philadelphicus, UNC, 192 mm SL. 8. (lower) Adult C. s. striatus, CU 22038, 159 mm SL. 44 Tulane Studies in Zoology Vol. 7 1. Material Examined Atlantic populations Centropristes striati/s striates (124 specimens): CNHM 7109 ( 1, 210) Buzzards Bay, Mass.; USNM 148150 (1, 146.8) Wood Hole. Mass.; ANSP 13676-77 (2, 50-187.2) Woods Hole, Mass.; USNM 163712 (4, 42.9-203.5) Woods Hole, Mass.; USNM 20618 (5, 33-66.4) Woods Hole, Mass.; TU 18125 (2, 37-59.7) Woods Hole, Mass.; TU 18126 (1, 78.4) Provincetown, Mass.; CU 18607 (1, 215) Sandy Neck, Cape Cod Bay, Mass.; CU 14372 (1, 91.4) Niantic R. at Waterford, Conn.; USNM 118242 (5, 58.3-88.5) Old Bare Shoal, N. J.; ANSP 18569-70 (2, 116.3-102.4) N. J.; ANSP 33062-63 (2, 107.8-114.7) N. J.; USNM 755 (2, 110.8- 162) Beesleys Pt., N. J.; USNM 49540 (1, 319) West End, N. }.; CU 22038 (7, 121- 158) Sandy Hook, N. J.; USNM 76703 (1, 154) Ocean City, Md.; TU 18127 (2, 116.5- 140.7) Ocean City, Md.; USNM 89815 (1, 147) Cobb Id., Potomac, Md.; FU 886 (4, 795-112.6) York R., 1 mi. off Tue. Pt. in Channel, York Co., Va.; CU 31015 (13, 67.7-101.3) Bogue Sd., N. C; TU 15057 (4, 78-149) 35" 07' N., 75° 13' W.; TU 17065 1, 129) 35" 07' N., 75° 14' W. CU 30940 (6, 115-179) Beaufort, N. C. CU 30747 (1,67.5) White Oak R., N.C. CU 30872 (3, 62.3-78.7) Bogue Sd., N.C. CU 31001 (2, 69.6-111) Bogue Sd., N.C. CNHM 40342 (6, 67-152.8) Beaufort Harbor, N.C; CNHM 37947 (2, 95.8- 123.2) Charleston Harbor, S.C; TU 14745 (1, 181) 33" 19' N., 78° 07' W.; TU 16327 (1, 145.2) 31° 55' N., 80° 19' W.; FU 5560 (1, 123.8) 3 mi. off St. Simons Is., Ga.; USNM 127486 (3, 120.8-132.3) Cumberland R. Sd., Ga.; USNM 127490 (5, 75.6-108.9) Cumberland Is., Cumber- land Sd., Brunswick, Ga.; CNHM 64295 ( 1, 176.2) Ga.; USNM 155182 (1, 97.7) Ga.; USNM 18033 (6, 91.1-133) Matanzas In- let, Fla.; USNM 53339 (1, 97.5) Hillsboro R., New Smyrna, Fla.; FU 678 (1, 129) 1 mi. S. Crescent Beach, St. Johns Co., Fla.; FU 3315 (1, 79.2) Vilano Beach, St. Johns Co., Fla.; TU 14791 (1, 139) 29° 03' N., 79° 59' W.; USNM 18033 (1, 114.4) Matanzas Inlet, Fla.; USNM 158084 (1, 145) S. of Miami, Fla.; USNM 155184 (2, 112.1-113.7) Cape Canaveral, Fla.; USNM 155183 (2, 82.8-109.2) Cape Canaveral, Fla.; USNM 155181 (1, 79.1) Cape Cana- veral, Fla.; USNM 62627 (5, 72.8-152.9) St. Augustine, Fla.; CNHM 64300 ( 1, 196) 30" 11' N., 81" 02' W.; TU 14722 (1, 113.5) 30° 20' N., 81° 06' W.; USNM 42383 (3, 38.2-45.7) Fla. Keys; ANSP 39934 (1, 100) Hawk's Park. Volusia Co., Fla. Gulf populations Centropristes striatus melanus (68 specimens ):' CNHM 50494 (1, 84.6) Manatee Bay, Englewood, Fla.; FU 1438 (1, 100.4) Clearwater, Fla.; CU 14059 (3, 233-270) Clearwater, Fla.; CU 12882 (1, 161) Clearwater Bay, Fla.; CU 26132 (7, 130.5-197) Gulf of Mex., Clear- water vie; TU uncat. (5, 54.5-83) mouth of Anclote R., Fla.; Ore. sta. no. 1721 (4, 161-190.6) 28° 33' N., 83" 08' W.; CNHM 64278 (2, 178-181) 28° 35' N., 83" 11' W.; FU 1452 (2, 135-156.2) Yankeetown ( Withlacoochie R.) Levy Co., Fla.; ANSP 21042-44 (3, 177-189) Bayport, Fla.; FU 2154 (1, 139) Cedar Keys (Gulf of Mex.); FU 832 (3, 86.6-103.6) Cedar Keys, Fla.; FU 674 (2, 97.5-97.6) Cedar Keys, Fla.; FU 2738 (4, 141-169.5) Cedar Keys, Fla.; FU 915 (3, 77.8-107.3) Cedar Keys, Fla.; FU 815 (3, 68.2-160.7) Cedar Keys, Fla.; FU 296 (11, 53.7-96.5) Cedar Keys, Fla.; TU 11938 (6, 49.7-69) Cedar Keys Islands, Fla.; FU 2124 (6, 42.5-88.1) Cedar Keys, Fla. 2. Description Dorsal X, 11; anal III, 7; caudal 17; pelvic I, 5; pectoral usually 16 to 19 (14- 20); vertebrae 24 [15}, rarely 23; pyloric caecae 6 (4-7) {9}. Scales large, 7 or 8 — 47 (46-49) — 14 to 17 (13-19); small scales at base of soft-dorsal membranes in two rows, scales at base of anal membranes in single row extending one-third length of membranes; caudal membranes scaled one- third to one-half of length from base; pec- torals scaled only near base; predorsal rows 11-15 (8-17). Total gill rakers plus tuber- cles usually 22-28 (20-29), lower limb usu- ally 14 to 18 (14-19); gill rakers rather long and slender at center of arch grading to small tubercles at both ends of arch. Body robust, the back somewhat elevated anteriorly. Head length usually 39-42 (37- 44) per cent of standard length; head large, thick, little compressed. Opercle drawn out to form broad, thin flap; middle opercular spine well developed, lower rather poorly No. 2 Miller: Review of Centropristes 45 Table 3. Frequency distribution of proportional characters, expressed as percent of standard length, in populations of Centropristes. Dorsal-fi n origin to snout Species 33 34 35 36 37 38 39 40 41 42 N s. striatus 1 1 11 12 12 8 12 7 1 1 66 s. melanus 1 5 8 6 5 14 4 1 44 ocyurus Gulf Atlantic 10 3 15 3 15 15 19 13 14 8 6 2 79 44 philadelphicus Gulf Atlantic 2 2 5 6 20 14 29 17 22 16 8 4 2 1 89 59 Anal-fin orig-in to caudal-fin base 38 39 40 41 42 43 44 45 46 47 48 N s. striatum 1 3 6 16 16 17 4 2 1 66 s. melanus 1 3 6 12 13 6 3 1 45 ocyurus Gulf Atlantic 2 2 1 9 3 12 3 20 11 20 15 12 5 3 4 2 80 44 philadelphicus Gulf Atlantic 1 1 2 3 11 11 14 10 22 18 15 12 5 8 1 2 72 64 Dorsal-fin origin to lateral line 8 9 10 11 12 13 14 N s. sti'iatu^ 6 27 25 5 1 64 s. melanus 1 19 17 7 44 ocyurus Gulf Atlantic 4 35 20 37 19 4 3 1 80 43 philadelphicus Gulf Atlantic 20 1 48 49 4 13 72 63 developed, upper in form of a small, blunt protuberance. Top of head, suborbital, maxillary, lower jaw, and snout naked; cheek scaled in about 11 rows; opercle with large scales in about 8 or 9 rows. Orbit usually 8 to 11 (6-12), about 4 in head. Eye some- what longer than deep. Upper jaw usually 16 to 19 (14-21); mouth large, terminal; lower jaw subequal to upper in front or slightly projecting. Teeth in jaws in wide bands, outer and inner teeth slightly en- larged; no canines; teeth only slightly de- pressible; vomerine teeth in crescent shape; patch on palatines long and narrow. An- terior naris with large dermal flap. Posterior border of preopercle finely serrate, the angle and lower border with larger teeth, some antrorse. Branchiostegals 7. Dorsal spines rather strong, first three evenly graduated, first about half as long as second, third al- most as long as fourth, the longest; follow- ing spines decrease gradually in length to tenth, which is slightly shorter than first dorsal ray; emargination between spinous and soft parts of dorsal slight; soft dorsal short, high; longest ray 18-34, males having noticeably higher fins; pectoral fin long, usually 26 to 29 (23-30), posterior margin nearly truncate, rounded at angles, more so below than above; pelvic long, usually 23- 28 (21-30), reaching to anus or just short of it, outer angle in front of lower pectoral angle; anal spines graduated, second spine a little shorter and stouter than third; soft anal short, high; fifth or sixth rays longest, usually 20-23 (17-27); caudal in small specimens asymmetrical, rounded for its greater and lower part, a moderate emargi- nation above, the second branched ray from the top moderately or slightly produced; larger specimens often three lobed, with middle and bottom rays produced; either middle or top lobes longest in tri-lobed specimens, top lobe usually longest in speci- 46 Tulane Studies in Zoology Vol. 7 ^^'^ ^g? Fi{i;ures 9-11. 9. (upper) Young- C. s. striatum, USNM 163712, .59.6 mm SL. 10. (mid- dle) Adult C. s. rnelanns, UF 815, 161.1 mm SL. 11. (lower) Young C. s. mclanus, CHML, 59 mm SL. mens with rounded caudal; longest ray 18-29. Color pattern in alcohol. — General ground color dark to nearly black; often with faint traces of wide darker cross-areas, separated by narrower, slightly lighter interspaces; scales with a lighter colored area on exposed part anteriorly, surrounded peripherally with dark pigment, presenting in gross appear- ance effect of bead-like longitudinal lines of light spots along rows of scales; dorsal dark with several series of elongate whitish spots forming oblique light stripes; pectoral light to dusky; pelvic and anal dark to nearly black, sometimes edged with white; caudal almost uniformly dusky or dark, sometimes with faint spots. Young with a black longi- tudinal band, which later breaks up, sides No. 2 Miller: Review of Centropristes Al variously mottled or with vertical cross shades; belly area paler than adults; large black spot at bases of last three dorsal spines. Color of adult in life (Hildebrand and Schroeder. 1929). — "Blue black, centers of scales pale to white, forming longitudinal lines; several bluish streaks in front of or below eye present or absent; dorsal blue- black, with pale stripes more numerous on soft part, caudal dusky or dark blue, streaked or mottled with pale markings; anal almost entirely pale or streaked with dark blue; ventrals bluish gray, the rays pale; pectorals grayish. Females are lighter than males, usu- ally gray-blue instead of blue-black." Males clevelop an adipose hump on the nape which is bright blue in the breeding season. Color is somewhat variable in this species, with individuals that have bronze or reddish brown hues variously placed on the fins and body being taken commonly. In general the basic color of the species varies from dark reddish brown to blue to almost black. Accounts of color differences in spawn- ing striatus are found also in Breder ( 1932 ) and Lavenda ( 1949). No notes are available on color of live specimens from the Gulf, but presumably it is similar to that of Atlantic forms. 3. Distribution Centropristes striatus striatus is found commonly from Cape Cod south to Cape Canaveral. There are occasional summer records from the Gulf of Maine and more frequent occurrences south to Miami, Flor- ida. In addition I have examined three small specimens which I identified as stri- atus from the Florida Keys. This form ap- pears to be rare south of Miami, but per- haps small concentrations exist in suitable habitats about the Keys ( fig. 2 ) . Centropristes striatus melanus is found on the west coast of Florida between Pensacola Table 4. Frequency distribution of pro^oi-tioifil characters, exvressed as percent of sta)idard length, in rorulat'ions of Cen+'>'o^^ri''t^earing at the angle of the preopercle; lateral bars are more interrupted, with the lower one-third becoming scarcely more than a series of inky spots. The first two bars in the occipital region differ con- siderably. The center of each scale in the upper four to six scale rows in these bars has an inky black spot in it. Some females have a vague suggestion of these spots, but none are developed to the intensity found in males. Possibly these spots are associated in some way with the courtship and breed- ing behavior of these fish and may be analogous to the nuchal hump in breeding male striatus. In the larger male the lateral bars are almost completely broken up, and the upper part of the body is a slightly mottled dark gray. The lateral spot is very dark and extends forward to the end of the opercle. This gives the appearance of a dark lateral bar on the anterior part of the body. Ventral and anal fins of males are darker and the caudal fin is darker than that of the female, and the spots in the caudal fin membranes are larger and more pro- nounced. The dorsal fin of the smaller male is similar to the typical female pattern, but the larger male's is not. Here the spots on the spines are lamost absent, whereas the bases of membranes between them are much darker. This darkening of the membrane bases extends back over the entire length of the soft dorsal as well as the spinous dor- sal. Dusky rows are absent on the mem- branes of the soft dorsal, and only a light uniform duskiness prevails. There are more melanophores along the mid-ventral area of the males; the belly thus appears to have a light dusky stripe from the anus to a point anterior to the pelvic fins. Sexual dimorphism becomes more pro- nounced with the approach of the spawning season. Though males are larger and have much longer fin rays, they also appear to assume a distinctive color pattern (see above) as they become reproductively moti- vated. After spawning, color differences diminish and eventually all but disappear. 3. Distribution Centropristes ocyurus ranges from Cape Lookout, North Carolina, in the north to the Yucatan banks of the southern Gulf of Mexico. These fish appear to be most com- mon in areas where there are coral patches or other types of hard bottom in rather deep water (figs. 1, 4). Although they are taken at times over mud and sand bottoms, this habitat is not typical. This form differs from the others in that it thrives in tropical waters off southern Florida and Yucatan. It is interesting to note that the species is un- recorded from most of the western Gulf, an area with extensive mud or blue-mud bot- tom. Although I have no records of this form from the east coast of Florida south of Cape Canaveral, the probability seems slight that a fish which thrives in tropical waters around the Tortugas would be excluded by temperature (as Ginsburg suggests) from living in or moving through the waters off the southeast coast of Florida. The Florida peninsula probably does not form a barrier 58 Tulane Studies in Zoology Vol. 7 between Gulf and Atlantic populations of this species. This subject will be discussed more fully later in this paper. 4. Population Analysis Meristic data for the Gulf and Atlantic populations of ocyurus are presented in Tables 1 and 2. These data clearly indicate the uniformity of the two populations, and in no case is there any material differentia- tion. The highest average divergence occurs in the number of scale rows between the anal-fin origin and the lateral line (Table 2 ) , and that is far less than a racial level of differentiation. Comparison of proportional measurements (Tables 3-10) shows the two populations to be nearly identical. On the basis of counts and body measurements, there is no reason for separating the Atlantic and Gulf popu- lations at any level. In comparing Gulf and Atlantic popula- tions of ocyurus with respect to color pat- tern, the total variation present was neces- sarily investigated. It was found that a selection of nine series included a high per- centage of the different patterns found in the complete sample examined. One series (CNHM 64089, described above) was chosen as a standard with which the others were compared. The following series were examined intensively and are representative of the kinds of variations found within the species: Ore. sta. no. 417; Ore. sta. no. 892; TU 12999; TU 17061; TU 17049; CNHM 45761; TU 16969; and CU 12869. Most differences are due to varying in- tensities of pigmentation. Small specimens, both male and female, tend to have rela- tively diffuse lateral bars, and there is usu- ally little spotting on the body. There is, however, considerable variation in this basic pattern, as can be seen in Figs. 13-15. Spotting on the head varies as does the intensity of the color pattern of the fins. Although there is much variation, Gulf and Atlantic populations cannot be separated on the basis of coloration observed in preserved speci- mens. In summary, comparison of the two popu- lations shows them to be similar, if not identical, in observable characters. The dis- tribution map shows a hiatus off southeast- ern Florida, but this gap may be closed with additional collecting in deep water. Fishes capable of enduring summer water tem- Fig'ure 16. Occipital region of C. ocyurus, showing- scaleless area on nape. Figure 17. Occipital region of C. philadel- phicas showing completely scaled nape. peratures on the Campeche banks or at the Dry Tortugas hardly would find the water temperature off the southeastern tip of Flor- ida a barrier to migration. Bottom types or other ecological conditions perhaps asso- ciated with the strong push of the Gulf Stream may have some effect in limiting the distribution of this species off southern Florida, but this seems unlikely. I conclude, therefore, that the two pop- ulations are not isolated from each other and that the genetic interchange between them is primarily responsible for their ob- servable similarities. 5. Biology Spawning. — No information about spawn- ing in this form is available in the literature. Specimens collected between December and March in the Gulf have well developed ovaries. By July, Gulf specimens are spawned out, and gonads in many cases have become small and shrivelled. There are no records for May or June, but it appears that No. 2 Miller: Review of Centropristes 59 spawning occurs sometime between late March and early June. Atlantic specimens appear to be spawned out by late June, though the males still have rather well developed testes at that time. Young of this species are never found in shallow water, so the entire life of the fish, from egg to spawning adult, probably is spent in deep water, usually where the bot- tom is rocky or covered with patches of coral. Grotvth and Body Proportions. — Measure- ments discussed under striatus are illustrated for ocytmis in Figs. 21-29. In many cases the measurements change relative to size in the same manner as they did in striatus. though the actual percentages may be dif- ferent. It is profitable to compare ocyurus to striatus in regard to some of these char- acters. Rate of increase in width of the inter- orbital is somewhat greater than in striatus (fig. 21), as is the rate of increase in the snout length (fig. 22). Orbit length (fig. 23) variation is comparable to that in striatus. Upper jaw length (fig. 24) seems to increase more sharply in ocyurus. as does dorsal fin length (fig. 25). The anal fin in ocyurus is relatively shorter (fig. 26) and does not show a very clear increase in length in larger specimens. The pectoral (fig. 27) and pelvic fins (fig. 28) are shorter in larger fishes, but the caudal-fin length (fig. 29) varies greatly. Few speci- mens under 90 mm. standard length have very long caudal rays, but those larger have fins that range in size from 21.7 to 56.2 per cent of the standard length. Either the top or the middle lobe can be longest, but the four longest fins recorded were from the middle lobes. C. Centropristes philadelphicus (Linnaeus) Rock Sea Bass (Figures 11-12, 17) 1. Material Examined Atlantic populations (67 specimens): USNM 142814 (2, 183-191) 40-60 mi. E.S.E. Cape Henry, Va.; UNC uncat ( 10, 110-188) off Cape Lookout, N.C.; USNM 126814 (2, 70.5-79.6) Beaufort, N.C.; USNM 101560 (1, 88.8) 25° 23' N 75° 24' W.; USNM 53404 (1, 83.6) Beau- fort, N.C.; TU 17057 (1, 144) 35° 07' N, 75° 13' W.; TU 17066 (2, 142-175) 35° 07' N, 75° 14' W.; UNC uncat (9, 105.8-195) 71/2 mi. E. Knuckle Buoy, Cape Lookout, N.C.; UNC uncat. (3, 172.4- 184.4) 18 mi. E.S.E. Cape Lookout, N.C.; UNC uncat. (1, 192.5) 13 mi. S.E. Cape Lookout, N.C.; UNC uncat (4, 113.8- 188) 14 mi. E.S.E. Cape Lookout, N.C.; USNM 25959 (1, 168.2) Charleston, S.C; USNM 25305 (2, 109.5-124.2) Blackfish Banks, 12 mi. off Charleston, S.C; USNM 33161 (1, 169) Charleston, S.C; ANSP 13638-40 (3, 132.2-185) S.C; USNM 127491 (3, 106.1-108.5) St. Simon's Sd., Brunswick, Ga.; USNM 127492 (3, 104.1- 115.2) Cumberland R. Sound, Ga.; USNM 92202 (1, 102.7) Savannah, Ga.; FU 5424 (6, 95.3-121) off mouth St. Simon's Sd., Ga.; FU 5561 (3, 101-109.4) 3 mi. off St. Simon's Is., Ga.; USNM 155159 (1, 173.1 ) 28° 57' N, 80° 43' W.; USNM 155139 (1, 108.5) 28° 03' N, 79° 55' W.; USNM 155145 (1, 130.7) 28° 12' N, 81° 31' W.; USNM 124391 (1, 164) off Cape Ca- naveral, Fla.; USNM 53349 (1, 118) Hills- boro R., New Smyrna, Fla.; TU 14803 (1, 113.1) 27° 57' N, 80' 04' W.; USNM 155158 (1, 172) Cape Canaveral, Fla.; CNHM 50766 (1, 189.2) Palm Beach, Fla. Gulf Populations (111 specimens ) : TU 10641 (1, 152.5) 26° 10' N, 96° 40' W.; Ore. sta. no. 1086 (3, 46.9-124.4) 26° 10' N, 96° 54' W.; Ore. sta. no. 1083 (1, 41) 26° 13' N, 96° 45' W.; TU 12954 (2, 112-113.8) 26° 10' N, 97° 00' W.; TU 13225 (2, 93-164) 26° 10' N, 96° 40' W.; TU 12916 (3, 125-163.3) 26° 13' N, 96° 45' W.; TU 12914 (1, 123.5) 26° 11' N, 96° 52' W.: CNHM 45133 (4, 138.8-170.2) 28° 20' N, 96° 40' W.; CNHM 40262 (5, 65.5-124.6) less than 50 mi. off Corpus Christi, Tex.; USNM 142813 (9, 76.8-95) Corpus Christi, Tex.; CNHM 45140 (2, 147.9-175.4) 27° 34' N, 96° 43' W.; CNHM 45134 (2, 146.2-184) 27° 37' N, 96° 39' W.; CNHM 45132 (1, 171) 27° 20' N, 96^ 40' W.; TU 16227 (2, 130-177.6) 28° 08' N, 93° 35' W.; TU 13006 (8, 133.2-154) 28° 41' N, 89° 32' W.; TU 12124 (1, 171.6) 28° 46' N, 89° 58' W.; TU 8821 (17, 52.2-81.7) Pass-A-Loutre, 140° from old light tower. La.; TU 6093 (1, 56.8) Bayou Taylor to Pass des Isle, Gulf of Mex., La.; TU 5347 (2, 76.5-99.2) mouth Oyster Bayou, La.; 60 Tulane Studies in Zoology Vol. 7 TU 9038 (1, 117.8) Lake Borgne, La.; TU 4075 (1, 134.5) mouth Oyster Bayou, La.; TU 9328 (1, 84.2) Lake Pontchartrain at Rigolets, La.; TU 8398 ( 1, 59.5) S.W. Pass, outside Vermillion Bay, La.; TU 16295 (1, 137.7 j 29" 04' N., 88° 43' W.; TU 11658 (4, 102.5-119.2) 29° 17' N., 88° 51' W.; TU 5404 (2, 86-147) 29° 15.5' N., 88° 55.8' W.; TU 5410 (11, 92-186) 29° 10' N., 88° 52' W.; TU 4217 (5, 94.5-138) mouth of Bayou Creole, La.; CNHM 45767 (4, 137-161) 29° 27' N., 87° 45' W.; CNHM 31830-31 (2, 100-103.8) Horn Is., Miss.; CNHM 45763 (3, 67.8-128.1) 30° 00' N., 88° 07' W.; CNHM 45768 (4, 173-209) 29° 30' N., 86° 10' W.; CNHM 33737 (1, 101.6) La.; TU 11768 (1, 187) 29° 50' N., 86° 30' W.; TU 13207 ( 1, 152) 29° 50' N., 86° 30' W.; TU 13198 (1, 150) 29° 50' N., 86° 30' W. The following 148 specimens were ex- amined and are included in discussions of species but are not represented in the tabu- lated data: AMNH 16332 (6, 161-183) Near Chavin, La.; AMNH 16385 (2, 82.3- 93) La.; AMNH 16333 (24, 39.8-117.3) near Chavin, La.; AMNH 16478 (1, 101.6) La.; ANSP 55864-69 (5, 93.2-112.3) Bre- ton Is., La.; ANSP 65160-61 (2, 88.2-177) Corpus Christi, Tex.; ANSP 75259 d, 93) 1 mi. off St. Aug., Fla.; TU 4067 (1, 85.3) mouth of Oyster Bayou, La.; TU 5566 (3, 86.5-98) Gulf Mex. near Oyster Bayou, La.; TU 5390 (45, 66-129.7) 29° 15' N., 88° 55' W.; TU 10007 (1, 68.1) Lake Pelto, La.; TU 4182 (10, 87.5-122) 4 mi. W. Atchafalaya R., La.; TU 13745 (4, 49.6- 66.5) Bayou Hero, La.; MCZ 2920 (3, 151.5-175) Charleston, S.C; TU 6046 (3, 49-54) Lake Pelto, La.; TU 9004 (11, 86.8- 117.7) off mouth Oyster Bayou, La.; TU 10620 (15, 36.7-138) 26= 10' N., 96° 54' W.; MCZ 3067 (6, 156-178.3) Charleston, S.C; TU 2774 (1, 71.4) 28° 44' N., 90° 09' W.; USNM 155162 (1, 32.4) (Poor specimen; may be ocyurus), Apalachicola Bay, Fla.; USNM 155138 (1, 72.5) New Smyrna, Fla.; USNM 155142 (1, 123) Cape Canaveral, Fla.; USNM 155147 (1, 128) New Smyrna, Fla. 2. Description Dorsal X, 1 1 ; anal III, 7; caudal 17; pel- vic I, 5; pectoral usually 18 (15-20); ver- tebrae usually 24 [17], rarely 22 or 23; pyloric caecae 6 [4]. Scales large, 7 or 8 — 47 (46-49) — 17 to 20 (15-22); small scales at base of soft dorsal membranes in single rows, scales at base of anal mem- branes in single rows extending one-half or less the length of membranes; caudal mem- branes scaled one-third to one-half of length from base; pectorals scaled only near base; pelvic rays finely scaled on underside for one-third length from base; predorsal rows usually 13-16 (11-19). Total gill rakers plus tubercles usually 19-21 (17-22), lower limb about 13; gill rakers long and slender at center of arch, grading evenly to small mbercies at ends of arch. Body robust, back elevated less than in C. striatus. Head length usually 38-41 (35-43). Head large, not as deep as in C. striatus, little compressed. Opercle drawn out into thin flap; middle opercular spine well developed, lower rather poorly developed, and upper barely visible. Top of head, suborbital, maxillary, lower jaw and snout naked; cheek with small scales in 9-11 rows; opercle with larger scales in 8 or 9 rows. Orbit usually 9-11 (7-13), about 4 in head. Eye longer than deep. Upper jaw usually 17-19 (16-21); mouth large, terminal; lower jaw projecting slight- ly beyond tip of upper. Teeth in jaws in bands, outer and inner teenth slightly to moderately enlarged, though none can be called canines; teeth only slightly depressi- ble; patch on vomer V-shaped, patch on palatines long and narrow. Nares very close to eye, anterior naris with large dermal flap. Posterior border of preopercle finely serrate, serrae on ventral border and at angle of pre- opercle somewhat larger and farther apart. Branchiostegals 7. Dorsal spines moderate, first three evenly graduated; first about one- half as long as second; third about as long Fig:ure 18. Anal fin of adult female ( up- per) and male (lower) C. philadelphiciis. No. 2 Miller: Review of Centropris(es 61 as fourth, one or the other being longest in fin; following spines decrease in length to tenth, which is shorter than first dorsal ray; spines with fleshy filaments attached to tips, filaments in large specimens often as long as spines; emargination between spinous and soft parts of dorsal moderate; soft dorsal short, high; longest ray 17-37, large males with noticeably higher fins; pectoral long, usually 25-28 (22-29), posterior margin nearly truncate, rounded at angles, more so below than above; pelvic long, pointed, usu- ally 20-22 (19-24), seldom reaching anus; outer angle in front of lower pectoral angle; anal spines graduated, second spine some- what shorter and stouter than third; soft anal short, high; fourth to seventh rays longest, usually 16-18 (15-29), mature males with obviously longer rays; caudal in small speci- mens rounded, sometimes with upper rays slightly produced; middle rays also some- what produced in medium-size ( approx. 100 mm.) specimens, large specimens dis- tinctly three lobed, with upper and middle lobes greatly produced; longest ray 21-57, mature males with much longer rays. Color pattern in alcohol. — General ground color pale to white; seven dark vertical bars along side of body, usually diffuse, never breaking up into series of dark spots, some- times interrupted by a pale longitudinal streak just above lateral line. Top of head and snout spotted and streaked with dusky color; dusky spot at ventral posterior angle of eye, scattered melanophores present on opercle including flap, and on tip of lower jaw; dorsal fin with several series of elongate dusky spots forming oblique dark stripes; large black spot at base of last three dorsal spines; pectoral pale; pelvic dusky, edged with white; anal clear, dusky color at base of membranes and at ends of fin rays, tips of rays edged with white; caudal spotted. Young (40 mm. or larger) similar to adults in color pattern. Color in life. — From Jordan and Ever- mann (1896): Olivaceous above, whitish below; 7 broad brown bars from dorsum obliquely forward to level of middle of pectorals, these almost obsolete along lateral line; the color of the bars is not intense and is formed by shadings along the base and margins of the scales; the anterior bar crosses the nape and is very indistinct; snout and upper part of head with numerous brownish- red spots and lines, three or four of these parallel, stretching from eye to snout, the interspace usually light blue; upper lip red- dish brown; tip of lower jaw broadly pur- plish; a dark blotch on opercle anteriorly and sometimes a small, dark spot behind eye; lining of opercle and throat lemon yellow; a large black blotch behind pseudo- branchiae; spinous dorsal translucent, with indistinct whitish and dusky longitudinal streaks; a large dark blotch on membrane of last spines immediately above fourth verti- cal bar of sides; some dark spots on the spines form two irregular lengthwise series; dorsal filaments bright scarlet, the fin usu- ally with light-bluish shading; soft dorsal with a series of blue-white spots near margin (one between each two rays), one or more incomplete series above and below this; the fin is margined with reddish brown, and has usually several series of reddish-brown spots, these most numerous posteriorly; some irregular olive-brown spots toward base; a small black spot on base of mem- brane between eighth and ninth and one between tenth and eleventh rays, the former frequently absent; caudal translucent, with irregular series of round brown-red spots, the space between them often with bluish- white spots; the fin margined above with brownish-red; lower lobe whitish, unspotted; anal white, with a median sulphur-yellow streak and a terminal dark bar; ventrals whitish, with dusky areas, often uniform black; pectorals translucent; peritoneum silvery. 3. Distribution In the Atlantic philadelpbicus is found from Cape Henry, Virginia, in the north to Palm Beach, Florida, in the south. In the Gulf it is limited to the waters between Brownsville, Texas, and Cape Haze, Florida ( fig. 5 ) . Although it occurs frequently in deep water, philadelpbicus is very common in shallow water at times and has been known to enter harbors, bays, and shallow brackish lakes. That preference for hard bottom is not so obvious in this form is supported by the large numbers found in the Mississippi delta region. In general philadelphici/s appears to be a more temper- ate water form than ocyurus and seldom, if ever, is able to round the tip of Florida. The paucity of records from the west coast of Florida is perplexing and is probably due 62 Tulane Studies in Zoology Vol. 7 to a combination of environmental condi- tions rather than water temperature alone. 4. Population Analysis Comparison of Gulf and Atlantic popu- lations of philadelphiais shows divergence in two meristic characters ( Table 2 ) . In number of scale rows between the dorsal origin and lateral line, average divergence of populations is 80.6 per cent, a differ- ence considered by some (Ginsburg, 1938) to merit subspecific recognition. In the number of scale rows between anal origin and lateral line, there is an average diver- gence of 75.4 per cent which, according to standards mentioned above, is on the border- line between subspecies and race. Body and fin measurements (Tables 3- 10) show population differences in a num- ber of cases. Distance between the dorsal- fin origin and the caudal base ( Table 7 ) is slightly greater in Atlantic specimens, as is the distance between the dorsal origin and the occiput (Table 7). Dorsal-fin origin to lateral line ( Table 3 ) shows a fairly marked divergence, with measurements of Atlantic fish averaging higher. The same condition is found with regard to depth of caudal peduncle ( Table 4, and fig. 20 ) which is greater in most Atlantic specimens. The head ( Table 4 ) is slightly longer in Gulf fish; the orbit (Table 5) is also somewhat longer in Gulf fish, but the anal fin ( Table 8) tends to be shorter in smaller Gulf specimens. The color pattern given in the general description is typical of most specimens. Although many differences found are due to fading of the pattern, there are minor vari- ations in the intensity of pigmentation about the head and snout. In philadelphiaa, as in ocyurus, there are no pigmentary differences in preserved specimens which separate Gulf and Atlantic populations. C. philadelphicus has a much smaller range of variation in color pattern and apparently exhibits less sexual dimor- phism in color than do the other forms. Taking all characters into consideration, the two populations appear to be well- defined races that have not diverged to sub- species level. C. philadelphicus seems to be a more temperate-water form than ocyurus, for the southernmost record for it is on the Texas coast, where it is found offshore almost to the Mexican border. In the Atlantic it is apparently absent south of Palm Beach, Florida. Only one specimen ( TU 18158) was available from the Florida west coast, although the Oregon Reports ( Fish. Rept. No. 196) list the species from three stations in this area. These specimens were not available for examination. Judging by the large number of collections that have been made in the area, it seems improbable that this species is very common there. Why philadelphicus should be almost absent here and the other two species present in perplex- ing, for all three forms are known to prefer hard bottom and coral patches similar to those found along this coastline. I have been unable to locate specimens or verified records of philadelphicus from the Florida Keys, and I believe that the peninsula forms a barrier between Gulf and Atlantic populations. The resulting isola- tion and subsequent differentiation probably accounts for the minor, though consistent, differences in measurements and counts noted above. The contrast expressed by the uniformity of ocyurus populations and the divergence of philadelphicus populations is due to a difference in the relationships between each population pair. In the more heat-tolerant ocyurus the gene flow between populations is essentially unhindered, whereas in phila- delphicus it is almost, if not completely, prevented. The fact that populations of philadelphicus are isolated and populations of ocyurus are not could be responsible for the situation explained above. 5. Biology Spawning. — Gulf specimens collected be- tween November and March usually have mature gonads, though there must be some variation in the season at which gonads de- velop, since I found markedly undeveloped specimens in December, February, and March. I am unable to correlate this with staggering of spawning dates or distribution in the Gulf. Spawning probably occurs from late March to May or possibly June. Atlantic fishes have well developed gonads from February to June and possibly spawn at a later date than their Gulf counterparts. Spawning probably occurs from May to early No. 2 Miller: Review of Centropristes 63 Figure 19. Internal anatomy of C. philadelphicns. A. Air bladder. B. Stomach. C. Right lobe of testis. D. Left lobe of testis. E. Urinary bladder. F. Kidney. G. U. G. papilla. H. Testis. I. Fat. J. Caecum. K. Gut. L. Liver. M. Pericardial cavity. June and may be partially influenced by water temperatures. Young are common in shallow (2-20 fms. ) waters of the Gulf during the entire summer. This may indicate that spawning is not entirely restricted to deep waters ( 50- 100 fms.) and could explain the apparent difference in spawning times in the Gulf and Atlantic. If philadelphicus spawns in shallower water, the water temperatures there would vary more than in deep water, and an optimum temperature for spawning would probably occur earlier in the Gulf than in the Atlantic. Growth and Body Proportions. — Scatter- grams reveal three characteristics which can be contrasted with those of the other two species: Interorbital width (fig. 21) is smaller in philadelphicus than in both of the other species and does not greatly in- crease with size. Neither the increase in snout length (fig. 22) nor the decrease in orbit length (fig. 23) are as sharp as in other forms. Other scattergrams show that the caudal peduncle (fig. 20) is definitely deeper in Atlantic than in Gulf populations; the dorsal fin (fig. 25) is much longer in larger fishes, with the longest fins on males; the anal fin (fig. 26) is the shortest of the three forms in smaller fishes, but shows a marked increase in length in mature males; great elongation of one or two of the last anal rays is very conspicuous in spawning males (fig. 18); the pectoral (fig. 27) does not vary with increased size, but the pelvic (fig. 28) tends to be slightly shorter in larger fishes; and the caudal fin increased in length quite sharply in specimens over 90 mm. standard length. As in ocyurus, either the top or the middle caudal lobes may be the longest with most of the very longest being middle lobes. VIL Internal Anatomy Very few preserved specimens are useful in studying internal anatomy of these fishes because of damage caused by handling and cutting. In 16 specimens examined, how- ever, the internal organs seemed to be simi- lar in all three species. Fig. 19 is a drawing of the body cavity and viscera of a speci- men of philadelphicus.. Except for minor differences in shape and relative placement, internal organs of the three forms conform to the plan presented in the drawing. 64 Tulane Studies in Zoology Vol. 7 22 ~ 0 a 20 « o o \'S 18 . ° ' ' . r ' ° '° 16 o o o o ^ o°o « ° O ° Oo o o„ o°o*>o /" - **° °° .»»/ 14 °= • ^ oo .^ orf> ° 12 o o 1 ° Atlantic > ° Gulf 14 - 12 10 lilt 50 100 150 SL 200 12 10 8 12- 10- 8 12 10 8 o o ~ o 0 o o " eo o o' = o ° I o * » °o ^ ' 0 o o^ a) o o »„ o o o" o to * 50 100 SL 150 200 Figure 20. Scatter diagrrams for various bodv measurements. Percent of Standard r^. .^.^ c. i. i ^.i. • Lengi^h (ordinate) plotted against Standard f^^Yv Vi .^""''^ ^^1^-K '" ' *• ^^2/«'*"«, Length (abscissa), a. Dorsal origin to oc- ^- Phtladelphtcus, c. striatus. ciput in striatus. b. Caudal peduncle depth in philadelphiciis. o*^1i_»%°° o _ o« o» gco » - „ B aji " o°" .1 °*o "*» o o o Bo °° o""""" 50 100 SL 150 200 Figure 21. Interorbital width in: a. ocyii- riis, b. philadelpliicus, c. striatus. 12 10 8 12 10 (/t 6? 12 10 h 8 ! ^ O oS ?A . „o „o o o o o o o J 8* o o 50 100 SL 150 200 Figure 23. Orbit length in : a. ocyrirus, b. philadelphicxs, c. striatus. No. 2 Miller: Review of Centropristes 65 65 o o o 20 - , " "Vo g 18 - 16 'a o Q o o o O o ^ 18 - » no oo o * • ^° o V " ° ■° o 16 - 14 b 1 > 18 O »° 0 0° ° O o o 16 o Op • o", o 8 #5~ o o S 0 o °o o " oo 14 " C o 1 1 1 1 50 100 SL 150 200 Figure 24. Upper jaw length in: a. ocyu- 1-21S, h. s. melanus, c. striatus. 26 22 18 30 26 22 18 6R 26 22- 18 e • « - %o , . < 'I '" w • • B « - a « • • 0 • • "■ e • • • • • • • • « b 0 ° • • - • • • • • » . • " . . * • °. . » • ., . » * ' A . . • - oo Joo«^ ""'">, ^% o o«* «■ - -o\<.. •fee ^* ' c 1 ..,-i 1 1 1 ■ 50 100 SL 150 200 250 Figure 25. Dorsal fin length in : a. ocyu- rus, b. philadelphicus, c. striatus. Z2 18 26 22 18 tfi 26 22 18 - • : ■Gulf oAtlantic ' s. Striatus ' s. melanus 50 100 150 SL 200 250 Figure 26. Anal fin length in : a. ocyurus, h. philadelphicus, c. striatus. 30 28 26 24 30 28 26 24 6? o ' o° ° o'o, o - o o8o ° S o ° " ' 0°%^ if^ °§° - .„ \' f°o° °o\°"S „ ° o o o o o o o o o "a - o _ o o o ^ O o o o o ° oo o° ° ^-oo:\;;% o ; o ' o" o °„ ooo tf * o o'o O . » » o o^ o » "• o e o b s. striatus- ~ • •• • •• • • . . melanus-° • o .•» . . ..*.•• • .• -•o ■■■■:■': o m * <• .oo_^ o « 1 1 1 50 100 SL 150 200 Figure 27. Pectoral fin length in: a. ocyu- rus, b. philadelphicus, c. striatus. 66 Tulane Studies in Zoology Vol. 7 26 241- 22 o -so 24 _ ■> o O o • o 0 22 - Otf>0 O o Oo ©0 0 0° o / -oo* „ 0 o ° ° 0° o o oO 0 o o 20 . ° s o o^? » '"'■'■ Oo O * ,.0 0 o o b o o 28 - o o o o o 28 - o o o 0 o o o^ <^ o - o o o o o o 24 ~ » t I "o * ° » o 0 0 o 22 - ° <• o J ° o o o "«> 20 "c , r 1 1 50 100 SL 150 200 Figure 28. Pelvic fin length in: a. ocyu- riis, h. philadelphicus, and caudal fin length in: c. striatus. 54 4S 38 30 • 1 • • • 1 1 • top lobe ° middle lobe 1 50 100 SL 150 200 22 Figure 29. Caudal fin length in ocyurus. 62 54 46 38- 6S 30 22- o - • - o • - o - 3 o " • • - o - 3 - » o _ o - o 0 o <«> '^ - o o ° . ' o — "% • • - o o ° • • - o 0 • • °* 0 • ~ • • • • o • • top lobe • .V • • • • o middle lobe • 1 • 1 1 1 50 100 SL 150 200 Fi^re 30. Caudal fin length in philadel- phicus. VIII. Summary Three species of Centropristes {striatus, philadelphicus and ocyurus) are recognized. Gulf and Atlantic populations of these have been compared. The absence of infra- specific differentiation in ocyurus is prob- ably due to the lack of isolation. Certain constant, but minor, differences were found in philadelphicus which were judged to represent a low degree of divergence, and the populations were designated as races. The amount of overlap in all characters of melanus and striatus was judged to be more No. 2 Miller: Review of Centropristes Gl representative of the subspecies level of dif- ferentiation than the species level. They were reduced, therefore, to subspecies, with Gulf forms designated C. striatus melanus Ginsburg and Atlantic forms designated C striatus striatus ( Linnaeus ) . Food habits were found to be similar in all forms with fish and crustaceans forming the major staples in the diet. Spawning occurs from early to late spring in all species. IX. Acknowledgements For the loan of specimens I wish to thank Dr. James Bohike of the Academy of Nat- ural Sciences of Philadelphia, Drs. Charles M. Breder, Jr. and Vladimir Walters of the American Museum of Natural History, Drs. John C. Briggs and Daniel Cohen of the University of Florida, Dr. Eugenie Clark of the Cape Haze Marine Laboratory, Drs. Earl E. Deubler, Jr. and William E. Fahy of the Institute of Fisheries Research of the Uni- versity of North Carolina, Drs. Ernest A. Lachner and Leonard P. Schultz of the United States National Museum, Dr. Giles W. Mead and Mr. Harvey R. Bullis of the United States Fish and Wildlife Service, Dr. Edward C. Raney of Cornell University, Mr. Loren P. Woods of the Chicago Natural History Museum, and Dr. Henry B. Bigelow and Mrs. M. Dick of the Museum of Com- parative Zoology, Harvard University. This investigation was conducted at Tu- lane University as a partial requirement to- ward the degree of Master of Science. I am indebted to Dr. Royal D. Suttkus of the Department of Zoology, for suggesting this problem and for encouragement and assist- ance during the study, and to Dr. Edward C. Raney of Cornell University for helpful suggestions on the manuscript. I also thank my wife, Helen C. Miller, for her patient help and encouragement at all times. X. References Cited Bailey, Reeve M. 1951. The authorship of names proposed in Cuvier and Valenci- ennes' "Histoire NatiireUe des Poissons." Copeia, 1951 (3) : 249-251. Bean, Tarleton H. 1903. Catalogue of the fishes of New York. Bull. 60, N. Y. State Mils., Ann. Rept. N. Y. State Mus., 3: 1-784. Block, Marc Elieser and Johann Gott- LOB Schneider 1801. Syste^na Ichthijolo- giae iconibus ex illustratum. Post obitum anctoris opus inchoatum absolvit, correx- it, interpolavit Jo. Gottlob Schneider, Saxo. Berolini, Ix: 1-584, 110 col. pis. BoULENGER, George Albert 1895. Cata- logue of the Fishes in the British Mn- senm. Ed 2, London, 394 pp. 15 pis. Breder, Charles M., Jr. 1932. A record of Sarda velox and notes on other Block Island fishes. Copeia, 1932 (1) : 32. CuviER, Georges L.C.F.D. 1829a. Le Regne Animal. Ed. 2, 5 vols., Paris, Illust. Cuvier, Georges (in: Cuvier and Valienci- ennes) 1829b. Histoire Naturelle des Poissons. 22 vols., Paris, 1828 ?-49; 650 pis. DeKay, James E. 1842. New York Fauna. Part 1, Zoology; part 4, Fishes, Albany, 1-415, 102 pis.* Earll, R. Edward 1885. Blackfish eggs im- pregnated. Bull. U. S. Fish. Comm., 5: 91. Gill, Theodore 1861. Catalogue of the fishes of the eastern coast of North America, from Greenland to Georgia. Pi^oc. Acad. Nat. Sci. Phila., (suppl.) 1-63. 1873. Catalogue of the fishes of the east coast of North America. Rept. U. S. Fish Comm., 1871-72, 1: 779- 882. Ginsburg, Isaac 1938. Arithmetic defini- tion of the species, subspecies, and race concept, with a proposal for a modified nomenclature. Zoologica, 23 : 253-286. 1952. Eight new fishes from the Gulf coast of the United States, with two new genera, and notes on geo- graphic distribution. .Jour. Wash. Acad. Sci., 42(3) : 84-101, 9 figs. Goode, G. Brown 1884. The Fisheries and Fishery Industries of the United States. Washington, Section 1, part 3: 162-682, 218 pis. 1888. American Fishes. New York, 14: 1-496, illust. Hildebrand, Samuel F. and William C. Schroeder 1928. Fishes of Chesapeake Bay. Bull. U. S. Bur. of Fish., XLIII (Part 1) : 1-366, 211 figs. HOLBROOK, John Edwards 1860. Ichthy- ology of South Carolina. Ed. 2, Charles- ton, S. C, 205 pp., pis. HuBBS, Carl L. 1943. Criteria for subspe- cies, species, and genera, as determined by researches on fishes. Annals N. Y. Acad. Sci., XLIV (Art. 2) : 109-123. HuBBS, Carl L. and Karl F. Lagler 1947. Fishes of the Great Lakes Region. Cran- brook Inst. Sci., Bull. 26: 1-186, 26 pis., 251 figs. Jordan, David Starr 1885a. A catalogue of the fishes known to inhabit the waters of North America, north of the Tropic of Cancer. Ann. Rept. Comm. of Fish and Fisheries for 188Jt, Washington, 789-973. 68 Tulane Studies in Zoology Vol. 7 1885b. Supplementary notes on North American fishes. Proc. U. S. Nat. Mns., 7: 545-548. 1886. Notes on t., pical speci- mens of fishes described by Cuvier and Valenciennes and preserved in the Musee D'Histoire Naturelle in Paris. Proc. U. S. Nat. Mas., 9: 525-546. Jordan, David Starr and Barton Warren EvERMANN 1886. Description of six new species of fishes from the Gulf of Mexico, with notes on other species. Proc. U. S. Nat. Mns., 9: 466-476. .^ 1896. Fishes of North and Middle America. Bull. 47, U. S. Nat. Mus., 1896-1900, 4 parts, 3313 pp., 392 pis. Jordan, David Starr, Barton Warren EvERMANN, and Howard Walton Clark 1930. Check list of the fishes and fish- like vertebrates of North and Middle America north of the northern boundary of Venezuela and Colombia. Rept. U. S. Bur. Fish., 670 pp. Jordan, David Starr and Charles Henry Gilbert 1882a. Synopsis of the fishes of North America. Bull. 16, U. S. Nat. Mus., pp. Ivii, 1-1018. 1882b. Notes on fishes observed about Pensacola, Florida, and Galveston, Texas, with descriptions on new species. Proc. U. S. Nat. Mus., 5: 241-307. 1882c. Notes on a collec- tion of fishes from Charleston, South Carolina, with descriptions of three new species. Proc. U. S. Nat. Mus., 5: 580- 620. Lacepede, Bernhard Germaine Etienne 1803. Histoire Naturelle des Poissons. 14 vols., Paris, (1799-1804). LAVENDA, Nathan 1949. Sexual differences and normal protogynous heraiaphroditism in the Atlantic sea bass, Centropristes striatus. Copeia, 1949 (3) : 185-194. Linnaeus, Carl 1758. Systema Naturae. Ed. 10, 2 vols., Holmiae, pp. 285, 291. 1766. Systema Naturae. Ed. 12. pp. 485, 489. MiTCHiLL, Samuel Latham 1815. The fishes of New York, described and ar- ranged. Trans. Lit. and Phil. Soc. New York, 1: 355-492, pis. 1-6. Nichols, J. T. and Charles M. Breder, Jr. 1927. The marine fishes of New York and southern New England. Zoologica, 9(1) : 1-192. PvEiD, George K., Jr. 1954. An ecological study of the Gulf of Mexico fishes, in the vicinitv of Cedar Kev, Florida. Bull. Mar. Sci. Gulf and Carlh., 4(1) : 1-94, 13 figs., 8 tables. Smith, Hugh M. 1898. Fishes found in the vicinity of Woods Hole. Bull. U. S. Fish Comm., 17: 85-111. 1907. The Fishes of North Carolina. N. Car. Gcol. and Econ. Surv., Vol. 2, Raleigh, 11: 1-453, 21 pis., 187 figs. Storer, David Humphreys 1846. A Synop- sis of the Fishes of No7~th America. Met- calf and Co., Cambridge, Mass., 298 pp. Sumner, Francis B., Raymond C. Osborn, and L. J. Cole 1913. A catalogue of the marine fauna of Woods Hole and vicinity. Bull. U. S. Bur. Fish., (1911), 31 (part 2) : 549-794. Walbaum, Johann Julius 1792. Petri Ar- tedi renovati, pars i et ii (and iii-v) i.e. Bibliotheca et philosophia ichthyologica, cura lohannis Iiilii WCilhaumii edidit. 5 5 parts in 3 vols., Gr^peswaldiaee, 4 pis. Weed, Alfred C. 1937. Notes on sea basses of the genus Centropristes. Zool. Ser., Field Mus. Nat. Hist., Chicago, 20: (23) : 291-320, 2 figs. Abstract Gulf and Atlantic populations of the three species of Centropristes are com- pared. The absence of infraspecific differentiation in ocyurus is attributed to lack of isolation between Gulf and Atlantic populations. Certain constant differences, which were judged to rep- resent a low degree of divergence, are found in philadclphicus, and the popu- lations are designated as races, dehot- ing distinct but minor differences. Di- vergence of melanus and striatus was found to be greater than in the other forms, but overlap in all characters is large enough to dispute their specific distinctness. They are consequently re- duced to subspecies. Additional biologi- cal data may indicate their exact status. Food habits are similar in all forms, with fish and crustaceans forming the major staples in the diet. Spawning probably occurs from early to late spring in all species, with some possible variations between Gulf and Atlantic populations. TULANE STUDIES IN ZOOLOGY VOLUME 1, 1953-54 Number Price 1 On a new genus and species of mysid from Louisiana (Crustacea, Malacostraca), by Albert H. Banner, pp. 1-8 (June 1, 1953) . .1$0.25 2 A contribution on the life history of the lizard Scincella laterale (Say), by Richard M. Johnson, pp. 9-27 (July 3, 1953) 75 3 An outline for the study of a reptile life history, by Fred R. Cagle, pp. 29-52 (July 28, 1953) .75 4 A population of Ilolbrook's salamander, Eurycea longicauda guttolineata (IIol- brook), by Robert E. 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Simpson and Gordon Gunter, pp. 113-134 (December 31, 1956) .35 5 Dominancesubordinance relationships in the crawfish Cambarellus shufcldtii, by Mildred Eileen Lowe, pp. 137-170 (Decem- ber 31, 1956) 45 6 Propogatlon of the white shrimp, Penaeus aetiferus (Linn.) in captivity, by Malcolm C. Johnson and J. R. Fielding, pp. 173-190 (December 31, 1956) 30 Complete volume, including title page, ta- ble of contents and index (unbound) $2.50 VOLUME 5, 1957 1 Oatniusia heterochir, a new poeciliid fish from Texas, with an account of its hybrid- ization with O. affinis, by Clark Hubbs, pp. 1-16 (March 18, 1957) I$0.30 2 New calanoid copepods of Pontella Dana and Labidocera Lubbock with notes on the distribution of the genera in the Gulf of Mexico, by Abraham Fleminger, pp. 17-34 (March 18, 1957) 80 3 Three new crayfishes Crom Alabama and Mississippi (Decapoda: Astacidae), by Horton H. Hobbs, Jr. and Margaret Wal- ton, pp. 37-52 (March 18, 1957) 30 4 Heat death and associated weight loss of the oyster Crassostrea vtrginica, by Milton Fingerman and Laurence D. Fairbanks, pp. 53-62 (April 1, 1957) 30 5 The Odonata of Louisiana, by George H. Bick, pp. 69-135 (May 15, 1957) 1.00 6 Endocrine control of the red and white chromatophores of the dwarf crawfish, Cambarellus shufeldti, by Milton Finger- man, pp. 137-148 (June 7, 1957) 30 7 Hormones controlling the chromatophores of the dwarf crawfish, Cambarellus shu- feldti : their secretion, stability, and sepa- ration by filter paper electrophoresis, by Milton Fingerman and Mildred E. Lowe, pp. 149-171 (June 7, 1957) 40 8 Cyprinid fishes of the subgenus Cyprinella of Notropis. III. Variation and subspecies of Notropis venustus (Girard), by Robert H. Gibbs, Jr. pp. 173-203 (August 7, 1957) .50 9 The early development of Rana capita sevo- aa, by E. Peter Volpe, pp. 205-225 (Septem- ber 12, 1957) 35 Orconectes palineri (Faxon) ( Decapoda, Astacidae), by George Henry Penn, pp. 229-262 (September 12, 1957) .60 11 The skeleton shrimps (Crustacea: Caprel- lidae) of the Gulf of Mexico, by Joan E. Steinberg and Ellsworth C. Dougherty, pp. 265-288 (December 30, 1957) 40 12 The systematic status of the suckers of the genus Aloxostoma from Texas, New Mexico, and Mexico, by C. Richard Robins and Edward C. Raney, pp. 289-318 (Decem- ber 30, 1957) 45 Complete volume, including title page, ta- ble of contents and index (unbound) ....$5.00 VOLUME 6, 1958 1 The systematics and ecology of the Sterno- thaerus carinatua complex (Testudinata, Chelydridae), by Donald W. Tinkle, pp. 1-56 (March 31, 1958) $1.25 2 The butterflies of Mississippi, by Bryant Mather and Katharine Mather, pp. 61-109 (June 6, 1958) 1.00 3 Aquatic and semiaquatic Hemiptera of Mis- sissippi, by Clifton A. Wilson, pp. 113-170 (September 5, 1958) 1.00 4 The copepod genus Halicyclops in North America, with description of a new species from Lake Pontchartrain, Louisiana, and the Texas coast, by Mildred Stratton Wil- son, pp. 176-189 (December 31, 1958). Ontogeny of the first and second pleopods of the male crawfish Orconectes clypeatus (Hay) (Dedapoda, Astacidae), by Joe B. Black, pp. 190-203 (December 31, 1958) ... .60 Complete volume, including title page, table of contents and index (unbound) $3.50 VOLUME 7, 1959 1 An illustrated key to the crawfishes of Louisiana with a summary of their distri- bution within the State (Decapoda, Asta- cidae), by George Henry Penn, pp. 3-20 (April 23, 1959). Comparison of the chromatophorotropns of two crayfishes with special reference to electrophoretic behavior, by Milton Finger- man, pp. 21-30 (April 23, 1959) $0.60 Orders should be addressed to Meade Natural History Library, c/o Department of Zoology, Tulane University, New Orleans, 18, La., USA Please make remittance payable to "Tulane University" SEP -8 1959 HAHVARQ UHiVERSin Volume 7, Number 3 August 24, 1959 DIGENETIC TREMATODES OF MARINE FISHES FROM THE GULF OF PANAMA AND BIMINI, BRITISH WEST INDIES FRANKLIN SOGANDARES^BERNAL, FLORIDA STATE BOARD OF CONSERVATION MARINE LABORATORY, MARITIME BASE, BAYBORO HARBOR, ST. PETERSBURG, FLORIDA TULANE UNIVERSITY NEW ORLEANS TULANE STUDIES IN ZOOLOGY is devoted primarily to the zoology of the waters and adjacent land areas of the Gulf of Mexico and the Caribbean Sea. Each number is issued separately and contains an individual study. As volumes are completed, title pages and tables of contents are distributed to institutions exchanging the entire series. Manuscripts submitted for publication are evaluated by the editor and by an editorial com- mittee selected for each paper. Contributors need not be members of the Tulane Univer- sity faculty. Manuscripts should be submitted on good paper, as original typewritten copy, double- spaced, and carefully corrected. Two carbon copies in addition to the original will help expedite editing and assure more rapid publication. An abstract not exceeding three percent of the length of the original article must accom- pany each manuscript submitted. This will be transmitted to Biological Abstracts and any other abstracting journal specified by the writer. Separate numbers or volumes may be purchased by individuals, but subscriptions are not accepted. Authors may obtain copies for personal use at cost. Address all communications concerning manuscripts and editorial matters to the editor; communications concerning exchanges, and orders for individual numbers to the Librarian, Meade Natural History Library. When citing this series authors are requested to use the following abbreviations: Tulane Stud. Zool. Price for this number: $1.00. George Henry Penn, Editor Meade Natural History Library, Tulane University, New Orleans, U. S. A. Assistant to the Editor: Robert K. Chipman TULANE STUDIES IN ZOOLOGY Volume 7, Number 3 SEP- 8 195! — UNIVERSITY August ^24, 19^1 9 DIGENETIC TREMATODES OF MARINE FISHES FROM THE GULF OF PANAMA AND BIMINI. BRITISH WEST INDIES ^ FRANKLIN SOGANDARES-BERNAL, Florida State Board of Conservation Marine Laboratory, Maritime Base. Bayboro Harbor, St. Petersburg. Florida CONTENTS 1. Introduction 71 II. The Digenea of the Gulf of Panama and Bimini, B.W.I 71 A. Family Aspidogastridae 71 B. Family Bucephalidae 7 1 C Family Paramphistomatidae 74 D. Family Pronocephalidae 74 E. Family Lepocreadiidae 74 F. Family Megaperidae 87 G. Family Acanthocolpidae 87 H. Family Waretrematidae 92 I. Family Haplosplanchnidae 93 J. Family Opecoelidae 95 K. Family Cryptogonimidae 98 L. Family Fellodistomatidae 101 M. Family Monorchidae 102 N. Family Zoogonidae 102 O. Family Bivesiculidae 102 P. Family Gorgoderidae 102 Q. Family Hemiuridae 103 R. Family Prosogonotrematidae 106 S. Family Accacoelidae 106 III. Measurements of pH of the Stomach and Intestine of Fish Hosts 107 IV. Exact Locations of Digenea in Their Hosts 108 V. Geographical Distribution 108 VI. Summary 114 VII. Acknowledgements 115 VIII. References Cited ■- 116 Abstract 117 1 Studies from the Department of Zoology, University of Nebraska, No. 315. EDITORIAL COMMITTEE FOR THIS NUMBER Raymond M. Cable, Professor of Parasitology, Purdue University, Lafayette. Indiana Robert F. Hutton, Biologist-in-Charge, Florida State Board of Conservation Marine Laboratory, Maritime Base, Bayboro Harbor, St. Petersburg, Florida H. W. Manter, Professor of Zoology, University of Nebraska, Lincoln, Nebraska No. 3 Sogandares-Bernd: Ttrematodes of Marine Fishes 71 I. Introduction This paper is based on collections of rrematodes made during the summer of 1956 in the Gulf of Panama and the summer of 1957 at Bimini, British West Indies. The trematode faunas of both these regions are little known. Manter (1940a) reported a few species from Panama, a region later studied by Caballero and colleagues (Cabal- lero, et al, 1952, 1953, 1956). The nearest well studied region to Bimini is the Tor- tugas, Florida region (Linton, 1910; Manter, 1947). A total or 484 specimens representing 208 species of fishes were examined. Eighty- eight species of Digenea were collected and studied. These included two new genera, 17 new species, and 29 new host records. Ob- servations were made on the exact location of the trematodes in the digestive tract of the host. II. The Digenea of the Gulf of Panama and Bimini, B.W.I. The following account includes almost all of the species of Digenea collected. A few were too immature or too macerated to identify with certainty. A few agreed well with some early descriptions which, how- ever, probably represented more than one species. These specimens will need to be compared with holotypes of the species in question. The exact location of a trematode along the intestine is indicated as from the pyloric junction; for example, "anterior 1/3 intes- tine" means within the distance from the pyloric valve to 1/3 the distance to the anus. All measurements are in millimeters un- less otherwise stated. Holotypes of new species have been de- posited in the United States National Mu- seum Helminthological Collection. All figures, except diagrams, were made with the aid of a camera lucida. Each pro- jected scale has the value indicated in milli- meters. A. Family ASPIDO CAST RID AE Poche, 1907 Lobatostoma ringens (Linton, 1907) Eckmann, 1932 Host. — Calamus bajonado (Bloch & Schneider), jolt-head porgy. Location. — Rectum. Locality — W. side (opposite Lerner Lab- oratory), N. Bimini, B.W.T. Discussion. — La Rue (1957) considered the non-reproduction by larvae of aspido- gastrids sufficient to separate this group from the Digenea. Although Le Rue stated (p. 310) that the Aspidogastrea "belong to a separate class", he apparently intended to follow Faust and Tang (1936) and others in separating them from the subclass Di- genea. They are clearly members of the class Trematoda. Although Aspidogaster conchicola has a direct life cycle, (Williams, \9A1) and its larvae do not reproduce, there is a possibility that this freshwater species maturing in bi- valves may be atypical of the group. Adult aspidogastrids have so many morphological similarities to certain Digenea that they are here retained as a family of Digenea. B. Family BUCEPHALIDAE Poche, 1907 Bucephalus varicus Manter, 1940 Hosts. — Palometa media (Peters), palo- meta {new host record}; Caranx crysos ( Mitchill ) , hard tailed jack [new host rec- ord]; Caranx hippos (Linn.), common jack; Caranx latus Ag^ssiz, horse-eyed jack; Car- anx ruber ( Bloch ) , skip jack. Location. — Anterior 1/4 intestine of P. media: pyloric ceca and length of entire in- testine of C. latus; mid-intestine of C. ruber. Locality. — P. media from Taboga Island, Panama Pacific. The other hosts from Bim- ini, B.W.I. Discussion. — B. varicus is a common para- site of carangids in the region of the Tropi- cal American Pacific, Tortugas, Florida; Bimini, B.W.L; the Red Sea, and the Oki- nawan area. Manter (1955) has already noted faunal similarities of trematodes of these areas. Prosorhynchus gonoderus Manter, 1940 (Figure 1) Host. — Epinephelus analogies Gill, ca- brilla pinta [new host record}. Location. — Intestine. Locality.. — Taboga Island, Panama Pacific [new locality record}. Discussion. — P. gonoderus was named and described from a specimen from a "yellow- 72 Tulane Studies in Zoology Vol. 7 spotted grouper" from the Galapagos Islands by Manter (1940a). Manter found P. go- noderus associated with P. ozakii Manter, 1934. I have examined paratype specimens of P. gonoderus and have found one speci- men which agrees almost exactly with my specimen from Ephinephelns analogus. Prosorhynchus ozakii Manter, 1934 Host. — Epinephelus analyogus Gill, ca- brilla pinta [new host record}. Location. — Intestine. Locality. — Taboga Island, Panama Pacific [new locality record}. Ddisci/ssion. — P. ozakii was originally de- scribed from Epinephelus niveatus (Cuv. & Val.) at Tortugas, Florida by Manter (1934). Later, Manter (1940a) reported the same species from a "yellow-spotted grouper" in the Galapagos Islands. Prosorhynchus pacificus Manter, 1940 Host. — Mycteroperca venenosa venenosa (Linn.). Location. — Pyloric ceca. Locality. — Bimini, B.W.I, [new locality record}. Discussion. — Mycteroperca v.. venenosa has 13 pyloric ceca. One cecum, which had a surface area of approximately 320 square millimeters, contained 6 P. pacificus and 3 immature lepocreadiids. Some ceca had no trematodes. If trematodes were found, there were more than one per cecum. P. pacificus has been reported from Myc- teroperca olfax and M. xenarcha in the Gala- pagos Islands by Manter (1940a) and from M. venenosa and M. microlepis in Tortugas by Manter (1940); and from Sebastopyr ruherrivius in Bermuda by Hanson (1950). This record from Bimini, extends the South- eastern range of P. pacificus in the Atlantic. Hanson (1950) considered P. atlanticus Manter, 1940 a synonym of P. pacificus Manter, 1940. Dollfustrema muraenae, sp. nov.- (Figures 2-5) Host. — Gymnothorax vicinus (Castelnau), brown moray. Location. — Proximal to pyloric junction in one host and in 3/4 intestine of another. Locality. — N. Bimini, B.W.I. - The name muraenae indicates the host which is in the family Muraenidae. Holotype.—\].S:NM. Helm. Coll. No. 38860. Diagnosis (based on 2 specimens; 4 speci- mens lost). — Body completely spined, elon- gate, rounded at anterior end, tapering to a point posteriorly, 1.273 to 1.35 long by 0.399 to 0.47 wide. Rhynchus terminal, well developed, elliptical in shape, with 3 equa- torial rows of spines. Spines difficult to measure due to their position. Dorsal spines in linear arrangement, ( fig. 5 ) approxi- mately equal in length, from 12 to 24 mi- crons long, (fig. 3A). Ventral spines, alter- nately arranged, (figs 3, 4), unequal in length; anterior spines shortest, from 10 to 11 microns long by 5 microns at their base; middle usually longest, 20 to 22 microns long by 3 microns wide at their base and tapering only very gradually to a point; bottom row of spines approximately 3/4 length and same diameter of spines in mid- dle row. Mouth postequatorial, level with anterior tip of cirrus sac. Pharynx imme- diately following mouth, globular, 0.101 to 0.114 long by 0.121 to 0.126 wide. Cecum club-shaped, extending from pharynx ap- proximately 2/3 distance to posterior edge of rhynchus. Gonads immediately post- equatorial, side by side in a transverse row. Testes spherical, on each side of body, separated by cecum and ovary in one speci- men; ovary to right of sinistral testis and cecum, and intertesticular in another speci- men. Cirrus sac dextral, in posterior 1/3 body; 0.295 to 0.37 long; internal seminal vesicle with one ascending one one descend- ing loop in anterior half of sac; pars pro- statica in posterior half of sac; genital lobe single in a genital atrium which is approxi- mately 1 10 length of cirrus sac; genital pore ventral 0.075 to 0.167 from posterior end of body. Vitellaria foUiailar, beginning at anterior level of cecum on each side of body, confluent at mid-body immediately posterior to edge of rhynchus. Ovary inter- testiailar or dextral to right testis, but al- ways at level of testes. Uterus descending posteriorly beyond ovary for a short distance, ascending to fill area between rhynchus and anterior edge of ceamn, descending again to end in genital atrium. Eggs 24 to 33 by 19 to 27 microns. Excretory vesicle not ob- served. Discussion. — Five species of Dollfustrema Eckmann, 1934 (syn. Dollfusina Eckmann, No. 3 Sogandares-Bernal: Ttrematodes of Marine Fishes 73 Figrures 1-10. 1. Prosorhynchus gonoderus Manter, ventral view. 2. DoUfiistreama mu- raenae, sp. nov., ventral view. 3,4. D. tmiraenae, portion or rhynchus showing' spines, ventral view. 5. D. muraenae, portion of rhynchus showing spines, dorsal view. 6. Pseu- docreadium biminensis, sp. nov., ventral view. 7,8. Pseudocreadimn scaphosonuim Man- ter, schematic diagrams showing position of ovary between testes. 9. Pseudocreadium lameUiforme (Linton), ventral view. 10. Lepidapedon triincatum, sp. nov., ventral view. 74 Tulane Studies in Zoology Vol. 7 1932) have been named. These species are: D. californiae Montgomery, 1957; D. echi- natuni (Komiya & Toyami, 1941) Yama- guti, 1953 (described from its metacercaria); D. gravidnm Manter, 1940; D. macracan- thum Hanson, 1950; and D. laneyi (Tseng & Shen, 1930) Eckmann, 1934 (type species ) . In D. californicum. D. gravidum. D. ma- cracantbum, and D. vaneyi, the testes are tandem, whereas in D. echinatum and D. viuraenae, the testes are side by side. D. muraenae differs from D. echinatum by having a postequatorial mouth, pharynx op- posite anterior edge of cirrus sac, and secum extending only two-thirds distance from pharynx to rhynchus as compared with a preequatorial mouth, pharynx distant from anterior edge of cirrus sac, and cecum almost in contact with rhynchus. All of the species of DoUfustrema re- ported from the coasts of the American Continent have been from Gymnothorax. One record (D. vaneyi) from the China coast has been from a fish other than eel. The other record ( D. echinatum ) is of a metacercaria in the muscle of a fish. Bucephaloides arcuatus ( Linton, 1900 ) Hopkins, 1954 Host. — Sphyraena barracuda (Linn.), bar- racuda. Location. — Pyloric ceca of 2 hosts and rectum of 1 host ( the one with only one trematode ) . Locality. — N. shore of S. Bimini [new locality record}. Discussion. — Manter (1940b) discussed the systematic status of this species in detail. The hosts examined in this study were caught over sandy bottom in water no deeper than one fathom. Host stomachs and intestines were either empty or contained only fish remains. B. arcuatus has been re- ported from Woods Hole, Mass., Beaufort, N.C., and Tortugas, Fla. Sparks ( 1957) re- ported trematodes from S. barracuda col- lected in the vicinity of Nassau but did not list B. arcuatus. C. Family PARAMPHISTOMATIDAE Fischroeder, 1901 Cleptodiscus reticulatus Linton, 1910 Host. — Pomacanthus aureus (Bloch), black angelfish. ■ Location. — Immature specimens in pyl- oric ceca and mature specimens in rectum. Locality.— -Lerner Fish Pens and N. shore, N. Bimini, B.W.I, [new locality record}. Cleptodiscus kyphosi Sogandares, 1959 Host. — Kyphosus sectatrix (Linn.), ber- muda club. Location. — Mid intestine. Locality. — Bimini, B.W.I. D. Family PRONOCEPHALIDAE Looss, 1902 Barisomum erubescens Linton, 1910 Host. — Pomacanthus aureus ( Bloch ) , black angelfish. Location. — Rectum. Locality.— N. shore, N. Bimini, B.W.I. [new locality record}. Disciission. — Linton (1910) and Manter (1947) reported B. erubescens from differ- ent fishes, all in the family Holacanthidae, from Tortugas, Florida. In addition, Linton reported specimens from a parrot fish, per- haps an accidental infection. E. Family LEPOCREADIIDAE Nicoll. 1933 Lepocreadium trulla (Linton, 1907) Linton, 1910) Hosts. — Ocyurus chrysurus (Bloch), yel- lowtail; Lutjanus buccanella (Cuv. & Val. ) , black-finned snapper [new host record}. Location. — Pyloric ceca and 14 intestine of O. chrysuriis and 1/3 intestine of L. buc- canella. Locality. — 0. chrysurus from Lerner Fish Pens, Bimini and near Cat Cay, B.W.I. ; and L. buccanella from 1 mi. W. of N. Bimini (20 fathoms), B.W.I. Discussion. — L. trulla has been reported from O. chrysurus in Bermuda by Linton (1907) and from the same host by Linton (1910) and Manter (1947) in Tortugas, Florida. Manter noted that Linton's ( 1910) record of one specimen of L. trulla in Cala- vius calamus in Tortugas, probably repre- sents an accidental infection. The speci- men of L. trulla foimd in L. buccanella from Bimini is also believed to be accidental in this host. Pseudocreadium biminensis, sp. nov.^ (Figure 6) Host. — Balistes capriscus Gmelin, ocean tally. •"' The name biiiiincvfiis indicates the type locality. No. 3 Sogandares-Bernal: Ttrematodes of Marine Fishes 75 Location. — 1/3 intestine in one host and 1/3 intestine (mature specimen) and 2/3 intestine (immature specimen). Locality. — W. side of N. Bimini and near Cat Cay, B.W.I. Holotype.—\].S.N.U. Helm. Coll. No. 38861. Diagnosis (based on 2 mature speci- mens).— Body elongate elliptical, spined; 1.577 to 2.294 long by 1.007 to 1.596 wide. Forebody 0.779 to 1.003 long. Posterior body 0.608 to 1.007 long. Oral sucker sim- ple, subterminal, 0.208 to 0.281 long by 0.241 to 0.348 wide. Acetabulum equa- torial, 0.208 to 0.255 long by 0.174 to 0.214 wide. Sucker ratio 1:0.51 to 0.89. Pre- pharynx as wide as, and half length of, pharynx which measures 0.161 long by 0.181 to 0.255 wide. Esophagus approxi- mately half length of pharynx. Cecal bifur- cation in anterior 1/3 body. Ceca arch laterally on each side of body, completely encircling testes, ending a short distance from posterior end of body. Genital pore sinistral, almost reaching inner edge of left cecum midway between acetabulum and anterior end of body. Gonads postequatorial. Testes round to irregular, side by side, in the posterior 1/3 body; right testis 0.348 to 0.779 long by 0.295 wide; left testis 0.235 to 0.295 long by 0.201 to 0.318 wide. Cir- rus sac extending from genital pore to slightly posterior to acetabulum, entirely to left of acetabulum, cirrus in anterior third of sac, followed by narrow coiled prostatic vesicle which tubular anteriorly and almost bulbular posteriorly; internal seminal vesicle spherical, in posterior 1/^6 sac. External seminal vesicle voluminous, extending from cirrus sac transversally between ovary and acetabulum. Ovary indistinctly tri-lobed, median, in contact with right testis, 0.134 to 0.181 long by 0.147 to 0.181 wide. Seminal receptacle club-shapde, between left testis and posterior tip of cirrus sac in one specimen and transverse, near midbody, im- mediately posterior to external seminal ves- icle in another. Vitellaria extending from level of posterior edge of oral sucker on each side of body, confluent medianly at level of cecal bifurcation and genital pore, separating anterior to acetabulum and over- lapping ceca on each side of body for their posterior extent to terminate a short dis- tance from posterior end of body, confluent posterior to testes. Uterus anterior to ovary, entering muscular metraterm to left of and slightly longer than cirrus sac. Eggs 64 to 72 by 32 to 36 microns. Excretory pore dorsal, median, between inner aspect of tips of ceca and testes. Excretory vesicle partly visible in immature specimen; tubular, ex- tending to mid-testicular level where two cornua branch off, one on each side of body; giving vesicle a Y-shaped appearance; they are not visible anterior to seminal receptacle. Discussion. — Layman (1930) named the genus Pseudocreadium for Pseudocreadium fuonacanthi. Ozaki ( 1936) named the genera Leptocreadium and Hypocreadium.. Lepto- creadium Ozaki, 1936 proparte and Hypo- creadium Ozaki, 1936 were considered syn- onyms of Pseudocreadium Layman, 1930 by Manter (1940a, 1945). Yamaguti (1953) considered Hypocreadium distinct from Pseudocreadium on the basis that the ovary is between the testes in the former genus and in advance of the testes in the latter genus. Bravo and Manter (1957) accepted Yamaguti's ( 1953) two genera on the basis of intertesticular ovarian position and posterior extent of the uterus. Exami- nation of paratypes of Panamanian Pseudo- creadium scaphosomum Manter, 1940, re- veals variation in intertesticular ovarian position (figs. 7, 8). Furthermore, if a series of the species P. scaphosomum Man- ter, 1940, P. spinosum Manter, 1940, P. ga/apagoense Manter, 1945 and P. mona- canthi Layman, 1930 are studied, they will clearly show intergradation from inter- testicular to pretesticular ovarian position. Extent of uterus posterior to ovary is also not constant. A study of a large series of P. scaphosomum showed the uterus extend- ing from mid-ovary to posterior to ovary, to posterior level of testes and slightly beyond. This observed variation did not appear to be correlated with size of individual speci- mens. For this reason, the genus Hypo- creadium is considered a synonym of Ps eudo creadiu m . Manter ( 1945 ) transferred four species of Pseudocreadium. to Lepocreadium. These were: P. balistes Nagaty, 1942, P. elonga- tum Nagaty, 1942, P. sohali Nagaty, 1942, and P. vitellosum (Ozaki, 1936) Manter, 1940 (syn: Leptocreadium vitellosum Oza- ki, 1936). Other species in the genus Pseudocreadium are: P. anandrum Manter, 76 Tulane Studies in Zoology Vol. 7 1947 (syn: H. anandrum (Manter, 1947) Yamagiiti, 1953); P. dampieriae ( Yama- guti, 1942) n. comb., (syn: H. dampieriae Yamaguti, 1942); P. galapagoensis Manter, 1945; P. lamellifor?ne (Linton, 1907) Man- ter, 1945, (syn: Dislofnu??i lartudliforme Linton, 1907 pro parte); P. monacanthi Layman, 1930, (syn: Leptocreadium skarja- bini Ozaki, 1936); P, ovale Yamaguti, 1942; P. patellare (Yamaguti, 1938) Man- ter, 1940, (syn: H. patellare Yamaguti, 1942); P. scaphosomum Manter, 1940; (syn: H. scaphosomum (Manter, 1940) Yamaguti, 1942; H. myohelicatu?n Bravo & Manter, 1947, new synonymy); P. spi- nosuyn Manter, 1940, (syn: H. spinosum (Manter, 1940) Yamaguti, 1942); P. sym- metrorchis (Ozaki, 1936) Manter, 1940, (syn: H. symmetrorchis Ozaki, 1936). Pseudocreadium himinensis is most close- ly related to P. galapagoensis. Both differ from all other species of Pseudocreadium by having the metraterm and cirrus sac on the left side of the acetabulum. P. himi- nensis differs from P. galapagoensis by hav- ing vitellaria which extend only to posterior edge of oral sucker as compared with vit- ellaria extending anteriorly as far as an- terior border of oral sucker; sucker ratio of from 1:0.51 to 0.89 as compared with sucker ratio 1:1 to 1.13; transverse, median external seminal vesicle as compared with a lateral diagonal external seminal vesicle; cirrus sac extending posteriorly beyond acetabulum as compared with cirrus sac extending to mid-acetabulum or slightly beyond, median position of acetabulum as compared with acetabulum more anterior to equator, and locality Atlantic rather than Pacific Ocean. Pseudocreadium scaphosomum Manter, 1940 Host. — Balistes naufragium Jordan & Starks, cochino [new host records}; Balistes verres Gilbert & Starks, pez puerco. Location. — Intestine, near pyloric junc- tion. Locality. — Taboga Island, Panama Pacific. Discussion. — Caballero et al (1953) re- ported P. scaphosomum from Balistes poly- lepis (type host) in Panama. Bravo and Manter (1957) named Hypocreadhnn myo- helicatum from the intestine of Balistes capistratus (=zBalistes verres). They dif- ferentiated H. myohelicatum from H. scap- hosomum (=zPseudocreadium scaphosomum) on the basis of vitellaria overlapping the ceca in the former but not the latter species. I examined the paratype series of P. scap- hosomum and found that at least a few vitellaria always overlap the ceca. H. myo- helicatuni is here considered a synonym of Pseudocreadium scaphoso ynu m . Pseudocreadium lamelliforme ( Linton, 1907) Manter, 1945 Host. — Balistes vetula Linn., queen trig- gerfish [new host record]. Location. — 1 4 intestine and 1 2 intes- tine. Locality. — Lerner fish pens, and N. shore, N. Bimini, B.W.L Dermadena lactophrysi Manter, 1945 Synonym. — Distomum lamelliforme (Lin- ton, 1907) pro parte. Host. — Lactophrys trigonus ( Linn. ) ; Lac- tophrys tricornis (Linn.), trunkfish. Location. — Mid-intestine of L. trigonus and 3/4 intestine of L. tricornis^ Locality. — L. trigonus from Lerner fish pens and L. tricornis from off Lerner Lab- oratory pier, Bimini, B.W.I. Lepidapedon truncatum, sp. nov.^ (Figure 10) Host. — Holocentrus ascencionis (Osbeck), squirrelfish. Location. — Pyloric ceca ( immature speci- mens) and 1/3 intestine (mature speci- mens ) . Locality. — Lerner fish pens, Bimini, and from near Cat Cay, B.W.L Holotype.—U.SN.U. Helm. Coll. No. 38862. Diagnosis (based on 3 mature speci- mens ) . — Body elongate, sides almost paral- lel, widest in testicular area, tapering an- teriorly and truncate posteriorly; cuticle spined; 1.976 to 2.261 long by 0.475 to 0.513 wide at testicular level. Forebody 0.551 to 0.646 long. Posterior body 1.330 to 1.501 long. Oral sucker subterminal; 0.107 to 0.121 long by 0.101 to 0.121 wide. Acetabulum 0.094 long by 0.074 to 0.094 wide. Sucker ratio 1:0.73 to 0.77. Pre- pharynx very short, almost absent. Pharynx ^ The ni'nie ti-nncatitin is for the charac- teristic truncate appearance of the posterior end of body. No. 3 Sogandares-Bernal: Tirematodes of Marine Fishes 77 ovoid, 0.094 to 0.101 long by 0.074 to 0.081 wide. Esophagus from equal to, to IY2 times length of, pharynx. Cecal bi- furcation approximately in middle of fore- body; ceca extending to posterior end of body, ending blindly. Genital pore sinistral to anterior border of acetabulum. Gonads postequatorial. Testes diagonal, in posterior l/'3 body, oval and smooth; anterior testis to left, posterior to right of midline; an- terior testis 0.194 to 0.215 long by 0.134 wide; posterior testis 0.201 to 0.248 long by 0.134 wide. Cirrus sac extends from genital pore to approximately 1/3 distance from acetabulum to ovary; cirrus and pro- static vesicle occupy anterior 5 6 sac; an- terior seminal vesicle in posterior 1/6 sac. Posterior seminal vesicle approximately as long as cirrus sac with one to three slight constrictions; surrounded by prostatic cells which are in turn surrounded by membrane originating from cirrus sac. Cirrus sac and posterior seminal vesicle extending approxi- mately 3 4 distance from acetabulum to ovary. Trilobed ovary median, separated from or in contact with anterior testis, 0.101 to 0.134 long by 0.134 wide. Club-shaped seminal receptacle to left of ovary and in contact with anterior testis. Vitellaria ex- tending from anterior border of acetabulum posteriorly, overlapping ceca, to end slightly beyond tips of ceca. Uterus anterior to ovary; entering muscular metraterm which may be as long as 1/2 length of cirrus. Un- collapsed eggs 67 to 72 microns long by 48 microns wide, (collapsed eggs 53 to 68 by 24 to 40 microns ) . Excretory pore pos- terior and terminal. Excretory vesicle tubu- lar, dorsal, beginning at excretory pore where it is constricted by a bulbulomuscular sphincter, ending at level of cecal bifurca- tion. Discussion. — There are six species of Lepidapedon that possess an excretory vesi- cle extending to the cecal bifurcation; these are: L. congeri Manter, 1954; L. epinepheli Bravo & Manter, 1957; L. hancocki Manter, 1940; L. levenseni (Linton, 1907) Manter, 1947; L. nicolli Manter, 1934; and L. tra- chinoti Hanson, 1950. L. truncatum differs \rovi L. epinepheli, L. levenseni and L. tra- chinoti by possessing vitellaria which ex- tend to the acetabulum as compared with vitellaria never extending to the acetabulum. L. truncatum further differs from L. tra- chinoti by lacking an accessory lateral sucker, possessing a trilobed ovary and testes diagonal as compared with tandem testes. L. truncatufu also differs from L. epinepheli and L. levenseni by possessing diagonal in- stead of tandem testes and trilobed instead of smooth ovary. L. truncatum differs from L. congeri by having vitellaria which are not confluent preacetabularly, genital pore sinistral as compared with median or only slightly submedian, and posterior seminal vesicle far posterior to acetabulum as com- pared with posterior seminal vesicle dorsal and extending only slightly posterior to acetabulum. L. truncatum is most closely related to L. nicolli and L. hancocki, differ- ing from these species by having separated diagonal testes as compared with contiguous testes in tandem, distinctly trilobed ovary as compared with smooth oval ovary, and trun- cate posterior end of body as compared with more pointed posterior end. One abnormal mature specimen (not de- scribed ) possessed only one testis which was median in posterior 13 of body. This specimen agreed in all other respect with L. truncatum. Lepidapedon parepinepheli, sp. nov.^ ( Figure 1 1 ) Host. — Epinephelus tigris (Cuv. & Val.), tiger grouper. Location. — 2 in pyloric ceca and 9 in 1/4 intestine. Locality. — 1 mile W. of N. Bimini, B.W.I.; in 18 fathoms. Ho/o/j/?^.— U.S.N.M. Helm. Coll. No. 38863. Diagnosis (based on 11 specimens). — Body elongate, sides of body almost parallel, tapering to a rounded end anteriorly and posteriorly; cuticle spined; 3.857 to 5.263 long by 0.380 to 0.627 wide. Forebody 0.950 to 0.969 long. Posterior body 2.774 to 4.18 long. Oral sucker terminal; 0.047 to 0.087 long by 0.101 to 0.121 wide. Acetabulum approximately in anterior 1/4 body; 0.114 to 0.134 long by 0.114 to 0.148 wide. Sucker ratio 1:1.11 to 1.29. Pre- pharynx from 14 to equal in pharynx length. Pharynx 0.067 to 0.081 long by ■"' The name pare pin eplxeli indicates the close relationship of this species with Lepi- dapedon epinepheli; para (= alongside) , epinepheli {^Lepidapedon epinepheli). 78 Tulane Studies in Zoology Vol. 7 0.054 to 0.067 wide. Esophagus from 2 to ending blindly. Genital pore sinistral to 4 times as long as pharynx. Cecal bifurca- anterior border of middle of acetabulum tion approximately in mid-forebody, ceca Gonads postequatorial. Testes tandem in extending to near posterior end of body, posterior 14 body, oval and smooth; sepa- • o" ^^. »s> 0m '^W- 0&S* ••« ••5/ ii* 11 \i^ Figures 11-24. 11. Lcpidapcdon ])arepincpheli, sp. nov., ventral view. 12. Diploproc- toduenm plicitnm (Linton), ventral view. 13-15. D. plicitnni, ratio of metraterm length to cirrus sac length. 16-18. D. plicitnm, vaiiation of anterior extent of vitellaria. 19-21. D. j)licifu)n, variation in shape of anterior end of body. 22. Enoitcnoii aiirenm Linton, ventral view. 23. E. uureain, cirrus sac. 24. E. aaretou, oral sucker. No. 3 Sogandares-Bernal: Ttrematodes of Marine Fishes 79 rated from each other by a distince from 1 to IVi times length of anterior testis; an- terior "testis 0.168 to 0.268 long by 0.147 to 0.023 wide; posterior testis 0.181 to 0.318 long by 0.168 to 0.268 wide. Cirrus sac extending approximately halfway from genital pore to ovary; 0.57 to 0.804 long; cirrus and prostatic vesicle occupying an- terior 6 7 sac; anterior seminal vesicle in posterior 1 7 sac. Convoluted posterior seminal vesicle approximately 1'2 length of cirrus sac, 0.268 to 0.482 long, sur- rounded by prostatic cells which are in turn surrounded by a membrane originating from rear portion of cirrus sac. Cirrus sac and posterior seminal vesicle extending approximately 2/3 distance from acetabul- um to ovary. Median, smooth, oval ovary approximately halfway between anterior testis to posterior tip of posterior seminal vesicle; 0.134 to 0.188 long to 0.127 to 0.168 -wide. Club-shaped seminal receptacle to left of ovary, tapering to the Mehlis, gland complex which is median and imme- diately anterior to ovary. Laurer's canal ex- tending from ootype laterally on the left side of body level with anterior border of ovary to open dorsally between left cecum and edge of body. Vitellaria extending from anterior to posterior edge of cirrus sac or posterior tip of posterior seminal vesicle towards posterior end of body over- lapping ceca along posterior extent, con- fluent between testes, filling posttesticular space to end near posterior end of body. Uterus anterior to ovary entering muscular metraterm which is half length of cirrus sac. Eggs from 56 to 69 by 24 to 32 microns. Excretory pore posterior and terminal. Ex- cretory vesicle tubular, dorsal, extending to near cecal bifurcation. Discussion. — There are seven species of Lepidapedon which possess an excretory vesicle extending to the cecal bifurcation, these are: L. congeri Manter, 1954; L. epinepheli Bravo & Manter, 1957; L. han- cocki Manter, 1940; L. levenseni (Linton, 1907) Manter, 1947; L. nicolli Manter, 1934; L. trachinoti Hanson, 1950 and L. truncatum (this paper). L. par epinepheli differs from L, congeri, L. hancocki, L. ni- colli and L. trtmcatuni in possessing vitel- laria which never extend to acetabulum as compared with vitellaria extending to acetabulum or beyond. L, parepinepheli dif- fers from L. trachinoti by lacking an ac- cessory lateral sucker as compared with lateral sucker present. L. parepinepheli is most closely related to L. levenseni and L. epinepheli, differing from L. levenseni in sucker ratio, 1:1.11 to 1.29 as compared with 1:0.75, posterior edge of external seminal vesicle lying 2/3 distance from ace- tabulum to ovary as compared with approxi- mately 1/3 distance from acetabulum to ovary, seminal receptacle sinistral to ovary as compared with between ovary and an- terior testis, and vitellaria confluent between the testes. L. parepinepheli is very closely related to L. epinepheli. I have examined the paratypes of L, epinepheli and except for two details — my specimens agree in almost all characters. The posterior seminal vesicle of L. parepinepheli is approximately half length of cirrus sac and never longer as compared with a length almost always twice as long as cirrus sac in L. epinepheli. In addition, the posterior tip of the posterior seminal vesicle lies 2/3 the distance from acetabulum to ovary in L. parepinepheli while it is never more than 1/2 way from acetabulum to ovary in L. epinepheli. Diploproctodaeum haustrum (McCallum, 1918) La Rue, 1926 Host. — Ceratacanthus scripta ( Osbeck ) , scrawled filefish. Location. — 1/3 intestine. Locality. — Bimini Vicinity; B.W.L Discussion. — Sogandares and Hutton (1958) have indicated that Bianium lecano- cephalum Perez-Vigueras, 1955 is a syn- onym of Diploproctodaeum haustrum.. D. haustrum and D. plicitum are very closely related. Further studies of larger series of specimens may reveal the species to be synonymous. At present, the constantly more sagittate forebody and longer esopha- gus of D. haustrum may be used to separate that species from D. plicitum. Diploproctodaeum plicitum (Linton, 1928) Sogandares & Hutton, 1958 (Figures 12-22) Host. — Sphoeroides annulatus (Jenyns), tamboril; Diodon hystrix (Linn.). Location. — 1/3 intestine. Locality. — Bella Vista Beach and off mouth of Chiman River, Gulf of Panama. Discussion. — Some observed variations in 80 Tulane Studies in Zoology Vol. 7 D. pUcitinn from Panama are illustrated in figs. 13-21. Enentermn aiireum Linton, 1910 (Figures 22-24) Hosts. — Kypbosus sectatrix (Linn.), Ber- muda chub; Kypbosus elegans (Peters), chopa. Location. — Mid and 3 '4 intestine of K. sectatrix and 3, 4 intestine of K. elegans. Locality. — K. sectatrix from W shore, Lerner Fish Pens, and 1 Mi. W. Entrance Point, N. Bimini, B.W.L; and K. elegans from Taboga Island, Gulf of Panama. Discussion.— Uvimtx (1947) redescribed and figured E. aureum. I have examined specimens of E. aureum from K. sectatrix and K. incisor from Tortugas, Florida. My specimens from Bimini agree in all details with the Florida specimens. E. aureum from K. elegans in Panama (fig. 22) differ only in that the vitellaria do not overlap the ceca. Live specimens of E. aureum from Bimini were observed for this detail. The vitellaria overlap of ceca is variable and is dependent upon contraction and flatten- ing pressure. Manter examined 2037 fishes of 237 species in Tortugas, Fla. and found E. aureum only in Kypbosus. Examinations of 224 species of fishes from Bimini and the Panama Pacific also yielded E. aureum from Kypbosus. Winter (1957) reported E. aureum from K. elegans in the Mexican Pacific. Two specimens of K. sectatrix measuring 15 and 45 mm respectively were taken from Sargassum weed 1 mi. W. of Entrance Point, N. Bimini, but did not harbor E. aureum. Jeancadenatia hrumpti Dollfus, 1946 (Figure 25) Host. — Kypbosus sectatrix (Linn.), Ber- muda chub. Location. — 1/4 intestine. Locality. — W. shore (opposite Lerner Laboratory), N. Bimini, B.W.I, [new lo- cality record}. Discussion. — L. hrumpti was described by Dollfus (1946) from Kypbosus sectatrix in Dakar, Africa. The species has not since been reported in the literature. Dollfus' .specimens were in poor condition and his description of the species was a provisional one. Dollfus was in doubt as to the presence of a seminal receptacle as he stated, "Le receptaculum seminis n'a pas ete reconnu, peut-ctre est-il represente par un masse volumnieuse, un peu en S, situee en avant de la glande de Mehlis et de I'ovaire: sur ce point, je teste dans I'incertitude ..." The seminal receptacle in my two speci- mens was clearly visible and is as described by Dollfus except that in one specimen it was C-shaped. The ceca and excretory vesi- cle open to the outside as in Enenterum aureum Linton, 1910. There is a single testis in both my specimens of /. hrumpti. Dollfus (1946) believed two testes to be present and stated, " ( Par suite de la mace- ration, la limite entre les deux testicules n'est pas observable ) ", and his figure shows one testis with a deep constriction at the equator. My specimens do not show this constriction. That a single testis may be- come divided into two testes with age has been demonstrated by Montgomery ( 1957) for the genus Scorpidicola (Waretrematidae). Thus there is a possibility that the testes in Dollfus' material were almost separated. Dollfus does not mention the presence of an external seminal vesicle in his description of /. hrumpti. An external seminal vesicle is present; after leaving the short cirrus sac it coils twice and ends approximately mid- way between the acetabulum and seminal receptacle. Fig. 25 shows that the accessory suckers, are 15 and 17 in number in each specimen respectively, and are surrounded by gland cells. Cableia trigoni, gen. nov., sp. nov.*' (Figure 26) Host. — Lactopbrys trigonus (Linn.), com- mon trunkfish. Location. — Intestine, next to pyloric junc- tion. Locality. — Lerner fish pens, Bimini, B.W.I. Holotype.—\M^:^M. Helm. Coll. No. 38864. Diagnosis (based on 1 specimen). — Body elongate; widest between oral sucker and acetabulum; spined; 1.596 long by 0.399 wide. Two lateral, papilla-like elevations, one on each side of body, between posterior end of posterior testis and posterior end of 6 The genus is named in honor of Dr. R. M. Cable, Purdue University, in recog- nition of his work on the life cycles of trematodes. The name trigoni indicates the host species Lactophrys trigoni. No. 3 Sogandares-Bernal: Ttre?7iatodes of Marine Fishes 81 body; another vesicular swelling ventral, on Oral sucker 0.147 long by 0.161 wide. Ace- dextral side of body posterior to lateral tabulum in anterior 1/3 body 0.19 long by dextral swelling, directly over right cecum. 0.201 wide. Sucker ratio 1:1.25. Pre- forebody 0.475 long; hind-body 0.931 long. pharynx approximately 1/2 length of phar- 45s>: »«.«. k«* • •a 3 0'^ 2.S •••/ M) b Figxires 25-30. 25. Jeuncadenatia brumpti Dollfus, accessory suckers, lateral view show- ing^ gland cells surrounding suckers. 26. Cableia trigoni, gen. nov., sp. nov., ventral view. 27. Apocreadiuni balistis Manter, ventral view. 28. Apocreadiutn colli, sp. nov., ventral view. 29. Apocreadium angustum, sp. nov., ventral view. 30. A. angustum: a, ventral view of oral sucker to show lateral fleshy lobes; b, dorsal view of oral sucker to show band of circular muscles at connection with prepharynx. 82 Tulane Studies in Zoology Vol. 7 ynx. Pharynx 0.081 long by 0.074 wide. Esophagus approximately as long as phar- ynx. Cecal bifurcation between pharynx and acetabulum. Ceca extending to pos- terior end of body where they join excretory vesicle to form a uroproct. Genital pore to left of midbody at level of cecal bifurcation. Gonads postequatorial, intercecal. Testes tandem, in contact, oval, smooth, longer than wide; anterior testis 0.235 long by 0.201 wide; posterior testis 0.302 long by 0.188 wide. Cirrus sac extending sinistrally from dextral side of posterior 1/4 acetabulum, obhquely across body to genital pore; in- ternal seminal vesicle in posterior l/'2 of sac, cirrus and ejaculatory duct surrounded by prostate cells in anterior 1/2 of sac. External seminal vesicle absent. Ovary me- dian, deeply 6-lobed, in contact with an- terior testis; 0.121 long by 0.268 wide. Club-shaped seminal receptacle extending along the left posterior edge of acetabulum from posterior to mid-acetabulum to the median ootype. Mehlis' gland large, occu- pying most of intercecal area between ovary and acetabulum. Laurer's canal not visible. Vitellaria of massive follicles, extending from posterior edge of acetabulum to pos- terior end of body, overlapping ceca. Vitel- line ducts fusing ventral to ovary to form median vitelline receptacle. Uterus coiling between ovary and acetabulum, passing along right side of acetabulum. Eggs 27 to 32 by 12 to 16 microns. Excretory pore at posterior end of body. Excretory vesicle not visible anteriorly beyond posterior testis. Generic diagnosis of Cableia. — Elongate, medium sized Lepocreadiidae. Cuticle spined. Oral sucker terminal. Acetabulum in anterior 1/3 body. Ceca joining excre- tory vesicle to form a uroproct. Genital pore sinistral to cecal bifurcation. Testes tandem, postequatorial. Cirrus sac with in- ternal seminal vesicle, and ejaculatory duct surrounded by prostate cells. External semi- nal vesicle absent. Ovary median, deeply lobed. Seminal receptacle present. Mehlis' gland massive, from ovary to acetabulum. Vitellaria follicular, overlapping ceca, ex- tending to posterior end of body. Uterus restricted to intercecal area between genital pore and ovary. Eggs small, operculate, thick .shelled. Excretory pore terminal. Excretory vesicle not observed anterior to testes. Type species: Cableia trigoni. There are only two genera of Lepo- creadiidae which lack an external seminal vesicle; these are: Stegodexamene Mac- Farlane, 1951; and Froenenterum Manter, 1954; Cableia differs from both genera by possessing a uroproct as compared with ceca ending blindly in Stegodexamene, and ceca joining but not forming a uroproct in Froenenterum. In addition, Cableia differs from Stegodexamene by having tandem postequatorial testes as compared with oblique preequatorial testes; and from Pro- enentertim by possessing an elongate in- stead of an almost round cirrus sac. The large vitelline follicles are common in the family Lepocreadiidae, but the small, thick- shelled eggs are unusual. Rhagorchis odbneriMamet, 1931 Host. — Ceratacanthus scripta (Osbeck), scrawled file fish [new host record}. Location. — 1/3 intestine. Locality. — Lerner fish pens, Bimini, B.W.I, [new locality record}. Discussion. — One immature specimen agreed in almost all details with R. odbneri described by Manter (1931) from Cerata- canthus schoepfi in Beaufort, N.C and later reported from the same host and Mono- canthus ciliatus in Tortugas by Manter (1947). Homalometron elongatum Manter, 1947 Hosts. — Gerres cinereus (Walbaum); Eucinostomus calif orniense (Gill), mojarra [new host record}. Location. — Entire length of intestine in G. cinereus and mid-intestine of £. cali- forniense. Locality. — G. cinereus from off Lerner Laboratory Pier, Bimini, B.W.I, [new lo- cality record}; and £. calif orniense from Taboga Island, Panama Pacific [new locality record}. Discussion. — Manter ( 1947 ) reported H. elongatum from Gerres cinereus in Tortu- gas, Florida. The Bimini specimens of H. elongatum agree in all details with the paratypes. The single specimen from Pana- ma appears mature in all respects except in lacking eggs, and is a little shorter than the minimum range given by Manter for H. elongatum. Bravo and Manter (1957) described papillae on the oral sucker of H. elongatum. The immature Panama speci- No. 3 Sogandares-Bernal: Ttrernatodes of Marine Fishes 83 men and the Bimini specimens have these papillae. Crassicutis marina Manter, 1947 Host. — Gerres cinereus (Walbaum), mo- jarra. Location. — Rectum. Locality. — Off Lerner Laboratory Pier, N. Bimini, B.W.I, [new locality record}. Discussion. — One mature specimen of C. marina agreed in all essential details with the paratypes of C. marina. Manter ( 1947) reported this species from Eucinostomus lefroyi and Gerres cinereus at Tortugas, Florida. Apocreadium longisinosum Manter, 1937 Host. — Sphoeroides annulatus ( Jenyns ) , tamboril. Location. — Intestine. Locality.. — Bella Vista Beach, Panama City Panarna Pacific. Discussion. — A. longisinosum in the col- lection possessed eggs whose maximum measurements were 105 microns. I have compared my specimens with paratypes and they agree in all details. Manter ( 1937 ) described the species in detail and aside from egg size nothing more need be men- tioned here regarding its morphology. Caballero, et al, (1952) reported an im- mature specimen of this species from Sphoeroides annulatus in Panama. Apocreadium balistis Manter, 1947 Host. — Balistes capriscus Gmelin, ocean tally [new host record}. Location. — 1/3 intestine. Locality. — Near Cat Cay, B.W.I, [new locality record}. Discussion.. — One specimen believed to be A. balistis was collected. It differed from A. balistis in that the acetabulum is in the anterior one-fourth body as compared with the anterior one-third body in the holotype. Collapsed eggs measured from 67 to 75 by 32 to 40 microns as compared with eggs 73 to 78 by 49 microns in the holotype. In addition, the testes of the holotype have a peculiar equatorial ridge which was not present in my specimen. Since A. balistis was described from a single specimen, the significance of the testicular ridge in the holotype is not evident. A. balistis is hitherto known from Balistes vetula. Apocreadium, colli, sp. nov.''^ (Figure 28) Hosts. — Balistes capriscus Gmelin, ocean tally; Balistes vetula Linn., queen trigger- fish. Location. — In rectum of all host speci- mens examined. Locality. — B. capriscus from near Cat Cay, B.W.I, and B. vetula from N. Shore, N. Bimini, B.W.I. Ho/o^'pe.— U.S.N.M. Helm. Coll. No. 38865. Diagnosis (based on one mature speci- men ) . — Body elongate, almost equally wide along its length, rounded at anterior and posterior ends, completely spined, 2.975 long by 0.225 wide. Oral sucker sub- terminal, elliptical, with two lateral fleshy lobes and a .sphincter muscle at junction of prepharynx and sucker, 0.429 long by 0.335 wide. Acetabulum immediately preequa- torial, 0.268 long by 0.335 wide. Sucker ratio 1:1. Prepharynx half as long as pharynx. Pharynx with a band of circular muscles in its anterior half, 0.268 long by 0.255 wide. Esophagus absent, ceca branch- ing immediately extending parallel to pos- terior end of the body. Genital pore me- dian, immediately preacetabular. Testes two, postequatorial, tandem, intercecal, smooth; anterior testis 0.220 long by 0.275 wide; posterior testis almost in contact with an- terior testis, 0.320 long by 0.247 wide. The posttesticular region about half as long as forebody. Seminal vesicle curved, club- shaped, extending from genital atrium, dor- sal to right half of acetabulum, almost to ovary. Cirrus and cirrus sac absent. Globu- lar ovary dextral, intercecal, between ace- tabulum and anterior testis; 0.150 long by 0.147 wide. Mehlis' gland sinistral, be- tween anterior testis and ovary. Seminal receptacle club-shaped, dorsal to ovary, com- ples and between ovary and anterior testis. Vitellaria follicular, from cecal bifurcation to posterior end of body, confluent anterior to acetabulum, posterior to testes, and pos- terior to tips of ceca. Uterus extending an- teriorly from ootype, with few coils. Col- lapsed eggs 80 to 88 by 20 to 40 microns. Excretory vesicle visible as a narrow tube extending at least to posterior testes. A ' This species is named in honor of Dr. William H. Coil. 84 Tulane Sti/dies in Zoology Vol. 7 lymphatic vessel extends along the inner side of each cecum, at least between anterior testis and acetabulum. Discussion. — Previously five species of Apocreadium have been named: A. mexi- canum Manter, 1937 ( type ) ; A. longisino- sum Manter, 1937; A. balistis Manter, 1940; A. synagris Yamaguti, 1953; and A. cabcd- leroi Bravo, 1954. A. coili differs from all in its short post- testicular region, acetabulum far back near midbody, and vitellaria confluent anterior to the acetabulum. Most similar to A. coili J , t? 6J is a species to be described tpr-690-) from Balistes naujragiuvi in the Panama Pacific. Apocreadium angustum, sp. nov.^ (Figures 29-30) • Host.. — Lactophrys trigonus (Linn.), com- mon trunkfish. Location. — Intestine, near pyloric junc- tion. Locality. — Lerner fish pens, Bimini, B.W.I. Holotype.—U.SNM. Helm. Coll. No. 38866. Diagnosis ( based on one mature speci- men).— Body elongate, almost equally wide along its entire length, tapering to rounded ends; spined to level of posterior testis; 7.750 long by 1. wide. Oral sucker sub- terminal, elliptical, 0.670 long by 0.0389 wide; with two lateral fleshy lobes and a set of sphincter muscles at junction of pre- pharynx and sucker. Acetabulum about one- third body length from anterior end, 0.4 long by 0.5 wide. Sucker ratio 1:1.29. Pre- pharynx four-fifths length of pharynx. An- terior fourth of pharynx composed of cir- cular muscles, 0.375 long by 0.4 wide. Short esophagus about one-sixth length of phar- ynx. Ceca extending to posterior end of body, ending blindly. Genital pore median, immediately preacetabular. Genital atrium very short, cannot be accurately measured because of its vertical position. Testes sepa- rated from each other by a short distance, postequatorial between acetabulum and pos- terior end of body, intercecal, tandem, with but slight superficial Jobation; anterior tes- tis 0.625 long by 0.26 wide; posterior testis 1.0 by 0.26 wide. Posttesticular space about six-sevenths as long as forebody. Seminal vesicle extending from genital atrium, curv- ^ The name a)ig)(sfniii (L. iiarroiv) refers to the slender body. ing around left side of acetabulum, continu- ing posteriorly past acetabulum to come nearly in contact with ovary. Globular, dextral, intercecal ovary between anterior testis and acetabulum, 0.26 long by 0.25 wide. Mehlis' gland and ootype median, between anterior testis and ovary. Seminal recepticle tubular, dorsal to ovary, Vitellaria beginning posterior to and confluent an- terior to acetabulum, overlapping ceca to posterior end of body, filling posttesticular space. Uterus preovarian with few coils. Eggs 60 to 74 by 11 to 14 microns. Ex- cretory pore terminal, vesicle not observed. One pair of lymphatic vessels extending along inner side of each cecum, visible be- tween anterior testis and acetabulum. Discussion. — This species is like A. coili in that the vitellaria are confluent anterior to the acetabulum, but its body is much narrower, the testes are much longer than wide, and the eggs considerably smaller. Apocreadimn uroproctof erum, sp. nov.*^ (Figure 31) Host. — Balistes vetula Linn., queen trig- gerfish. Location. — 2/3 intestine. Locality.— N. Shore, N. Bimini, B.W.I. Holotype.—\J.S.N.M. Helm. Coll. No. 38867. Diagnosis (based on 8 mature speci- mens).— Body lanceolate, widest at mid- posttesticular region, completely spined; 2.019 to 4.009 long by 0.270 to 1.045 wide at acetabular level and from 0.703 to 1.254 at widest point. Forebody 0.532 to 0.798 long. Hindbody 1.501 to 2.495 long. Oral sucker simple, subterminal, 0.181 to 0.248 long by 0.211 to 0.268 wide. Acetabulum at end of anterior fourth of body, recessed into body, 0.261 to 0.369 long by 0.268 to 0.362 wide. Sucker ratio 1:1.18 to 1.41. Prepharynx one-half to three-fourths length of pharynx. Anterior fourth of pharynx with a weak band of circular muscles, 0.141 to 0.181 long by 0.067 to 0.147 wide. Esophagus approximately one-fourth length of pharynx. Ceca ending in a uroproct a short distance from posterior end of body. Genital pore median, immediately preace- tabular. Genital atrium extending from genital pore posteriorly dosal to anterior '■* The name lo-oprocfofcnnii indicates that the species bears a uroproct. No. 3 Sogandares-Bernal: Ttrematodes of Marine Fishes 85 M ."^^ %• -«1 ». m 33 ^ii 34 32 Figures 31-34. 31. Apocreadmm uroproctoferum, sp. nov., ventral view. 32. Apocrea- dium bravoi, sp. nov., ventral view^. 33. Stephanostomnm proviteUosum, sp. nov., ventral view. 34. S. proviteUosHm, oral sucker and spines, ventral view. half of acetabulum. Testes equatorial, tan- dem, in contact or slightly overlapping each other; edges indented; anterior testis almost square, usually half length of posterior tes- tis, 0.268 to 0.402 long by 0.215 to 0.567 long; posterior testis widest in its anterior region, tapering posteriorly rather sharply in some specimens to produce an almost triangular appearance, 0.318 to 0.643 long by 0.328 to 0.556 wide at its anterior end. Posttesticular space approximately equal with distance from anterior testis to anterior tip of body; 0.798 to 1.520 long. Seminal vesicle club-shaped, extending from genital atrium, dorsal to acetabulum, to near or in contact with ovary. Ovary globular, dextral, intercecal, usually in contact with anterior testis; 0.168 to 0.369 long by 0.141 to 0.261 wide. Mehlis' gland median, imme- diately anterior to anterior testis, posterior to ovary. Club-shaped seminal receptacle dorsal to ovary. Vitellaria extending from 86 Tulane Studies in Zoology Vol. 7 anterior edge of acetabulum or sometimes lateral fleshy lobe on each side of mouth. from level of ovary, mostly lateral to ceca, Sucker 0.46 to 0.48 long by 0.32 to 0.36 not confluent anterior to testes, filling post- wide. Forebody 0.93 to 1.03 long. Hind- testicular space. Uterus anterior to ootype, body 1.96 to 2.34 long. Acetabulum one- coiling a few times, to enter genital atrium third from anterior end of body. 0.25 to at mid-acetabular level. Uncollapsed eggs 0.28 long by 0.25 to 0.27 wide. Sucker ratio 64 to 80 by 40 to 45 microns. Excretory from 1:0.7 to 0.81. Prepharynx from one- vesicle Y-shaped, extending anteriorly from fourth to one-half length of pharynx. Phar- terminal excretory pore to posterior testis ynx 0.25 to 0.29 long by 0.23 to 0.25 wide where it forks, sending one branch on each with prominent anterior circular muscle side of body, along inner border of ceca band. Esophagus short, about one-fifth crossing ceca anterior to acetabulum, along length of pharynx. Cecal bifurcation one- outer borders of ceca, branching profusely fourth to one-half to three-fourths distance at level of oral sucker; branches extending from oral sucker to anterior border of aceta- anterior to oral sucker. Lymphatic system bulum. Ceca extending to posterior tip of present; vessels extending along inner side body; ending blindly. Genital pore median, of ceca. A lymphatic sinus surrounding immediately preacetabular. Testes tandem, dorsal side of acetabulum, extending an- intercecal, posterior to midbody, in contact teriorly at least as far as cecal bifurcation, with one another or separated by about one- One vessel ( expecially visible in immature fifth length of posterior testes. Seminal specimens) forks from main stem vessel at vesicle sac-like from mid-dorsal region of posterior end of body next to uroproct, acetabulum extending posterior to acetabu- curves outwards from main stem on each lum one-half way to ovary. Genital atrium side of body to come in contact with each a slender tube dorsal to anterior half of side of body at level of posterior testis. acetabulum. Prostatic cells around anterior Discussion.— This species differs from all part of seminal vesicle and posterior part of others in the genus Apocreadium by pos- atrium. sessing a uroproct but otherwise is a typical Ovary ovoid, unlobed, dextral, just an- member of the genus. The presence of a terior to right edge of anterior testis and uroproct is considered a generic character near right cecum, 0.17 to 0.21 long by 0.15 in certain Opecoelidae but not in the to 0.19 wide. Mehlis' gland large, median, Acanthocolpidae (genus Stephanostomum) . immediately anterior to anterior testes to Among the species of Apocreaditim, A. left of ovary. Seminal receptacle claviform, uroproctojerum has a relatively wide body, extending forward from oviduct dorsal to Apocreadiuvi\iX2iyQ\,s^.no^y^ ovary, overlapping right side of seminal cc-.r^ ^^:ii\ vesicle. Vitellana from cecal bifurcation to (figure 32) • J £ u J n J J _, ,. , . ^ , , posterior end of body, confluent anterior to HosL-Bahstes naufragmm Jordan and acetabulum and posterior to testes. Uterus Starks, cochino. ^-^^^ ^^^ ^^^^^ anterior to Mehlis" gland. Io.^//o,^.-Intestine. , , , , Eggs 72 to 91 by 41 to 51 microns. Excre- Lo.^///)/.— Saint Bartholomew Island, Ar- ^^^j^.,^ thin-walled, extending from ex- ch.pie ago de las Perlas, Panama Pacific. ^ .^^ posterior end of body to Holotype.— U. S.N. M. Helm. Coll. No. posterior edge of rear testis where it sends •^ . ■ . one excretory branch on each side of body Diagnosis (based on five specimens).— anteriorly to vicinity of oral sucker. Ex- Body eOngate, rounded at both ends, cy- ^^ branches overlapping ceca in a tor- lindrical,^spined at least to level of testes, .^.^^^ ^^^,3^ Lymphatic vessels two, one 3.21 to x7l long by 0.7 to 0.93 wide at „j^ ^^^^ side of body, following excretory level of acetabulum. Oral sucker subtermi- branches, extending at least from anterior nal^jacking papillae but with one large ^„ ^..^j,^^, ^^.^ium to posterior to testes where 1" The species is named for Miss Marga- ^'^^T ^"^""^^ laterally to come close to each rita Bravo-Hollis of the Instituto Nacional side of body. de BioloKia, Mexico, in recoju-nition of her Discussioii.~A. bravoi is most similar to contributions to the knowledge of marine > .,.,.,. u 1 ■ rr c fish trematodolof-y from the Pacific Coast ^- ''^"'' ^'""^■''' ." resembles in confluence of of Mexico. vitellaria anterior to the acetabulum and in No. 3 Sogandares-Bernal: Trre?naiodes of Marine Fishes 87 arrangement of the gonads. However, the sucker ratio is different (1:0.7 to 0.8 as compared with 1:1); the posttesticular re- gion is much longer. The Genus Apocreadium The genus Apocreadium is closely re- lated to Homalometron, differing in the presence of lymphatic vessels. The four new species named above raise the total to nine. All are from plectognath fishes with the exception of A. mexicanum which is from Labrisomus xanti (family Clinidae), and A. synagris which is from Synagris taeniopterus (family Lutjanidae). Of the nine species, five are from Balistes (trigger fishes), one from puffers, and from trunk- fishes. The genus is predominantly American; 4 species are from Bimini; 1 Tortugas; 3 from the American Pacific from Mexico to the Galapagos Islands. The only non-American species is A. synagris from the Celebes Islands. The nine species may be segregated into three groups: (1) those with oral sucker possessing two lateral, fleshy lobes {A. bra- voi, A. coili, A. angustum)\ (2) those with simple oral sucker and without a uro- proct {A. mexicanum, A. balistis, A. cabal- leroi. A. longisinosum, A., synagris); (3) one with simple oral sucker and with a uroproct {A. uroporctoferum) . As the genus grows it may prove advisa- ble to recognize three genera, or at least three subgenera on the basis of these differ- ences. Another grouping based on the anterior confluence of vitellaria is possible. This character is more easily observed than a uro- proct. Three {A. coili, A. angustmn, A. bravoi) have vitellaria confluent anterior to the acetabulum; the other six do not. F. Family MEGAPERIDAE Manter, 1934 Megapera gyrina (Linton, 1907) Manter, 1934 Hosts. — Lactophrys tricornis (Linn.); Lactophrys bicaudalis (Linn.), trunkfish. Location. — Middle intestine of L.. tricor- nis and 2/3 intestine of L. bicaudalis. Locality. — N. shore N. Bimini, B.W.I. and near Cat Cay, B.W.I. Thysanopbarynx elongatus Manter, 1933 Host. — Lactophrys tricornis (Linn.), trunkfish. Location. — Middle intestine. Locality.— N. Shore of N. Bimini, B.W.I. G. Family ACANTHOCOLPIDAE Liihe 1909 Stephanostomum provitellosum, sp. nov.^^ (Figures 33-34) Hosts. — Balistes naujragium Jordan & Starks; Balistes polylepis Steindachner, trig- ger fish, cochino. Location. — Rectum. Locality^ — B. naujragium from Taboga Island, Panama; and B. polylepis from Gala- pagos Islands (Manter's collection). Holotype.—U.SN.U. Helm. Coll. No. 38869. Diagnosis (based on ten specimens; meas- urements on nine). — Body 2.622 to 3-572 long by 0.513 to 0.665 wide, spined to pos- terior end. Cuticle heavily spined, with anteriormost row of spines (proximal to oral spines) larger than other body spines. If the second and third row of body spines are missing, the oral crown appears to have three rows of spines of which the posterior row is separated by some distance; this lat- ter row of spines is to be interpreted, I be- lieve, as the anterior row of body spines. Forebody 0.893 to 1.116 long. Posterior body 1.425 ot 1.950 long. Oral sucker ter- minal, 0.285 to 0.361 long by 0.241 to 0.342 wide; with two rows of 28 spines each; spines in both rows approximately 40 microns long. Acetabulum immediately an- terior to midbody from 0.285 to 0.361 long by 0.268 to 0.323 wide. Sucker ratio from 1:0.94 to 1.12. Prepharynx 0.285 to 0.4 long. Pharynx pear-shaped, with a band of circular muscles around its anterior third; 0.133 to 0.190 long by 0.133 to 0.19 wide. Esophagus a little shorter than pharynx. Ceca extend to posterior end of body where they join the excretory bladder to form a uroproct. Genital pore median, close to an- terior edge of acetabulum. Gonads post- equatorial, tandem, intercecal. Testes in con- tact with each other or slightly overlapping, globular; anterior testis 0.304 to 0.435 long by 0.235 to 0.342 wide; posterior testis 0.380 to 0.494 long by 0.228 to 0.323 wide. 11 The name proviteUosnm is from the latin pro-, anterior and viteUus, >oIk glands, in reference to the anterior band of vitelline follicles which characterize the species. 88 Tulane Studies in Zoology Vol. 7 Cirrus sac extending posterior to acetabulum from 1/2 way to almost in contact with ovary; internal seminal vesicle in posterior 1/2 to 13 sac; cirrus in anterior 1/2 to 2/3 sac, spined. Ovary globular, pretesticu- lar, median, usually in contact with anterior testis, 0.190 to 0.266 long by 0.152 to 0.266 wide. Vitellaria follicular, filling posttes- ticular space, overlapping dorsally and ven- trally and extending across body immedi- ately anterior to acetabulum, interrupted at acetabular level. Uterus intercecal between ovary and genital pore; entering a thin- walled metraterm which is as long as cirrus sac and unspined. Eggs thin-shelled. Col- lapsed eggs in utero, 57 to 75 microns long. UncoUapsed eggs in utero proximal to ovary 58 to 74 by 46 to 53 microns. Excretory vesicle not observed anterior to posterior testis. Discussion. — At least 40 species have been named in the genus Stephanostomum Looss, 1899. S. proiitellosum differs from all by possessing a transverse band of vitel- line follicles anterior to acetabulum and with follicles interrupted at the level of the acetabulum. The species is probably of the S. dentatinn (Linton, 1901) type, but differs in details of vitellaria and in having a uroproct. It resembles S. casum (Linton, 1910) in possessing a uroproct but differs in distribution of vitellaria and number of oral spines. Stephanostomum hispidum ( Yamaguti, 1934) Manter, 1940 Host. — Seriola mazatlana Steindachner, bohala [new host record}. Location. — Rectum. Locality. — Saint Bartholomew Island and Galera Island, Archipielago de Las Perlas, Panama Pacific. Discussion. — Yamaguti (1934) described this species from Seriola quinqueradiata in the Japanese Pacific and Manter (1940a) recorded it from Seriola dorsalis and Seriola sp. "not dorsalis" in Mexico, and from Seri- ola dorsalis in Panama. I have studied the specimens from Seriola spp. and these agree with the S. hispidum in my collection and with some specimens sent by Yamaguti to Manter. Two specimens of the S. hispidum of Manter (1940a; from Elagatis bipinnu- latus in Panama were also studied. One was decapitated, the other intact. The crown spines of the intact specimen reveal that it is probably S. ditrematis. Stephanostoynum ditrematis (Yamaguti, 1939) Manter, 1947 Hosts. — Caranx latus Agassiz, horse-eyed jack; Elagatis bipinnulatus (Quoy & Gain- ard ) , blue runner. Location. — Rectum of C. latus and in- testine other than rectum in E. bipinnulatus.. Locality. — C. latus from Lerner fish pens, and off Lerner Laboratory Pier, N. Bimini, B.W.I, [new locality record}; and E. bi- pinnulatus from Niagara Rock, between San Jose and Pedro Gonzales Islands, Archi- pielago de Las Perlas, Panama Pacific. Discussion. — This species was described by Yamaguti (1939) from Ditrema tem- nicki in Japan. S. ditrematis has been pre- viously reported from Caranx caballus at Woods Hole by Linton (1940) and from Caranx ruber and C. latus at Tortugas. These fishes are related to Elagatis bipinnu- latus.. Perez-Vigueras ( 1955 ) described Stephanostomum manteri and Stephanosto- mum cubanum from Caranx ruber in Cuba. S. manteri was described from one mature specimen and 5'. cubanum from an imma- ture one. Neither species can be differenti- ated from Stephanostomum ditrematis, and are here considered synonyms of it. The other species of Stephanostomum described by Perez-Vigueras (1955), are also syno- nyms of one species or another, I believe. S. mediovitellarum and S. lopezneyrai agree in detail with the description of 5'. sentum, and in addition have been collected from the same hosts by Manter (1947) at Tor- tugas and by myself from Calamus at Bi- mini. S. microcephalum was described from one specimen in which some of the peri- buccal spines were missing. The descrip- tions and figure of S. microcephalum agree in morphological detail with both 5'. di- trematis and S. hispidum. The true specific nature of i\ ?mcrocephalum cannot be under- stood until the peribuccal spines and ar- rangement are studied in the holotype. 5". admicrostephanufn from Epinephelus mys- tacinus in Cuba is believed to be a synonym of S. microstephanum Manter, 1934 from the same host at Tortugas. Perez-Vigueras (1955) compared the two species and stated the differences as being ( 1 ) sucker ratio 1:1.33 as compared with 1:1.49, (2) No. 3 Sogandares-Bernal: Ttrematodes of Marine Fishes 89 "the separation between the esophageal bi- furcation and the acetabulum"; and (3) the number and disposition of the peribuccal spines. Manter (1934) stated that the sucker ratio was "about" 4:3; — the figure given by Manter and his description of S. microstephanus show no difference of posi- tion of the esophageal bifurcation from S. admicrostephanus: and Manter stated that the peribuccal spines in his specimens were easily lost which probably accounts for the dorsal and ventral bare spots of the oral crown in Vigueras" specimen. Stephanostomum hispidum ( Yamaguti, 1934) Manter, 1940 and Stephanostomum ditrematis (Yamaguti, 1939) Manter, 1947 are very closely related and may prove to be one species. Stephanostom.um sentum (Linton, 1900) Manter, 1947 Host. — Calamus bajonado (Bloch & Schneider), jolthead porgy; Cynoscion albus (Gunther), corvina {new host record] Mala- canthus plnmieri (Bloch), sand fish \_new host record] Location. — Rectum. Locality. — C. banjonado from 1/2 mi. W. (off Lerner Laboratory grounds) of N. Bi- mini, B.W.L; C. albus from Taboga Island, Panama Pacific; and M. plumieri from be- tween S. Bimini and Cat Cay, B.W.L Discussion. — These specimens were iden- tified as S. sentum for the following rea- ( 1 ) the vitellaria extend anteriorly sons: only to the middle of the cirrus sac; (2) the cirrus sac extends less than halfway between acetabulum and ovary; (3) the oral spines are 36 in number in two alternate rows of 18 spines each; (4) the pre-pharynx is more than twice the length of the pharynx; (5) the sucker ratio is from 1:1.0 to 1.3; (6) the testes are in contact with each other in the posterior half of the body; and (7) some vitelline follicles come between the ovary and anterior testis. S.. casum (Linton, 1910) is very closely related to S. sentum, differing mainly in that the cirrus sac does not extend halfway between acetabulum; the vitellaria are less extensive anteriorly, and extend between the anterior testis and ovary. StephanostOTUum dentatum (Linton, 1910) Manter, 1947 Host. — Mycteroperca v. venenosa (Linn.), yellow-fin grouper. Location. — Rectum. Locality. — Lerner fish pens, N. Bimini, B.W.L [new locality record]. Stephanostomum minutum (Looss, 1901) Manter, 1940 Host. — Malacanthus plumieri (Bloch), sandfish [new host record]. Location. — Rectum. Locality. — Between S. Bimini and Cat Cay, B.W.L [new locality record]. Discussion. — Caballero (1952) considered S. sentum (Linton, 1910) Manter, 1947 a synonym of S. minutum. I do not agree with such synonymy because S. minutum is fully mature at a much smaller size than S. sentum from the same host. The host from which S. minutum is reported in this paper also possessed two immature speci- mens of what I have considered 5". sentum. These immature specimens of S. sentum are approximately twice the size of the fully mature S. minutum. In addition, the oral spines of S.. minutum are more delicate than those of S. sentum. I prefer to retain S. minutuyn at least until life cycles of the two species are studied. Morphological dif- ferences may be found in the cercariae of the two species. Stephanostomum coryphaenae Manter, 1947 Host. — Coryphaena hippurus Linn., dol- phin. Location. — Rectum. Locality. — Lerner fish pens, Bimini, B.W.L [new locality record]. Discussion. — Yamaguti ( 1934) reported collections of trematodes from Coryphaena hippurus in Japan, but apparently never found S. coryphaenae in that area. Both Manter (1940a) and I have examined a number of dolphins from the tropical Amer- ican Pacific but have not found Stephanos- tomum coryphaenae. S. coryphaenae has been reported only from the American At- lantic. So far as is known, this appears to be a case where the host is widespread but harbors S. coryphaenae in only one ocean. This may indicate that the life cycle of 5'. coryphaenae may be completed only in the American Atlantic, and that the dolphin does not migrate easily from one ocean to the other. 90 Tulane Studies in Zoology Vol. 7 Stephanostomuni-psQ\idoc^.VAr\^\s, sp. nov.*- ( Figures 35-36) Host. — Holocentrus ascensionis (Osbeck), squirrel-fish. Location. — Immature in pyloric cecum, mature forms in rectum. Locality. — Lerner fish pens, N. Bimini; and near Cat Cay, B.W.I. Holotype.—\].SNM. Helm. Coll. No. 38870. Diagnosis (based on 3 mature speci- mens).— Body elongate, widest at testicular level, spined, 3.267 to 3629 long by 0.513 to 0.646 wide at acetabulum. Two eye- spots present at level of mid-prepharynx. Forebody 0.969 to 1.045 long. Hindbody 1.976 to 2.067 long. Oral sucker terminal; 0.107 to 0.167 long by 0.167 to 0.201 wide; with 36 peribuccal spines in alternate rows of 18 each, measuring from 47 to 54 mi- crons long. Acetabulum approximately in anterior 1/3 body; 0.241 to 0.255 long by 0.249 to 0.308 wide. Sucker ratio 1:1.27 to 1.53- Prepharynx approximately 2.25 times longer than pharynx. Pharynx pyri- form, almost equidistant between oral sucker and acetabulum; 0.160 to 0.214 long by 0.167 wide. Esophagus about 1.25 times longer than pharynx. Cecal bifurcation ap- proximately midway between pharynx and acetabulum; ceca joining excretory vesicle to form a uroproct. Genital pore median, immediately preacetabular. Genital atrium to about mid-acetabulum on left side. Gon- ads intercecal ,tandem in posterior third of body. Testes slightly separated from each other or juxtaposed, roundish and smooth; anterior testis 0.274 to 0.295 long by 0.228 to 0.282 wide; posterior testis 0.308 to 0.362 long by 0.227 to 0.261 wide. Cirrus sac from genital atrium to approximately 2/3 distance from acetabulum to ovary; in- ternal seminal vesicle bulbular in posterior 1/4 sac; cirrus licavily spined, in anterior 3/4 sac. Ovary rounded, immediately an- terior to testes and juxtaposed or separated from testes by a short distance. Vitellaria forming a transverse band preacetabularly between cecal bifurcation and acetabulum. 1- The name j)scHdoca raiiyia indicates the close relationship of the species with Sfei)h- nnoafonriDn cayaiigiH (Yamaj2:uti, 19.")1) Ca- ballero, 1952. extendmg to posterior end of body, over- lapping ceca dorsally and ventrally, filling posttesticular space. Uterus coiling a few times between ovary and cirrus sac; metra- term spined, to left of cirrus sac, almost as long or longer than cirrus sac — depending on whether the cirrus is restracted or pro- truded. Genital atrium short. Eggs thin shelled, 51 to 59 by 32 to 35 microns. Ex- cretory pore terminal; excretory vesicle not visible anterior to posterior testis. Discussion. — Only three species of Steph- anostomurn possess vitellaria that are con- fluent anterior to the acetabulum; these are: S. carangis (Yamaguti, 1951) Caballero, 1952; S. microcephalum Manter, 1934; and S. provitellosum (this paper). S. pseudo- carangis differs from all three by possessing a spinous metraterm. It further differs from 5'. carafigis by possessing a cirrus sac which extends posteriorly 2 3 distance from the acetabulum to the ovary, as compared with an extent midway from acetabulum to ovary. S. pseudocarangis differs from S. micro- cephalum by possessing 36 peribuccal spines in 2 rows of 18 each as compared with 150 spines in 3 concentric rows. S. pseudocaran- gis further differs from S. provitellosu^n by possessing vitellaria which are not inter- rupted opposite the acetabulum. Tormopsolus orientalis Yamaguti, 1934 (Figure 37) Host. — Seriola mazatlana Steindachner, amber jack, bohala [new host record}. Location. — Intestine. Locality. — Saint Bartholomew Island, Ar- chipelago de las Perles, Panama. Discussion. — This species was described from Seriola quinqueradiata in Japan. Ya- maguti ( 1934) illustrated the type speci- men with the cirrus sac reaching approxi- mately 3 4 distance from the acetabulum to beginning of the vitellaria, the hermaphro- ditic duct about 1 4 length of cirrus sac, and the genital pore immediately preace- tabular. Yamaguti stated that the posterior limit of the cirrus sac may sometimes reach the level of the vitellaria or even beyond. My specimen of T. orientalis does not agree with the type specimen with respect to ex- tent of the cirrus sac and length of the hermaphroditic duct. The hermaphroditic duct is almost as long as cirrus sac, the junction of uterus with hermaphroditic duct No. 3 Sogandares-Bernal: Ttrematodes of Marine Fishes 91 \% I* J* I m • ♦ • •. >• m* •% :••' T Figures 35-39. 35. Stephanostomum pseudocarangis, sp. nov., ventral view. 36. S. pseu- docarangis, oral sucker and spines, ventral view. 37. Toiuiopsolns orientalis Yamaguti, ventral view. 38. Mgodera t)iagna, sp. nov., latero-ventral view. 39. M. magna, herma- phroditic sac. is approximately 1 /3 distance from aceta- bulum to vitellaria (as compared with im- mediately behind acetabulum in the holo- type). I have examined Hanson's (1950) specimens of T. orientalis reported from a bonito in Bermuda. These specimens agree with mine in all details. The differences between my specimen and the figured holo- type apparently are similar to the variations in T. orientalis described by Yamaguti. Measurements of my specimen are as fol- lows: Body 11.286 long by 0.475 wide at acetabular level. Forebody 0.665 long. Hind- body 10.298 long. Oral sucker 0.209 long by 0.209 wide. Acetabulum 0.323 long by 0.361 wide. Sucker ratio approximately 92 Tulane Studies in Zoology Vol. 7 1:1.55. Pharynx 0.152 long by 0.114 wide. Ovary 0.21 long by 0.152 wide. Anterior testis 0.475 long by 0.190 wide. Posterior testis 0.490 long by 0.247 wide. Posttesticu- lar space 0.665 long. Eggs 54 to 87 by 26 to 60 microns. H. Faymly WARETREMATIDAE Srivastai'a. 1939 Myodera magna, sp. nov.'-^ (Figures 38-39) Host. — Kypbosus elegans Peters, chopa, chub. Location. — Intestine. Locality. — Taboga Island, Gulf of Panama. Holotype.—U.S^.M. Helm. Coll. No. 38875. Diagnosis (based on one specimen). — Body cylindrical, spined anteriorly, 5.237 long by 0.77 wide. Oral sucker terminal, 0.228 long by 0.228 wide. Forebody 0.737 long, with ventrolateral parenchymal muscle bands. Posterior body 4.5 long. Acetabulum 0.282 long, retractile, distally located on a peduncle; with one muscular, conical, equa- torial papilla on each side of longitudinal aperture. Acetabular peduncle 0.335 long. Sucker length ratio 1:1.22. Prepharynx ap- proximately 1/7 length of pharynx. Phar- ynx 0.282 long by 0.282 wide, with anterior and posterior circular bands 1/3 and 17 length of pharynx respectively. Esophagus approximately twice longer than pharynx, muscular, bifurcating for 1/4 its posterior length before connecting with ceca. Ceca voluminous, extending to posterior end of body, joining excretory vesicle to form a uroproct. Testis single, in posterior 1/4 body, intercecal, more than 3 times longer than wide, 0.740 long by 0.201 wide. Her- maphroditic sac 0.310 long, between aceta- bulum and oral sucker, surrounded by pro- static cells which are in turn surrounded by a thin membrane. Hermaphroditic sac (fig. 39) contains an internal seminal vesi- cle which occupies posterior 3 /4 of the sac, a small prostatic vesicle, and a tube which opens on a genital lobe separated from the hermaphroditic sac by a weak muscular band. Also within the sac is a muscular metraterm which opens on the genital lobe separately from the male duct. External seminal vesicle tubular, entering '•'' The name magna indicates the laiRe size of the body. right side of posterior end of hermaphro- ditic sac after first making one anterior loop at level of mid-hermaphroditic sac, extend- ing posteriorly to near ovary. Ovary globu- lar, 0.268 long by 0.268 wide, intercecal, widely separated from testis by a space al- most equal to posttesticular space. Mehlis' gland dorsal to ovary. Uterine seminal re- ceptacle present. Laurer's canal opening dor- sally at level of Mehlis' gland. Vitellaria extending from about 0.5 posterior to cecal bifurcation to posterior end of body, dorsal and ventral to ceca. Uterus intercecal, an- terior to ovary, entering left side of posterior end of hermaphroditic sac to connect with muscular metraterm. Eggs with polar fila- ments approximately 3 times longer than egg, 60 to 64 by 20 to 36 microns. Excre- tory vesicle not visible anterior to testis. One pair of lymphatic vessels on each side of body visible a short distance posterior to cecal bifurcation. Discussion. — Myodera magna differs from M. medialunae Montgomery, 1957, the only other species in the genus, as follows: ( 1 ) the anterior portion of the uterus is non-muscular; ( 2 ) the acetabular peduncle is not as long; ( 3 ) the posterior body is about 4 times longer than forebody as com- pared with only half as long; (4) external seminal vesicle extends almost to ovary as compared with extending only 2 3 distance from hermaphroditic sac to ovary; ( 5 ) the ovary is separated from the testis by a dis- tance of almost 3 '4 length of testis as com- pared with ovary and testis almost in con- tact; (6) papillae absent from posterior end of body; and, ( 7 ) polar filaments 3 times longer than egg as compared with filaments 4 to 5 times longer than egg. Although Montgomery ( 1957 ) did not figure or describe a membrane around the prostatic cells surrounding the hermaphro- ditic sac, a study of a paratype specimen of M. medialunae shows that there is a very delicate membrane present. The paratype of M. medialunae at my disposal is very darkly stained and details of a lymphatic system cannot be accurately observed. If Myodera medialunae does not possess a lymphatic system, a new genus may be erected for M. magna. Manter (1947) has already used this character to separate Hofualometron from Apocreadium in the Lepocreadiidae. No. 3 Sogandares-Bernal: Ttreniatodes of Marine Fishes 93 Megasolena kyphosi, sp. nov.^* ( Figures 40-42 ) Host. — Kyphoses analogus (Gill), salema. Location. — Intestine. Locality. — Taboga Island, Gulf of Panama. Ho/o/)/7^.— U.S.N.M. Helm. Coll. No. 38876. Diagnosis (based on four specimens). — Body unspined, elongate, cylindrical, 2.790 to 2.83 long by 0.480 to 0.660 wide, bearing eyespots slightly lateral to, or overlapping, dorsal posterior edge of oral sucker. Oral sucker subterminal, 0.320 to 0.399 wide, posterior portion of sucker with circular muscle band 1/14 to 1/24 length of sucker (fig. 42). Acetabulum 0.893 to 0.912 from anterior end of body, 0.17 to 0.228 wide. Sucker ratio 1:0.53 to 0.58. Prepharynx ap- proximately 1 4 length of pharynx. Phar- ynx 0.230 'to 0.275 long by 0.023 to 0.260 wide, with anterior circular muscle band 1/4 to 1/3 length of pharynx. Esophagus approximately same length as pharynx. Cecal bifurcation slightly posterior to or dorsal to acetabulum. Ceca voluminous, in- dented on inner aspect at level of each testis, extending to posterior end of body, ending blindly. Genital pore slightly sinis- tral, usually at level with left posterior edge of pharynx. Testes intercecal, immediately postequatorial, round in shape, tandem; an- terior testis separated from posterior testis by approximately half its length, 0.228 to 0.320 long by 0.190 to 0.220 wide; pos- terior testis 0.266 to 0.322 long by 0.190 to 0.247 wide. Posttesticular space 0.684 to 0.799. Hermaphroditic sac immediately to left of midline, 0.270 to 0.370 long, partly ventral to pharynx, partly dorsal to aceta- bulum, posterior end extending to mid- or posterior-edge of acetabulum or slightly beyond. Internal seminal vesicle connecting with metraterm near middle of hermaphro- ditic sac. Genital sinus leads to a short genital atrium which opens to outside through genital pore. Prostatic cells sur- rounding internal seminal vesicle and geni- tal sinus within hermaphroditic sac. Ex- ternal seminal vesicle from halfway between acetabulum and ovary from anterior edge of ovary, to hermaphroditic sac. Ovary 1/4 to 1/2 distance from anterior testis to aceta- 1-1 The name kyphosi is for the host genus Kyphosus. bulum, globular in shape, 0.13 to 0.174 long by 0.131 to 0.152 wide. Vitellaria ex- tending from about 1/2 way from anterior testis to acetabulum, to posterior end of body, overlapping ceca along posterior ex- tent, fusing medially between ovary and anterior testis, between testes, to fill post- testicular area. Uterus preovarian, intercecal entering genital sinus on left side of herma- phroditic sac, basal portion sometimes swol- len with sperm. Eggs 64 to 74 by 43 to 47 microns. Excretory vesicle Y-shaped, bi- furcating immediately behind ovary, arms extending laterally and dorsally to level with posterior edge of acetabulum. Lymphatic system present, but vessels not clearly visible. Immature specimens. — Measurements (3 specimens): body 2.280 to 2.740 long by 0.532 to 0.665 wide. Oral sucker subtermi- nal, 0.320 to 0.360 wide. Acetabulum 0.836 to 0.391 from anterior end of body, 1.311 to 1.824 from posterior end of body, 0.171 to 0.190 wide. Sucker ratio 1:0.49 to 0.56. Pharynx 0.210 to 0.370 long by 0.230 to 0.240 wide. Anterior testis 0.108 to 0.25 long by 0.055 to 0.103 wide. Posterior testis 0.250 to 0.228 long by 0.114 to 0.130 wide. Ovary 0.090 to 0.190 long by 0.090 to 0.209 wide. Vitelline glands apparently just differentiating in a specimen 2.74 long (fig. 41). Discussion. — M. kyphosi is closely related to M. estrix Linton, 1910 (figs. 43, 44). Manter (1935) redescribed the latter. I have studied specimens from Kyphosus in- cisor and K. sectatrix from Tortugas, Florida. M. kyphosi has vitellaria confluent between the ovary and anterior testis and between testes. In all six specimens of M. estrix observed, the testes are always in close con- tact and no vitelline follicles come between them. This character appears to be constant and easily separates the two species. I. Family HAPLOSPLANCHNIDAE Poche,1923 Haplosplanchnus acutus (Linton, 1910) Manter, 1937 Hosts.. — Thyrinops pachylepis (Gunther) [new host record}; Tylosurus raphidoma ( Ranzani ) , needle fish. Location. — Intestine, next to pyloric junc- tion. Locality. — T. pachylepis from Bella Vista Beach, Panama City, Panama Pacific; and 94 Tulane Studies in Zoology Vol. 7 42 Figures 40-46. 40. Meganoleva kyphosi, sp. nov., ventral view of mature specimen. 41. M. Icyphosi, ventral view of immature specimen. 42. M. kyjihoxi, ventral view, show- ing circular muscle bands at base of oral sucker and anterior border of pharynx. 43. Megasolena estrix (Linton), dorsal view. 44. M. estrix, dorsal view, showing circular muscle bands at base of oral sucker and anterior border of pharynx. 45. Hclicometrina mmia Linton, ventral view, from LHtjauas synagris. 46. H. )iiiiiia, ventral view, fi'om Haeniulon sciurus. No. 3 Sogandares-Bernal: Ttrematodes of Marine Fishes 95 T. raphidoma from off Lerner Laboratory pier, N. Bimini, B.W.I, [new locality record}. Discussion. — H. acutus has been reported from Tylosurus rhaphidoma and T. acus in Bermuda by Linton, and from T. marinus at Tortugas by Manter (1947). Caballero, Bravo and Grocott ( 1953) reported it from Tylosurus jodiator in the Panama Pacific. I examined one specimen each of Tylosurus stolzmani and T. pacificus but did not find H. acutus. The record from Thyrinops pachylepis is believed to represent an acci- dental infection. Manter (1940a) reported a single specimen of H. acutus from Ky- phosus. His record from this host may also represent an accidental infection. The tre- matode is typically a parasite of needle fishes. J. Family OPECOELIDAE Ozaki, 1923 Helicometrina nimia Linton, 1910 (Figures 45-46) Hosts. — Epinephelus analogus Gill, group- er [new host record}; Haemulon albu?n Cuv. & Val., margatefish [new host record}; Hae- mulon sciurus (Shaw), blue-striped grunt [new host record}; Lutjanus synagris (Linn.), lane snapper. Location. — Pyloric ceca. Locality.. — E. analogus from tidepool in Bella Vista, Panama City, Panama Pacific; H. album from Lerner Fish Pens, N. Bimini, and H. sciurus and L. synagris from N. shore, N. Bimini, B.W.L Discussion. — H. nimia has been reported from 16 and 7 host species from the Atlantic and Pacific coasts of the continent respec- tively. Twelve families of fishes serve as definitive hosts for H. nim-ia. The only fish family which H. nimia has in common as a host from both oceans is the Scorpaenidae. Helicometra torta Linton, 1910 ( Figure 47 ) Host. — Epinephelus striatus (Bloch), Nassau grouper. Location. — 1/4 intestine. Locality. — Lerner Fish Pens, N. Bimini, B.W.L [new locality record}. Helicometra execta Linton, 1910 Host. — Haemulon sciurus ( Shaw ) , blue- striped grunt; Gymnothor'AX vicinus (Castel- nau), brown moray [new host record}. Location. — 1/4 intestine of H. sciurus; and mid intestine of G. vicinus. Locality.. — H. sciurus from N. shore, N. Bimini; G. vicinus from opposite Lerner Laboratory Pier, N. Bimini, B.W.L [new locality record}. Stenopera equilata Manter, 1933 ( Figure 48 ) Host. — Holocentrus ascencionis (Osbeck), squirrel-fish. Location. — Pyloric ceca. Locality. — Lerner Fish Pens, N. Bimini, and near Cat Cay, B.W.L [new locality record}. Neonotoporus yamagutii Manter, 1947 Host.. — Selar crumenopthalmus ( Bloch ) , goggle-eyed scad. Location. — 1/2 intestine. Locality. — Off Lerner Laboratory Pier, Bimini, B.W.L [new locality record}. Pre- viously known from Tortugas. Pseudopecoeloides gracilis Manter, 1947 Host. — Selar crumenopthalmus ( Bloch ) , goggle-eyed scad; Apogon bmotatus (Poey), cardinal fish. Location. — Mid-intestine of S. crumenop- thalmus and 3/4 intestine of A. binotatus. Locality. — Off Lerner Laboratory Pier, N. Bimini, B.W.L Podocotyle mycteropercae, sp. nov.^*"^ ( Figure 49 ) Host. — Mycteroperca falcata ( Poey ) , scamp. Location. — Pyloric ceca. Locality. — "Bimini vicinity", B.W.L Holotype.—U. S.N.M. Helm. Coll. No. 38871. Diagnosis (measurements on one mature specimen). — Body elongate, widest at level of acetabulum and tapering to a rounded end; cuticle unspined; 2.360 long by 0.540 at greatest width. Forebody 0.750 long. Hindbody 1.210 long. Oral sucker sub- terminal, 0.201 long by 0.201 wide. Aceta- bulum short pedunculated, 0.402 long by 0.369 wide. Sucker ratio 1:1.83. Prephar- ynx absent. Pharynx 0.101 long by 0.094 wide. Esophagus about equal to pharynx length. Ceca extending to posterior end of body, ending blindly. Genital pore slightly sinistral, midway between cecal bifurcation i"' The name mycteropercae is for the host genus, Micteroperca. 96 Tulane Studies in Zoology Vol. 7 ■ \r 49 Figures 47-52. 47. Helicometra torta Linton, ventral view. 48. Stenopera eqnilata Man- ter, ventral view. 49. Podocotyle mycteropercae, sp. nov., ventral view. 50. Pachycrea- dium lerneri, sp. nov., ventral view. 51. Opecoeliis sphaericiis Ozaki, ventral view. 52. Pseudopecoeloides equesi Manter, latero-ventral view. No. Sogandares-Bernal: T(re??iatodes of Marine Fishes 97 and acetabulum. Gonads tandem, postequa- torial, intercecal. Testes roundish, smooth, almost in contact with each other; anterior testis 0.21 long by 0.2 v/ide; posterior testis 0.2 long by 0.2 wide. Cirrus sac extending posterior to acetabulum about halfway to ovary ( in contact with ovary in immature specimens); cirrus in anterior 16 sac; in- ternal seminal vesicle surrounded by prostate cells. Ovary smooth, roundish, median, slightly overlapping anterior testis; 0.15 long by 0.13 wide. Seminal receptacle club- shaped, dextral and anterior to ovary. Vitel- laria extending laterally from level of an- terior end of ovary to posterior end of body, overlapping ceca and portions of testes, fill- ing posttesticular space. Uterus preovarian. Eggs (collapsed) 53 by 24 microns. Excre- tory pore terminal, excretory vesicle saccu- lar, extending to level of ovary. Discussion. — There are 38 species in the genus Podocotyle Dujardin, 1845. Of these only 6 possess a smooth ovary and cirrus sac extending posteriorly past the aceta- bulum; these are: P. epinepheli Yamaguti, 1942; P. gracilis Yamaguti, 1952; P. meco- pera Manter, 1940; P. pedicillatuni (Stos- sich, 1887) Stossich, 1898; P. petallophal- lus (Yamaguti, 1934) Park, 1937, and P. serrani Yamaguti, 1952. P. mycteropercae differs from P. gracilis and P. pedicillatum by possessing vitellaria which do not reach the acetabulum, and an almost median genital pore as compared with vitellaria reaching acetabulum and a definitely sinistral genital pore. P. myc- teropercae differs from P. pet all op hall us by possessing a smooth cirrus, short stalked acetabulum^ and vitellaria ending at the level of the anterior edge of the ovary, as com- pared with a cirrus with petaloid append- ages, a longer stalked acetabulum and vitel- laria extending anteriorly far beyond level of ovary. P. mycteropercae differs from P. epinepheli and P. mecopera in sucker ratio (1:1.83), an almost straight seminal vesicle, and genital pore almost median and between cecal bifurcation and acetabulum, as com- pared with sucker ratio of about 1:2.3, a convoluted internal seminal vesicle, and genital pore on outer side of cecum at level of cecal bifurcation. P. mycteropercae dif- fers from P. serrani in that the vitellaria do not extend anteriorly much past the ovary, a more posterior acetabulum, almost straight cirrus sac extending only a short distance posterior to acetabulum, genital pore almost median on inner aspect, as compared with vitellaria clearly extending far beyond ovary, more anterior acetabulum, sigmoid-shaped cirrus sac extending for at least half its length posteriorly beyond acetabulum, and genital pore to the left of cecum. Pachycreadium lerneri, sp. nov.^*^ (Figure 50) Host. — Gerres cinereus (Walbaum), mo- jarra. Location. — Pyloric cecum. Locality. — Off Lerner Laboratory Pier, Bimini, B.W.I. Ho/o/>/?e.— U.S.N.M. Helm. Coll. No. 38872. Diagnosis. — Body robust, elongate; cuti- cle unspined and thick; broadly rounded at posterior end; 1.406 long by 0.646 wide at acetabular level. Forebody 0.482 long. Hind- body 0.637 long. Oral sucker subterminal; 0.161 long by 0.141 wide. Acetabulum pre- equatorial; 0.26 long by 0.328 wide. Sucker ratio 1:2.33. Prepharynx absent. Pharynx 0.1 long by 0.074 wide. Esophagus approxi- mately 1/2 length of pharynx. Ceca ending blindly near posterior end of body. Genital pore slightly sinistral, between cecal bifurca- tion and pharynx. Gonads postequatorial, intercecal. Testes diagonal, sinistral testis anteriormost, smooth, twice longer than wide, 0.228 by 0.114 wide; dextral testis 0.228 long by 0.1 wide. Cirrus sac median, overlaping anterior 1/4 of acetabulum; cir- rus in anterior 1/5 sac; internal seminal vesicle in posterior 4/5 sac. Ovary to right of midline, overlapping dextral testis slightly, smooth, globular, 0.134 long by 0.101 wide. Seminal receptacle not observed. Vitellaria extending from level of genital pore to pos- terior end of body, dorsal and ventral to ceca. Uterus from mid-ovarian level to geni- tal pore. Collapsed eggs 56 to 69 by 32 to 40 microns; one egg with an antiopercular filament 21 microns long; other eggs with- out filament. Excretory pore ventral, at level of ends of ceca; anterior extent of excretory bladder not observed. Discussion. — Manter (1954) erected the genus Pachycreadium for Plagioporus gas- 16 The name lerneri is in honor of Mr. Michael Lerner in appi-eciation for his aid in this study. 98 Tulane Studies in Zoology Vol. 7 trocotylum Manter, 1940. Aside from P. gas- trocotyluni (Manter, 1940) Manter, 1954 (type species) the genus contains one other species, Pachycreaclinm crasugidum ( Linton, 1910) Manter, 1954, (syn: Plagioporus cras- signlns (Linton, 1934) Price, 1934). P. lerneri differs from P. crasslguliim by pos- sessing a longer hindbody and more unequal sucker ratio. P. lerneri appears to be most closely related to P. gastrocotylum differing in sucker ratio (1:2.33) as compared with 1:2), smaller pharynx, vitellaria not con- fluent anterior to the acetabulum, and testes longer than wide rather than wider than long. P. gastrocotylum is from Calamus brachy- somus in the Galapagos Islands; P. crassigu- lum from Calamus and other hosts is known from Tortugas, Florida. Pseudopecoelus barkeri Hanson, 1950 Host. — Holocentrus ascensionis (Osbeck), squirrelfish. Location. — Pyloric cecum. Locality. — Lerner Fish Pens, Bimini, B.W.I, [new locality record]; formerly known from the same host at Bermuda. Hamacreadium mutabile Linton, 1910 Hosts.. — Epinephelus striatus (Bloch), Nassau grouper [new host record}; Haeniu- lon sciurus (Shaw), bluestriped grunt [new host record}; Lutjanus synagris (Linn.), lane snapper; Petrometopon cruentatus (Lace- pede), graysby [new host record]. Location. — Pyloric ceca of E. striatus, H. sciurus and P. cruentatus, and pyloric ceca (immature) and whole length of intestine except rectum in L. synagris. Locality. — £. striatus from Lerner Fish Pens, H. sciurus from N. shore N. Bimini; L. synagris, N. shore N. Bimini; and P. cru- entatus from "Bimini vicinity", B.W.I. Discussion. — I have studied the genital pore position of 23 specimens of Al. muta- bile from Lutjanus griseus from Tortugas ( Manter Collection ) and Lutjanus synagris from Bimini and find that 56.6 percent pos- sess a sinistral genital pore and the rest, 43. 5 percent, possess a median genital pore. Nagaty ( 1941 ) studied 34 specimens of this species and found a sinistral genital pore in 21, median genital pore 9, while 4 possessed a dextral genital pore. This widely distributed trematode is found in many species of fishes. Opecoelus sphaericus Ozaki, 1925 (Figure 51) Host. — Scorpaena mystes Jordan & Starks, lapon [new host record]. Location. — Intestine. Locality. — Taboga Island, Panama Pacific [new locality record}. Discussion. — The one specimen collected was cut halfway through the body at a point posterior to the acetabular stalk, between the ovary and anterior testis. The cut may have been made accidentally in opening the host intestine for examination. However, the tissue around the cut appeared regener- ated in part and the accident may have oc- curred under natural conditions. The speci- men differs from Ozaki's (1925) original description of O. sphaericus in having a smooth ovary; in (9. sphaericus the ovary is slightly trilobed. The ovary could be viewed only from the side in my specimen; perhaps it would appear trilobed were it viewed from the ventral side. Other small differ- ences such as vitelline distribution were con- sidered negligible. Pseudopecoeloides e que si Manter, 1947 (Figure 52) Host. — Stellifer sp. [new host record}. Location.. — Intestine. Locality. — Bella Vista, Panama City, Pan- ama. Discussion. — The paratype specimens of P. equesi show the posterior testis to be larger than the anterior testis but this differ- ence is not considered important enough to warrant two species although it may repre- sent geographical variation. P. equesi has been reported heretofore only from Eques lanceolatus and Eques acuminatus at Tortu- gas, Florida. It is another example of a species occurring in related hosts in the two oceans. K. Family CRYPTOGONIMIDAE dure a, 1933 Par aery ptogonimus americanus Manter 1940 ( Figures 54-56) Host. — Lutjanus novemjasciatus (Gill), pargo. Location. — Pyloric ceca. Locality. — Taboga Island, Gulf of Panama. Discussion. — This species was reported from Lutjanus novemjasciatus "or L. cyanop- No. 3 Sogandares-Bernal: Ttrernatodes of Marine Fishes 99 Figures 53-62. 53. Nagmia pacifica (Caballero), ventral view. 54. Paraci-yptogonimus americanus Manter, ventral view. 55. P. americamis, ventral view, oi'al sucker in normal position. 56. P. americanus, ventral view, oral sucker expanded. 57. Mefadena crassu- lata Linton, ventral view. 58. Gonacanihella lutjanl, gen. nov., sp. nov., ventral view. 59. Sterrhurns tahoganas, sp. nov., ventral view. 60. S. taboganns, terminal genital structures. 61. S. taboganns, sagittal section through preoral lobe showing musculature. 62. S. taboganns, sagittal section showing musculature of preacetabular pit. 100 Tidane Studies in Zoology Vol. 7 terus" from Mexico and Panama, by Manter (1940). The original description of P. americanus stated only that the acetabulum is usually about 2 '3 width of oral sucker. Twelve specimens of P. americanus in my collection had a sucker ratio from 1:0.54 to 1.0. The gland cells of the forebody of P. americanui were studied as follows: specimens fixed in A.F.A. and preserved in 70*^ ethanol were sectioned (10 microns) and stained with 0.05% Toluidin-Blue in citric acid ( pH 3.5) buffer. The sections were observed in water mounts and later dehydrated and mounted in balsam. The Toluidin-Blue staining indicated that these gland cells are probably not mucus producing. Since the glands are near the pharynx, they may pro- duce some enzyme. Body shape of P. americanus is extremely variable; specimens vary from elongate to almost completely round in outline. Figures 55 and 56 show that the anterior end of the body may be retracted and the oral sucker may be flared out until the oral spines appear to be on the body surface rather than on the oral sucker. Careful fo- cusing revealed that the spines were actually on the oral sucker. Once the spines are at- tached to the host tissue, the body surface which surrounds the spines may act as an additional retainer in keeping the host tis- sues attached to the spines. This arrange- ment would facilitate protrusion of the pharynx through the sucker, the anterior gland cells of the pharynx possibly having a lytic action on the host tissue. Metadena crassulata Linton, 1910 (Figure 57) Host. — Lutjanus analis (Cuv. & Val.), muttonfish. Location. — 1/4 intestine. Locality.. — Lerner fish pens, N. Bimini, B.W.I, [new locality record}. Metadena globosa (Linton, 1910) Price, 1940 Host. — Ocyurus chrysurus (Bloch), yel- lowtail [new host record}. Location. — Pyloric ceca and 1/4 intestine. Locality. — Close to Cat Cay, B.W.L [new locality record}. Siphodera vinaledwardsi (Linton, 1899) Linton, 1910 Host. — Lutjanus synagris (Linn.), spot snapper [new host record}. Location. — Pyloric ceca and mainly in in- testine excepting rectum. Locality. — Lerner fish pens and N. shore, N. Bimini, B.W.L [new locality record}. Gonacanthella lut jani, gen. nov., sp. nov.^' (Figure 58) Host. — Lutjanus jordani{?) Gilbert, pargo hilguero. Location.. — Pyloric ceca. Locality. — Taboga Island, Gulf of Panama. Holotype.— U.S.N. U. Helm. Coll. No. 38873. Diagnosis (based on two specimens, meas- urements on one favorable specimen ) . — Body 0.630 long by 0.429 wide; ellipsoidal in outline; spined to posterior end; with numerous gland cells in parenchyma. Oral sucker 0.074 long by 0.040 wide; terminal and retractile. Acetabulum 0.030 long by 0.040 wide, in anterior 1/3 body, recessed into body. Prepharynx absent. Pharynx 0.035 long by 0.043 wide. Esophagus as long as pharynx. Cecal bifurcation immedi- ately preacetabular. Ceca extending to pos- terior 3 4 body; ending blindly. Gonads im- mediately postequatorial, side by side and intercecal. Testes globular, separated from each other by ovary and seminal receptacle; dextral testis 0.107 long by 0.107 wide; sinistral testis 0.121 long by 0.114 wide. Gonotyl immediately preacetabular; protrud- ing from body; bulbular, 0.067 long by 0.{)67 wide, provided with spines 16 microns long by 3 microns wide which surround gonotyl in three rows. Seminal vesicle forming a sigmoidal-like curve between ovary and gonotyl, extending anteriorly to pharynx, then making a sharp turn pos- teriorly to insert dorsally into gonotyl. Ovary 0.072 long by 0.096 wide; globular; intertesticular, slightly overlapping both tes- tes ventrally. Seminal receptacle pyriform, bent sharply at its narrow anterior end, half its posterior length between testes, over- lapping anterior 1/5 ovary dorsally. Vitel- laria of large follicles dorsal to ceca, extend- 1" The name Gonacanthella indicates the spines on the gonotyl; and lut jani indicates the host genus Li(tj(tnus. No. 3 Sogandares-Bernal: Ttrematodes of Marine Fishes 101 ing across body between gonotyl and testes. Uterus filling most of hindbody to aceta- bulum in one specimen and to posterior level of pharynx in another. Eggs 16 to 23 by 11 to 15 microns. Excretory vesicle ob- scured by numerous eggs in uterus. Generic diagnosis of Gonacantbella. — Cryptogonimidae; body elliptical in outline, spined to posterior end. Forebody with nu- merous gland cells in parenchyma. Oral sucker terminal and retractile. Acetabulum in anterior 1/3 body, recessed into body. Pharynx spherical. Esophagus present. Ceca extending slightly posterior to testes. Gon- ads postequatorial, intercecal, side by side. Testes globular, separated from each other by ovary. Gonotyl immediately preacetabu- lar, protrusible, globular in shape, spined. Seminal vesicle extending from between ovary and gonotyl to region of pharynx, bending sharply backward to insert into gonotyl dorsally. Ovary round, between tes- tes. Vitellaria follicular, extending across the body between gonotyl and testes. Semi- nal receptacle median, between testes. Uter- us filling most of hindbody. Eggs opercu- lated, yellow, thin-shelled. Excretory vesicle not observed. Type species: Gonacantbella lutjani. Discussion. — The family Cryptogonimidae Ciurea, 1933 contains at least 21 genera. Gonacantbella is most closely related to Metadena Linton, 1910, differing only by possessing a spined gonotyl as compared with spines lacking on gonotyl; and smooth instead of follicular or deeply lobed ovary. The genus Proneocbasmus Szidat, 1954 is actually a synonym of Paraspina Pearse, 1920, agreeing in all important details. Both genera are known only from pimelodid cat- fishes of South America. Proneocbasmus argentinensis Szidat, 1954 becomes Para- spina argentinensis (Szidat, 1954) new comb. Paraspina bagre Pearse, 1920, type and only other species, differs from Para- spina argentinensis in egg size (l6 by 18 microns as compared with 30 by 13 mi- crons) and in possessing 20 instead of 26 peribuccal spines. L. Family FELLODISTOMATIDAE Nicoll, 1933 Proctoeces subtenue (Linton, 1907) Hanson, 1950 Host. — Calamus bajonado (Bloch & Schnieder), jolt-head porgy. Location. — Intestine. Locality. — Lerner fish pens, N. Bimini, B.W.I, [new locality record}. Discussion. — P. subtenue was reported from Calamus at Bermuda by Linton (1907) and Hanson (1950) and from Calamus spp. at Tortugas by Manter ( 1947 ) under the name of P. erythraeus Odhner, 1911. Odhner described his material from ]ulis lunaris and Cbrysopbrys bifasciatus in the Red Sea. Hanson considered P. erytbraeus a synonym of P. subtenue. which is found almost ex- clusively in the family Sparidae from Ber- muda, Tortugas, Fla., Bimini, B.W.I, and the Red Sea. Julis lunaris is a Labrid fish. P. subtenue is also reported from New Zea- land (Manter, 1954) from Latridopsis cili- aris, family Latridae. Antorcbis urna (Linton, 1910) Linton, 1911 Host. — Pomacantbus aureus ( Bloch ) , black angelfish. Location. — Pyloric ceca and entire length of intestine but mainly in pyloric ceca and 1/4 intestine. Locality.. — Lerner fish pens and N. shore N. Bimini, B.W.I. [new locality record}; known also at Tortugas. Bacciger barengulae Yamaguri, 1938 Host. — Harengula clupeola (Cuvier), big- eyed sardine. Location. — Pyloric ceca. Locality. — Off Lerner Laboratory pier, N. Bimini, B.W.I, [new locality record}; other localities are Tortugas, Florida, and Japan. Discussion. — There were usually two B. barengulae per cecum. The worms were usually located in the middle of the cecum and were separated from each other by at least one body length. The worms were easily observed in situ as the host's ceca were almost transparent. Tergestiapectinata (Linton, 1905) Manter, 1940 Hosts. — Caranx crysos (Mitchill), hard- tailed jack; Caranx latus (Agassiz), horse- eyed jack; Caranx ruber (Bloch), skip jack; Selar crumenopthalmus (Bloch), goggle eyed scad. Location. — Rectum of C. crysos; mid- intestine of C. ruber and S. crumenoptbal- mus: and 1/4 and mid-intestine of C. latus. 102 Tulane Studies in Zoology Vol. 7 Locality. — C. crysos. from N. shore, N. Bimini; C. latus from Lerner fish pens, N. Bimini; C. ruber from W. side of N. Bimini; and 5". crumenopthalmus from off Lerner Laboratory pier, N. Bimini, B.W.L [new locaUty record}; known also from Woods Hole, Massachusetts; Beaufort, North Caro- lina; and Tortugas. M. Vamily MONORCHIIDAE Odhner, 1911 Genolopa ampullacea Linton 1910 Hosts. — Haemulon album Cuv. & Val., margate-fish; Haemulon parra (Desmarest), sailor's choice [new most record}; Haemulon plumieri (Lacepedej, white grunt; Haemu- lon sciurus ( Shaw ) , blue-striped grunt. Location. — Pyloric ceca. Locality. — H. album and H. sciurus from Lerner fish pens, N. Bimini; H. sciurus from N. shore, N. Bimini; H. parra and H. plu- mieri from 12 mi. S. of S. Bimini. Proctotrema truncatum (Linton, 1910) Manter, 1940 Hosts.. — Caranx ruber (Bloch), skip-jack [new host record} ; Haemulon parra ( Des- marest) , sailor's choice; Haemulon plumieri (Lacepede), white grunt; Haemulon sciurus ( Shaw ) , blue-striped grunt. Location. — Pyloric ceca. Locality. — C. ruber from W. shore of N. Bimini; H. parra and H. plum-ieri from 1/2 mi. S. of S. Bimini; and H. sciurus from Lerner fish pens and N. shore, N. Bimini, B.W.L [new locality records}. N. Family ZOOGONIDAE Odhner. 1911 Diplangus paxillus Linton, 1910 Hosts. — Haemulon album Cuv. & Val., margatefish; Haemulon parra (Desmarest), sailor's choice; Haemulon plumieri (Lace- pede), white grunt; Haemulon sciurus (Shaw), blue-striped grunt. Location. — Predominantly in 2/3 intes- tine but sometimes ( 2 cases ) in pyloric ceca and then always both in ceca and in 2 3 intestine. Locality. — Lerner Fish Pens, Bimini, B.W.I. Steganoderma atherinae (Price, 1934) Yamaguti, 1953 Host. — Hepsetia stipes ( Muller & Tros- chel), hardhead, silverside [new host rec- ord}. Location. — 1/4 intestine and mid-intestine (1 specimen in 1 host specimen). Locality. — Opposite Lerner Laboratory Pier, Bimini, B.W.L Discussion. — The Bimini report is the northernmost record for S. atherinae which was described from the Dominican Repub- lic. Manter (1947) examined 66 specimens of Hepsetia stipes in Tortugas, Florida, but did not find S. atherinae. O. Family BIVESICULIDAE Yamaguti, 1939 Bivesicula hepsetiae Manter, 1947 Host. — Hepsetia stipes (Muller & Tro- schler), silverside. Location. — Mid-intestine. Locality. — Off Lerner Laboratory Pier, Bimini, B.W.L P. Family GORGODERIDAE Looss, 1901 Nagmiapacifica (Caballero, 1945) Markell, 1953 Host. — Car char hinus natator Meek & Hil- debrand, black-tipped shark [new host rec- ord}. Location. — Coelom and intestinal mesen- teries. Locality. — Opposite mouth of Chiman River, near Pelado Island; Cocos Point, Isla del Rey and Playa Grande, San Jose Island, Archipielago de las Perlas; and Pihas Bay; all localities in the Gulf of Panama. Discussion. — Caballero (1945) described Petalodistomum pacificum from 20 speci- mens taken from the coelom of an unidenti- fied shark from the coast of Manzanillo, Mexico. Markell (1953) transferred P. pacificu7)i to the genus Nagmia and the name became Nagmia pacifica (Caballero, 1945), Markell, 1953. Markell differenti- ated Nagfuia from Petalodistomum John- ston, 1934 on the basis of whether or not there is branching of the efferent ducts of the testes such as is found in Probolitrema Looss, 1902 and in Nagmia Nagaty, 1930, but not in Petalodistomum. (Dollfus) 1937, Johnston (1934), Caballero (1945) and Caballero, et. al. ( 1956) considered Nagmia a synonym of Petalodistomum. When more species are known, the possible generic value of the vasa efferentia can be judged better. N. pacifica is highly host-specific. In one night's collection several species of sharks over five feet in length were exam- No. 3 Sogandares-Bernal: Ttrematodes of Marine Fishes 103 ined for this parasite. Only Carcharhinus natator harbored N. pacijica. Other similar collections in other parts of the Gulf of Panama and south west of Pinas Bay did not yield N. pacijica from any shark-like fish other than Carcharhinus natator. I have also studied three specimens of Nagmia pacijica collected by Miss Margarita Bravo Hollis at Punta Mita, Puerto Vallarta, Ja- lisco, Mexico, from an unidentified shark. Caballero's (1945) specimens are 9.355 to 14.647 mm. long. My Panama specimens were 6 to 15 mm long and one of Miss Bravo's specimens was 16.5 mm long. A specimen 6 mm long had eggs in the uterus. Caballero recorded a sucker ratio of 1:1.25 to 1.86 although his figure shows a sucker ratio of 1:1.12. Sucker ratios of specimens from my collection are from 1:1.09 to 2.34 (average about 1:1.44). Eggs of N. pa- cijicuni were described as 59 to 63 by 36 to 38 microns. Eggs from specimens in my col- lection measured 54 to 67 by 27 to 40 mi- crons. Acetabulum position varies from an- terior 14 to 13 of body, agreeing with Caballero's description "hacia delante del ecuador del cuerpo, por delante del ovario." (His plate shows the acetabulum in the an- terior 1/4 body.) Recently Caballero, Bar- roeta, & Grocott (1956) reported N. pa- cijica from Carcharias in Panama. These authors apparently collected one specimen because they redescribed the species from one specimen and stated that the specimens from Mexico are larger in size than those from Panama and Alta California, U.S.A. As has been indicated above, N. pacijica appears to be host-specific to Carcharhinus natator in the Gulf of Panama, large num- bers always being found in this shark. The record of a specimen of N. pacijica from Carcharias sp. in the Gulf of Panama may represent an accidental infection in this host or a host misidentification. Q. Family HEMIURIDAE Lilhe, 1901 Ectenurus virgulus Linton, 1910 Hosts. — Harengula clupeola (Cuvier), big- eyed sardine; Selar crumenopthalmus (Bloch), goggle-eyed scad. Location. — Stomach. Locality. — Off Lerner Laboratory pier, N. Bimini, B.W.l. [new locality record}. Brachadena pyrijormis Linton, 1910 Hosts. — Haemulon album Cuv. and Val., margate fish; Haemulon plumieri (Lace- pede ) , white grunt; Haemulon sciurus (Shaw), blue-striped grunt; Haemulon parr a (Desmarest), sailor's choice; Malacanthus plumieri ( Bloch ) , sandf ish. Location. — Stomach. Locality. — H. album and H. sciurus from Lerner fish pens, N. Bimini; H. parra. H. plumieri, from 1/2 mi. S. of S. Bimini; and M. plumieri, from between Cat Cay and Bimini, B.W.L [new locality records}. Discussion. — This trematode is known from numerous hosts at Tortugas, Florida, and Beaufort, North Carolina. It also occurs at La Jolla, California. Leurodera decora Linton, 1910 Hosts. — Haemulon album Cuv. and Val., margate fish; Haemulon plumieri ( Lace- pede), margate fish; Haemulon sciurus (Shaw), blue-striped grunt; Haemulon parra ( Desmarest ) , sailor's choice. Location. — Stomach. Locality. — H.. sciurus and H. album from Lerner fish pens and N. shore N. Bimini; H. plumieri and H. parra, from 1/2 mi. S. of S. Bimini, B.W.l. [new locality record}; known from Tortugas. Hysterolecitha rosea Linton, 1910 Host. — Acanthtirus caeruleus Bloch and Schneider, blue tang. Location. — Stomach. Locality. — Off Lerner Laboratory pier, N. Bimini, B.W.I, [new locality record}; known from Tortugas. Dinurus longisinus Looss, 1907 Host. — Coryphaena hippurus Linn., dol- phin. Location. — Stomach. Locality. — Two hosts from 12 mi. N.W. San Jose Island, Archipielago de las Perlas, Panama Pacific; and 1 host from the Lerner fish pens, N. Bimini, B.W.L [new locality record}. Discussion. — D. longisinus is a common parasite of dolphins from the Panamanian Pacific. Manter (1947) reported D. longi- sinus from 2 of 6 hosts in Tortugas. Florida. It also occurs in Japan and in the Red Sea, and is probably almost world wide in dis- tribution. Dinurus tornatus (Rud., 1895) Looss, 1907 Host, — Coryphaena hippurus Linn., dol- phin. 104 Tulane Studies in Zoology Vol. 7 Location. — Stomach. Locality. — One host from Lerner fish pens, N. Bimini, B.W.I, [new locaUty rec- ord}. Discussion. — This species was not col- lected by me in Panama, but was found among specimens of D. longismus collected and reported by Manter (1940a). D. tor- natus may be separated from D. longisinus in that the cuticular annulations begin dor- sally about level with the posterior border of the acetabulum, and the hermaphroditic duct extends past posterior border of ace- tabulum. D. longisinus differs by having transverse annulations of the soma cuticula beginning dorsal to oral sucker and the hermaphroditic duct never extends posterior to the acetabulum. Approximately 200 speci- mens of D. longisinus from five hosts in Panama Bay indicate that these are constant characteristics of this species. D. tornatus has been reported from the Red Sea, Massa- chusetts, Tortugas, and the South Atlantic. Opisthadena dimidia Linton, 1910 Hosts. — Kyphosus elegans (Peters), chopa [new host record]; Kyphosus sectatrix ( Linn. ) , Bermuda chub. Location. — Stomach. Locality. — K.. elegans from North Re- stinga Beach, Taboga Island, Panama Pacific; and K. sectatrix from Bimini Vicinity, B.W.I, [new locality records}. Discussion. — 0. dimidia was reported from Kyphosus incisor (Cuv. & Val. ) and Kyphosus sectatrix (Linn.) from Tortugas, Florida by Manter ( 1947). The occurrence of O. dimidia from the Pacific Ocean is the first on record for this species. 0. dimidia may have been a parasite of Kyphosus spp. when the two oceans were continuous dur- ing the last continental gap. Tubulovesicula lindbergi (Layman, 1930) Yamaguti, 19.34 Hosts. — Unidentified eel and Synodus sp. Location. — Stomach. Locality. — Off mouth of Chiman River near Pelado Island, Panama Bay, Panama. Discussion. — Manter (1947) discussed at some length the status of the species of the genus Tuhulovesicula. He considered T. californica Park, 1936; T, pseudorhombi Yamaguti, 1938; and T. muraenasocis Ya- maguti, 1934 synonyms of T. spari Yama- guti, 1934. T. madurensis Nigrelli, 1940 was considered a synonym of T. lindbergi (Layman, 1930) Yamaguti, 1934. Manter later (1954) considered the following spe- cies as valid: T. lindbergi (Layman, 1930) Yamaguti, 1934; T. pinguis (Linton, 1940) Manter, 1947; T. angusticauda (NicoU, 1915); T. ?nagnacetabulu?n Yamaguti, 1939; T. anguillae Yamaguti, 1934, and T. spari Yamaguti, 1934. Nagaty (1956) described another species, T. serrani. A study of the Panama specimens leads me to conclude that not only are T. cali- fornica, T. pseudorhombi, and T. murae- nasocis synonyms of T. spari. but the latter in turn appears to be a synonym of 7\ lindbergi. as does T. anguillae. The only ap- parent difference between T. lindbergi and T. spari is egg size and this character is rather variable in Tubulovesicula. Manter (1954) found the only difference between T. spari and T. anguillae to be the length of the ecsoma. In my material from Pana- manian fishes, the ecsoma varied from shorter than body length to as much as three times the body length, probably de- pending upon contraction and flattening pressure during fixation. Since T. spari ap- pears to be a synonym of T. lindbergi the latter becomes the type species of the genus. T. serrani Nagaty, 1956, which was de- scribed from one specimen, also appears to be a synonym of T. lindbergi. I believe only four species of Tubulo- vesicula are valid: J .. lindbergi, T. angusti- cauda. T. pinguis and T. magnacetabuhwi. The species may be separated by the follow- ing key which is modified from Manter (1954): 1. Pars prostatica extending: pos- teriorly past anterior ^/s of acetabulum 2 Pars prostatica not extending: posterior to anterior edge of acetabulum T. angusticauda 2. Acetabulum 3 times width of oral sucker T. magnacetabuhun Acetabulum less than 3 times width of oral sucker — 3 3. Prostatic gland interrupted op- posite acetabulum T. pinguis Prostatic gland not interrupted opposite acetabulum T. lindbergi Varahemiurus merus (Linton, 1910) Woolcock, 1935 Host. — Harengula clupeola (Cuvier), big eyed sardine. Location. — Stomach. No. 3 Sogandares-Bernal: Ttrematodes of Marine Fishes 105 Locality. — Off Lerner Laboratory Pier, N. Bimini, B.W.I, [new locality record}; known from Tortugas, Florida and from coast of Ecuador. Lecithochirium microstomum Chandler, 1935 Host. — Sarda velox Meek & Hildebrand, bonito. Location.. — Stomach. Locality — Morro Sentinela, Pinas Bay, Panama Pacific. St err hums taboganus, sp. nov.^^ (Figures 59-62 ) Host. — Gymnothorax dovii (Gunther), morena. Location. — Stomach. Locality. — Taboga, Panama Pacific. Ho/o/}'/?^.— U.S.N.M. Helm. Coll. No. 38874. Diagnosis. — Sterrhurus (measurements on six favorable specimens); body 3-610 to 4.360 long by 1.159 to 1.387 wide; maxi- mum width between acetabulum and oral sucker. Cuticle smooth. Forebody 1.045 to 1.425 long; with a deep concavity of which the posterior rim is provided with a heavy bundle of circular muscle fibers ( fig. 62 ) ; lateral borders of concavity usually extend anteriorly from posterior rim, parallel with edges of acetabulum for about 12 to 3/4 distance between acetabulum and oral sucker. Hindbody 1,520 to 2.128 long. Ecsoma usu- ally retracted but may be extended a dis- tance of 0.133. Preoral lobe present. Oral sucker 0.266 to 0.361 long by 0.361 to 0.418 wide; subterminal; lacking any type of projection into its lumen. Acetabulum 0.655 to 0.912 long by 0.665 to 0.703 wide; preequatorial or equatorial; aperture ovoid, slightly transverse or longitudinal. Sucker ratio 1:1.68 to 1.95. Prepharynx absent. Pharynx 0.114 to 0.152 long by 0.095 to 0.133 wide. Esophagus very short or absent. Ceca first transverse from pharynx then bending sharply towards posterior end of body at pharyngeal level, continuing a sinu- ous course on each side of body ending at the base of the retracted ecsoma. Testes 0.152 to 0.266 long by 0.171 to 0.285 wide, symmetrical one on each side of body im- mediately posterior to acetabulum, over- lapping ceca or not; never extracecal. Geni- i** The name taboganus is for the type lo- cality. tal pore median, opposite posterior half of pharynx, followed by a genital atrium and a short muscular hermaphroditic duct which is surrounded by prostatic cells enclosed in a sinus sac. Prostatic vesicle connects with hermaphroditic duct and is also completely enclosed in the sinus sac. Metraterm enters the sinus sac and joins the male duct im- mediately anterior to prostatic vesicle. Pro- static cells surround the bipartite, external seminal vesicle near its junction with the sinus sac. Seminal vesicle extends posterior- ly to overlap anterior half of acetabulum. Ovary 0.152 to 0.247 long by 0.152 to 0.285 wide, median or sinistral sometimes overlapping left cecum. Uterus intercecal or slightly overlapping ceca, mostly between testes and ecsoma, descending posterior to ovary and ascending to join hermaphroditic duct, not entering ecsoma. Vitelline glands lobed, lobes as long as wide, immediately posterior or slightly overlapping ovary, me- dian or sinistral, left gland with two to four digitform lobes, right gland with one to three lobes. Eggs collapsed, lacking pro- jections of any kind, thin-shelled, 16 to 22 by 8 to 10 microns. Discussion. — Numerous species have been named, added and deleted in the genus Ster- rhurus Looss, 1907. Sterrhurus taboganus differs from all other species by possessing a large bundle of semi-circular muscle fibers along the pos- terior rim of the concavity (fig. 62). Such muscular fibers of the concavity have not been described for any other species of Sterrhur/is. A forebody concavity has been described or figured for at least S. flori- densis, S. grandiporus. S. imocavus. S. mag- nus. and S. musculus. In most of these species the concavity seems to be formed by con- traction of the forebody and thus may be temporary. The concavity of S. taboganus appears to be permanent. Of approximately 26 specimens of S.. taboganus examined, all showed this concavity clearly. In S. tabo- ganus there are also some longitudinal mus- cular fibers which connect the base of the concavity with the acetabulum (fig. 62). These muscle fibers apparently are for further deepening the concavity. These two sets of muscles would allow the concavity to act as an additional holdfast organ. In fact, when the stomach of the host was opened and laid flat for examination, most of the 106 Tulane Studies in Zoology Vol. 7 trematode specimens exhibited an angular posture of about from 15 to 30 degrees with the plane of the intestine. S. taboganus appears to be most similar to S. musculus. S. mitsculus differs from S. taboganus as follows: Body size 1.0 to 1.5 long as compared with 3.572 to 4.370 long in S. taboganus. In S. musculus the body is more rounded posteriorly, the maximum width being at midbody or behind the ace- tabulum as compared with forebody widest in S. taboganus. The wall of the herma- phroditic duct is cuticular in S. musculus and muscular in 5'. taboganus. Looss (1907) named the genus Sterrhurus for the species S. musculus ( type species ) ; S. grandiporus ( Rudolphi, 1819) Looss, 1907, S. imocavus Looss, 1907; and S. fusi- formis (Liihe, 1901) Looss, 1907. Since that time, there has been much confusion between Lecithochiriu7n Liihe, 1906, Ster- rhurus Looss, 1907, and related genera. The Russian parasitologists (Skrjabin and Guschanskaja, 1955) proposed entirely new criteria for genera of this group of hemiur- ids. They believe that details of the "herma- phroditic bursa" {i.e.. sinus sac) and ducts within it should be the chief basis of genera. Many transfers of species were made. Pending appraisal of such new criteria by other parasitologists, earlier concepts of these genera are followed here. Sterrhurus floridensis Manter, 1934 Hosts. — Abudefduf saxatilis (Linn.), ser- geant-major [new host record {; Conger con- ger (Linn.), conger eel [new host record}; Holocentrus ascensionis (Osbeck), squirrel fish; Labrisomus nuchipinnis Quoy & Gai- mard, hairy blenny [new host record]; Malacanthus plumieri ( Bloch ) , sand eel [new host record]; Ocyurus chrysurus (Bloch), yellowtail; Ogcocephalus vesper- tilio (Linn.), batfish [new host record]; Harengula clupeola (Cuvier), big-eyed sar- dine [new host record]; Scorpaena plu- mieri Bloch, scorpion fish. Location. — Stomach of all hosts. Locality. — All hosts are from near N. Bimini, (C. conger from 120 fathoms, W. side of N. Bimini), except for two speci- mens of each H. ascensionis. M. plumieri and one specimen of O. chrysurus from near Cat Cay, B.W.I. Discussion. — Manter (1934, 1947) noted the pronounced lack of host specificity of S. floridensis. Sparks (1957) reported S. floridensis from several fishes of the Nassau vicinity, B.W.I. Hirudinella clarata (Menzies, 1791) Blainville, 1828 Host. — Sarda velox Meek & Hildebrand, bonito [new host record]. Location. — Stomach. Locality. — Morro Sentinela, Pinas Bay, Panama Pacific. Hirudinella marina (Garcin, 1730) Host. — Acanthocybium solandri (Cuv. & Val. ), wahoo. Location. — Stomach. Locality. — 15 miles S.E. Taboguilla Island, Gulf of Panama. R. Family PROSOGONOTREMATIDAE Vigueras, 1940 Prosogonotrema bilabiatum Perez Vigueras, 1940 Host. — Ocyurus chrysurus (Bloch), yel- lowtail. Location. — Stomach. Locality. — Near Cat Cay, B.W.I. [new lo- cality record]. Discussion. — Manter (1947) examined 47 O. chrysurus from Tortugas, Florida, but did not find P. bilabiatum. This species has been reported only from 0. chrysurus in Cuba by Perez Vigueras (1940). The fam- ily Prosogonotrematidae is of doubtful va- lidity. At most it probably deserves sub- family status within the Hemiuridae. S. Family ACCACOELIDAE Looss. 1912 Tetrochetus coryphaenae Yamaguti, 1934 Host. — Coryphaena hippurus Linn., dol- phin. Location. — Rectum. Locality. — Lerner Fish Pens, Bimini, B.W.I., and from 12 mi. N.W. San Jose Island, Archipielago de las Perlas, Panama Pacific [new locality records]. Discussion. — This species was described from Coryphaena hippurus in Japan by Yamaguti ( 1934 ) and reported from that host in Tortugas, Florida by Manter ( 1947 ). Coryphaena hippurus is widely distributed and it is not surprising that its parasites are well established in at least Japan, the tropi- cal American Pacific, the Gulf of Mexico, and neighboring areas. No. 3 Sogandares-Bernal: Ttrematodes of Marine Fishes 107 III. Measurements of pH of the Stomach and Intestine of Fish Hosts The ecology of adult helminths of fish has been little studied. My studies at Bimini included some effort to observe certain de- tails of host-parasite relationships. Some of these details are included in the preceding papers. For example, each fish was exam- ined separately, and the exact location and numbers of trematodes present were noted, as well as the number of uninfected hosts. Most species of trematodes occur in a fairly definite region of the digestive tract. If two or more species occur simultaneously, they usually, but not always, live in separated portions of the tract. Since trematodes are usually at least part- ly immersed by intestinal fluids, the writer thought their different locations may be due to differences in pH of the intestinal con- tent. pH measurements were made with a Beckman pH meter after stunning the fish. The entire operation took less than three minutes after some practice. Measurements were made at the following points: stomach, pyloric ceca, plyoric junction of the stomach and intestine, 1/3 intestine, 1/2 intestine, 3/4 intestine, 4/4 intestine (middle of rectum ) . Most complete measurements were made on 10 specimens of Haemulon sciiiriis { Table 1 ) . A surprising variation in pH occurs from time to time in the same organ. The range in the stomach was 3.0 to 7.7. Hemiurid trematodes occurred in these stomachs, hence are able to withstand such a range. Stomachs containing food were acid (3-0 to 6.9); those of specimens starved for five days were alkaline (7.3 to 7.7). The pH of the pyloric ceca varied from 5.0 to 7.8, a range endured by living trema- todes there. The pH of the intestine varied from 5.3 to 8.2, the more acid readings being from specimens which had recently fed. In all four fishes which had not fed for five days, the intestine was alkaline. The content of the intestine seems to affect the pH. Large bolli of crustacean remains, probably containing some gastric fluid, re- sulted in acidic spots in the intestine. While the acid conditions in the intestine are probably localized and temporary, the parasites are exposed to them at least briefly. There is a large probably rather rapid shift in pH along the intestine, depending chiefly on recency of feeding and type of food. The environment of an intestinal parasite is not as monotonous as might be assumed. Read (1950) working with warm-blooded verte- brates stated: "considerable evidence has accumulated to show that there is a certain constancy of pH in various sections of the small intestine." My measurements of the pH of the intestine of fishes indicates it is less constant than that of warm-blooded vertebrates. In fact, trematodes of the di- gestive tract of marine fishes probably ex- perience a greater variation in pH than do free-living marine animals. Sea water is naturally buffered and offers a more stable pH environment except perhaps very locally. Even if the substrate pH of a free living marine animal shifts periodically during the day, this shift in pH would not be as rapid as that caused by a large crustacean boUus passing through the intestine of a marine fish. Table 1. ipH measurements of 10 Haemulon sciurus. ( — =no measurement possible; + = lire trem- todes were present; *= host starved for 3 days). Site of pH Reading pH Specimen No. 1 2 3 4* 5* 6* 7* 8 9 10 Stomach 3.0 + 6.7 + 6.7 7.3 + 7.7 + 7.5 + 7.5 + 5.1 + 4.6 + 6.9 Pyloric Ceca 6.5 + 7.0 + 6.9 + 7.3 7.2 + 7.8 + 7.4 + 7.8 + 7.2 + 5.0 + Pyloric Jet. 6.7 7.2 7.4 7.2 7.5 7.6 7.6 8.1 6.8 7.5 1/3 Intestine 7.2 + 7.8 + 7.4 7.7 + 7.9 + 7.9 + 7.0 8.0 + 6.3 8.2 1/2 Intestine 7.2 + 7.6 + 8.2 8.2 7.9 7.9 8.2 8.2 7.2 8.0 3/4 Intestine — + + 8.0 7.6 7.7 7.9 8.4 7.3 7.9 4/4 Intestine 6.9 7.6 7.8 7.5 7.5 7.6 7.2 7.7 7.4 7.5 108 Tulane Studies in Zoology Vol. 7 IV. Exact Locations of Digenea IN Their Hosts Most collectors of parasites indicate only the organ infected, not the exact location in the organ. Laboratory facilities at Bimini usually permitted observation of the exact location of the parasites. The location with- in the intestine has been indicated in the descriptions above. A particular species of trematode is al- most always found only in a limited area of the intestine. Exceptions are usually either immature individuals or cases of un- usually intense infection. Other exceptions may be due to movement after death of the host. Under natural conditions trematodes appear to have a rather localized location along the intestine. Consequently, several species may occur in a host, each occupying its own ecological site which may not ap- preciably overlap that of other species with- in the same intestine. On the other hand, specimens of as many as three different genera may occupy the same pyloric cecum; specimens of the genera Aiouorchis. Procio- rema, and Genolopa (all Monorchiidae) have been observed in a single cecum of Haemulon plumieri. Specimens of Procto- trenia and Genolopa observed in Bimini were practically in contact with one another; the tip of a needle could not be inserted between them. Table 2 shows the localization in the di- Table 2. The location of three species of Apocrea- dium, within the digestive tract of Batistes capriscus and B. vetula. Trematode Species Location of Trematodes in Digestive Tract Host Species A. ba list is 1/3 intestine B. capriscus A . coin Rectum B. capriscus B. vetula A. uroprocto- 2/3 intestine B. vetula feyiim gestive tract of three species of Apocrea- dium in species of Balistes. A. balistis and A. coili may occur widely separated in a single B. capriscus; or A. coili may occur with A. uroproctoferm in B. vetula. Even in the cases of single infections, the species of Apocreaclium live in a particular part of the intestine. No triple infections were found but these are theoretically possible without overlapping. Ecological isolation, so common among free living animals, is probably the rule among parasites. Speciation would be fav- ored by this occupation of different parts of the intestine by different groups of tre- matodes. Stunkard (1957) suggested that the type of genetic origin of species occur- ring in bisexual animals may not apply to trematodes because of their hermaphrodit- ism. However, cross fertilization is known to occur in some species of Digenea and may be more-or-less common. Area isola- tion within the intestine of a species of host might operate much as isolation of free- living animals by a valley, river, or mountain range. In the case of related species or genera found intermingled in the same spot, two conditions are possible. Most likely the spe- cies would not cross-fertilize one another because of differences in the reproductive organs. For example, the morphology and spination of the cirrus and metraterm of Monorchis, Proctotrema and Genolopa found together in Haemulon could prevent copulation either mechanically or act as a recognition character between genera. The other possibility is that two very similar species might cross and result in hybrids. Price (1953) suspected that hybridization between Fasciola hepatica and F. gigantica eixsted in cattle in the southern United States. Very heavy infestations involving closely related species would favor such hy- bridization. V. Geographical Distribution The fauna of Bimini is typical of the Caribbean and West Indian region. The fishes and invertebrates generally are the same as occur at Tortugas, Cuba, and the West Indies. Much of this fauna extends to Bermuda. The trematodes of these re- gions, as expected, are also similar although they are distinct from the northern Atlantic coast of America and Europe. Of the 62 species collected at Bimini, 48 (about 80 percent) are also known from Tortugas, Florida; 34.5 percent are known from Bermuda. Twelve of these species (about 20 percent) were also collected by me in the Gulf of Panama. In addition six species {Ha??iacreadium mutabile, Heli- No. 3 Sogandares-Bernal: Ttrematodes of Marine Fishes 109 Table 3. Parasite-host list, showing trematodes from the Gulf of Panama and Bimini, and their hosts. Hosts and trematodes marked (P) are from Panama, ( B) are from Bim,ini. FAM I LY ACANTIKK -OLIMDAE Hteplidiwstoniinii coriiphdenae Manter (B) t'oriii)h(unii liiiiiiinux ( B, P) 8tei)li i /, ( P ) Metadeiia crass uhttd Linton (B) Lutjaiius aiialis, (B) Metadeiia globosa (Linton) (B) Ocpurus chri/surus, (B) Paracriiptoi/oiiiiiius diiiericdiius Manter (P) Lutjaiius iioreiiifdsciatiis, (P) Siphodera riualedirdrdsi (Linton) (B) Lutjaiius sijnayris, (B) FAMILY FELLODISTOMATIDAE Antorchis iirna (Linton) (B) Poiiiacaiithiis aureus, (B) Bucciiier liarciifiulac Yamaguti (B) Harenijula ciupeula, (B) Proctoeces subtciiue (Linton) (B) Calamus bujoiiado, (B) Tergcstia pectinata (Linton) (B) Caraiix crysos, (B) Caranx latus, (B) Carunx ruber, (B) tielar crumcnoptlialiiius, (B) FAMILY GOIKiODEKIDAE JSIagiiiia pacifica (Caballero) (P) Carcfiarhinus natator, (P) FAMILY HAPLOSPLANCHIDAE Hapldsplanchnus acutus (Linton) (B, P Thiirinops pachi/lepis, (P) Tylosiirus rhaphidoma, (B) FAMILY IIEMIURIDAE Brachddi na piiriforiiiis Linton (B) Haeiiiuloii nUium, (B) Haeiiiiiloii parra, (B» Haemiilon pluinicri, (B) Haciii uiijii yciurus, (B) MuUii-diiUius iiliiiiiieri, (B) Diiiurus loiii/iKiiius Looss (B. P) Coriiplidi lid liippurus (15, P) Diiiuriix toriidtus (Rurtoliihi) (B, P) Coriiphdciid hipjiurus, (B, P) Ectenurus vin/ulus Linton (B) Hdrciifiula clupeoUi, (B) Selar cruiucuoptlialmus, (B) Hirudincllii clarata (Menzies) (P) t'Sardd relox, (P) HirudincUd iiidriiid (Bhiinville) (P) Acaiitliiiciihiu III Niilinidru lu , (P) Hystcriilccithd rosea Linton (B) AcdiitJiiirus cdcrulcus, (B) Lecitliochiriuiii iiiicrostoiiiuiii Chandler (P) Sarda velox, (P) Leurodcra decora Linton (B) Haeinuloii iilbuiii, (B) Haeiiiulnn purrd, (B( Harm iiloii phiiiiivri, (B) Haemulon sciurus, (B) Opisthadi lid diiiiidid Linton (B. P) Kyphnsus elci/diis, (P) KpiihiiKUK sri-tdtrix. (B) PardJiciiiiurus lueriis (Linton) (B) Hun 111,1 iild rlupcolii, (P.) Sterrhurus floridciixis Manter (B) Abiidrfduf saxatilis, (P.) Coiif/cr roiifier. (I?) Holoieiitrus dsccnsionis, (B) Ldbrisdiii u'< inicliipiiinis. (B) Mdldcdiithiis pJuiiiieri, (B) Ocyuriis i-lirysiirus, (B) Of/eocrplial us respcrtilio, (B) Harcniiula ilupeula. (B) Scorpaciia pluiuicri, (B) Stcrrtiurus tdboiiaii us (this paper) (P) (linmiothordx dorii, (I'l Tubulori'siculd liiidberyi (Layman) (P) Eel, unidentified, (P) Synodus sp., (P) FAMILY LEPOCREADIIDAE Apocreddiuiii balistis Manter (B) Balistrs rdjirisriis, (B) Apocrcadiuiii liniroi (this paper) (P) Bulistcs Hdufriifiium, (I') Apocrcadiuiii to(7i (this paper) (B) Balistes capriscus, (B) Balistes vetula, (B) Apocreadium linifiisinosuiii Manter (P) tSphoeroides aiiuiildtus, (I') Apocreadium aiii/ustum (this paper) (B) LactopJiri/s triipnnis. (P.) Apocreadium uroprortoferuin (this paper) (B) Balistes retuld, (Bt Cdbleid trifjoiii (this ijaper) (B) Lactoplirys triyonus, (B) Crassicutis marina Manter (B) Gerres cinereus, (B) Dermadena lactophrysi Manter (B) Lactophrys tricornis, (P.) Lactophrys trigonus. (B) Diploproctodaeum yVu-itinii (Linton) (P) hiodon liystri.r, ( I'l Syhoemides diniiildtus. (1') Dililoproctoddcum lidiistrum (McCallum) (B) Ccratacanthus scripta, (B) Encntcrum aureuni Linton (B, P) Kyphosus elegans, (P) Kiiphosus sectatrix. (B) Honialometrou elongatuiit Manter (B, P) Eucinostomus calif orniensis, (P) Gerres cinereus, (Bi Jeancadciiatid brumpti DoUfus (B) Kiipliosiis sectdtrix, (B) Lepidapedon parepinepheli (this paper) (B) Epinephelus tigris. (B) Lepidapedon truncutum (this paper) (B) Holocentrus ascensionis, (B) Lepocreadiuin trulla (Linton) (B) Lutjaiius liuccaneUa, (B) Ocyurus chrysurus, (B) Pseudocn adiuiii biminensis (this paper) (B) Balistes cayriscus. (B) Pseudocreadium lamelliforme (Linton) (B) 110 Tulane Studies in Zoology Vol. 7 Table 3. — (Continued) lialistes retiiht. (I!) I.tictoiiJir/ix hiinnilalifi. (B> J'scKilocn ailiiim KraphosiDinim Maiitcr (!') ISiilisIrs iiinifriiiiiiiiK. ( I'l lUtlistcx rem K, (T) Ceratucaiithus scripta, (H) FAMILY MKUArKUIKAK Alci/apeni niniiiii (Linton) di) Lnctophriis bicoiKldlis. (H) Lttctojthriis trici)))iis, (Bi ThiisaiioiiJiitnjnx cjonnittiix >LTntor (B) Liictdphriis iriconiis, ilJ) FA M ILY M(1X0K( 1 1 1 II >A K Geiioltipii iimpiilhtirn Linton (III IlitrmiiUm nlbum, (I'l Iliieiii iiliiit piiyni . ( Rl lltit iimhni iiliniiicri, (Iti IliHiiiiiloii sciiints. {]'>) I'roi-totrciiiii tniiiciitiOK (Linloiil (I!) ('(intii.r miter. (It ) ]{(ieiiiiihiii purra. (B) lliteuiuhin plumieri, (B) Uaeiiiiiloti sciuriis, (B) I A.MILY ()l'i:C()ELII)AK lldiniirretiiliiiiii iiiiitiibile Linton (P.) E pine iilirl tits strintil-s, (]'<) llneiii iihiii xeiiinis. (Iti Lilt inn IIS s/iniii/ris. (11) I'etriiiiii tiiptin enii iitiit lis, (I!) Ihlieniiietrii e.reetii Linton (B) (liimiiiitJinrui- rieinus. (B) lliieiiiiihiii sciiinis, (B) ileHeoiiietni tortfi Linton (B) i'.piiieplieliis strifitiis. (Ii) Helieoiiietriiiii iiimiu Linton (B. P) IliietiiuJiin (ilhiiiii. (B) lliieiinihiii sciiiriis, (P.) I.iil iniiiis siniiii/ris. (I!) Epiiieplieliis iiiiiiliKjiis (P) 2\eoiiotiiiioriis un inniiiitii Mantcr (I!) Seliir eriniienojilliiiliii Its. (li) Opevneliis spliiiericus Ozaki (P) Seorpiieiiii niiistes, (I'» PdehiieriiiiUinn lernerl (this paper) (B) (rcrres einereiis. (B) J'ltiloeotiile injieteriipereae (this paper) (B) M !/rteroper) rueuilopceoeloiiles i/rnrilis Manter (B) Ap'K.ion hiiiiiliitiis. (Bi Seliir enimenopthiiliiiiis, (B) P.<.) atherinne (I'licu) (B) HepHctia fttipis, (P.) (P.1 cometra torta. Metadena globosa, Farahemi- uriis merus, Proctotrema truncaturn. an(d Siphodera vinaledwardsi) are previously known from the tropical American Pacific, a total of 18 species. In other worcis, based on presently known collections, the Bimini trematodes are much the same as those at Tortugas; and are more like those of the American Pacific than like those of Ber- muda. Manter (1940a, 1947, 1955) noted this similarity of Atlantic and Pacific Tre- matodes. He found 28 (19-1 percent) of 147 shallow water species at Tortugas also occuring at Bermuda; 21 of the 28 in the same species of hosts. Twenty-six shallow water species (17.6 percent) occurred in the Pacific but with two exceptions in dif- ferent species of host. He interpreted this approximately equivalent position of Ber- muda and the warm Pacific as reflecting the former continuity of the two oceans together with the greater persistence of parasite species as compared with hosts. My collections in the Gulf of Panama in- crease by five the number of Atlantic spe- cies also known from the Pacific. Manter (1955) noted a total of 32 species known from Tortugas, or probably occurring there, and also known from the American Pacific, as compared with 28 species also known from Bermuda. Bravo-Hollis (1954, 1956) added Siphodera vinaledwardsi. Proctotrema truncaturn, Metadena glohosa, and Haplos- phlanchnus sparisomae. My collections add the following to this list: Stephanostomum sentuin, Opisthadena dimidia. Enenteru7n aureum, Homalo??ietron elongatum, and Dinurus tornatus. Thus, at present 41 spe- cies are common to fishes of the American Atlantic and American Pacific. Present col- lections still indicate a stronger similarity of Caribbean trematodes to the Pacific re- gion than to Bermuda. The proximity of Bimini to Tortugas and the West Indies does not mean these regions have identical trematodes faunas. Even when many more collections are made some species will probably be found to have local distributions. Several species common at Tortugas and whose host species ocair at Bimini were not found in my collections; for example, Stephanostomum casum and Sterrhurus fusiformis. Furthermore, some 12 species (20 percent; at Bimini are not yet known from Tortugas. However, fewer col- No. 3 Sogandares-Bernal: Ttrematodes of Marine Fishes 111 lections have been made ar Bermuda. Table 3 lists the Digenea collected by me at Bimini and in the Gulf of Panama with indication of the locality. These species are arranged alphabetically according to family. Table 4 lists the hosts examined at Bimini and in the Gulf of Panama, and the Digenea found. The hosts are arranged alphabetically without regard to family. Table 5 lists the Digenea in my collections which are also known from Bermuda; Tor- tugas, Florida; Nassau, B.W.I.; and the Gulf of Panama. This list does not include all species known from the Caribbean region and in these other regions. Table 4. Alphabetical list of fishes from, the Gulf of Panama (P) and Bimini, B.W.I. (B) indi- cating trematodes found. Numbers of speci- mens listed after each fish and trematodes. Al)ndef(hif saa-ullUx. ( Pomacentridae), 3 B Sterrhunts floridcnsis. (Hemluridae). 1. in 1 of 3 B Acanthori/hium solandrinn, (AeanthocvbiidaeK 1 P HinidiueUd inarinn. ( Hcmiuridap). 2. in 1 P Acaiifhiinis huhianus. ( Aeantluiridae). 1 B Negative Acanthurus crestonifi. (Acanthuridae). 1 P Negative Acanthurus caerulcus, (Aeantliuridae), 4 B Hysterolecitha rosea, (Hemiuridae), 2, in 1 of 4B Albitla vulpes, (Albulidae). 2 P. 1 B Negative Alectis crinitus, (Carangidae), 1 B Negative Anchovia sp., (Engraulidae), 1 P Negative Anisotremus taeniatus, (Pomadasyidae), 1 P Negative Apopon binotatus. (Ctieilodipteridae), 3 B Pseudopecoeloides gracilis, (Opecoelidae), 1, in 1 of 3 B Apocionichthys steUntus, (Cheilodipteiridae) 2 B Negative Artos macropthalma, (Gobiesocidae), 1 B Negative Aulostomus maculatus, (Aulostomidae). 1 B Negative Balistes capriscus, (Balistidae), 4 B Apocrradinm balistes, (Lepocreadiidae), 1, in 1 of 4 B Apocreadium colli, (Lepocreadiidae), 5. in 1 of Pseudocreadium biminensis, (Lepocreadiidae) 1 2. in 2 of 4 B Balistes naufrugium. (Balistidae). 2 P Apocreadium braroi, (Lepocreadiidae), 8, in 1 of Pseudocreadiitm seaphosomum, (Lepocreadiidae). 30 + , in 1 of 2 B Stephanosttjiaum provitellosiim. (Acantliocolni- dae). 2(1, in 1 of 2 B Balistes verres, (Balistidae), 2 P Pseudocreadium seaphosomum , (Lepocreadiidae) 30 + , in 1 of 2 P " Balistes vetula, (Balistidae). 7 B Apocreadium colli, (Lepocreadiidae), 3, 1. in 2 of 7 B Apocreadium uroproctoferum, (Lepocreadiidae), 18, In 1 of 7 B Pseudocreadium lamelliforme, (Lepocreadiidae) 1, 1, in 2 of 2 of 7 B Bathi/gobius soporator, (Gobiidae), 1 B Negative Brachi/geni/s chrysagerus, (Pomadasyidae), 1 B Negative Viilamus hdjunudo. (Sparidae). 4 B Lobutostoma ringens. (Aspidogastridae), 1. in 1 of 4 B Proctoeces suhtenuc, (Fellodistomatidae), 1. in 1 of 4 B Stephanostomum seiitum, (Acanthocolpidae), 12, in 1 of 4 B Canthrriiies pullus, (Monacanthidae), 3 B Neg;i tive Crtiitliigaster rostratus, (Canthigastridae). 3 B Negative Curatij- criixos, (Carangidae), 2 B Bucephalus varicus, (Bucephalidae), 300 + . in 1 of 2 B Tergestia pectinata, (Fellodistomatidae). 4 in 1 of 2 B Caran.r hippos, (Carangidae), 1 B. 2 P Negative Caranx latus. (Carangidae). 4 B Bucephalus raricus. (Bucephalidae). 12. 30, 75, 300 + . in 4 of 4 B Stephanostomum d it re mat is, (Acanthocolpidae), 2. 3. 5, in 3 of 4 B Terqestia pectinata, (Fellodistomatidae), 1, 6, 1, in 3 of 4 B Caranx lugubris, (Carangidae), 1 B Negative Caran.r marginatus, (Carangidae), 1 P Nega five Caranx ruber, (Carangidae). 4 B Bucephalus varicus, (Bucephalidae). 1. in 1 of 4 B Tergestia pectinata, (Fellodistomatidae). 1, in 1 of 4 B Carapus bermudensis, (Carapidae), 1 B Negative Carchurhinus acronotus, (Carcharhinidae). 1 B Negative Carcharhinus cerdale, (Carcharhinidae), 1 P Negative Carcharhinus natator, (Carcharhinidae). 8 P Nagmia pacifica, (Gorqoderidae, 100 + , 100+, 100 + . 10O+, 100 + , 100 + . 100 + , 100+, in 8 of 8 P Carcharhinus obscurus, (Caracharhinidae), 2 B Negative Centropomus undecimalis, (Centropomidae), 1 P Negative Cephalopholis fulvus, (Serranidae), 1 B Negative Ceratacauthus scripta, (Monacanthidae), 1 B Diploproctodaeum haustrum (Lepocreadiidae), 32, in 1 of 1 B Cestracion tiburo, (Cestraciontidae). 2 P Negative Cestracion tudes, (Cestraciontidae). 1 P Negative Chaetodipterus faber, (Epihippidae), 1 P Neagtive Chaetodon humeralis, (Chaetodontidae), 2 P Negative Clupeid (unidentified). (Clupeldae), 1 P Negative Conger conger, (Congi-idae), 1 B Sterrhurus floridcnsis (Hemiuridae), 6, in 1 of 1 K Corgphaeno hipi)urus, (Coryphaenidae). 1 B. 2 P Binurux loni/iKinus. (Hemiuridae). 200 + . in 1 of 1 B : 200+. 200 + . in 2 of 2 P Dinurus tornatus, (Hemiuridae), not counted, in 1 of 1 B Stephanostomum con/phaenae, (Acanthocolpi- dae). 1 in 1 of IB Tctrochactus coruphaenae. (Accacoelidae), 4. in 1 of 1 B : 6. in 1 of 2 P Corgphapterus glaucofraenum, (Gobiidae), 1 B Negative Cynoscion albus, (Sciaenidae). 3 P Stephanostomum senium. (Acanthocolpidae). 3, in 1 of 3 P Cyno.^cion sp., (Sciaenidae). 1 P Negative Dasyatis americana, (Dasyatidae), 1 B Negative Diodon holocanthus, (Diodontidae), 1 P Negative Diodon hystrijr. (Diodontidae). 2 P: 1 B Diploproctodaeum plicitum. (Lepocreadiidae), not counted, in 1 of 2 P : in 1 of 1 B Echidna catenata, (Muraenidae), 1 B Negative 112 Tulane Studies in Zoology Vol. 7 Table 4. — ( Continued ) Eel (nniflcntififMl). 2 P Tubiiloicsicula liiidbrnii. (Ileinluridae). 5. in 1 of 2 P Elagatis liipinnulatus. (Carangidae), 3 P StephanostoiiiiiDi ditiematis. (Acanthocolpiilae), 6 in 1 of 3 P Epinepheliix aiuilofius, (Serranirtae). r> P HcHromt triiia iihiiia, (Opocoi'lidac ). 1. in 1 of •5 P ProiiOthiinc1n(>< i/onoderus, (Bucephalidae). 1, in 1 of « P Prosorhiinchii.s ozakii, (Bucephalidae), 1. in 1 of 6 P Epinrpheliis (iitttutufi, (Serranidae). 1 P: 4 B Negative Epiiirphelus labriformix, (Serranidae), 3 P Negative Epin< uioritt. (Serranidae). 1 B Iniiicntifii'd Icpocri-adiid trematodes Epiiii iilirlim sp., (Serranidae), 1 P Negative Ephieplieliis striatus, (Serranidae). 1 P : 6 B HdiiKicrcdiJiuin mutabile, (Opecoelidae), 1,1,4,8,5 in .J of 6 B IhlUoinctni torto. (Opecoelidae). 1, in 1 of 6 B Eiiiiii phihis tu/rix. ( Serranidaei. 1 B Lepidtilirilon ixinphiepheli. (I^epocreadiidae), 11. in 1 uf 1 B Equcx iiciniiiiititiix or phichcr, (Sciat-nidae), 1 B rnidentificd liemiurid. 1. in 1 of 1 B Eucinotttotitus riiliforiiirsis, ( Ccrridae), 2 P Homalometron eloinidtimi . ( Lepocreadiidae). 1. in 1 of 2 P Eucino-sloiiiiix ypp., (deniihu) . 1 B : ."> P Negative Eupeni'iis (jraiHlisqiKiuiis, (Mvillidae). G P Negative Felkhthiis piiniiacuhitiis, (Silniidac). 1 P Negative Fifituluriti corneta, (Fistulariidae). 5 I' Negatve Gah'ichtlnjs seemani, (Siluridae). 8 P Negative Galeocerdo anticus, ((jaleidae). 1 P Negative Qerres vinereus, (Gerridae). 4 P. Crunsicutis marina, (Lepocreadiidae). 1. in 1 of 4B Homalometron rloiK/ntiim. (I-epocreadiidae). 4,.j, in 2 of 4 B Pachinrendium leriicri. (Lepocreadiidae), 1, in i of 4 B Oerres sp.. ((Jerridae). 1 P Nejiative GiKitholritix thoiiiiixoiii, ( ( joUiidae). 1 B Negative Girmiiothorii.r dorii. ( .Muraenidae). 1 P SterrJniniK tnhoiidiiiis. ( llemiiiridae). 2r>. in 1 of 1 Giniiiiotliora.r iiioridi/n, (Muraenidae). 1 It Negative Gindiiothnruj- iioIii I! Genol'jpd diiiijiilldirn. (.Monoicliiiil.ie i. '.).l(i.l2, in 3 of .". B Helicomctrina iiimiu, (Opecorlidaei, 2. in 1 of 5 B Leurodera decora, 30,7,8.7. in 4 of ."> B. . . . IJaemiilon flavolineatum, (Pomadasyidae). 1 P. Negative llditiiidon ixirrd. (Pomadasyidae). ;> B Jlrdcliadend pijriforniis, ( I Iiiniiiiidae ). !'.•, in 1 of 3 B J>iplan DiithnKiuf />«Ji7/i<.v. (Zoogonidae). 2. in 1 of SB Genolopd nmpuUacea, (Monorchiidae). 30,70. in 2 of 3 B . ^, Leurodera decora. (Hemiuridae). 2.4. in 2 ot :, I. Proctotrema tniitcatinn. i Monorchiidae). 2. in 1 of .•', P. Ildciiiiihiii sriurus, (Pomadasyidae). 14 B linicliddt ltd piiril'ormi>i, (Hemiuridae), 1.3,6,2.5, in 5 of 14 B Diplamiux pd.riUus, (Zoogonidae), 3,2,4,3,10. in 5 of 14 B Gettoloiid iinniuUacea, (Monorchiidae), ol.21.14. 14. 4. 4. 7".>.. •?.-.. 25. l.^. 5. 10. 12. in 13 of 14 B Hamdcreadiuiii nuitabile. ( Opecoelidae) . 1. in 1 of 14 B Helicoiiiitra execta. ((Jpecoelidae), 2, in 1 of 14 B „ , Helicometriud iiiiiiia. (Opecoelidae). 1.2, in 2 of 14 B Leurodera decora. (Hemiuridae), 1,2,2, 2,3, 4, o. in 7 of 14 B Proctotrema truncatum (Monorchiidae). l.l..->. in 3 of 14 B Haemulon steindachneri, (Pomadasyidae), 1 1 Negative Haemulon sp., (Pomadasyidae). 5 P Ni 'native Hulichoerus bivitatta, (Labridae), 2 B Negative Halichoerus poeyi, (Labridae), 1 B Negative Hareni/ula clupeola. (Clupeidae). 6 B Bac'ciqer harenfjulae, (Pellodistomatidae). 40, in i of 6 B Ectenurus vinjulus, (Hemiuridae). 12, in 1 of (i B Paraheniiurus menis. (Hemiuridae), 8,10.3, in 3 of 0 B Hepsetia stipes, (Atherinidae), 16 B BivesicuJa hepsetia, (Bivesiculidae), 1. in 1 of 16 B Stet/anoderma atherinae, (Zoogonidae). 2.1,1.1. in 4 of 16 B Histrio histrio, (Antennariidae), 1 B Negative Holocanthus ciUaris, (Chaetodontidae), 1 B Negative Holocanthus isabelita, (Chaetodontidae), 1 B Negative Holocentrus ascensionis. (Holocentridae). 4 B Lepidapedon truncatum, (Lepocreadiidae). 5.1.1, in 3 of 4 I;> Pseudopecoelus harkeri, (Opecoelidae), 1. in 1 of 4 B Ktenopera equilata, (Opecoelidae). 8,8, in 2 of ^ K , , ■ Stcphauostomum pseudocarangi. (Acanthocolpi- dae), 2,1.1. in 3 of 4 B tliistomatidae), 14. in 1 of 4 B En( iiliriim aureum. (Lepocreadiidae), S,]4'.t, in 2 of 4 P. .IcdiKdiliiidtia hruinpti, (Lepocreadiidae), 2. in 1 uf 4 B No. 3 Sogandares-Bernal: Ttrematodes of Marine Fishes 113 Table 4. — ( Continued) Oi>ixth(uUna (Ihnidia, (Ilemiuridae). 88. in 1 of 4 B Labri.soniiis niicliipinnis, (Rlonnidac). 4 P. Sterrlnoiix florideiisis (Hemiuridao). 1, in 1 of 4 P. Lactopliriin biraiidnlis. (Ostraclidae). 1 P. Meriaprra (/i/r'na. (Megapcridne). 1. in 1 of 1 B LactophriiK iricornix. (Ostraciidae). .S B Dermailriia lactophrysi, (Lepocreadiidae), 1. in 1 of .S B Mcfiniif" o.iniini, (Meganeridao). 1. in 1 of 3 B Thiixuii/iiilnirinu' eloin/iitus, (Megapcridae), 1, in 1 of :: B Laciiijihiiix triiinniis, (Ostraciidae). 3 P. Apocrciiiliinii anijitstum, (Lepocreadiidae). 1. in 1 of 3 B Cahlein frifjoni. (Lepocreadiidae). 1, in 1 of 3 B Dermadena lactophrysi, (Lepocreadiidae). 1. in 1 of 3 B Lari III lift (irt/entatus, (Seiaenidae). 2 P Negative Lutiaiiiis analis, (Lutjanidae), 1 B Mrtiuhtiii rrassiilnta, (Cr.vptogoniniidae). '_'. in 1 of 1 Lutjanus apodus, (Lutjanidae). 2 B Negative Lutjamis biiccaiielhi. (Lutjanidae). 1 B Lepocriudiuin tnilla. (Lepocreadiidae), 1, in 1 of 1 Lutjanus fiiiseus, (Lutjanidae). 1 B Negative LuijdHus yuttaius, (Lutjanidae). 1 I' Negative Lutjunus iorduiii. (Lutjanidae). 1 P Oonacautlnlki lutjani, (Crjptogonimidae), 2. in 1 of 1 P Lutjaniis iiovemfasciatus, (Lutjanidae). 1 P Paracniptoi/otiimus americaiius, (Crvptogoninii- daei. 10U + , in 1 of 1 P Lutiniiiis spp., (Lutjanidae). 4 P Negative Lutjiinux sjinnifris, (Lutjanidae), 5 B Haiinirrciidiiiiii mutahile, (Opecoelidae). 3.1.127, 11.7. in 5 of 5 B Helicoiiu'trina nimia, (Opecoelidae), 2.3, in 2 of 5 B Siphodcra riiialedivardsi. (Cryptogonimidae) , 3,411.4.21.78. in 5 of 5 B Makaira marliini, (Istiophoridae), 2 P Negative Mukuiru nUjricana, (Istiophoridae), 1 B Negative Makaini sp., (Silver Marlin), (Istiophoridae), 1 P Negative Microspatliodon dorsalis, (Pomacentridae), 2 P Negative MaUicuHthus plumieri, (Malacanthidae), 8 B BriuliiideiKi pi/riformis, (Hemiuridae), 1. in 1 of S B StepliuiioNtuiiiinii mil) lit inn. (Acanthocolpidae). 1, in 1 of 8 B i^tephiiiio.ytoiinim sentum, (Acanthocolpidae), 2, in 1 of s B HtirrlinruK floiidensis, (Hemiuridae), 1,1, in 2 of 8 B Malacoctriiiis (iiiroliiieatiiSj (Clinidae), 1 B Negative MalacoctcniiK iiiiicropus, (Clinidae), 1 B Negative MelichtJnix pisi-eiis. (Balistidae). 5 P. Unidentified Lepocreadiid trematode. in 1 of 5 B MoiHiiuiithux riliatiis, (Monacanthidae). 1 P. Negative Mufjil sp.. (Mugllidae). 6 P Unidentified luiploporid trematode, 2 in 1 of G P Myctcropi nil faJcdtu, (Serranldae), 1 B Podocoti/lc iiiycteropercae, (Opecoelidae), 12. in 1 of 1 B Myctrropercii v. veneiiosa, (Serranidae). 1 B Prij.sorln/nchux pacificiis. ( Bueephalidae). 28. in 1 of 1 B titepJiuiioxtoiii inn dnitiitinii, ( Acanthocoli)idae). 2. in 1 of 1 B Nef/apHoH hreriiostris. (Carcharhinidae), 2 B Negative OcyuriiK rJirysurus, (Lutjanidae), 3 B Lepocrriidiinn triillo, (Lepocreadiidae) . 10')-]-, ."),."). in 3 of 3 B Mctudena ijlubosa, (Cryptogonimidae), 21, in 1 of 3 B Prosonnniii renin bilabintum, (Prosogonotremati- daet, 1. in 1 of 3 B ..,,,.,, Kterrhnnis floridcnsis, (Hemiuridae). 1. in 1 ot 3 P. Or/cocephalux respertilio, (Ogcocephalidae). 2 B ' Sterrliiiruft floridensis, (Hemiuridae). 6,6, in 2 of 2 B Opxaniis V sp.. (Batrachoididae). 3 P Negative Palometa media. (Carangidae). IP , ^^ . , . Bucephalus rariciis, (Bueephalidae), in, in 1 ot 1 P Petromitopon ernentatus. (Serranidae), IB. Hamaereadiuin miitabile. (Opecoelidae). 1. in 1 Pfjmaeauthux aureus. (Il'olaeanthidae), 6 B AntoichiK lima, (Fellodlstomatidae), 14.1,..1(), 7,2!). in ."> of (j B Barisomum enibc^eens. (Pronocephalidae). 1. in 1 of 6 ^. . ^. , ^ Cleptodi.< Negative Prionotns sp.. (Triglidae). 6 P Negative Psenes sp.. (Nomeidae), 1 B Negative Reinora reiiiora, (Echeneidae), 6 P Rliinobatiis leucorhynchus, (Rhinobatidae). 2 P Negative .^ . , r. Rypticus saponaceous, (Serranidae). 1 1» Rjiptfcus saponaceous bicolor, (Serranidae). 1 P Negative t Negative lninrus sierra, (Cybiidae), P Negative iicorpacna mystes. (Scorpaenidae), 1 P ^ . , , Opeeoelus sp]iaericus, (Opecoelidae), 1, in 1 ot 1 I' l^corpacna iiluniieri. (Scorpaenidae), 2 B {■fterrliiiriis jloridensis. (Hemiuridae), .3. in 1 of 2 B Selar crumenoptlialmus. (Carangidae). IB Ectenurus viryulus, (Hemiuridae), 4, in 1 ot 1 B Neonotoporus yamagutii, (Opecoelidae), 1, in 1 of 1 B Pseudopecoeloides i/raeilis. (Opecoelidae), 4, in 1 of 1 B Terriestia peclinatn. ( Fellodistomatidae), 2, in 1 ' of 1 B Helene breroorti, (Carangidae), 1 P Negative f^eriola dorsalis. (Carangidae), 1 P Negative t^criola iniizatlaiia. (Carangidae). .5 P i p ^ocrc:i(liiil.-ic i. not coiintcMl. In t> of !• 1' ili!>'i<1'i<''. !• i" 1 of 2 P StelUfcr sp., (Sciaenidae). 1 V Negative stc'ilifrr sp.. (Sciaenidae). 1 1' Pseuihjijeroeloides ci/uesi, (<)peco(>lidae), 7. in 1 of 1 .finifiniithiix rosseau, (Syngnathidae). 1 B Negative iijiiiodiix sp.. (Synodontidae), 5 V ' Tubulovesiciila lindhergi, (Hemiuridae), 9, in 1 of 5 Trtnuidon liifiiiidits. (Tetraodontidae), 3 I' Negative ThdUixstjina hifasciatii in . (Labridae). 1 B Negative Thiirinojix fiiichiilciiix, (.\tlierinidae). 3 P Hiiiiloxfihinchnnx nculux. ( Ilaplosplanclinidae), 1 in 1 of 3 P Tracliinocephalux wiio/ix, (Synodontidae), 1 B Negative Trachinotux falcatiix. (Carangidae). 1 B Negative Trarliinotiis f/lauciix. (Carangidae). 2 B Unidentified lepoereadid trematodes, not counted, in 2 of 2 B Trichiiinix leiitiniix. (Trieliiuridae), 2 P Negative l^iiloxunis raiiliidoniti . ( I'.elonidae). 7 B ' HuiiJdxpJinicluuix ariitux. ( llaplosplanehnidae), 1,3. in 2 of 7 B Tyloxunix iiacifiriix, (Belonidae), 1 P Negative Tyiosurux Ntol::ni(niiii, (Belonidae), 1 P Negative Vpcneux muculatux, (Mullidae), 1 B Negative Urohatix jamaicensis, (Dasyatidae), 2 B Negative Urotri/i/on asterias, (Dasyatidae), 1 P Negative Voinir declirifrons, (Carangidae I. <'« I' Negative Xi/richthys psittacus, (l^abridae). 1 B Negative Table 5. Bimini trematodes also known from Ber- muda; Tortugas, Fla.; Nassau, B.W.I.; and the Gulf of Panama. lier- Tor- Nas- Pana- Trematodo muda tugas sau ma Antorcliin urn II A porifddium biilixtix lUn-ciiitr liamifjiilue Hinixomum cm hescens ItiicxivuUi licpxctia Unirliiidciiu pinifiifinis liUCCpllllluX niriciix Clcplodixcus reticulatus CrnssicutiK marina Dcrmadciia lacioplirysi Dill II nix lonyixinuH + + + + + + + + + + + + + + — — + Ber- Tor- Nas- Pana- Trematode muda tugas sau ma Diniirnx toriintiix — + — + Dililiiiiiinx pa.rill iix — + — — IHphipnictiiiliii u III pliiit II III — + — + Ectcniinix rininliis — + + — Encii t( III III aiin II III — + — + (Icniihipa a III piillaicii + + + — lliniKicrcadin III iniitiiliili — ■ + + — Ilaplosiiliniclnius ai-iitiix + + — + Hcliciiinctni iwcctn — + + — Hcliciiinct ni titrtii — + — — IIcliciniK 1 1 i nil II i mill — + + + Homiil'iini Iron ihiniiatnin — + — + Hyxtcrolccitlia roxea — + — Lepocrcadiii III triiUa + + + Lcnniilcra decora — + — Lobatoxtoiiia rinf/ois + + — Mcf/apcra (/yrinu + + — Metadeiia craxsiilata — + — Metadciia fllohoxa — + — yeoiiotoporiis yainiit/utii — + — OpixtJiadi iia dimidia — + — + Purahcmiuriix incrux — + — + Pructocces xiihtenue + + — — Prfjctofrciiia triincatiiin — + — — Proxorliinicfiiix paiificiix + + — — P xc iidoc rciidi u III laincUiforine -f + — — Pxeiidopccocloidcs !/racilis — + + — Pxeudopccoelus barki ri + • — - — — h'liiinorcliix odlincri — + ■ — — Sipliodcrn riniilrdinirdxi + + + — i^tciiopcru cquiiatA — + + — Stepliaiioxtoiiiuiii coritphaciiac — + — — Stcpli a n ox to III II III denial II in + + — — Stcpliaiioxtoinuin dil rcinal i>i . — + — + Stcpliaiioxtoinuin xcntiDii — + — + t^tcrrlniriix floridvnxis — + + — Tcri/cxtia pcctiiiata — + + — Tctroclnictiix coriiplntcnac — + — + Thiixanopharynx eloiujatus ~ + - + VI. Summary 1. During the summers of 1956 and 1957 the writer collected trematodes from fishes in the Gulf of Panama (1956) and at Bimini, British West Indies (1957). In No. 3 Sogandares-Bernal: Ttrematodes of Marine Fishes 115 Panama, 234 specimens of 121 species of fishes were individually examined; at Bi- mini, 250 specimens of 109 species. Totals were 484 specimens of 209 species. 2. Tremarodes were found in 19 percent of the species of fishes in the Gulf of Pan- ama; in 56 percent of the species of fishes at Bimini. A total of 88 species of digenetic trematodes were collected and studied. Of these, 62 were at Bimini; 31 in the Gulf of Panama; 5 species occurred in both regions. 3. Two new genera are named: Cableia (Lepocreadiidae) and Gonacanthella (Cryp- togonimidae ) . 4. Seventeen new species, distributed in 7 families are named. These are Acantho- colpidae: Stephanostomum provitellosum, Stephanostomum pseudocarangis: Bucephali- dae: Dolljustrema niuraenae; Cryptogoni- midae: Gonacanthella lutjani; Hemiuridae: Sterrhufus taboganus; Lepocreadiidae: Apo- creadiuni bravoi, Apocreadiuni coili, Apocre- adium angustum, Apocreadium nroprocto- ferum, Cableia trigoni, Lepidapedon parepi- nepheli, Lepidapedon truncatum, Pseudo- creadimn biminensis; Opecoelidae: Pachy- creadium lerneri, Podocotyle mycteropercae; Waretrematidae: Megasolena kyphosi, Myo- dera magna. 5. The names of ten species and one genus are considered synonyms. These are Bianium lecanocephalum Perez-Vigueras, 1955, synonym of D. haustrum (McCallum, 1918) La Rue, 1926; Proneochasmus Szidat, 1954, synonym of Paraspina Pearse, 1920; Psendocreadium myohelicatmn Bravo & Manter, 1951, synonym of P. scaphosomum Manter, 1940; Stephanostomum cubanus Perez-Vigueras, 1955 and S. m,anteri Perez- Vigueras, 1955, synonyms of S. ditrematis (Yamaguti, 1939) Manter, 1947; Stepha- nostomum mediovitellarium Perez-Vigueras, 1955, and S. lopezneyrai Perez-Vigueras, 1955, synonyms of S. sentum (Linton, 1910) Manter, 1947; Stephanostomum admicro- stephanum Perez-Vigueras, 1955, synonym of S. microstephanus Manter, 1934; Tubulo- vesicula anguillae Yamaguti, 1934; T. spari Yamaguti, 1934 and T. serrani Nagaty, 1956; synonyms of T. lindbergi (Layman, 1930) Yamaguti, 1934. 6. Twenty-nine new host records are noted. 7. Some observations regarding the ecol- ogy of these parasites within the host in- cluded the exact location of the trematodes in the digestive tract, and, in some cases, measurement of the pH of these locations. The pH was surprisingly inconstant even in the same region of different individuals depending chiefly on recency of feeding. 8. The Digenea of Bimini are very simi- lar to those known from Tortugas, Florida and from Bermuda. However, the known resemblance of those of Tortugas to the American Pacific is increased by five species found in both oceans, making an impressive total of 41 species. 9. A list of Digenea and their hosts, as well as a list of hosts found infected, are included. VIL Acknowledgements A number of people and institutions were very generous in providing aid in one way or another. The trip to Panama was made posisble by a Wolcott Scholarship from the Zoology Department, University of Nebras- ka. The following persons aided in collect- ing hosts and/or lending laboratory facili- ties: Messrs. Martin Burkenroad, FAO, Laboratorio Nacional de Pesca, Panama City; Paul A. Gambotti, Panama City; Antonio Landa, Inter-American Tuna Commission Laboratory, Taboga Island, Panama; Antonio Marciacq, Panama City; John and Louis Schmidt, Panama Canal Zone; Anastasio Sogandares, my father; Dr. Carl Johnson, Gorgas Memorial Laobratory, Panama City who supplied me with alcohol. The work at the Lerner Marine Laboratory on Bimini, B.W.I. was made possible by the American Museum of Natural History, New York City. Gratitude is expressed to Mr. John Bergmann, maintenance engineer and other employees of the Lerner Marine Laboratory for collection of hosts and in- numerable other favors; to Mr. and Mrs. Michael Lerner who many times caught fish hosts for me and aided in many other ways; to Mr. C. L. Smith, Museum of Zo- ology, University of Michigan, and Dr. Vladimir Walters of the American Museum of Natural History, who caught and identi- fied some hosts. Appreciation is also extended to my wife, Judithe Stanley Sogandares, who directly assisted in the work at Bimini, to Mr. E. S. Robinson, Zoology Department, University 116 Tulane Studies in Zoology Vol. 7 of Nebraska for aid in integrating some Faust, E. C. and C. C. Tang 1936. Notes on L J r „ -a-^:^: „^A r^ r^r u \Y/ new aspidog-astrid species, with considera- host data from Bmimi, and to Dr. H. W. ^.^^^ of the phvlogenv of the group. Para- Manter under whose direction this woric g{i^ 28: 487-501. was done. Hanson, Mary Louise 1950. Some digenetic _ trematodes of marine fishes of Bermuda. VIII. References Cited proc. Helm. Soc. Wash., 17: 74-88. Bravo-Hollis, Margarita 1954. Tremato- Johnston, T. H. 1934. Some Australian dos de peces marinos de aguas Mexicanas. anaporrhutine trematodes. Tra)it<. & Proc. VII. Ann. Inst. Biol. (Mexico), 25: 219 Roy. Soc. So. Anstralia, 5S: 139-148. 252. La Rue, George R. 1957. The classification 1956. Trematodos de pe- Digenetic Trematoda: A review and a ces marinos de aguas Mexicanas. XI. new system. Exper. Parasit., 6: 306-349. Estudio de 17 digeneos de las del Pacifico, layman, E. M. 1930. Parasitic worms from incluyendo seis especies nuevas y un ^.j^^ fishes of Peter the Great Bay. Bull. nuevo genero. Ibid.. 27: 245-277. Pacif. Sci. Fish. Res., 3: 1-120. (Russian Bravo-Hollis, Margarita and H. W. Man- with German summary ) . TER 1957. Trematodes of marine fishes Linton, E. 1907. Notes on parasites of Ber- of Mexican Waters. X. Thirteen Digenea, i-^u^ia fishes. Proc. U. S. Nat. Mns., 33: including nine new species and two new 85-126. genera from the Pacific Coast. Proc. ^^^^ Helminth fauna of Helm. Soc. Mash., 24: 35-48. p^.^ Tortugas. 2. Trematodes. Pap. Tor- Bravo-Hollis, Margarita and Franklin tugas Lab., Dept. Marine Biol., 4: 11-98. Sogandares-Bernal 1956. Trematodes of ^94q_ Trematodes from marine fishes of Mexican Watei-s. IX. fishes mainly from the Woods Hole re- Four gasterostomes from the Pacitic ^j^^^ Massachusetts. Proc. U. S. Nat. Coast. Jour. Parasit., 42: 536-039. Mas., 88: 1-172. Caballero y C, Eduardo 1945. Hallazgo de looss, A. 1907. Beitrage Zur Systematik una especie nueva del genero Petalodisto- ^^^. Distomen zur Kenntnis, der Familie HH(/« Johnston, 1913. (Trematoda: Gorgo- Hemiuridae. Zool. .Jahrb. Sijst. (Abt. deridae) en las costas de Manzanillo, g^^^ ) .?g. ^3.130 ^^I'fr^'i '^""' ^'''^' ^'''^' ^^^"■'''''^' ^^- Manter,' H. W. 1931. Some digenetic tre- 359-365. matodes of marine fishes of Beaufort, 1952. Revision de los gen- j^^j.^j^ Carolina. Parasit., 23: 396-411. eros y especies que integran la familia ^^3^ g^^^^ digenetic tre- Acanthocolpidae Luhe, 1909. Rev. Med. ^^atodes from deep-water fish of Tortu- Vet. & Parasit., 11: 1-Zdi. ^^^^ Florida. Carnegie Inst. Wash. Publ, Caballero y C, Eudardo, L. F. Barroeta 435. 257-345. and R. G. Grocott 1956. Helmintos de la -^^35 rpj^^ structure and Republica de Panama. V. Redescripciones 'taxoriomic' position of Megasoleua estrix de algunos trematodos ya conocidos pero Linton, 1910 (Trematoda) with notes on nuevos en la fauna helmintologica de este j-glated trematodes. Parasit., 27: 431-439. pais i^e^.. Biol. Trop. (Costa Rica), 4: ^^3^ ^ ^^^ ^^^^^^^ ^^ ■ distomes (Trematoda) with lymphatic Caballero y C, Eduardo, Margarita Bravo vessels. Rept. Allan Hancock Pacif. Ex- HoLLis and R. G. Grocott 1952. Helmm- ^^^^ 2- 11-22. tos de la Republica de Panama. III. Tres " ' ^g^^^ Digenetic trema- trematodos de peces marinos con descrip- --todes' of fishes from Galapagos Islands ^^'w ?L""^ ."ot''''i«7Pi'«T- '"• and neighboring Pacif ic. //,k/., 2 : 327-497. 5ioWM.xzco;, 23: 167-180. ^^^^^ Gasterostomes Caballero y C, Eduardo, Margarita Bravo (Trematoda) of Tortugas, Florida. Pap. HOLLis and R. C. Grocott 1953. Helmin- Tortugas Lab., 33: 1-19. tos de la Republica de Panama. VII De- ^9^5 Dermadena lacto- scripcion de algunos trematodos de peces ---^/^.^../nV-^enV, n- sp. (Trematoda: Lepo- mannos. Ibid., Z4: 9i-Ub. creadiidae) and consideration of the re- ChandleR, A. C. 1935. Parasites of fishes lated genus Psendocreadinm. Jour. Para- in Galveston Bay. Proc. U. S. Nat. Mus., sit., 31: 411-417. 83: 123-157. I947. The digenetic tre- Dollfus, R. Ph. 1937. Les trematodes Di- matodes of marine fishes of Tortugas, genea des selaciens (Plagiostomes). Cata- Florida. Amer. Midi. Nat., 38: 257-416. logue par botes. Distribution geograph- I954. Some digenetic tre- ique. A7in. Parasit., 15: 58-73; 164-281. matodes from fishes of New Zealand. 1946. Sur trois especies Trans. Roy. Soc. N. Z., 82: 475-568. de distomes, dont un a 17 ventouses (En- 1955. The zoogeography eyiterinn (Jeancadeyiatia) brumpti n. sp.-*. of trematodes of marine fishes. Exper. Ibid., 21: 119-128. Parasit., 4: 62-86. No. 3 Sogandares-Bernal: Ttrematodes of Marine Fishes 117 Markell, E. K. 1953. Nagmia floridensis 1938. Studies on the hel- n. sp., an anaporihutine trematode from minth fauna of Japan. Part 24. Tre- the coeloni of the stingray, Amphotistus matodes of fishes. V. Ibid., 8: 15-74. sabinus. Jour. Parasit., 39: 45-51. 1939. Studies on the hel- MoNTGOMERY, W. R. 1957. Studies on dige- minth fauna of Japan. Part 26. Trema- netic trematodes from marine fishes of todes of fishes. VI. Ibid., 8: 211-230. La Jolla, California. Trans. Amer. Mi- 1942. Studies on the hel- cros. Soc, 76: 13-36. minth fauna of Japan. Part 39. Trema- Nagaty, H. F. 1941. Trematodes of fishes todes of fishes mainly from Naha. Trans. from the Red Sea. Part 2. The genus Biogeogr. Soc. Jap., 3 : 329-398. Hamacreadium Linton, 1910 (Fam. Alio- 1953. Systema Hehnin- creadiidae) with a description of two new thum. Part I. Digenetic Trematodes of species. Jour. Egypt. Med. Assoc, 27: Fishes. Tokyo, 405 pp. ^^^-^1^- Abstract 1956. Trematodes of ^ „ . ABSTRACT fishes from the Red Sea. Part 6. On five Collections of digentic ti-ematodes of distomes including one new genus and marine fishes from the Gulf of Pan- four new species. Jour. Parasit., A2: 151- ^™a and Bimini, British West Indies, 155 were made during the summers of 1956 OzAxi, YOSHIDA 1925. Preliminary note on and 1957 respectively. A total of 234 a trematode. Jour. Parasit., 12: 51-53. specimens of 121 species of fishes were 1936. Two new genera of ^^^"^^"^d in the Gulf of Panama, and the trematode family Allocreadiidae. Zool. ^^ specimens of 109 species m Bimini Mag., 48: 513-519. "(Japanese with Eng- Trematodes were found in about 19 lish summary) percent of fish hosts in the Gulf of Perez-Vigueras,'l 1940. Prosogonotrema- Pamama and in about 56 percent of tidae n. fam. y Prosogonotrema hilabia- fish hosts in Bimini. A total of 88 turn n. gen. n. sp. (Trematoda: Disto- ^Pf"^% °^ trematodes are reported mata) parasito de Ocyurus chrysurus J^^^f. trematdoes are distributed in 19 (Bloch) (Pisces). Mem. Soc. Cubana families. Two new trematode genem in Hist. Nat.; 14 : 249-252. \^^ families Lepocreadiidae {i.e., Cab- 1955. Descripcion de seis ^^'""k ^J'^ ^ryptogonimidae (; e., Gona- especies nuevas de trematodes de la fam- IZttf'^l Zl^^f^J!^"^' ^'7f k ^^ •i- A „ ,1 1 • , J- • • J 1 species or trematodes are described m iha Acanthocolpidae y division del genero ^fj^ following families: Acanthocolpi- Stephanostomum en sub-generos. Rev. ^ Stephunostomum proviteUosum, Ibertca Parasit., Extraordinary Volume: ^„^ ^^ pseudocarangis ; Bucephalidae: Price, E. W. 1953. The fluke situation in ^^J Gonacanthella hit jaJ; tAmiuvi- nTlSr" '■"'"'"^"t^- '^^"'■- P»>-»^^t-' 39: dae: Sterrhurus taboganus; Lepocrea- T}^^^ f^, l^.~ r. ^(\r^(\ rpi j. u i. ii diidae : Pseudocreadium. biminensis, Le- Read, Clark P. 1950. The vertebrate small pidapedon truncatum, L. parajyinepheli, intestine as an environment for Parasitic ^ableia trigoni, Apocreadium coiU, A. helminths Rice Inst Pamphlet, 3/: 1-94. angustum, A. uroproctoferum, and A. ^^lZl'^^%f^r^\: ^r.''''^7.e"^T.^- ^- hZvoi; Opocoelidae: Podocotyle my- HUTTON 1958 The status of the Trema- cteropercae, and Pachycreadium ler- tode Eenns Bmnmm Stunkard, 1930, a „g„- Waretrematidae : Myodera mag- synonym of Diploproctodae^um La Rue, ^nd Megasoleua kyphosi. Ten spe- 1926. Jour^ Parasit 44: 566-567. ^.jes ^nd two genera of trematodes are Sparks, A K. 1957. Some digenetic trema- synonymized. Twenty-nine new host todes of marine fishes of the Bahama records are reported. Records regard- isiands bull. Mar. Sci. Gulf & Canb., jng. the ecology of these trematodes in- 7: 25.j-zb5. ^„_„ ^ , ... . elude, when possible, the exact loca. Stunkard, H. W. 1957. Intraspecific varia- tion in their respective hosts. Results tion in parasitic flatworms. Syst. Zool., of observations of the gastro-intestinal 6: 5-18. . tract of 10 specimens of 1 host species Williams, C. O. 1942. Observations on the seem to indicae that recency of feeding life histoiT and taxonomic relationships is a factor determing the pH of any of the trematode Aspidogaster conchicola. particular site in the digestive tract, Jo»r. Parasii., 28: 467-475. especially the stomach, and that tre- Winter, H. a. 1957. Trematodos de peces matodes lived there in spite of large marinos de aguas Mexicanas. XII. Dos shifts in pH. Certain ecological fac- generos de digeneos (Lepocreadiidae, in- tors possibly influencing speciation of cluyendo una nueva especie procedente trematodes are discussed. The known de Kyphosus elegans (Peters) de las resemblance of Digenea from marine Islas Tres Marias, en el Oceano Pacifico. fishes from Tortugas, Florida, with Ann. Inst. Biol. (Mexico), 27: 403-413. the American Pacific is increased by 5 Yamaguti, Satyu 1934. Studies on the hel- species, making a total of 41 species minth fauna of Japan, Part 2. Trema- known from both oceans. Host para- todes of fishes. I. Jap. joiage, table of contents and index (unbound) $3.50 VOLUME 7, 1959 1 An illustrated key to the crawfishes of Louisiana with a summary of their distri- bution within the State (Decapoda, Asta- cidae), by George Henry Penn, pp. 3-20 (April 23, 1959). Comparison of the chromatophoro tropins of two crayfishes with special reference to electrophoretic behavior, by Milton Finger- man, pp. 21-30 (April 23, 1959) $0.60 2 A review of the seabasscs of the genus Vciitropristes (Serranidae), by Rudolph J. Miller, pi). 33-68 (July 9, 1959) 0.75 Orders should be addressed to Meade Natural History Library, c/o Department of Zoology, Tulane University, New Orleans, 18, La., USA Please make remittance payable to "Tulane University" Jim ^(e)(DIL®(§l' LIDnAttl NOV -3 1959 uliivERsiTY X" Volume 7, Number 4 / r-,., .^ ^ 7~^ -LAA. tl A.. A /^<1 ' October 19, 1959 PARASITES OF THE COMMERCIAL SHRIMPS, PENAEUS AZTECUS IVES, P, DUO R ARUM BURKENROAD, AND P. SETIFERUS (LINNAEUS) DWAYNE NATHANIEL KRUSE, DEPARTMENT OF BIOLOGICAL SCIEJfCES, FLORIDA STATE UNIVERSITY, TALLAHASSEE, FLORIDA THE LARVA OF THE OAK TOAD, BUFO QUERCICUS HOLBROOK E. PETER VOLPE and JAMES L. DOBIE, DEPARTMENT OF ZOOLOGY, NEWCOMB COLLEGE, TULANE UNIVERSITY, NEW ORLEANS, LOUISIANA TULANE UNIVERSITY NEW ORLEANS TULANE STUDIES IN ZOOLOGY is devoted primarily to the zoology of the waters and adjacent land areas of the Gulf of Mexico and the Caribbean Sea. Each number is issued separately and contains an individual study. As volumes are completed, title pages and tables of contents are distributed to institutions exchanging the entire series. Manuscripts submitted for publication are evaluated by the editor and by an editorial com- mittee selected for each paper. Contributors need not be members of the Tulane Univer- sity faculty. 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George Henry Penn, Editor Meade Natural History Library, Tulane University, New Orleans, U. S. A. Assistant to the Editor: Robert K. Chipman TULANE STUDIES IN ZOOLOGY Volume 7, Number 4 October 19, 1959 CONTENTS PARASITES OF THE COMMERCIAL SHRIMPS, PENAEUS AZTECUS, IVES, P. DUORARUM BURKENROAD, AND P. SETIFERUS (LINNAEUS) Dwayne Nathaniel Kruse 123 Editorial Committee: Robert Ph. Dollfus, Museum National d'Histoire Naturelle, Paris, France Robert B. Short, Professor of Zoology, Florida State University, Tallahassee, Florida Franklin Sogandares-Bernal, Instructor in Zoology, Tulane University, New Orleans, Louisiana THE LARVA OF THE OAK TOAD, BUFO QUERCICUS HOLBROOK E. Peter Volpe and James L. Dobie 145 Editorial Committee: Fred R. Cagle, Professor of Zoology, Tulane University, New Orleans, Louisiana Kenneth L. Gosner, Science Department Staff, Newark Museum, Newark, New Jersey M. Graham Netting, Director, Carnegie Museum, Pittsburgh, Pennsylvania MUS. COWP. ZOOL NOV- 31959 HARVARD UNIVERSITY PARASITES OF THE COMMERCIAL SHRIMPS, PENAEUS AZTECUS IVES, P. DUORARUM BURKENROAD AND P. SETIFERUS (LINNAEUS) i- - DWAYNE NATHANIEL KRUSE, Department of Biological Sciences, Florida State University, Tallahassee, Florida Introduction The U. S. Fish and WildHfe Service Fish- ery Statistics of the United States for 1956 show that shrimp are by far the leading sea- food industry in the United States. The total catch for 1956 was worth $70,894,000. The Gulf of Mexico produced $62,499,000 of this amount. Though the importance of shrimp in the fishing economy has been in- creasing steadily since 1956, little is known about their parasites. Knowledge concern- ing deleterious effects, distribution and host specificity of shrimp parasites is desirable for both academic and economic interests. The specific goals of this preliminary sur- vey were: (1) description and identification of the parasites of P. aztecus, P. duorarum and P. setiferus from Alligator Harbor and Apalachicola Bay, Florida; (2) determina- tion of the incidence and site of infection of these parasites; and (3) recording observa- tions regarding host specificity and life his- tories of these parasites. After the present paper had been sub- mitted for publication Hutton, et al ( 1959) published a preliminary report dealing with parasites of shrimp in Florida waters. For a historical review of the subject the reader is referred to this work. Materials and Methods Collection of Hosts The shrimp hosts, Penaeus duorarum, P. aztecus and P. setiferus, were collected from three locations along the northwest Gulf coast of Florida between June and Novem- ber, 1958. Most P. duorarum were collected at approximately monthly intervals from July through October in Alligator Harbor, Franklin County, Florida. These shrimp were collected in the grassy flats on the sand bar in the mouth of the harbor with either a push net or a small beam trawl. One collection of ten specimens of P. duo- rarum was made in Apalachicola Bay, Frank- lin County, Florida in October; another col- lection of four shrimp was made in the grassy flats near the mouth of the St. Marks River, Wakulla County, Florida in Septem- ber. All specimens of P. aztecus and P. seti- ferus examined were collected from Apa- lachicola Bay with a large shrimp trawl. The collections were made at seven different stations in the bay that varied from shallow grassy flats near its head to a depth of 40 feet near St. George's Island. P. aztecus and P. setiferus were collected at approximately monthly intervals from June through Oc- tober. The shrimp were identified with the aid of the keys to the family Penaeidae by Anderson and Linder (1934) and Voss (1955). Examination of Hosts All of the shrimp caught were macro- scopically examined for evidence of a micro- sporidian infection locally known as "cotton shrimp", "blue shrimp" or "milk shrimp". The incidence of this infection in relation to the total catch was noted. The infected specimens were either preserved in 10% formalin or brought back alive to the labo- ratory at Florida State University for further study. A representive sample of normal "non-cotton shrimp" was taken from each station; these were transported alive in three gallon glass jars to the laboratory for com- plete examination. The shrimp were dissected alive in petri dishes in a small amount of sea water or physiological saline. Hosts which had been iced or preserved in formalin proved to be unsatisfatcory for examination. The diges- tive gland in iced shrimp liquefies rapidly and fills the hemocoel with cytolized yellow tissue making small translucent parasites 1 A portion of a thesis submitted to the g^raduate Council of Florida State University in partial fulfillment of the requirements for the degi-ee of Master of Science. - A contribution from the Department of Biological Sciences and the Oceanographic Institute (No. 120), Florida State University. 124 Tulane Studies in Zoology Vol. 7 difficult ro find or to determine their nat- ural position in the host. The smaller para- sires in preserved shrimp are difficult to de- tect due to the lack of movement. Also the parasites, especially the gregarines, in the gut and digestive gland deteriorate rapidly. In order to fix the parasites in good condi- tion one should cut open the shrimp in the field before preserving. Tissues of the shrimp studied were teased apart and ex- amined with a dissecting microscope. The intestines were ait open, placed on a micro- scope slide, covered with a coverslip, and examined with a compound microscope. Details of methods will be given with the descriptions of individual parasites. All measurements are in millimeters unless stated otherwise; averages usually precede the ranges in parentheses. All illustrations were made from living specimens unless stated otherwise and were drawn with the aid of a microprojecror or camera lucida. Observations Subphylum Sporozoa Leuckart, 1897 Class Telosporidea Schaudinn, 1900 Order Gregarinida Lankester, 1866 Suborder Eugregarinia Doflein, 1901 Tribe Cephalina Delage and Herouard, 1896 Cephalolobus penaeus, gen. nov., sp. nov. (Figures 1-5) These gregarines were first studied alive in their natural habitat by removing the stomach and mounting the entire structure on a glass slide in physiological saline. The specimens were then fixed in AFA and mounted in Damar without staining. A few gregarines were stained with Ehrlich's acid hematoxylin. All measurements are in mi- crons and were made from 20 living speci- mens. Description. — Trophozoites divided into protomerite and deutomerite by ectoplasmic septum; young, solitary trophozoites up to 171 long by 45 wide, protomerite 46 long, deutomerite 125 long. Trophozoite asso- ciations 328 to 660 long by 66 to 125 wide; primite 118 to 231 long by GG to 158 wide, usually wider than satellite, protomerite of primite 33 to 85 long, deutomerite 99 to 198 long. Satellite 211 to 429 long by 33 to 118 wide; protomerite of satellite 26 to 52 long, deutomerite 211 to 376 long. No true epimerite observed; anterior end of protomerite modified into holdfast, consist- ing of sub-cylindrical cone with 15 to 40 irregular lobe-like processes around edge. Holdfast region in mature associative tro- phozoites usually set off by a constriction from rest of protomerite; holdfast used to attach young individuals and mature asso- ciative trophozoites to chitinous sheet be- tween ventral surface of the base of the terminal lappets and the filter. Primite of many individuals bent at about 45 degree angle. Usually one satellite attached to pri- mite, sometimes two or three satellites; pro- tomerite of satellite with no special lobed holdfast ( anterior surface of protomerite of dislodged satellite only slightly irregular). Endoplasm in deutomerite of primite and satellite more densely granular than that in protomerite of primite and satellite. Ecto- plasm a thick, hyaline layer around peri- phery of organism and on both sides of ectoplasmic membrane that separates pro- tomerite from deutomerite. Nucleus 33 to 56, in deutomerite only, containing 3 to 12 sperical granules. Single individuals and those in association capable of locomotion when dislodged from chitinous sheet. Spores, sporozoites, free sporonts and cysts not ob- served. Hosts. — Penaeus aztecus and P. duorarum. Location. — Young individual and mature associative trophozoites attached to chitinous sheet that extends from ventral surface of the base of the terminal lappets to the filter of the stomach. Locality. — Northern Gulf coast of Florida. Types. — Holotype and paratypes in Hel- minthological Collection of the Department Biological Sciences, Florida State University. Generic diagnosis of Cephalolobus. — Young individual and mature associative trophozoites septate, trophozoites associative and mature while attached in gut of host; one, two or three satellites in syzygy with primite, syzygy caudofrontal, satellite small- er than primite; development extracellular; true epimerite absent, anterior end of pro- tomerite modified into holdfast consisting of sub-cylindrical cone with irregular lobe- like processes on anterior edge; protomerite of satellite without specialized holdfast. In gut of marine decapods. Type species. — Cephalolobus penaeus. The trophozoites of C. penaeus were No. 4 Kruse: Parasites of Commercial Shrimps 125 Figures 1-5. Cephalolohus peiiaens, gen. nov., sp. nov. 1,2. Illustrations showing the method and place of attachment of the organisms on the thin chitincus sheet between base of the lappet and filter. 3. A multiple association with one primite and two satel- lites; note constriction of the protomerite which sets off the holdfast. 4. The most com- mon type of association with one primite and satellite. 5. A solitary trophozoite. 126 Tulane Studies in Zoology Vol. 7 found in 26 or 15.2 percent of 170 shrimp examined. Eleven or 17.7 percent of 62 P. aztecus were infected with an average of 7.7 trophozoites per shrimp whereas 15 or 20.8 percent of 72 P. duorarum were in- fected with an average of 4.5 trophozoites per host. The number of trophozoites per shrimp ranged from one to 21. The 36 P. setijenis examined were all found to be negative. Only one other genus, Anisolobus Vincent 1924, has been reported as lacking an epi- merite and having the anterior end of the protomerite developed into a lobed holdfast. The genus Cephalolobus differs from Ani- solobus in the following points: (1) The protomerite of Anisolobus is expanded con- siderably to form a few large, flat lobes which function as a sucker-like organ for attachment of the trophozoite to the cells of the host's gut wall; the protomerite of Cep- halolobus is not expanded but is a sub- cylindrical cone and is composed of many digitform lobes that attach the trophozoite by holding the thin chitinous sheet of the stomach between the lobes; ( 2 ) The very young trophozoites of Cephalolobus have the special holdfast whereas the young of Anisolobus do not; ( 3 ) Cephalolobus oc- curs in syzygy while in the attached tro- phozoite stage; Anisolobus occurs in syzygy only in the free sporont stage; (4) Spora- dins of Anisolobus are biassociative; the primite of Cephalolobus may have one, two, or three satellites; ( 5 ) The host of Ani- solobus is a coleopteran whereas Cephalolo- bus parasitizes a marine decapod. Cephalolobus penaeus also differs from most gregarines in being attached to a thin chitinous sheet from the very young stages to mature associations and thus presumably obtains all of its nourishment by absorbing through its body wall food in the host's stomach. An epimerite in the young stages of most gregarines remains inside the h(\st's gut cell and serves for attachment and ab- sorption of the cell contents for food. These gregarines lose their epimerites, then mature and become associative while free in the host's gut where they presumably absorb food through their body surfaces. The nature of the holdfast prevents as- signing Cephalolobus to a family at the present time; however, it is most similar to members of the family Gregarinidae Labbe, 1899. Kamm ( 1922 ), Grasse (1953) and Kudo { 1954) stated that the presence of a simple knob-like epimerite is one of the characters of the family Gregarinidae; Cep- halolobus and Anisolobus. therefore, do not belong in the family. If the life cycle of C. penaeus is proven to be similar to that of Anisolobus, both genera should be placed in a new family. Cephalolobus is somewhat similar to members of the family Dactylophoridae Leger, 1892 except that the dactylophorids are solitary and they are described as having an epimerite. However, Grasse (1953) be- lieved that the true epimerite is replaced in the young stages with protomerite exten- sions, and that the complex holdfasts of the dactylophorids are not epimerites but actu- ally modifications of the protomerite. This may be what occurs in Cephalolobus^ Family Porosporidae Leger, 1899 Nematopsis penaeus Sprague, 1954 (Figures 6-13 ) The sporonts were removed from the lu- men of the intestine with a needle, placed in sea water on a glass slide and studied alive under the microscope. Young tropho- zoites were observed by removing the entire intestine, cutting it lengthwise, flattening it on a slide, teasing it apart and then flatten- ing it slightly with cover glass pressure. Gametocysts were removed from the wall of the rectum and studied alive. Gym- nospores were liberated from gametocysts by slight cover glass pressure which broke the cyst wall. Some of the gymnospores were stained with Ehrlich's acid hematoxylin. All measurements are in microns and were taken from 35 ro 50 living specimens. It is the author's opinion that this species is N. penaeus Sprague, 1954 in the family Porosporidae Leger; although, as Sprague, (1954: 249) indicated, definite generic de- termination cannot be made until stages in the intermediate host are known, since it is on the basis of these that the genera (Porospora Schneider and Nematopsis Schneider) in the family Porosporidae are differentiated. Sprague described N. penaeus from P. aztecus collected off the Louisiana coast and he also found what seemed to be the same species in P. setiferus. Sprague's description did not include illustrations. Krme: Parasites of Commercial Shrimps 111 5JL 50p 3. 11 ® y Fig-ure 6-13. Nematopsis pcuaeus Sprag-ue. 6. Young- gametocyst. 7. Mature gymno- spore; drawn from preserved material. 8. Solitary sporont. 9-13. Mature sporonts show- ing various types of bifurcated and linear syzygy. The data and illustrations here supplement by 19 wide. Older sporadins in multiple Sprague's account. linear and bifurcated syzygy. Ectoplasmic Description. — Young trophozoites at- wall disappears between protomerite and tached to intestinal cells by simple, spherical, deutomerite of satellites. In bifurcated epimerite; solitary sporadins up to 70 long syzygy nuclei of both individuals move to 128 Tulane Studies in Zoology Vol. 7 stem portion. Smallest biassociation 120 long by 26 wide; multiple associations up to 590 long by 40 wide. Up to five indi- viduals in linear syzygy. Gametocysts spher- ical, attached to wall of rectum, 158 (99- 240) in diameter; mature spherical gym- nospores 6.5 (6.3-9-8) in diameter. Hosts. — Penaeus aztecus, P. cbiorarnm and P. setiferus. Location. — Young forms attached to cells of intestinal wall, sporonts and associations free in lumen of intestine, gametocysts at- tached to wall of entire length of rectum. Locality. — Alligator Harbor and Apalachi- cola Bay, Florida. At least one stage (young trophozoite, sporont or gametocyst ) was present in all of the approximately 60 shrimp that the author examined thoroughly. All three stages were usually present in most individuals of all three species of shrimp; however, in some shrimp only the very young tropho- zoites or mature gametocysts could be found. Sprague (1954) discussed the compari- sons and affinities of N. penaetts with other members of the family Porosporidae. He also presented evidence that N. penaeus has an intermediate host in its life cycle as do N. ostrearurn and N. prytherchi. This idea has been substantiated by the writer's work. Shrimp kept in laboratory aquaria for a week or longer lose infections of N. penaeus even if they live in close association with each other and are fed infected shrimp. This suggests that the spores from the gymnos- pores must undergo development in an in- termediate host before they are infective for the shrimp. The intermediate host must be .some organism that the shrimp feed on frequently, because in nature the shrimp are usually heavily infected with young and older stages, and the parasite does not re- produce while inside the shrimp host. Class Cnidosporidea Doflein, 1901 Order Microsporidia Balbiani, 1883 Family Nosematidae Labbe, 1899 Thelohania sp. (Figures 14-20) This species was studied alive with the phase contrast microscope and in fixed smears and sections. Smears, and cross and longitudinal sections of infected abdominal muscles were fixed in Bouin's and stained with Erhlich's acid hematoxylin. Extrusion of polar filaments was accomplished by exerting cover glass pressure on live speci- mens. All measurements are in microns and were taken from 30 living specimens. Description. — Preponderance of mature pansporoblasts with eight spores in living muscle smears; pansporoblasts spherical with thin, clear limiting membranes through which spores are visible; mature pansporo- blasts 10.2 (8.8-13-6) in diameter; spores in mature pansporoblasts 5.4 (6.4-4.0) long by 3-4 (3.0-4.0) wide, highly refractile with large vacuole at one end. Free, solitary spores 5.7 (6.1-4.7) long by 3.7 (3.0-4.0) wide, posterior end slightly wider than an- terior end, with large posterior vacuole. Polar filament extruded from anterior end 109 (97-142) long, almost uniform in width for entire length, tapering slightly at distal end. In stained smears structure of the sporonts and spores visible in more detail. Young developing sporonts with 1, 2, 4 or 8 darkly stained nuclei, cytoplasm light and finely granular. Eight nucleated stage followed by partitioning of cytoplasm into eight sections with a nucleus in each section destined to form a spore; this developmental stage of pansporoblast slightly larger in diameter than mature pansporoblast. Stained free spores with large clear vacuole at posterior end and smaller clear area at anterior end; darkly stained nucleus near middle, almost as wide as spore; lighter grey cone-shaped polar capsule projecting from nuclear area toward anterior end; anterior tip of polar capsule darkly stained at opening, remaining portion of spore stained a homogeneous light blue color. Hosts. — Penaeus aztecus, P. duorarum and P. setiferus. Location. — Mainly in muscles but also in other organs. Locality. — Alligator Harbor and Apalachi- cola Bay, Florida. The musculature of "cotton shrimp" is white with intermingled blue-black areas and lacks the firmness of normal muscle. Of the 4816 examined, 553 (11 percent) were grossly identifiable as infected. The incidence of infection in each host species was as follows: P. aztecus. 310 (16 per- cent) of 1874 examined; P. duorarum. 15 (1.9 percent) of 784; P. setiferus. 228 (10 percent; of 2158. P. duorarum had the low- No. 4 Kruse: Parasites of Commercial Shrimps 129 5p 5p 20 Figures 14-22. All figui-es drawn from preserved material except fig. 20. 14-20. Tlielo- hania sp. 14-17. 1-, 2-, 4-, and 8-nucleated developing sporcnts. 18. Developing pansporoblast. 19. Mature pansporoblast. 20. Living tree spore with polar filament extruded. 21-22. Juvenile of Contracaecam sp. showing anterior and posterior ends. 130 Tulane Studies in Zoology Vol. 7 est incidence of infection probably because the specimens examined were much younger than those of P. aztecus and P. setijerus. These figures obviously do not represent the true incidence of infection because the early and light infections cannot be detected with the unaided eye; in fact, the writer noted many such light infections during dissection of shrimp under the microscope. In a heavy infection most of the muscles and other organs of the body were para- sitized. Histological sections showed that the organisms saturate the muscle tissue, lying in large sheets or masses in and around the muscle fibers. Apparently all stages of development take place in the muscles. A satisfactory specific identification of this microsporidian cannot be made until it can be compared with Tbelohania penaei Sprague, 1950, T. hunterae Jones, 1958, and T. duorara Iversen and Manning (in press). Sprague did not give a detailed description of T. penaei which he found in the gonads of P. setijerus collected in the vicinity of Grand Isle, Louisiana. According to his description T. penaei differs from the pres- ent species in the following ways: (1) The spores of T. penaei average 4 by 2.2 whereas spores of the organism herein reported aver- age 5.7 by 3.7; (2) The polar filament of T. penaei averages 70 long and has a thick proximal half with a thinner distal half; the polar filament of the present species averages 109 long and is essentially uniform in diameter for its entire length; (3) T. penaei was found in the gonads only; the present one is mainly in the muscles. Jones' (1958; report on T, hunterae is an abstract (according to a personal com- munication he has a detailed paper in press ) and has very little information. T. hunterae is described as primarily a coelozoic parasite which sometimes invades the ovary and nerve cord, while the present specimens were found mainly in the muscles. Iversen and Manning permitted the author to re?d a paper by them (in press) describ- ing Thelohania duorara from the muscula- ture of P. duorarum collected from Biscayne Bay, Florida Bay and the Dry Tortguas region, Florida. Their description of T. duorara agrees well with the author's speci- mens except for polar filament length. The polar filament of 7 . duorara averages 38 in length whereas the polar filament of the present specimens average 109- Class Trematoda Rudolphi, 1809 Order Plagiorchiida La Rue. 1957 Family Opecoelidae Ozaki, 1925 Metacercaria of Opecoeloides jimbriatus (Linton, 1934) Sogandares-Bernal and Hutton (Figures 23-27) Synony??iy. — Distomum vitellosum Linton, 1901; Cymbephallus jimbriatus Linton, 1934; Fimbriatus jimbriatus (Linton, 1934) Von Wicklen, 1946. The metacercariae used in this study were first observed alive and then fixed in an alcohol-formalin-acetic acid solution (AFA). A few specimens were fixed under slight coverslip pressure; most were fixed without flattening in hot AFA so the acetabulum would be fixed in an extended position. The whole mounts were stained in Semi- chon's acetocarmine or Ehrlich's acid hema- toxylin. Serial frontal sections were cut at ten microns and stained with Ehrlich's acid hematoxylin and eosin. All measurements of metacercariae (30 encysted, 30 excysted) were taken from living specimens; those of adults are from Linton's work ( 1934, 1942). The cyst wall is thin, clear, about 1.7 microns thick and easily broken by active metacercariae. The size and shape of the cysts depends on the age of infection; the young cysts are almost spherical, the older ones almost sausage shaped. In the more mature cyst the worm is usually folded on itself. The average size of the cysts is about 0.68 by 0.25, the more mature cysts attain- ing 1.2 by 0.5. Worms removed from cysts average 0.84 by 0.3, the larger ones attain- ing 1.9 by 0.48. The morphology (fig. 23) is very similar to that of the adult except that the metacercariae apparently do not become sexually mature while encysted. Of 137 P. duorarum from Apalachicola Bay, Alligator Harbor and near the mouth of the St. Marks River every one was in- fected with this metacercaria. In contrast, none of 120 specimens of P. setijerus and 36 of P. aztecus from Apalachicola Bay were infected with this parasite. The three spe- cies of shrimp live closely associated in the same habitat and have been collected to- gether in a single trawl haul. All three shrimp species are undoubtedly exposed to No. 4 Kruse: Parasites of Commercial Shrimps 131 the cercariae of O. fimbriatus, yet in the present work, only P. duorarum has been found infected. Hutton et al (1959) found a single specimen of O. fimbriatus in P. setiferus from Apalachicola Bay and Wood- not as favorable for morphological studies as some in the writer's collection. The spe- cies is therefore redescribed from the type specimens and from whole mounts and serial sections of metacercariae obtained in the burn et al (1957) reported Trachypeneus present work. Data are also included from sp. as a host in southwest Florida. For a dis- cussion of the hosts and distribution of metacercariae and adults of this trematode the reader is referred to papers by Linton (1901, 1905, 1940), Woodburne/d/( 1957), Sparks (1958), Hutton et al (1959) and Sogandares-Bernal and Hutton (1959). The metacercariae were found in all stages of development in young and old shrimp indicating that the shrimp are being Linton (1934, 1942) and Hopkins (1941). Opecoeloides fimbriatus (Linton, 1934) Sogandares-Bernal and Hutton Description. — Body elongate, broadly rounded posteriorly, 1.28 to 4.62 long by 0.60 to 0.84 wide. Oral sucker well devel- oped, 0.12 to 0.26 in diameter, located on antero-ventral extremity. Ventral sucker larger than oral, 0.21 to 0.35 in diameter, continually infected in the bays or the shal- pedunculated, surrounded by raised border low coastal waters. There is some correlation made up of four lobes each having from five between the number of encysted metacer- to nine papillae; main body of sucker pro- cariae and age (determined by the size) truding as two prominent papillae at the of the shrimp. Table 1 is an acairate record junction of the four lobes. Small accessory of the number of these parasites recovered sucker with limiting membrane, anterior to from 75 shrimp examined from June ventral sucker, 0.08 to 0.04 in diameter, through September. The data indicate an Pharynx broadly ovate, 0.09 by 0.14; esoph- increase in the number of metacercariae per agus longer than pharynx; intestinal ceca host as the shrimp increases in size. Other extending to the posterior end and empty- data in this author's possession suggest what ing separately into the excretory vesicle or appears to be an increase in the number of opening separately to outside depending metacercariae in shrimp of the same size upon the contraction of the body. Genital range as the season progresses from spring atrium common, muscular, genital pore im- to fall. mediately in front of accessory sucker. Tes- AU details of anatomy are not visible in tes two, tandem, usually lobed, post-equa- the immature specimens. For this reason as torial; sperm ducts long, uniting at seminal well as to be reasonably certain of the in dentification of this metacercaria, the writer examined the adult type specimens. ( U. S. National Museum No. 8266) from which Linton (1934) described Cymbephallus fim- briatus. This study revealed certain errors in vesicle, 1/3 to 1/2 the distance from ovary to bifurcation of gut. Seminal vesicle com- posed of short bulbous posterior portion and narrow, elongate anterior portion which con- nects to muscular cirrus. Ovary anterior to fore-testis, usually not lobed; ootype and Linton's description. His specimens were Mehlis' gland slightly anterior to ovary; Table 1. Intensity of parasitism by Opecoeloides fimbriatus in Penaeus duorarum. of various s'zes Size Range of Number of Infected Number of Par£ 1 sites Per Shrimp Shrimp in mm Shrimp Examined Average Range 10-19 1 2 2 20-29 6 2.3 1-3 30-39 13 4 1-8 40-49 19 7.8 1-26 50-59 10 10.6 4-21 60-69 10 10.5 4-14 70-79 9 12.7 4-30 80-89 3 15 12-20 90-99 3 31 14-37 100-109 1 97 97 132 Tulane Studies in Zoology Vol. 7 uterus moderately folded, lying mostly in direct line between ovary and genital pore; eggs about 0.06 by 0.03- Vitelline reservoir at the dorsal, anterior border of the ovary; vitellaria filling the body posterior to the testes; extending laterally along the margins to a point about half-way between the ovary and ventral sucker. Excretory vesicle a long sac-shaped structure that extends anteriorly to anterior border of first testis, with two main collecting tubules emptying into the antero-lateral corners. Hosts. — Adults in Menticirrhiis saxatilis, M. amencanus, Bairdiella chrysura, Micropo- gon undulatus and Sciaenops ocellata. Meta- cercariae in Penaeus duoramm. P. setiferus and Trachypeneus constrictus. Location. — Adults in intestine, metacer- cariae in coverings and other tissues sur- rounding the digestive gland, stomach, heart, gonads, and intestine; also in tissue of the head region and tissues next to the lateral and dorso-lateral portions of carapace. Number. — Average number of metacer- carial cysts per shrimp: 10; range: 2 to 97. Locality. — Atlantic and Gulf Coasts of United States. The writer's description differs from Lin- ton's ( 1934) in the following ways: ( 1 ) A more complete description is given of the ventral sucker which in Linton's type speci- ments was retracted; ( 2 ) The ceca are described as emptying separately into the excretory vesicle which empties to the out- side rather than ending blindly as Linton described; ( 3 ) A description is given of the excretory vesicle which Linton omitted and which agrees with the description by Hop- kins (1941) of the same species; (4) The ejaculatory duct is described as opening into a common genital atrium along with the uterus rather than opening through the ac- cessory sucker as Linton described. The fact that this trematode has an ac- cessory sucker, ceca which empty into the excretory vesicle, a pedunculated acetabulum with papillae and no cirrus poucli places it in the genus Opecoeloides Odhner, 1928. Von "Wicklen (1946) reported that each cecimi opened separately to the outside. Consequently she established a new genus for this trematode, giving it the name Fim- hriattis jimbriatus. In the type specimens examined by the writer the vitellaria ob- scured the posterior ends of the ceca. Frontal sections and mounts of metacercaria showed that the ceca empty into the excretory vesi- cle. Sogandares-Bernal and Hutton (1959) observed that the posterior tips of the ceca moved to positions where each cecum and the bladder emptied separately to the out- side when the metacercaria was extended and then returned to the normal position of emptying into the excretory vesicle, when the worm contracted. The writer also ob- served this phenomenon (figs. 25-27) and saw particles pass from the ceca to the ex- cretory vesicle and then to the outside when the worm contracted, and directly to the out- side when extended. The writer fixed a group of 55 metacercariae with a consider- able amount of coverslip pressure. One of the 55 was fixed in the extended position with the ceca emptying separately to the outside and several were fixed in intermedi- ate positions. A specimen fixed in a flat- tened and extended position with the ceca emptying directly to the outside could ac- count for a description of this species as having two ani; however, the writer did not observe this condition in the type specimens. The remaining life cycle stages for Ope- coeloides jimbriatus are not known. Ope- coelid cercariae possess a stylet, lack eye- spots and body spines, have a short glandu- lar tail, possess a thick-walled excretory vesicle that has a very uneven lumen due to protrusion of large wall cells into it, have a 2[(2 + 2) + (2 + 2)] excretory formula, and develop within sporocysts in proso- branch snails (Cable 1956a, 1956b). The life cycle of Anisopoms rnanteri Hunninen and Cable, 1940, which Von Wicklen ( 1946) synonymized with Opecoeloides vitellosus (Linton, 1900) has been worked out by Hunninen and Cable ( 1941 ) and is prob- ably similar to the life cycle of O. fi??ibriatus. The adult of 0. vitellosus has been reported in over 60 species of marine fishes (Linton, 1934; Hunninen and Cable, 1940); however, Von "Wicklen (1946) stated that some of the trematodes identified as 0. vitellosus are other species. Tlic cercaria of O. vitellosus ocairs in the marine gastropod Mitrella lunata (Say) and the metacercaria in the marine amphipods, Carinogammarus mucro- natus (Say) and Amphithoe longimanu Smith. No. 4 Kruse: Parasites of Commercial Shrimps 24 133 25 26 27 Figures 23-27. Metacercaria of Opecoeloides fimbriatus (Linton) Sogandf res-Bernal and Hutton. All illustrations from preserved specimens. 23. Excysted matacercaria. 24. Schematic drawing showing- morphology of acetabulum, oblique view of surface. 25-27. Stages of contraction and extension cf metacercaria showing change in posi- tion of cecal openings. 25. Contracted position with cecal openings emptying into ex- cretory vesicle. 26. Intermediate position. 27. Extended position with ceca and excre- tory vesicle opening separately to the outside. 134 Tulane Studies in Zoology Vol. 7 Class Cestoidea Rudolphi, 1809 Order Trypanorhyncha Diesing, 1863 Family Eutetrarhynchidae Guiart, 1927 Prochristianella penaei, sp. nov. (Figures 28-34) Most of the observations and illustrations recorded herein were made from live plero- cerci. The live specimens showed the ar- rangement of hooks, bulbs, tentacles, and de- tails of the excretory system much better than preserved material. A small needle was used to break the cyst wall and to free the holdfast end of the body. Protrusion of the tentacles for study of hook arrangement was accomplished by exerting slight cover glass pressure on living specimens or by relaxing the specimens in saline solution at 4° C overnight. Some larvae were fixed in AFA and stained with Semichon's acetocarmine or Ehrlich's acid hematoxylin. The measure- ments are based on 20 worms taken from the three species of shrimp collected at various times during the summer and fall. All hook measurements were made near the middle of the tentacle. Description. — Encysted plerocerci cylin- drical, 1.1 (0.62-2.30) long by 0.52 (0.33- 1.03) wide; cyst wall clear, thin, 1.8 microns thick; holdfast region not withdrawn into blastocyst, surrounded only by cyst wall. Excysted pleroceri 1.8 (0.82-3.29) long by 0.46 (0.21-0.68) wide; holdfast region 0.82 (0.39-1.24) long by 0.15 (0.12-0.19) wide; blastocyst 1.0 (0.39-2.27) long by 0.46 (0.21-0.68) wide. Bothridia two, patelli- form; bothridial surfaces parallel to holdfast region with posterior margins free, not notched, 0.15 (0.12-0.17) long. Bulbs four. 0.44 (0.37-0.56) long by 0.04 (0.03-0.05) wide with mass of red granules at anterior end of each bulb. Proximal part of sheath sinuous. 0.40 ( 0.27-0.64 ) long; tentacles 0.60 (0.48-0.74) long by 0.02 (0.015- 0.03 ) wide at mid-point, covered with hooks. Hook arrangement heterocanthous, heteromorphous; metabasal armature com- posed of a single longitudinal row of large, stout hooks on internal surface of each ten- tacle and a series of obliquely ascending, half-turn spiral rows of smaller hooks that start on internal surface and continue dis- tally around bothridial and antibothridial surfaces until they meet on external surface; last few hooks of each half-turn spiral row meet on the external surface in an off-set inverted V where the hook of one ascend- ing row meets the opposite row between the last and next to last hooks. Large basal hooks about 10 microns long, with base about 8.5 microns long, short toe, long heel and recurved point; smaller hooks long, slender, spinous, and diminishing in size as they pass to external surface; usually seven to nine hooks in each ascending row. Hooks in each row becoming fewer, closer together and smaller toward distal end of tentacle; toward base of tentacle hooks in each row becoming more numerous, farther apart and larger, but the same basic arrangement per- sisting except at base. Proximal or basal portion of tentacle swollen with special ar- rangement of hooks: few, large, recurved hooks; many rows of long, slender, spinous hooks. Tentacles protruding through open- ings in antero-lateral corners of external surfaces of bothridia. Postbulbosal region Table 2. Incidence of infection of shrimp by the plerocercus of Prochristianella penaei P. aztecus P. duorarum P. setiferns Number of Shrimp Examined 128 137 36 Number of Shrimp Infected 116 132 34 Percentajje of Shrimp Infected 90.6 97 94.4 Average Number of Parasites per Shrimp 6.2 7.0 14.7 Range of Numbei- of Parasites per Shrimp 0-42 0-37 0-26 Average Size of Shrimp Examined 110.4 mm 52.4 mm 114.7 mm No. 4 Kruse: Parasites of Commercial Shrimps 135 28 30 E E fife 9 #' • - i«4 o. » 32 a IT) CM Fig-ures 28-34. Plerocercus of Prochristianella penaei, sp. nov. 28. Excysted plerocer- cus. 29. Encysted plerocercus. 30-34. Portions of tentacles showing arrangement and morphology of hooks. 30. Metabasal armature of internal surface showing' ascending, oblique, half-turn spiral rows; numbers 1-9 represent one bothrial row, numbers 1-7 the opposite row; b is the large, recurved, basal hook. 31. Same as fig. 30, external sur- face. 32. Side view of metabasal armature drawn from a preserved specimen showing that preservation shrinks the tentacle, drawing the hooks closer together. 33. Armature of internal surface of basal swelling. 34. Armature of external surface of basal swelling. 136 Tulane Studies in Zoology Vol. 7 (between bulbs and blastoq^st) short. Blastocyst muscular, divided into two regions by constriction; anterior portion filled with numerous, dark, granular bodies, 50 microns in diameter; posterior portion filled with small homogeneous granules. Blastocyst usu- ally attached to host tissue or another cyst at constriction. Hosts. — Plerocerci in Penaeus aztecus, P. duorarurn and P. setiferus. Location. — Digestive gland and tissues cent infected with a high incidence in all three host species. The average number of parasites per shrimp was 8.1, and the range was 0 to 42. This trypanorhynchan larva was found in shrimp collected from each of the three collections areas. Alligator Harbor, near the mouth of the St. Marks River, and Apalachicola Bay. Table 3 presents data indicating an in- crease in the number of plerocerci as the age (indicated by size) of the shrimp in- surrounding digestive gland and stomach, creases. This is probably due to two reasons; blastocysts of larvae frequently penetrating first, the shrimp are being infected continu- wall of digestive gland. Locality. — Northern Gulf coast of Florida. Types. — Holotype will be deposited in the U. S. N. M. Helminthological Collection; paratypes in the Helminthological Collec- tion of the Department of Biological Sci- ences, Florida State University. Table 2 presents data on incidence of ally, thus building up the infection; second, the larger shrimp are usually collected dur- ing the fall when a higher rate of infection may be taking place. The second explana- tion is supported by data from shrimp of the same species and same size range in the early summer and late fall. Shrimp in the fall show a considerable increase in the aver- infection. The writer examined 301 shrimp age number of parasites per individual over for this cestode and found 282 or 93.6 per- those collected in early summer. However, Table 3. Inteiisity of parasitism by Prochristianella penaei in shrimp of various sizes Numbei • of Parasites Per Size Range of Shrimp in mm Number of Infected Shrimp Shrimp Species Averages Ranges P. aztecus 60-69 1 7 — 70-79 1 6 80-89 8 6 4-12 90-99 14 5.3 2-17 100-109 18 5.6 1-11 110-119 24 6.7 1-31 120-129 11 9 3-42 130-139 7 6 1-23 140-149 2 5.5 2-9 150-159 1 8 P. duorarurn 10-19 1 3 20-29 6 2.6 1-4 30-39 13 3.1 1-7 40-49 16 2.9 1-17 50-59 10 5.2 4-18 60-69 11 6.6 4-14 70-79 9 6.5 3-21 80-89 3 16.3 12-20 90-99 3 22.3 14-37 100-109 1 20 — P. setiferus 60-69 2 9 6-12 70-79 2 6.5 3-10 80-89 1 8 — 90-99 2 5 4-6 100-109 2 10 8-12 110-119 4 19.2 17-24 120-129 6 13.3 4-26 130-139 3 18.3 17-21 140-149 2 11 8-14 No. 4 Kruse: Parasites of Commercial Shrimps 137 F. aztecus, which live in the bay and shallow coastal waters mainly during the late spring and early summer, have a lower percentage and lighter intensity of infetcion than P. duorari/vi and P. setifems, which live in the bay and shallow coastal waters mainly dur- ing late summer and fall ( Table 2 ) . Possi- ble explanations of this difference are a higher resistance in P. aztecus than in the other two species or more mature procer- coids and first intermediate hosts during the late summer and fall. Woodburn et al (1937), Sparks and Mackin (1957), and Button et al (1959) give accounts of the known hosts and dis- tribution of this cestode. The genus Prochristianella was established by Dollfus (1946) to accommodate species in the family Eutetrarhynchidae which have a basal tentacular swelling and either hetero- morphic or hemoeomorphic armature. He included two species in the genus, the type species P. trygonicola Dollfus, 1946 [adults from spiral valve of Dasyatis pastinaca (L) and plerocercus from the digestive gland of Upogebia stellata ( Montagu ) ] and Rhyn- chobothrium tenuispine Linton, 1890 [adults from Dasyatis centroura (Mitchill)}. P. penaei differs considerably from P. trygoni- cola and P. tenuispinis (Linton 1890) in arrangement and morphology of hooks. The row of large, recurved hooks in P. penaei is lacking in the other two species; the metabasal armature of P. tenuispinis is com- pletely homeomorphic; and in P. trygonicola each half-turn spiral rows starts with short slender hooks on the internal surface, the hooks becoming long and slender on the bothridial and antibothridial surfaces and then becoming short and slender again on the external surface. Although no complete life cycle has been elucidated in the Order Trypanorhyncha, adults are known to occur in the spiral valve of selachians. The work of Ruzkowski (1932, 1934) {fide Dollfus, 1942) as well as other bits of evidence, indicate the fol- lowing course of events: the eggs are re- leased into the sea when the selachian defe- cates, a free-swimming coracidium hatches from the egg; the coracidium is eaten by the first intermediate host (copepod) and then develops into a procercoid; the pro- cercoid is eaten, along with the first inter- mediate host, by a second intermediate host ( fish, mollusk, or decapod ) in which the procercoid develops into a plerocercus; the plerocercus is eaten, along with the second intermediate host, by a selachian; the plero- cercus then develops into an adult in the selachian. The writer autopsied three specimens of Dasyatis sabina (LeSueur), two of Rhinop- tera bonasus (Mitchill) and one of Raja texana Chandler, all of which feed extensive- ly on shrimp. Their stomachs were crowded with shrimp that still contained live plero- cerci; however, no adult Prochristianella penaei were found in their intestines. Ex- amination of shrimp stomachs for parasites frequently revealed what appeared to be remnants of copepods, amphipods, and os- tracods, crustaceans which could very pos- sibly serve as first intermediate hosts for P. penaei. Parachristianellamoi\ovcieg3iCSini\\3., sp. nov. (Figures 35, 36) The plerocerci were first studied alive and then fixed in AFA and stained in Semi- chon's acetocarmine. Protrusion of the ten- tacles was accomplished by exerting slight cover glass pressure. Only two specimens were recovered, both from P. duorarinn. One was accidentally destroyed, therefore, all measurements were taken from the remain- ing specimen. The holdfast regions of both specimens were free and active by the time the parasites were recovered; therefore, no information is available on the nature of the encysted plerocercus. A total of 301 shrimp was examined, 137 of which were P. duorarnm. Description. — Plerocercus large, stout 3.82 long; body divided into distinct an- terior holdfast portion and posterior blasto- cyst region; holdfast sub-cylindrical, 1.13 long by 0.198 wide; bothridia two, patelli- form, 0.128 long, slanted toward mid-point anteriorly but not confluent, posterior mar- gins not free. Bulbs four, 0.56 long by 0.03 wide; with mass of red granules at anterior end of each bulb; sheath 0.46 long; ten- tacles protruding at antero-lateral corners of external surface of bothridia, 0.77 long by 0.028 wide at mid-point; armed with hooks which are heterocanthous and heteromor- phous. Metabasal armature a sequence of oblique, ascending, half-turn spiral rows of 138 Tulane Studies in Zoology Vol. 7 P^'ig'ures 35-40. 35-36. P. monomegacantha,sp. nov. 35. Excysted plerocercus. 36. Meta- basal armature of tentacle; hooks numbered 1-14 represent one ascending-, oblique, half- turn spiral row and 1-11 the opposite row. 37-40. P. dinwfjaccoitlKi, sp. nov. 37. Excyst- ed plerocercus. 38. Side view of tentacle from preserved specimen. 39. Metabasal arm- ature of internal surface; hooks numbered 1-6 represent proximal portion of half-turn spiral row; hooks 1 and 2 are large, basal, recurved hooks. 40. Metabasal armature of external surface; hooks numbered 3-11 repi'esent distal portion of half-turn spiral row. hooks which start on internal surface of ternal surface where the last hook of one tentacle and cross to external surface alter- nately around the bothridial and antibothri- dial surface; last hooks of each half-turn spiral making an off-set inverted V on ex- ascending spiral row ends at a level slightly below the second from last hook of the opposite spiral row. First hook of each half- turn spiral large, about 19 microns long, No. 4 Kruse: Parasites of Commercial Shrimps 139 with base about 14 microns long, short toe, mediate and the definitive hosts are differ- long heel and a recurved point; these first ent; however, the extremely low incidence hooks arranged in two alternate rows on of infection in shrimp with P. monomega- internal surface; other hooks very long, cantba indicates that shrimp may not be the slender and spiny; hooks gradually diminish- only, or even the usual, intermediate host ing in size as they pass to external surface, for this species, usually about 11 to 14 hooks in each half turn spiral at mid-region of tentacle. Same Parachnstianella dimegacantha, sp. nov. arrangement of hooks persisting for entire ( rigures 3/-4U) metabasal armature; toward base of the ten- One specimen of P. dimegacantha was re- tacle the hooks in each row becoming large, covered; it was first studied alive, then fixed more numerous and farther apart; toward in AFA and stained with Semichon's aceto- the distal end the hooks in each row becom- carmine. Protrusion of the tentacles was ing smaller, fewer and closer together. Base accomplished by placing the larva in saline of tentacle not swollen, basal armature re- solution at 4° C for 24 hours. All measure- duced, consisting of a few small, slender ments were made from this individual, hooks. Postbulbosal region very short, 0.112 P. dimegacantha was recovered from P. long; blastocyst large, granular, muscular, 2.8 duorarum. A total of 301 shrimps was ex- long by 0.70 wide at widest portion near amined, 137 of which were P. duorarum. anterior end. The holdfast was free and active when the Host. — Plerocercus in Penaeus duorarum.. specimen was recovered; therefore, no in- Location. — In digestive gland, with blasto- formation is available on the encysted stage, cyst protruding through wall of digestive Description.. — Plerocerus large, stout, gland. 3 28 long; body divided into distinct an- Locality. — Northern Gulf coast of Florida, terior holdfast region and posterior blasto- Type. — Holotype in author's collection. cyst region; holdfast sub-cylindrical, 1.05 Dollfus (1946) established the genus long by 0.211 wide. Bothridia two, small, Parachristianella to include the species in the patelliform, 0.132 long, slanted toward mid- family Eutetrarhynchidae Guiart, 1927 that point anteriorly but not confluent, posterior have heteromorphic metabasal armature and edges not free; bulbs shorter than sheath or no basal swelling of the tentacle. The only postbulbosal region, 0.297 long by 0.039 species he placed in the genus was the type wide; mass of red granules at anterior end species P. trygonis from the spiral valve of of each bulb; sheaths slightly sinous, 0.326 Dasyatis pastinaca (Linnaeus) from Con- long. Tentacles protruding at antero-lateral carneau, France. P. mono me gac ant ha is the borders of external surface of bothridia, second species and strengthens the validity 0.389 long by 0.021 wide at mid-point, of the genus. The plerocercus of P. trygonis armed with hooks. Hook arrangement he- was described (Dollfus, 1946) from Vpo- terocanthus, heteromorphic; metabasal arma- gebia stellata ( Montagu ) collected in Ar- ture a series of obliquely ascending, half- cachon, France. turn spiral rows that start on the internal P. monomegacantha differs from P. try- surface and pass alternatively across the gowij- in the following points: (1) P. tnono- bothridial and antibothrial surfaces to the megacantha has two rows of large, recurved external surface; last hooks of each half-turn hooks with long bases on the internal sur- spiral meeting on the external surface in an face of the tentacle whereas P. trygonis has off-set inverted V where the last hook of one four rows of such hooks; ( 2 ) The posterior ascending row meets the opposite row be- margins of the bothridia are not as free in tween the last and next to the last hooks. P. monomegacantha and the bothridia are The first two hooks of each ascending row more closely approximated anteriorly than about 14 and 10 microns long respectively, in P. trygonis: ( 3 ) The blastocyst in P. with bases about 9 and 7 microns long, with monomegacantha is not as well differenti- short toes, long heels and recurved points; ated into anterior and posterior regions as these first hooks arranged in two alternate in P. trygonis; (4) The overall size of the rows. Other hooks long, slender, spinous, plerocercus of P. trygonis is greater than and diminishing in size as they pass to ex- that of P. monomegacantha: (5) The inter- ternal surface; usually 9 to 12 hooks in each 140 Tulane Studies in Zoology Vol. 7 half-turn spiral at middle of tentacle. Same arrangement of hooks persists for entire metabasal armature; toward base of tentacle hooks in each row becoming larger, more numerous and farther apart; toward the dis- tal end hooks in each row becoming smaller, fewer and closer together. Base of tentacle not swollen, basal armature reduced, consist- ing of a few small, slender hooks. Post- bulbosal region relatively long, 0.415; blasto- cyst large, granular, muscular, 2.21 long by 1.37 wide at widest portion near anterior end. Host. — Plerocerais in Penaeus duor arum- Location. — In digestive gland, part of blastocyst protruding through wall of di- gestive gland. Locality. — Alligator Harbor ,Florida. Type. — Holotype in writer's collection. P. dimegacantha is the third species in the genus and differs from the plerocercus of P. trygonis in the following points: ( 1 ) P. dimegacantha has fewer hooks in each ascending half-turn spiral row; the rows are arranged differently and meet on the external surface in a different manner; ( 2 ) The bulbs of P. dimegacantha are less than a third of the length of the holdfast, whereas in P. trygonis they are almost one half the length of the holdfast; ( 3 ) The bothridia of P. dimegacantha are closer to- gether at the anterior end and the posterior margins are not as free; { 4 ) The postbul- bosal region is longer in P. dimegacantha; however, this distance may not be constant; ( 5 ) The blastocyst of P. dimegacantha is not sharply divided into an anterior and posterior region as in P. trygonis: { 6 ) The over-all size of the plerocercus of P. trygonis is much larger than that of P. dimegacantha: (1) The intermediate and definitive hosts are different; however, the extremely low incidence of infection in shrimp with P. dimegacantha indicates that shrimp may not be the only, or even the usual, intermediate host for this species. P. dimegacantha and P., monomegacantha (described above) differ considerably in hook arrangement ( figs. 36, 38-40 ) and in the relative sizes of the bothridia, bulbs, and postbulbosal regions. Cestode Larva The specimens were observed alive and then stained with Semichon's acetocarmine and mounted in Damar for further study. All measurements were taken from 20 pre- served specimens; other details of the fol- lowing description are from living speci- mens. Description. — Small, solid, pyriform, larva; body 0.125 (0.112-0.132) long by 0.066 { 0.052-0.085 ) wide. Well developed apical sucker, 0.033 (0.026-0.029) long, by 0.046 (0.039-0.052) wide. Four well developed glands immediately posterior to apical sucker, each with a separate duct leading forward, passing through sucker wall and emptying into sucker cavity. Small sac-like osmoregulatory bladder near posterior end of body emptying to outside through prom- inent pore; flame cells and excretory tubules asymmetrical in arrangement, three flame cells maximum number seen in one individ- ual. Body cuticle covered with closely ar- ranged rows of very fine spines. Thirteen to 19 large calcareous corpuscles scattered throughout body; rest of body filled with smaller homogeneous granules. Hosts. — Penaeus aztecus and P. duorarum. Location. — Internal lining of mid-intestine. Locality. — Alligator Harbor and Apalachi- cola Bay, Florida. Number. — 27 (14.3 percent) of the 188 shrimp examined were infected with this parasite. Twenty-one (16.4 percent) of the 128 P. aztecus and six (10 percent) of the 60 P. duorarum harbored this cestode. From 27 to 122 of these parasites were found in every infected shrimp. Lack of specific structures rendered fur- ther identification of this larva uncertain. According to Wardle and McLeod ( 1952), larval forms of this type have been reported from other marine invertebrates such as pearl oysters, other lamellibranchs, and me- dusae; they have also been recorded from fishes. None of these larvae have been cor- related with an adult. Hutton et al ( 1959) tentatively referred what appears to be the same larva to the Lecanicephala. Class Nematoda (Rudolphi, 1809) Cobb, 1919 Order Rhabditida Chitwood, 1933 Family Ascarididae Blancliard, 1896 Juvenile of Contracaecum sp. ( Figures 21, 22 ) These juvenile nematodes were studied alive and then fixed in hot ten percent for- malin for further study. All measurements No. 4 Kruse: Parasites of Commercial Shrimps 141 are based on ten preserved specimens, four specimens each from Penaeus duoranmi and P. aztecHs, and two from P. setijerus. Description. — Body slender, tapering gradually anteriorly and more abruptly pos- teriorly, 1.74 (1.32-2.44) long by 0.08 (0.06-0.10) wide at widest portion near middle. Cuticle finely annulated, annula- tions 5 to 9 microns wide. Lips three, about equal in size, 0.020 (0.016-0.023) long by 0.015 (0.014-0.018) wide, (dorsal lip often slightly shorter than ventro-lateral lips); large cuticular boring tooth on one ventro- lateral lip. Esophagus muscular 0.396 (0.315-0.462) long by 0.029 (0.028-0.030) wide at mid-point, no distinct bulb, en- larged slightly posteriorly, ventriculus 0.029 (0.021-0.035) long by 0.029 (0.022-0.037) wide; ventricular diverticulum well devel- oped, 0.277 (0.169-0.315) long by 0.028 (0.016-0.035) wide at mid-point. Intestine long, very thick walled, filling most of body cavity, 0.050 (0.042-0.065) wide at mid- point; intestinal diverticulum projecting an- terior to ventriailus, 0.118 (0.107-0.126) long by 0.020 (0.014-0.029) wide at mid- point; the intestine narrowing abruptly pos- teriorly; well developed sphincter muscle separating intestine from short, narrow rec- tum; rectum provided with three pairs of well developed rectal glands, two pairs dorsal to rectum, one pair ventral; anus 0.098 (0.083-0.113) from posterior end. Nerve ring 0.112 to 0.161 from anterior end; excretory pore 0.130 to 0.190 from an- terior end. Sex of individual juvenile nema- todes could not be determined. Hosts. — Penaeus aztecus. P. duorarum and P. setijerus. Location. — In digestive gland and tissues surrounding digestive gland and stomach, not encysted. Locality. — Alligator Harbor and Apalachi- cola Bay, Florida. Specimens on deposit. — U. S. N. M. Hel- minthological Collection No. 37269- The writer examined 255 shrimps col- lected from June to November, 1958, for nematodes and found 14 parasitized with an average of 3.1 per host. Table 4 gives data on the parasitism in each species of shrimp. Juveniles of the genus Contracaecum Railliet and Henry, 1912 have been found in fishes and a variety of invertebrates such as medusae, Sagitta, copepods, amphipods, and cephalopods ( Hyman, 1951). The adults are typically parasites of the digestive tract of fishes, fish-eating birds, and mam- mals. To the writer's knowledge there is only one record of an adult Contracaecum sp. infecting a prawn, Pandalus borealis Kroyer, and this was from the coast of British Columbia, Canada (Margolis and Butler, 1954). These authors concluded that the prawn was an abnormal definitive host and tentatively identified the nematode as Contracaecu7n aduncinn ( Rudolphi, 1802). Woodburn et al (1957) reported nema- todes in P. duorarum from the Tortugas region and in a "white shrimp" from Apa- lachicola Bay, but did not identify them further. Hutton et al (1959) identified a nematode of the genus Contracaecum in P, duorarum from southwest Florida. Summary 1. A total of 301 shrimps of the species Penaeus aztecus. P. duorarum and P. seti- jerus was examined for parasites. These were collected from Apalachicola Bay, Alli- gator Harbor and near the mouth of the St. Marks River, Florida during the summer and fall of 1958. 2. Cephalolobus penaeus, gen. nov., sp. Table 4. Incidence of parasitism in shrimp by juveniles of Contracaecum sp. P. aztecus P. duo)arum P. setiferus Number of Shrimp Examined 128 91 36 Number of Shrimp Infected 3(2.3%) 9(9.79r) 2(5.57.) Average Number Parasites per Shr of imp 4 1.7 4 Range of Number Parasites per Shr: of imp 1-10 1-3 2-6 142 Tulane Studies in Zoology Vol. 7 nov. ( Eugregarinida Dolfein ) is described from Penaeus aztecus and P. duorarum. It was found attached to a thin, chitinous sheet in the stomach of 17.7 percent of the P. aztecus (average of 1.7 parasites per host) and 20.8 percent of the P. duorarmn (aver- age of 4.5 parasites per host). This species differs from most cephaHne gregarines by the absence of a true epimerite; the anterior end of the protomerite is modified into a lobed holdfast. 3. Nematopsis penaeus Sprague, 1954 (Eugregarinida Doflein) is reported from Penaeus aztecus, P. duorarum and P. seti- ferus. Every shrimp examined was found to harbor at least one stage of this gregarine; the young trophozoites were attached to the gut cells, the sporonts were free in the gut and the gametocysts were encysted on the wall of the rectum. Additional details of morphology are added to Sprague's descrip- tion. 4. A microsporidian of the genus Tbelo- bania is described from the musculature of Penaeus aztecus, P. duorarum and P. seti- ferus. Of the 4816 shrimp examined, 553 (11 percent) were grossly identified as "cotton shrimp". The microsporidian ap- parently is not T. penaei Sprague, 1950 re- ported from P. setiferus; comparison with T. hunterae Jones, 1958 and T. duorara In- versen and Manning (in press) must be made before specific identification. 5. The metacercaria of the trematode Opecoeloides fimhriatus (Linton, 1934) Sogandares-Bernal and Hutton is reported and described from the pink shrimp, Pe- naeus duorarum. This parasite was found encysted in the tissue surrounding the di- gestive gland and stomach. Penaeus duo- rarum was the only shrimp that harbored this trematode, and all specimens examined were positive. The average number of para- sites per shrimp was ten, with a higher aver- age number during the fall than during the summer. O. fi?nbriatus is redescribed from the type specimens and from new informa- tion obtained during the study of the meta- cercariae. 6. Prochristianella penaei is described as a new species of Trypanorhyncha from plerocerci found in the digestive gland of Penaeus aztecus, P. duorarum, and P. seti- ferus. The plerocerci were found in 93.6 percent of the shrimp with an average of 8.1 per host. The number of parasites per shrimp increased with the age of the host and toward the fall of the year. P. penaei differs from the other two species in the genus, P. trygonicola Dollfus, 1946 and P. tenuispinis (Linton, 1890), chiefly in the arrangement and structure of hooks of the metabasal armature of the tentacles. 7. Paracbristianella monomegacantba and Paracbristianella dimegacantba are described as new species of Trypanorhyncha from plerocerci recovered from the digestive gland of P. duorarum. Only two specimens of P. monomegacantba and one of P. dime- gacantba were found in 137 P. duorarum. The two species differ considerably from each other and from P. trygonis Dollfus, 1946 (the only other species in the genus) in the arrangement and structure of hooks of the metabasal armature of the tentacles. 8. A small unidentified cestode larva from the intestinal lining of Penaeus aztecus and P. duorarum is reported. From 27 to 122 of these parasites were found in the infected shrimp; 14.3 percent of the hosts examined were infected. 9. An immature nematode in the genus Contracaecum is reported from Penaeus aztecus, P. duorarum and P. setiferus. This parasite was found in the digestive gland and tissues surrounding the stomach of 5.4 percent of the shrimp examined with an average of 3-1 per host. Acknowledgements The writer wishes to thank his major professor. Dr. Robert B. Short, for his un- selfish aid and many helpful criticisms dur- ing the course of this work. The writer also thanks Mr. Robert M. Ingle of the Florida State Board of Conservation for suggesting a study of the parasites of shrimp and for aid in collecting specimens. Dr. Franklin Sogandares-Bernal, Department of Zoology, Tulane University, offered several helpful suggestions. Mr. Allen Mcintosh, Dr. M. B. Chitw(X)d of the Animal Disease and Para- site Research Branch, ABS, USDA, Belts- ville, Md. and Dr. R. Ph. Dollfus of the Museum National D'Histoire Naturelle, Paris aided in identification of specimens. References Cited Anderson, W. W. and M. J. Lindner 1943. A provisional key to the family Penae- idae with -special reference to American No. 4 Kruse: Parasites of Commercial Shrimps 143 forms.. Trails. Amer. Fish. Soc, 73: 284: 319. Cable, R. M. 1956a. Opistholebes diodontis n. sp., its development in the final host, the affinities of some amphistomatous trematodes from mai'ine fishes and the allocreadioid problem. Parasitology, 46 (1-2) : 1-13. 1956b. Marine cercariae of Puerto Rico. Scientific Survey of Puerto Rico and the Virgin Islands. N. Y. Acad. Sci., 16(4) : 491-577. DOLLFUS, R. Ph. 1942. Etudes critiques sur les tetrarhynques du museum de Paris. Arch. Mus. Nat. Hist. Nat., Paris, 19: 7.466. 1946. Notes diverses sur des tetrarhynques. Mem. Mus. Nat. Hist. Nat., Paris, 22(5) : 179-220. Grasse, p. p. 1953. Traite de Zoologie. Anatomic, Systematique, Biologic, Tome I. Fasc. II. Protozoaircs: Rhizopodes, Ac- tinopodes, Sporozoaircs, C)iidosporides. Masson et cie, Paris. 1160 p. Hopkins, S. H. 1941. The excretory sys- tems of Helicometra and Cymhcphallus (Trematoda), with remarks on their re- lationships. Trans. Amcr. Mioos. Soc, 60(1) : 41.44. HuNNiNEN, A. V. and R. M. Cable 1941. Studies on the life history of Anisoporns manteri (Trematoda: Allocreadiidae) . Biol. Bull., 80(3) : 415-428. HuTTON, R. F., F. Sogandares-Bernal, Bon- nie Eldred, R. M. Ingle, and K. D. WooDBURN 1959. Investigations on the parasites and diseases of saltv^^ater shrimps (Penaeidae) of sports and com- mercial importance to Florida. Fla. St. Bd. Cons., Tech. Ser., No. 26: 1-38. Hyman, Libbie H. 1951. The Invertebrates: Acanthocephala, Aschelminthes, and En- toprocta. The Pseudocoelomate Bilateria. McGraw-Hill, New York. 572 p. Jones, E, E. 1958. Microsporidiosis in shrimp. Assoc. Southeastern Biol. Bull., 5(1) : 10. Kamm, Minnie (Watson) 1922. Studies on gregarines II. Synopsis of the polycystid gregarines of the world, excluding those from the Myi-iapoda, Orthoptera and Co- leoptera. Illinois Biol. Monogr., 7(1): 1-104. Kudo, R. R. 1954. Protozoology. 4th Ed. Charles C. Thomas, Springfield, 111. 966 p. Linton, Edwin 1901. Parasites of fishes of the Woods Hole region. Bull. U. S. Fish Comm. (1899), 19: 405-492. 1905. Parasites of fishes of Beaufort, North Carolina. Bull. U. S. Bur. Fish. (1904), 24: 321-428. 1934. A new genus of trematodes belonging to the subfamily Allocreadiinae. Jour. Wash. Acad. Sci., 24(2): 81.83. 1940. Trematodes from fishes mainly from the Woods Hole re- gion, Massachusetts. P)-oc. U. S. Nat. Mus., 88: 1-172. Margolis, Leo and T. H. Butler 1954. An unusual and heavy infection of a prawn, Pandalus borcalis Kroyer, by a nematode, Contracaecum sp. Jour. ParasitoL, 40(6) : 649-655. RuszKOWSKi, J. S. 1932. Etudes sur le cycle evolutif et sur la structure des cestodes marins. III. Le cycle evolutif du tetra- rhynque Grillotia erinaceus (Van Beneden, 1858) Acad. Polon. Sci. et Lett., Compt. Rend. Mens. CI. Sci. Math, et Nat., 9: 6. 1934. Etudes sur le cycle evolutif et sur la structure des cestodes de mer. III. Le cycle evolutif du tetra- rhynque Grillotia erinaceus (Van Beneden, 1858). Mew. Acad. Polon. Sci. et Lett., Cracovie, CI. Sci. Math, et Nat., s. B.: Sci. Nat., 6: 1.9. Sogandares-'Bernal, F. and R. F. Hutton 1959. Studies on helminth parasites from the coast of Florida. III. Digenetic tre- matodes of marine fishes from Tampa and Boca Ciega Bays. Jour. ParasitoL, 45: 337-346. Sparks, A. K. 1958. Some digenetic tre- matodes of fishes of Grand Isle, Louisi- ana. Proc. La. Acad. Sci., 20: 71-82. Sparks, A. K. and J. G. Mackin 1957. A larval trypanorhynchid cestode from com- mercial shrimp. Texas Jour. Sci., 9(4) : 475-476. Sprague, Victor 1950a. Notes on three microsporidian parasites of decapod Crus- tacea of Louisiana waters. Occ. Pap. Mar. Lab., Louisiaiia State Univ., 5: 1-8. 1950b. Studies on Nema^ topsis prytherchi Sprague and N. ostre- arum, emended. Mimeographed for pri- vate distribution by Texas A & M Re- search Foundation. 59 p. 1954. Protozoa, In : Gulf of Mexico, its origin, waters and marine life. Fish. Bull. U. S. Fish and Wildlife Sew., 55: 243-256. and P. E. Orr, Jr. 1955. Nematopsis ostreamm and N. prytherchi (Eugregarinida: Porosporidae) with spe- cial reference to the host-parasite rela- tions. Jour. ParasitoL, 41(1): 89-104. Vincent, Mary 1924. On a new gregarine Anisolobus dacnecola n. g., n. sp. A para- site of Dacne rufifrons, Fabr. (Coleop- tera). Parasitology, 16(1) : 44-47. Von Wicklen, Jane H. 1946. The trema- tode genus Opecoeloides and related gen- era, with a description of Opecoeloides polynemi n. sp. Jour. ParasitoL, 32(2): 156-163. Voss, G. L. 1955. Key to the commercial and potentially commercial shrimp of the family Penaeidae of the Western North Atlantic and the Gulf of Mexico. Fla. St. 144 Tulane Studies in Zoology Vol. 7 Bd. Cons., Tech. Ser., No. 14: 1-23. Wakdle, R. a. and J. A. McLeod. 1952. Zoology of Tapeworms. Univ. Minn. Press., Minneapolis. 780 p. WooDBURN, K. D., Bonnie Eldred, Eugenie Clark, R. F. Hutton, and R. M. Ingle 1957. The live bait shrimp industry of the West Coast of Florida (Cedar Key to Naples). Fla. St. Bd. Cons., Tech. Ser., No. 21: 1.33. Abstract A total of 301 shrimp of the species Penaeus aztecus, P. diiorarum and P. setiferiis was examined for parasites. The shrimp were collected in the sum- mer and fall of 1958 from the northwest Gulf coast of Florida. A new genus and species of gregarine, Cephalolobus penaeus, is described from P. aztecus and P. duorarnni. It was found at- tached to a thin chitinous sheet be- tween the filter and the base of the terminal lappets of the stomach. Nema- topsis penaeus Sprague, 1954 is re- ported from all three species of shrimp and additional details of morphology are added to Sprague's description. Every shrimp examined harbored at least one stage of this gregarine; the young trophozoites were attached to the gut cells, the sporonts were free in the gut and gametocysts were encysted on the wall of the rectum. A micro- sporidian of the genus Thelohania is reported from the musculature of all three species of shrimp. The metacer- caria of the trematode Opecoeloides fimhriatus (Linton, 1934) Sogandares- Bernal and Hutton is reported from the pink shrimp, Penaeus duorarum. These parasites were found encysted mainly in the tissue surrounding the digestive gland and stomach. This spe- cies is redescribed from type specimens and metacercariae. The Trypanorhyn- cha Prochristianella penaei, Parachris- tianella monomegacantha and Para- christianella dimegacantha are named and described from plerocerci recovered from the digestive gland. P. penaei from all three species of shrimp, P. monomegacantha and P. dimegacan- tha from P. duorarum. A small uniden- tified cestode lai^va is reported from the intestinal lining of P. aztecus and P. dtiorarum. Immature specimens of a nematode of the genus Contracaecum were found in the digestive gland and tissues surrounding the stomach of all three species of shrimp. The taxonomy, morphology, incidence, specificity and biology of each parasite is discussed. THE LARVA OF THE OAK TOAD, BUFO QUERCICUS HOLBROOK E. PETER VOLPE and JAMES L. DOBIE, Department of Zoology. Neivcomb College, Tulane University, New Orleans, Louisiana One aspect of systematic studies on anu- ran species is the establishment of criteria by which larvae can be unequivocally identi- fied, particularly when samples are collected from areas in which two or more species coexist. In southeastern Louisiana and Mis- sissippi, the genus Bufo is represented by four species, whose ranges partially or en- tirely overlap ( fig. 1 ) . Precise knowledge of the early development of each species should facilitate accurate diagnoses of larvae in mixed collections, even in samples de- rived from localities in which interspecific hybridization occurs (Volpe, 1956). Tadpoles of B. terrestns and B. foitleri are distinguished primarily on the basis of the relative distribution of melanophores in the tail musculature (Gosner and Black, 1958). Larvae of the Gulf Coast toad, B. valliceps, differ mainly from those of the Southern and Fowler's toads in labial tooth row proportions and pigmentation pattern of the tail ( Limbaugh and Volpe, 1957). The tadpole of the Oak toad, B. quercicns, has been reported to have a distinctive tail pattern. The pattern of "six or seven black saddles on the musculature, and with heavily marked upper tail crest" ( Wright and Wright, 1949, p. 199) should serve to dif- ferentiate larvae of B. quercicus from those of other species of toads. However, such a supposition proves untenable. The investigation reported here was under- taken to amplify the description of the lar- vae of B. quercicus, particularly with the view of establishing characters useful in identification. Embryonic Development The opportunity to study the early de- velopment of the Oak toad was afforded when several adults were collected, on May 27, 1958, from a rain pool in pine-oak wood- lands 1.4 miles northwest of the junction of State Highways 604 and 43, near Pearling- ton, Hancock County, Mississippi, 2.6 miles east of the Louisiana border ( fig. 1 ) . Am- plexing pairs were placed in a gravel- bottomed terrarium containing a small amount of pond water. Eggs were obtained from one mated pair. The eggs were laid in small groups, each consisting of a short strand or "bar" of three to six eggs. The entire complement deposited by the female was estimated at 500 eggs. The mean di- ameter of the vitellus was 1.09 mm (range of 21 eggs : 0.98 - 1.13 mm). Each egg was enclosed in a single layer of hyaline jelly; the diameter of this gelatinous en- velope averaged 1.25 mm (range of 20 measurements : 1.20 - 1.34 mm). A thin layer of jelly of loose consistency, compris- ing the jelly tube, united the individual eggs. The jelly tube was drawn out and pinched off between individual bars of eggs ( fig. 2 ) . Strands of fertilized eggs were transferred to finger bowls ( 350 ml capacity ) , each con- taining 200 ml of pond water. The develop- ment of the embryos was observed at approximately three-hour intervals. Stages in the embryological development were similar to those of valliceps, as illustrated by Limbaugh and Volpe (1957). As is char- acteristic of toads, the embryos hatched be- fore they were motile (tail bud stage 17). Embryonic development was extremely rap- id, in comparison with other species of toads known to the authors. At room temperature (27.5° - 29.5° C), the embryos hatched within 24 hours after fertilization. The embryos were noticeably smaller than those of valliceps. At stage 20, the onset of gill circulation, the average total length (snout to tip of tail) was 3.2 mm; at stage 25, the last of the prefeeding stages, 5.5 mm. Em- bryos of valliceps at equivalent stages meas- ure 3.7 and 7.3 mm, respectively. The mouthparts differentiated during the closing stages of the embryonic period. Cornified frameworks of the beaks and rudi- ments of the labial ridges were recognizable 146 Tulane Studies in Zoology Vol. 7 B. valliceps B. fowleri Fig'ure 1. Approximate rang'es of four species of toads in Louisiana and Mississippi. The black dot sig:nifies the source of origin of the B. qitercicns adults utilized in the in- vestigation. (Note: The status of B. fotvleri west of the Mississippi River is question- able. ) at embryonic stage 23. Teeth were evident at stage 24, arising in the first upper labial ridge or row. The inner edges of the horny beaks were pigmented and serrated. At stage 25, the lateral papillary fringes were emarginate and contained small papillae. Teeth were conspicuous on all labial ridges (two upper and three lower); however, no row possessed a full complement of teeth. During the period between the origin and resorption of the external gills ( stages 20- 25), melanophores and xanthophores arose in the dorsolateral surfaces of the tail muscu- lature. At stage 25, tlie last embryonic stage, the tail musculature was sharply bi- color; the dorsal half was darkened by me- lanophores, the ventral half was immaculate. Clusters of xanthophores occurred at irregu- lar intervals along the dorsal edge of the tail musculature. Larval Development At the onset of feeding activity (stage 26, defined in Limbaugh and Volpe, 1957), the larvae were transferred to 12 X 8 X 2 inch enamel pans containing pond water to a depth of one inch. Each enamel pan, hous- ing approximately 10 larvae, was kept at room temperature (27.5° - 29.5° C). The larvae were fed slightly boiled letture until the closing stages of metamorphosis. The larvae were examined at approxi- mately 24-hour intervals. Records were kept of tlie stages of development, distribution of chromarophores, and linear dimensions (snout to tail tip) of the larvae. Photo- graphs were taken of living larvae (nar- No. 4 Volpe and Dobie: Larva of Bufo quercicus Ul cotized in "M.S. 222" V) to reveal the dis- tribution of formalin-soluble yellow chro- matophores ( xanthophores ) . The following discussion of larval characteristics empha- sizes the structure of the oral disc, the size in relation to stage, and the pigmentation 1 Trade name for tricaine methanesul- fonate, prepared by the Sandoz Chemical Works, New York, New York. pattern of the tail. Mouthparts. — The configuration of the oral disc is illustrated in figure 2. The labial tooth formula is 2/3. The first tooth row in the upper labium is continuous; the second is divided medially. The median space of the second upper tooth row is nar- row, comprising 0.15 - 0.20 times the length of either lateral segment of teeth. The first -<^ ^^J'^^^SS^'^^^^^'^^I^ Figure 2. "Bars" of eggs (A); oral disc (B); oblique and lateril views of formalin- preserved larvae at stages 30 (C) p.nd 41 (D). (Illustrated by Mr:. C:rc,yn Thcrne Volpe.) 148 Tidane Studies in Zoology Vol. 7 lower tooth row is interrupted medially by a short space- and rises to a point in the midline in the direction of the broadly V- shaped lower beak. The second lower row corresponds in outline to the first but is slightly shorter. The third lower tooth row is approximately half the length of the sec- ond lower row. The oral disc is emarginate. Marginal papillae are present along the sides of the upper and lower labia. The relative lengths of the three lower tooth rows constitute the best criterion for distinguishing the oral discs of qiiercicus and ralliceps,. In the latter species, the three lower tooth rows are of nearly equal chord length ( Limbaugh and Volpe, 1957; Gosner and Black, 1958). In mouthparts of larval fowleri and terrestris, the lateral papillae of the lower labium extend ventrad to slightly below the outer edge of the short third lower row of teeth ( see fig. 10 in Volpe, 1956). The lower labial papillary fringe of quercicus extends to the outer edge of the second lower tooth row (fig. 2). In our sample, the third lower row is usually flanked on each side by a single, isolated papilla. We do not know whether this feature is generally characteristic of querci- cus larvae, or is peculiar to the particular group of larvae studied. Linear dimensions. — The size-stage rela- tionships during larval development are graphically represented in figure 3. The growth curve of quercicus is an approxima- tion based on small samples of two or three larvae at each stage. Obviously, greater numbers would result in a refinement of the curve. The present data, however, are of value in revealing the relatively small size of the tadpole of quercicus. The larvae of terrestris, fowleri, and valliceps are conspic- uously larger. Gosner and Black ( 1958 ) found that the size-staging curves of the latter three species are nearly equivalent. Data presented in figure 3, based on large samples of laboratory-reared larvae of ter- restris, fowleri, and valliceps from southeast- ern Louisiana, substantiate their findings. Tadpoles of quercicus reached a maxi- mum length of 19.4 mm at stage 41. The - The majority of larvae examined pos- sessed a divided first lower tooth row. A median break in this row occurs (or is con- sidered) as a variant condition in other species of toads. body lengths (snout to vent) of ten toads preserved at transformation averaged 8.2 mm (range: 7.2 - 8.9 mm). Transforma- tion sizes recorded by Wright (1932) for the Oak toad are between 7 and 8 mm. Pigmentation pattern of the tail. — Two types of chromatophores, the black (melano- phore) and the yellow ( xanthophore ) ,'' im- part a characteristic pattern to the tail. The distribution of xanthophores is shown in photographs of living larvae at stages 30, 37, 39, and 41 (fig. 4). The appearance of the tails of two of the larvae (stages 30 and 41 ) after formalin preservation is il- lustrated in figure 2. The striking contrast before and after preservation reflects the fact that the pigment of xanthophores is soluble in formalin. It is evident that larval descriptions based solely on preserved speci- mens are not only incomplete but may be misleading. The dorsal portion of the tail musculature is covered with melanophores; the greater part of the ventral half is devoid of melano- phores. The dividing line between the melanophore-darkened dorsal region and the immaculate ventral area may be referred to as the "melanophore border". This border tends to be more ventrad in advanced larvae (cf.. stages 30 and 41 in fig. 4). The dorsal tail fin in our sample can not be designated as "heavily marked" ( Wright and Wright. 1949). Rather, the dorsal fin is sparsely flecked with melanophores, the number vary- ing with the stage of development. In gen- eral, the intensity of mottling increases pro- gressively during development (fig. 4). The greatest concentration of xanthopores occurs along the dorsal edge of the tail mus- culature and along the melanophore border. In a formalin-preserved specimen viewed dorsally or obliquely (fig. 2), a series of light areas may be seen in the dorsal edge of the musculature. These light areas repre- sent the sites of the formalin-soluble xantho- phores. In like manner, the melanophore •^ The term "xanthophore" is used with reservation. At least two types of chro- matophores, xanthophores and leucophores, comprise the "lig-ht spots'' of the tadpole. The piedominant type appears to be the xanthophore ( = lipophore) , containing: a yellow pigment. Others fit the description of an iridescent leucophore ( ^iridophore, jl'uanophorc) , appeai'lnji' silvery with re- flected lifi'ht. Additional studies are needed. No. 4 Volpe and Dobie: Larva of Bufo quercicus 149 E E CD 24- 23- 22 - 21 20 19- 18 - 17 16 15 14 - 13- 12 - 11 ^ 10 9 8 7 X B. terrestris B. valliceps B. fowler i X X A X f— 26 — I— 27 -T— 28 — r— 29 30 — r- 31 32 -r- 33 — I— 34 I 35 — T" 36 37 38 — r~ 39 40 — I— 41 42 43 — r— 44 45 46 LARVAL STAGES Figure 3. Average dimensions in millimeters of the total length of tadpoles of four species of toads during development. Stages of larval development are defined and il- lustrated in Limbaugh and Volpe (1957). border in formalin-preserved specimens is irregular in outline, the indentations signi- fying those areas occupied by clusters of xanthophores. The scalloped appearance of the melanophore border may conceivably create an overall pattern effect of "six or seven black saddles on the musculature", as envisioned by Wright and Wright (1949). Comparison of Tail Patterns Xanthophores are prevalent also in the tail of the larva of valliceps; larvae of ter- restris and fowleri contain relatively insig- nificant amounts of yellow chromatophores ( fig. 5 ) . Although the disposition of xan- thophores in quercicus may be somewhat comparable, particularly along the dorsal edge of the musculature, to that of valliceps, larvae of the latter species can be distin- guished readily on the basis of the amount and distribution of melanophores. In sharp contrast to quercicus, melanophores are dis- tributed throughout the tail musculature in valliceps and form an elaborate, reticulate pattern in the dorsal tail fin. Formalin-preserved specimens of terrestris and fowleri resemble superficially the tad- pole of quercicus, especially when compari- sons are made utilizing larvae of terrestris that possess unspotted dorsal fins {cf., A in fig. 5 with C in fig. 2 ) . In each of these three species, the melanophore-darkened dorsal region contrasts sharply with the ventral area lacking melanophores. How- ever, a close examination of the dorsal edge of the musculature of larvae of quercicus 150 Tulane Studies in Zoology Vol. 7 Figure 4. Latei-al views of the tails of living- larvae of Bnfo qnercicus at stages 80 (A), 37 (B), 39 (C), and 41 (D). reveals light areas or patches representing outline in terrestris and jowleri than in the sites of formalin-soluble xanthophores, cjiarciciai. whereas the same region in larvae of ter- restris or fowleri is covered uniformly with Summary melanophores. Furthermore, the melano- The only previous description of the tad- phore border tends to be more regular in pole of the Oak toad, Bufo qnercicus, Volpe and Dobie: Larva of Bufo quercicus Figure 5. Lateral views of the tails of living larvae of Bufo terrestyis (A), B. quer- cicus (B), and B. valliceps (C). stemmed from studies of specimens collected from the Okefinokee swamp area in Georgia (Wright, 1932). A detailed description of laboratory-reared larvae derived from a sin- gle mated pair collected near Pearlington, Mississippi has been presented here. Em- phasis has been placed on those characters that serve to differentiate these tadpoles of B. quercicus from those of B. valliceps, B. terrestris, and B. fowleri. References Cited 'GosNER, Kenneth L. and Irving H. Black 1958. Notes on larval toads in the east- ern United States with special refei-ence to natural hybridization. Herpetologica, 14: 133-140. LiMBAUGH, Beverly A. and E. Peter Volpe 1957. Early development of the Gulf Coast toad, Bufo valliceps Wiegmann. Amer. Mus. Nointates, No. 1842: 1-32. Volpe, E. Peter 1956. Experimental Fi hybrids between Bufo valliceps and Bufo fowleri. Tulane Stud. Zool., 4(2) : 61-75. Wright, Albert Hazen 1932. Life-histories of the Frogs of Okefinokee Swamp, Georgia. Macmillan Company, New York, XV + 497 pp., fig. 1, pis. 1-45. Wright, Albert Hazen and Anna Allen Wright 1949. Handbook of Frogs and 152 Tulane Studies in Zoology Vol. 7 Toads. Comstock Publishing Company, Pearlington, Mississippi, have been de- New York, xii + 640 pp., figs. 1-7, pis. scribed and compared with those of B. 1-126, maps 1-37. vaUiceps, B. terrestris, and B. fowleri. As judged by this limited sample, lar- AbSTRACT vae of B. qiieycicHS may be distin- Laboratory-reared larvae of the Oak guished primarily on the basis of the toad, Bufo qHercicns Holbrook, derived distributional pattern of melanophores from a single mated pair collected near and xanthophores in the tail. TULANE STUDIES IN ZOOLOGY VOLUME 1, 1953-54 Number Price 1 On a new genus and species of mysid from Louisiana (Crustacea, Malacostraca), by Albert H. Banner, pp. 1-8 (June 1, 1953) . .I?0.25 2 A contribution on the life history of the lizard Scincella laterale (Say), by Richard M. Johnson, pp. 9-27 (July 3, 1958) 75 3 An outline for the study of a reptile life history, by Fred R. Cagle, pp. 29-52 (July 28, 1953) 75 4 A population of Holbrook's salamander, Eurycea longicauda guttolineata (Hol- brook), by Robert E. Gordon, pp. 53-60 (August 15, 1953) 25 5 A redescription of the crawfish Procam- barns hinei (Ortmann) (Decapoda, Astaci- dae), by George Henry Penn, pp. 61-68 (September 1, 1953) 25 6 A new burrowing crawfish of the genus Procambarus from Louisiana and Missis- sippi (Decapoda, Astacidae), by George Henry Penn, pp. 69-76 (September 15, 1953) 25 7 The life history of the crawfish Orconectes (Faxonella) clypeatus (Hay) (Decapoda, Astacidae), by Elsie Wayne Smith, pp. 77- 96 (October 23, 1953) 50 8 Ostrincola gracilis C. B. Wilson, a parasite of marine pelecypods in Louisiana (Cope- poda, Cyclopoida), by Arthur G. Humes, pp. 97-107 (December 21, 1953) 25 9 Hybrid inviabillty between L'ana pipiens from Wisconsin and Mexico, by B. Peter Volpe, pp. 109-123 (February 8, 1954) 35 10 The butterflies and skippers of Louisiana, by Edward Nelson Lambremont, pp. 125- 164 (April 30, 1954) 60 11 Two new species of the genus Oraptemys, by Fred R. Cagle, pp. 165-186 (August 26, 1954) 75 12 The taxonomlc status of the mid-Gulf Coast Amphiuma, by Imogene R. Hill, pp. 189-215 (August 26, 1954) 35 Complete volume, including title page and table of contents, (unbound) $4.25 VOLUME 2, 1954-55 1 A new Eulimnadia from the rice fields of Arkansas with a key to the American species of the genus (Conchostraca, Lim- nadiidae). by N. T. Mattox, pp. 1-10 (Sep- tember 14, 1954) $0.25 2 Studies in the ecology of the narrow- mouthed toad, Microhyla carolinenais caro' linensis, by Paul K. Anderson, pp. 13-46 (November 8, 1954) 50 3 A new species of Diaptomus from Louisi- ana and Texas with notes on the subgenus Leptodiaptotnus (Copepoda, Calanoida), by Mildred Stratton Wilson, pp. 47-60 (No- vember 29. 1954) 25 4 Three new species and new records of southern mllllpeds. by Nell B. Causey, pp. 61-68 (December 28. 1954) 25 5 Notropis baileyi, a new cyprinid fish from Pascagoula and Mobile Bay drainages of Mississippi and Alabama, bv Royal D. Snttkus and Edward C. Raney, pp. 69-86 (January 12. 1955) 30 VOLUME 2, 1954-55-Coiitinued Number Price 6 A reconsideration of the racer. Coluber constrictor, in eastern United States, by Walter Auffenberg, pp. 87-155 (February 25, 1955) 1.00 7 liotropis Jiypsilepis, a new cyprinid fish from the Apalachicola River system of Georgia and Alabama, by Royal D. Suttkus and Edward C. Raney, pp. 157-170 (Feb- ruary 25, 1955) 30 8 The fishes of two Gulf Coastal marsh areas of Florida, by John D. Kilby, pp. 171-247 (May 4, 1955) 1.00 Complete volume, including title page, table of contents and index (unbound) . . . .$3.75 VOLUME 3, 1955-56 1 Notropis asperifrotis, a new cyprinid fish from the Mobile Bay drainage of Alabama and Georgia, with studies of related spe- cies, by Royal D. Suttkus and Edward C. Raney, pp. 1-33 (July 8, 1955) I$0.50 2 A new Louisiana copepod related to Diap- tomus (Aglaodiaptomus) clavipes Schacht (Copepoda, Calanoida), by Mildred Strat- ton Wilson, pp. 35-47 (August 1, 1955) ... .30 3 A new species of Sternotherus with a dis- cussion of the i^ternotherus carinatus com- plex (Chelonia, Kinosternidae), by Don- ald W. Tinkle and Robert G. Webb, pp. 51-67 (August 30, 1955) 50 4 A new Cambarus of the Diogenes section from North Louisiana (Decapoda, A.«taci- dae). by George Henry I'enn, pp. 71-81 (September 30, 1955) 25 5 Notropis euryzontis, a new cyprinid fish from the Chattahoochee River system of Georgia and Alabama, by Royal D. Suttkus, pp. 83-100 (December 28, 1955) 50 6 Factors influencing the rate of oxygen con- sumption of the dwarf crawfish, Camba- relliis shufeldtii (Decapoda Astacidae), by Milton FIngerman, pp. 101-116 (December 28, 1955) 35 7 Identification and geographical variation of the cyprinodout fishes Fundulus oliva- ceus (Storer) and Fundulus notatus (Rafl- nesque), by Jerram L. Brown, pp. 117-134, (February 3, 1956) 50 8 The physiology of the melanophores of the isopod Idothea exotica, by Milton Finger- man, pp. 137-148 (April 12, 1956) .30 9 Osmotic behavior and bleeding of the oyster Crassostrea virginica, by Milton FIn- german and Laurence D. Fairbanks, pp. 149-168 (April 12. 1956) 50 10 Anatomy of the e.vestalk of the white shrimp, Penaeus setiferus (Linn. 1758), by Joseph H. Young, pp. 169-190 (June 22, 1956) 5a Complete volume. Including title page, ta- ble of contents and Index (unbound) S4.00 VOLUME 4, 1956 1 A study of the distribution and taxonomy of the percid fish Percinn nuirofasciata (Agassiz). by Ronald W. Crawford, pp. 1- 55 (August 1, 1956) $0.75 TULANE STUDIES IN ZOOLOGY VOLUME 4, 1956-Continued Number Price 2 Experimental Fi hybrids between Bufo vallicepa and Bufo fowleri, by E. Peter Volpe, pp. 59-75 (September 30, 1956) 40 3 An outline for the study of an amphibian life history, by Kred 11. Cagle, pp. 77-110 (October 31, 1956) 40 4 Notes on habitats, systematic characters and life histories of Texas salt water Cyp- rinodontes. by Don G. Simpson and Gordon Gunter, pp. 113-134 (December 31. 1956) .35 5 Dominance subordinance relationships in the crawfish Cambarellus shufcldtii, by Mildred Eileen Lowe. pp. 137-170 (Decem- ber 31, 1956) 45 6 Propogation of the white shrimp, Penaeua eetiferus (Linn.) in captivity, by Malcolm C. Johnson and J. U. Ifielding, pp. 173-190 ( December 31. 1956 ) 30 Complete volume, including title page, ta- ble of contents and index (unbound) $2.50 VOLUME 5, 1957 1 Oatnhusia heterochir, a new poecilild fish from Texas, w^ith an account of Its hybrid- ization with a. ajfinia, by Clark llubbs, pp. 1-16 (March 18. 1957) 50.30 2 New calanoid copepods of Pontella Dana and Labidocera Lubbock with notes on the distribution of the genera in the Gulf of Mexico, by Abraham Fleminger, pp. 17-34 (March IS, 1957) 30 3 Three new crayfishes from Alabama and Mississippi (Decapoda: Astacidae), by iix>rton U. Hobbs, Jr. and Margaret Wal- ton, pp. 37 52 (March 18, 1957) 30 4 Heat death and associated weight loss of the oyster Crasaostrea virginica, by Milton Ftngerman and Laurence D, Fairbanks, pp. 53-62 (April 1. 1957) 30 6 The Odonata of Louisiana, by George H. Blck. pp. 69-135 (May 15, 1957) 1.00 6 Endocrine control of the red and white chromatophores of the dwarf crawfish, Cambarellua ahujeldti, by Milton Finger- man, pp. 137-148 (June 7, 1957) 30 7 Hormones controlling the chromatophores of the dwarf crawfish, Cambarellua ahu- jeldti : their secretion, stability, and sepa- ration by filter paper electrophoresis, by Milton Fingerman and Mildred E. Lowe, pp. 149-171 (June 7, 1957) 40 8 Cyprlnld fishes of the subgenus Cyprinella or Notropia. III. Variation and subspecies of Notropia venuatua (Girard), by Robert H. Glbbs, Jr. pp. 173-203 (August 7, 1957) .50 9 The earlv development of Rana capita aevo- aa, by E." Peter Volpe, pp. 205-225 (Septem- ber 12, 1957) 35 11 The skeleton shrimps (Crustacea : Caprel- lidae) of the Gulf of Mexico, by Joan E. Steinberg and Ellsworth C. Dougherty, pp. 265-288 (December 30, 1957) 40 10 Variation and subspecies of the crawfish Orconectea palmeri (I<'axon) ( Dccapodu, Astacidae), by George Henry Penn, pp. 229-262 (September 12, 1957) 60 VOLUME 5, 1957-Continued Number Price 12 The systematic status of the suckers of the genus Moxostoma from Texas, New Mexico, and Mexico, by C. Richard Robins and Edward C. Raney. pp. 289-318 (Decem- ber 30. 1957) 45 Complete volume, including title page, ta- ble of contents and index (unbound) ....$5.00 VOLUME 6, 1958 1 The systematics and ecology of the Sterna- thuerua carinntua complex (Testudinata, Chelydridae), by Donald W. Tinkle, pp. 1-56 (March 31, 1958) $1.25 2 The butterflies of Mississippi, by Bryant Mather and Ivatharine Mather, pp. 61-109 (June 6, 1958) 1.00 3 Aquatic and semiaquatic Hemiptera of Mis- sissippi, by Clifton A. Wilson, pp. 113-170 (September 5, 1958) 1.00 4 The copepod genus Halicyclops in North America, with description of a new species from Lake Pontchartrain, Louisiana, and the Texas coast, by Mildred Stratton Wil- son, pp. 176-189 (December 31, 1958). Ontogeny of the first and second pleopods of the male crawfish Orconectea clypeatus (Hay) (Decapoda, Astacidae), by Joe B. Black, pp. 190 203 (December 31, 1958) ... .60 Complete volume, including title page, table of contents and index (unbound) $3.50 VOLUME 7, 1959 1 An illustrated key to the crawfishes of Louisiana with a summary of their distri- bution within the State (Decapoda, Asta- cidae), by George Henry Penn, pp. 3-20 (April 23. 1959). Comparison of the chromatophorotropins of two crayfishes with special reference to electrophoretic behavior, by Milton Finger- man, pp. 21-30 (April 23, 1959) $0.60 2 A review of the seabasses of the genus Centropristea (Serranidae), by Rudolph J. Miller, pp. 33-68 (July 9, 1959) 0.75 3 Digenetic trematodes of marine fishes from the Gulf of Panama and Bimini, British West Indies, by Franklin Sogandares-Ber- nal, pp. 69-117 (August 24, 1959) 1.00 4 Parasites of the commercial shrimps, Pet^^ aeua aztecua Ives, P. duorarum Burkenroad, and P. aetiferus (Linnaeus), by Dwayne Nathaniel Kruse, pp. 123-144 (October 19, 1959). The larva of the oak toad, Bufo quercicus Holbrook, by B. Peter Volpe and James L. Dobie, pp. 145-152 (October 19, 1959) 60 Orders should be addressed to Meade Natural History Library, c/o Department of Zoology, Tulane University, New Orleans. 18, La., USA Please make remittance payable to "Tulane Unlyerslty" TULANE STUDIES IN ZOOLOGY VOLUME 7 INDEX TO AUTHORS AND SCIENTIFIC NAMES (New species and genera in boldface) Abndefduf saxatilis, 106 Acanthocolpidae, 87-92 AcanthocijhiHm solandri, 106 Acantlutrus caernleus, 103 Accacoelidae, 106 Amphibia, 145-152 Antorchis arna, 101 Apocreadiuin, 83-87 angustum, sp. nov., 84 balistis, 83, 108 bravoi, sp. nov., 86-87 coili, sp. nov., 83-84, 108 longisiiiosinn, 83 uroproctoferum, sp. nov., 84-86, 108 Apo(/o)i hi)iotatHS, 95 Ascarididae, 140-141 Aspidogastridae, 71 Astacidae, 3-20, 21-30 Bacciger harengulae, 101 BairdieUa chrysura, 132 Balistes caprisctis, 74, 83, 108 naufragium, 76, 86, 87 polylepis, 87 verres, 76 vetula, 76, 83, 84, 108 Barisonmm erubescens, 74 Bevesicula hepsetiae, 102 Bevesiculidae, 102 Brachadena pyriformis, 103 Bucephalidae, 71-74 Bucephaloides arcuahis, 74 Bucephalus varicits, 71 Bnfo fowleri, 146 qiiericicus, 145-152 terrestris, 146, 151 valliceps, 146, 151 Cableia, gen. nov., 80-82 trigoni, sp. nov., 80-82 Calirilla pi)ita, 71 Calamus bajonado, 71, 89, 101 Cambarellus puer, 14 shufeldti, 14,21-30 Cambarus dio genes diogenes, 14 ludovicianus, 14 dissitus, 16 hedgpetJti, 14-16 oryktes, 16 Caranx crysos, 71, 101 hippos, 71 latus, 71, 88, 101 ruber, 71, 101, 102 Carcharhinus natator, 102 Centropristes, 33-68 ocyurus, 50-59 philadelphicus, 59-63 striatus, 43-50 melanus, 47-48 striatus, 47 Cephalolobus, gen. nov., 124-126 penaeus, sp. nov., 124-126 Ceratacanthus scripta, 79, 82 Cestoidea, 134-140 Cleptodiscus kyphosi, 74 reticulatus, 74 Cnidosporidia, 128-130 Conger conger, 106 Contracecum sp., 140-141 Coryphaena hippurus, 89, 103, 106 Crassicutis Diariiia, 83 Crustacea, 3-20, 21-30, 123-144 Cryptogonimidae, 98-101 Cy)io.^cio)i albus, 89 Dasyutis sabinu, 137 Decapoda, 3-20, 21-30, 123-144 Dermadena lactophyysi, 76 Dinm-ns lougisinus, 103 tornatus, 103-104 Diodon hystnx, 79 Diplaiigus paxillus, 102 Dipluproctodaeuni haustrum, 79 plicitum, 79-80 Dobie, James L., article, 145-152 Dollfnstrema muraenae, sp. nov., 72-74 Ectcnurus virgnlus, 103 Elagaiis bipinnulatus, 88 Eyienterum aurewm, 80 Epitiephelus analogus, 71, 72, 95 striatus, 95, 98 tigris, 77 Eucinostomus calif or niense, 82 Eutetrarhynchidae, 134-140 Fellcdistomatidae, 101-102 Fingerman, Milton, article, 21-30 Genolopa anipullacea, 102 Gerres cinereus, 82, 83, 97 Gonacanthella, gen. nov., 100-101 lutjani, sp. nov., lOO-lOl Gorgoderidae, 102-103 Gregarinida, 124-128 Gymnothorax dovii, 105 vicinus, 72, 95 Haemulon album, 95, 102, 103 parra, 102, 103 plumieri, 102, 103 sciiirus, 95, 98, 102, 103, 107 Hamacreadium mutabile, 98 Haplosplanchnidae, 93-95 Haplosplanchnus acutus, 93-95 Harengulaciupeola, 101, 103, 104, 106 Heliconietra execta, 95 nimia, 95 torta, 95 INDEX TO AUTHORS AND SCIENTIFIC NAMES— Continued Hemiuridae, 103-106 Hepsetia stipes, 102 HiriidineUa clarata, 106 tuariiia, 106 Holoceiitnis ascoiciottis, 76, 90, 95, 98 106 Homalometron elongatiim, 82 Hi/sferolecithn rosea, 103 JrdiicadeiKifid hrii)Hj)ti, 80 Kruse, Dwayne Nathanial, article, 123-144 Kil])hosi(s analog i(S, 93 elegans, 80, 92. 104 sectatrix, 74,80, 104 [Athrisoinus niichipiviiis, 106 Lactophrijs hicandalis, 87 tricornis, 76, 87 trigonis, 76, 84 Lecithochirinui mio-ostnmuDi, 105 Lcjjidapedou parepinepheli, sp. nov., 77-79 truncatum, sp. nov., 76-77 Lepocreadiidae, 74-87 Lepocreadinm trnlla, 74 Leurodera decora, 103 Lohatosoma ringens, 71 analis, 100 bnccanella, 74 jordaiii, 100 iiovemfasciafiis, 98 synagris, 95, 98, 100 Malacavfhus plumieri, 89, 103, 106 Megapera gyriiia, 87 Megraperidae, 87 Megasolena kyphosi, sp. nov., 93 americamm, 132 saxatilis, 132 crassidatu, 100 globosa, 100 Micropogov ini.diilatns, 132 Microsporidia, 128-130 Miller, Rudolph J., article, 33-68 Monorchiidae, 102 falcata, 95 venenosa venenosa, 72, 89 Myodera magna, sp. nov., 92 Nagniia pacifica, 102 Nematoda, 140-141 Nematopsis penaeus, 126-128 Neonotoporus yaniagiifii, 95 Ocyurus chrysurus, 74, 100, 106 Ogcncephahis resperfilio, 106 Opecoelidae, 95-98, 130-133 Opecoclnides fiwhriatiis, 130-133 Opecoelns sphacricits, 98 Opisthadena dimidia, 104 OrcoMectes heyeri, 16-18 clypeatus, 16, 21-30 difficilis, 19 haihaivayi, 18-19 hohhsi, 18 lancifer, 18 creolauHs, 18 lotigimanus, 18 pahneri, 18 Osteichythyes, 33-68 Fachycreadiion lerneri, sp. nov., 97-98 Pachycryptogonimns americanns, 98-100 Palo))ieta media, 71 Paracliristianellu dimegacantha, sp. nov., 139-140 monomegacantha, sp. nov., 137-139 Paralieniinriis nicras, 104-105 Paramphistomatidae, 74 argeiitinensis, 101 bagre, 101 aztecns, 123-144 dnorarnm, 123-144 setiferns, 123-144 Penn, George Henry, article, 3-20 Petrometopov cruenfutus, 98 Podocotyle mycteropercae, sp. nov., 95-97 Poniacanfhns anrcns, 74, 101 Prochristianella penaei, sp. nov., 134-137 Procambariis birittatus, 10 blatidingi aciitas, 10 darki, 10 dupratzi, 13 hiuei, 11 iiatchitochae, 14 planirostris, 10 penui, 13 shermani, 11-13 simidans simnUuis, 13 tulanei, 13 riaeviridis, 10 vioscai, 13 Proctoeces subtenne, 101 Proctotrema truncatum, 102 Proneochasmu.s, 101 Pronocephalidae, 74 Prosogo)wtre>ua bilabiat)i))i. 106 Prosog-onotrematidae, 106 Prosorhynchns gonoderiis, 71-72 ozak-ii, 72 l)acificiis, 72 Psendocrcadium biminensis, sp. nov., 74-76 lamellifo))ne, 76 scaphosomam, 76 Pseudopecoeloides equesi, 98 gracilis, 95 Pseiidnpecnchis ha rkeri, 98 /I'r'/f/ fcxana, 137 Rhagorchis odiineri, 82 Rhinoptera bonasus, 137 Sarda velox, 106 Sciaenops ocellata, 132 INDEX TO AUTHORS AND SCIENTIFIC NAMES — Continued Scorpaena mystes, 98 phimieri, 106 Selar cnimenopthalmus, 95, 101, 103 Seriola hispidum, 88, 90 Serranidae, 33-68 Siphodera vinaledwardsi, 100 Sogandares-Bernal, Franklin, article, 69-117 Spiwei-oides annulatus, 79, 83 Sphyraena barracuda, 74 Sporozoa, 124-130 Steganoderma atherinae, 102 Stellifer sp., 98 Stenopera equilata, 95 Sfephanostomum coryphaenae, 89 dentatum, 89 difrematis, 88-89 hispidum, 88 minutum, 89 provitellosum, sp. nov., 87-88 pseudocarangis, sp. nov., 90 senium, 89 Sterrhurus florideiisis, 106 taboganus, sp. nov. Synodus sp., 104 105-106 Telosporidia, 124-128 Tergestia pectinata, 101 Tetrochetus coryphaenae, 106 Thelohania sp., 128-130 Thyrinops pachylepis, 93 Tormopsolus orientalis, 90-92 Trachypeneus constrictus, 132 Trematoda, 69-117, 130-133 Tubulovesicula angusticauda, 104 lindbergi, 104 magnacetabulum, 104 pinguis, 104 Tylosurus raphidoma, 93 Volpe, E. Peter, ai-ticle, 145-152 Waretrematidae, 92-93 Zoogonidae, 1C2 1