CHAPTER X PARASITIC WORMS Blank page retained for pagination PARASITIC HELMINTHS

By ASA C. CHANDLER, Rice Institute. and HAROLD W. MANTER, University of Nebraska

The helminth parasites of the Gulf of Mexico, Chandler (1935a), in a study of 23 species of with the exception of the trematodes, have been fish from various parts of Galveston Bay, pointed studied very little. The bulk of the collecting out a great excess of immature over adult stages done so far has been at the former Biological of parasites, 15 of 25 of the species identified Laboratory of the Carnegie Institution at Dry being larval or immature stages; a scarcity of Tortugas, Florida, at the east!'lrn side of the Gulf. flukes and adult tapeworms (no elasmobranchs A. few desultory investigations or reports of one were included in the study); a predominance of or a few species have been made at other points Acanthocephala as common adult parasites; and on the United States coast and from the north very marked differences in the helminth fauna in coast of Cuba, but with the exception of one new local areas, even in the very limited region in­ species of trematode reported from a manatee no cluded in the 'study. These differences are prob­ studies have yet been made along the extensive ably associated with the local distribution of Mexican coast where many new helminths will definitive hosts in the case of immature species probably be found, particularly in the non­ and of intermediate hosts in the case of adult lUigratory fishes of the shallower waters and species. Of 10 species of adults and 15 of im­ bays. Practically all the work done so far has mature species, 9 of the former and 11 of the latter been taxonomic and morphological, dealing either were previously undescribed species. This is in with adult or larval forms. Of more than 200 contrast to the findings of Manter in oceanic species of trematodes from Gulf animals only 3 species of fish from Dry Tortugas and suggests, life cycles have been studied, and the life cycles of as remarked above, that many new species of none of the cestodes, acanthocephalans, or nem­ fish helminths will be found when the parasites atodes have been studied at all. of nonmigratory species of the shores and bays of The only parasitic helminths of Gulf animals the western and southern sides of the Gulf are that have thus far been studied comprehensively studied. are the trematodes of oceanic fishes and these only In preliminary studies on marine fishes from at Dry Tortugas where the pioneer work was done Corpus Christi Bay, Tex., Aaron Seamster (1950) by Linton (1907-10) followed by an extensive reported in a personal communication that of 30 study by Manter from 1930-49. Linton also did fish examined, belonging to 16 species, parasites Some pioneer work on the other parasitic helminths were found in 27. The heaviest load was in the of Dry Tortugas, but this work has not been fol­ yellow-tail, Bairdiella chrysura. However, only lowed up as has the work on the trematodes. It 1 specimen, Menticirrhus americanus, was infested Was observed by Linton, however, that Acantho­ with an acanthocephalan, Illiosentis furcatus, and cephala and Nematoda, both in adult and larval only 1 specimen, Synodusfoetens, with a nematode. stages, are relatively scanty in fishes of Dry A few cestode infections were reported. Tortugas and in fishes of other southern regions In addition to the trematodes, cestodes, acantho­ (Bermuda and North Carolina) as compared with cephalans, and nematodes considered in the fol­ their occurrence in fishes at Woods Hole, Mas­ lowing sections a few other parasitic or semi­ sachusetts. Larval cestodes, also, were relatively parasitic helminths have been reported from the scanty except adult Tetraphyllidea and Try­ Gulf. Several polyclad turbellarians are as­ Panorhyncha in selachians which compared favor­ socipted with particular marine organisms on ably with the numbers found in the Woods Hole, which they are predatory in a semi-parasitic Massachusetts, area. fashion. Stylochus frontalis is common in Gulf 333 334 FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE oysters. Other species are Stylochus ellipticus Another helminth recorded from the Gulf is commonly attacking barnacles, and Hoplo­ a nemertean, Carcinonemertes carcinophila (Kol~ plana inquilina, attacking the oyster drill (Thais liker 1845) Coe, 1902, a variety of which, C. c. jloridana). Hoploplana inquilina was studied at imminuta, was found living on the egg masses of Barataria Bay, Louisiana, by Schechter (1943), Callinectes sapidus, blue crab, at Grand Isle, and the others were reported from Apalachicola Louisiana. Bay, Florida, by Pearse and Wharton (1938). NOTE.-Bibliography follows chapter on Nematoda, p. 358. TREMATODA OF THE GULF OF MEXICO

By HAROLD W. MANTER, University of Nebraska

Trematodes of the Gulf of Mexico have been 3. Hexastoma macracanthum Fujii, 1944. studied chiefly at the former Biological Laboratory Host: Euthynnus alletteratus, Little tuna. of the Carnegie Institution at Dry Tortugas, Reported by Fujii (1944). Florida. The rich fauna there of fishes, mollusks, 4. Heterocotyle jloridana (Pratt, 1910) Price, 1938. and Crustacea would suggest a great variety and Host: Aetobatus freminvillii, Stingaree. abundance of these parasites, and such has been Reported by Pratt (1910). proved to be the case. Early in the history of the 5. Udonella caligorum Johnston, 1835. Laboratory, Linton (1910) and Pratt (1910-16) Host: On a copepod on gills of Lutjanus described species of trematodes from Tortugas. griseus, Gray snapper. Reported by Linton (1910). Linton's work in particular indicated the unusual \7ariety of trematode species occurring within 6. Protomicrocotyle mirabile (MacCallum, 1918) fishes there. Later, Miller (1925-29) studied Johnston and Tiegs, 1922. Host: Caranx latus, Horse-eye jack. larval stages (cercariae) of trematodes from New record. MacCallum described the spe­ lnollusks; McCoy (1929, 1930) studied the life cies from Caranx hippos, Jack crevalle, CYcles of two species; and the author (Manter, from the New York Aquarium. 1930-49) made extensive taxonomic studies of his bee~eported collections at Tortugas during 1930-32. As a The following four species have result, the trematodes vf fishes, both surface water from other parts of the Gulf region: sPecies and species down to a depth of about 500 7. Capsala poeyi (Vigueras, 1935) Price, 1938. fathoms, are relatively well known. Little study Host: Skin of Makaira ampla, Blue marlin, has been made of these parasites elsewhere in the from Havana, Cuba. Gulf. Vigueras (1935, 1940) has reported a few Reported by Vigueras (1935). sPecies fro'll Cuba; while Chandler (1935), Von 8. Encotyllabe monticelli Vigueras, 1940. Wicklen (1946), and Read (1947) described several Host: Calamus bajonado, Jolt-head porgy, sPecies from Galveston Bay. Faust and Tang from north coast of Cuba. (1936) described one species from Biloxi Bay, Reported by Vigueras (1940). Mississippi. 9. Entobdella squamula (Heath, 1902) Johnston, I am indebted to Dr. Luis R. Rivas for checking 1929. and correcting the scientific names of the fish Host: Undetermined fish "presumably from Gulf of Mexico." hosts and also for suggestions for the common Reported by Price (1939). llallles of these fishes. 10. N eoheterobothrium affine (Linton, 1898) Price, MONOGENEA 1939. The following species of Monogellea have been Host: Paralichthys sp., Flounder, from Grand ~ported Isle, La. from the gills of fishes at Key West or Reported by Melugin (1940). This species ortugas: is known from two species of Paralichthys 1. Cyclocotyla neomaenis (MacCallum, 1917) Price, at Woods Hole, Mass. 1943. Host: Lutjanus analis, Gray snapper. This small number of species of Monogenea Reported by MacCallum (1917). reported does not mean they are uncommon para­ 2. Cyclocotyla hysteroncha Fujii, 1944. sites, but rather that they have not been collected Hosts: Bathystoma striatum; Brachygenys and studied. I have a number of species still in­ chrysargyreus; Haemulon jlavolineatum, French grunt. completely studied collected from fishes at Tortu­ Reported by Fujii (1944). gas. Most of them are undescribed species. 335 336 FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE

ASPIDOGASTREA 2. Bucephalopsis "bennetti" Melugin, 1940. Host: Paralichthys lethostigma, Flounder. 1. Lobatostoma ringens (Linton, 1907) Eckmann, Reported by Melugin (1940) from Gra 1932. Isle, Louisiana. The name "bennetti" w Hosts: Calamus calamus, Saucer-eye porgy; printed in this abstract by Melugin, b Calamus bajonado, Jolt-head porgy. the trematode was not described, hen Reported by Linton (1910). the name is a nomen nudum. 2. Stichocotyle cristata Faust and Tang, 1936. 3. Bucephalopsis longoviferus Manter, 1940. Host: Rhinoptera bonasus, Cow-nosed ray Host: Sphyraena barracuda, Barracuda. from Biloxi Bay, Miss. Reported by Faust and Tang (1936). 4. Bucephalus kathetostomae (Manter, 1934) MB ter, 1940. A large aspidogastrid, Lophotaspis vallei, occurs Host: Kathetostoma albigutta, Stargazer. in the loggerhead turtle, Caretta caretta. It will 5. Bucephalus priacanthi Manter, 1940. be discussed with the trematodes of turtles. Host: Priacanthus arenatus, Glass-eyed sna DIGENEA per. 6. Bucephalus scorpaenae Manter, 1940. Approximately 200 species of digenetic trema­ Host: Scorpaena plumieri, Scorpion fish. todes are known from fishes of the Gulf of Mexico, 7. Bucephalus varicus Manter, 1940. mostly from its eastern region. Linton (1910) Hosts: Caranx bartholomaei, Yellow jack; reported 53 species from fishes at Tortugas. All latus, jack; C. rubel', Runner. except one of these were also collected by the 8. Dolljustrema gravidum Manter, 1940. author. Manter (1947, 1949) lists 190 species of Host: Gymnothorax moringa, Moray. digenetic trematodes, including 1 species of aspi­ 9. Prosorhynchus atlanticus Manter, 1940. dogastrid, from the fishes at Tortugas. This num­ Hosts: Mycteroperca microlepis, Gag; M~ ber is relatively large compared with the number tel'operca venosa, Yellow-fin grouper. known from marine fishes of New England, Great 10. Prosorhynchus ozakii Manter, 1934. Britain, or the Mediterranean, although Yamaguti Host: Epinephelus niveatus, Snowy grouper has reported a larger number from Japanese fishes. 11. Prosorhynchus promicropsi (Linton, 1905) Ec The richness of the trematode fauna of the Gulf mann, 1932. Host: Promicrops itaiara, Jewfis of Mexico is associated with the great variety of fishes mollusks, and Crustacea in that region. 12. Rhipidocotyle angusticolle Chandler, 1941. Thes~ Host: Sarda sarda, Bonito. four groups are ecologically associated in Reported from the Texas coast near Freepo that trematodes require a molluscan host for their by Chandler (1941). larvae and frequently use Crustacea as second 13. Rhipidocotyle adbaculum Manter, 1940. intermediate hosts. Thus, trematode parasites Host: Scomberomorus regalis, Cero. constitute a very common and widespread element 14. Rhipidocotyle baculum (Linton, 1905) Ec in the ecological picture of a region such as mann, 1932. Tortugas. 15. Rhipidocotyle barracudae Manter, 1940. Apparently only 13 other records of trematodes Host: Sphyraena barracuda, Barracuda. from Gulf of Mexico fishes have been published 16. Rhipidocotyle longleyi Manter, 1934. (Chandler, 1935, 1941; Melugin, 1940; Vigueras, Host: Synagrops bell-us. 1940, 1940a, 1940b; Von WickleJ4. 1946; Read, 1947). Of these species, four are known at Tor­ 17. Rhipidocotyle nagatyi Manter, 1940. Host: Euthynnus alletteratus, Little tuna. tugas. In the list following, the record is from Tortugas or Key West unless otherwise noted. 18. Rhipidocotyle transversale Chandler, 1935. Host: M enidia menidia, Silverside. b GASTEROSTOMATA Reported from Galveston Bay, Texas, Chandler (1935). Family BUCEPHALIDAE 1. Bucephalopsis longicirrus Nagaty, 1937; syn­ PROSOSTOMATA onym: Bucephalopsis arcuatus (Linton, Family PARAMPHISTOMATIDAE 1900) Eckmann, 1932. New synonymy. Also known from Red Sea. 19. Cleptodiscus reticulatus Linton, 1910. 1fisl Host: Sphyraena barracuda, Barracuda. Host: Pomacanthus arcuatus, Black ange GULF OF MEXICO 337

20. M acrorchitrema havanensis Vigueras, 1940. 37. Postporus mycteropercae (Manter, 1947) Manter, Host: Holacanthus tricolor, Rock beauty. 1949. Reported from north coast of Cuba by Host: Mycteroperca venenosa, Yellow-fin Vigueras (1940). grouper. Family PRONOCEPHALIDAE 38. Lepidapedon levenseni (Linton, 1907) Manter, 21. Pleurogonius candibulus (Linton, 1910) Ruiz, 1947. 1946. Host: Epinephelus morio, Red grouper. Hosts: Angelichthys ciliaris, Queen angelfish; 39. Lepidapedon elongatum (Bebour, 1908) Nicoll, Pomacanthus arcuatus, Black angelfish. 1915. 22. Barisomum erubescens Linton, 1910. Hosts: Coelorhynchus carminatus, Grenadier; Hosts: Angelichthys ciliaris, Queen angelfish; Epigonus occidentalis; Laemonema bar­ A. isabelita, Angelfish; Pomacanthus arcu­ batulum; Urophycis chesteri. atus, Bla"k angelfish; 8carus croicensis, 40..Lepidapedon lebouri Manter, 1934. Parrotfish. Host: Unidentified Macrouridae. Family MEGASOLENIDAE 41. Lepidapedon nicolli Manter, 1934. 23. Hapladena varia Linton, 1910. Host: Epinephelus niveatus, Snowy grouper. Hosts: Acanthus coeruleus, Blue tang; A. hepatus, Doctorfish. 42. Lepidapedon rachion (Cobbold, 1858) Stafford, 1904. 24. Hapladena ovalis (Linton, 1910) Manter, 1947. Host: Coelorhynchus carminatus, Grenadier. Hosts: Sparisoma pachycephalum, Parrotfish; Pseudoscarus coelestinus, Parrotfish. 43. M yzoxenus lachnolaimi Manter, 1947. Host: Lachnolaimus maximus, Hogfish. 25. Hapladena leptotelea Manter, 1947. Host: Pomacanthus arcuatus, Black angelfish. 44. M yzoxenus vitellosus Manter, 1934. Hosts: Calamus calamus, Saucer-eye porgy; 26. M egasolena estrix Linton, 1910. Decodon puellaris, Cuban hogfish. Hosts: Kyphosus incisor, Yellow chub; K. sectatrix, Bermuda chub. 45. Bianium plicitum (Lint-on, 1928) Stunkard, 1931. Family OPISTHOLEBETIDAE Hosts: 8phoeroides spengleri, Southern puf­ 27. Opistholebes adcotylopho1'1ts Manter, 1947. fer; 8phoeroides sp., Puffer; Monacanthus Host: Diodon holocanthus, Balloonfish. hispidus, Filefish. Family LEPOCREADIIDAE 46. Multitestis chaetodoni Manter, 1947. 28. Lepocreadium trulla (Linton, 1907) Linton, 1910. Hosts: Chaetodon ocellatus, Butterfly fish; Host: Ocyurus chrysurus, Yellowtail. C. capistratus, Butterfly fish. 29. Lepocreadium bimarinum Manter, 1940. 47. Rhagorchis odhneri Manter, 1931. Host: Lachnolaimus maximus, Hogfish. Hosts: Alutera schoepfii, Orange filefishj M onacanthus ciliatus, Fringed filefish. 30. Pseudocreadium anandrum Manter, 1947. Host: Calamus calamus, Saucer-eye porgy. 48. Enenterum aureum Linton, 1910. Hosts: Kyphosus sectatrix, Bermuda chub; 31. Dermadena lactophrysi Manter, 1946. K. incisor, Yellow chub. Hosts: Lactophrys tricornis, Cowfish; L. trigonus, Trunkfish; L. triqueter, Trunkfish. 49. Cadenatella americana Manter, 1949. Host: Kyphosus incisor, Yellow chub. 32. Opechona gracilis (Linton, 1910) Manter, 1947. Host: Harengula clupeola, Pilchard. Family OPECOELIDAE 50. Opegaster synodi Manter, 1947. 33. Apocreadium balistis Manter, 1947. Host: Synodus foetens, Lizardfish. Host: Balistes vetula, Queen triggerfish. 34. Homalometron elongatum Manter, 1947. 51. Opecoeloides brachyteleus Manter, 1947. Host: Gerres cinereum, Broad shad. Host: Upeneus martinicus, Yellow goatfish. 35. Crassicutis marina Manter, 1947. 52. Opecoeloides elongatus Manter, 1947. Hosts: Eucinostomus lefroyi, Mojarra; Gerres .Host: Upeneus maculatus, Red goatfish. cinercus, Broad shad. 53. Opecoeloides polyfimbriatus Read, 1947. 36. Postporus epinepheli (Manter, 1947) Manter, Host: 8ynodus foetens, Lizardfish. 1949. Reported from Galveston Bay, Texas, by Host: Epinephelus morio, Red grouper. Read (1947). 338 FISHEPoY BULLETIN OF THE FISH AND WILDLIFE SERVICE

54. Opecoeloides polynemi Von Wicklen, 1946. 69. Hamacreadium mutabile Linton, 1910. Host: Polynemus octonemus, Thread-fin fish. Hosts: Lutjanus analis, Muttonfish; L. Reported from Galveston Bay by Von apodus, Schoolmaster; L. griseus, Gray Wicklen (1946). snapper; L. jocu, Dog snapper; L. synagris, Lane snapper; Anisotremus virginicus, 55. Pseudopecoeloides gracilis Manter, 1947. Porkfish; Ocyurus chrysurus, Yellowtail. Host: Trachurops crumenophthalma, Goggle­ eye jack. 70. Hamacreadium oscitans Linton, 1910. Hosts: A nisotremus virginicus, Porkfish; 56. Pseudopecoeloides equesi Manter, 1947. Brachygenys chrysargyreus, Bronze grunt; Hosts: Eques Ian ceolatus, Ribbonfish; E. Haemulon carbonarium, Caesar grunt; H. acuminatus, Ribbonfish. plumieri, White grunt; H. sciurus, Yellow 57. Pseudopecoeloides carangis (Yamaguti, 1938) grunt. Yamaguti, 1940. 71. Helicometra execta Linton, 1910. Hosts: Caranx bartholomaei, Yellow jack; Hosts: Doratonotus megalepis; Eques acumina­ C. ruber, Runner. tus, Ribbonfish; Haemulon plumieri, White grunt; H. sciurus, Yellow grunt; Hali­ 58. Pseudopecoelus priacanthi (MacCallum, 1921) choeres bivittatus, Slippery dick; H. poeyi; Manter, 1947. H. radiatus; Labrisomus bucciferus; L. Hosts: Priacanthus arenatus, Glass-eye snap­ haitensis; Lachnolaimus maximus, Hogfish; per; P. cruentatus, Big-eye. M ycteroperca venenosa, Yellow-fin grouper; 59. Pseudopecoelus tortugae von Wicklen, 1946. Thalassoma bifasciatum, Slippery dick. Host: Coelorhynchus carminatus, Grenadier. 72. Helicometrafasciata (Rud., 1819) Odhner, 1902. 60. Pseudopecoelus vulgaris (Manter, 1947) von Hosts: Prionotus alatus, Sea robin; Prionodes Wicklen, 1946. sp.; Bellator militaris. Hosts: 16 species of fishes collected from 73. Helicometra torta Linton, 1910. depths of 50 to 315 fathoms (see Manter, Hosts: Epinephelus morio, Red grouper; 1934, p. 293). E. striatus, Nassau grouper. 61. Neopecoelus scorpaenae Manter, 1947. 74. Stenopera equilata Manter, 1933. Hosts: Scorpaena grandicornis, Scorpion fish; Host: Holocentrus ascensionis, Squirrelfish. S. brasiliensis. 75. Helicometrina nimia Linton, 1910. 62. Neopecoelus holocentri Manter, 1947. Hosts: 14 species of fishes including cardinal Host: Holocentrus coruscus, Squirrelfish. fish, porgies, snappers, and scorpion fishes. 63. Genitocotyle atlantica Manter, 1947. 76. Helicometrina parva Manter, 1933. Hosts: Carapus affinis, Pearlfish; Haemulon Host: Halichoeres bivittatus, Slippery dick. flavolineatum, French grunt; M alacoctenu8 macropus, Blenny; Opisthognathus sp., 77. Podocotyle pearsei Manter, 1934. Jawfish; Syngnathus robertsi, Pipefish. Host: Urophycis chesteri. 78. Eurycreadium vitellosum Manter, 1934. 64. Opecoelina helicoleni Manter, 1934. Host: Laemonema barbatulum. Host: Helicolenus rnaderensis. Subfamily Horatrematinae 65. Opecoelina scorpaenae Manter, 1934. Host: Scorpaena cristula/a, Scorpion fish. 79. Horatrema crassum Manter, 1947. Hosts: Odontoscion dentex, Corvina; Eques Subfamily Plagioporinat; acuminatus, Ribbonfish; E. lanceolatuS, er 66. Plagioporus crassigulus (Linton, 1910) Price, Ribbonfish; Haemulon carbonarium, Caes 1934. grunt. Hosts: Calamus bajonada, Jolt-head porgy; Subfamily Notoporinae C. calamus, Saucer-eye porgy; Decodon puellaris, Cuban hogfish; Diplodus hol­ 80. Neonotoporus yamagutii Manter, 1947. brookii, Spot-tail pinfish. Host: 'l'rachurops crumenophthalmus, Goggle­ eye. 67. Hamacreadium consueturn Linton, 1910. Family ACANTHOCOLPIDAE Host: Haemulon plumieri, White grunt. 81. M onorchistephanostomum gracile Vigueras, 1942. 68. Hamacreadium gulella Linton, 1910. Host: Sphyraena barracuda, Barracuda. Hosts: Lutjanus analis, Muttonfish; L. Reported from north coast of Cuba bY griseus, Gray snapper. Vigueras (I942). GULF OF MEXICO 339

82. Stephanostomum sp. 97. Diplangus paxillus Linton, 1910. Host: Galeichthys felis, Sea catfish. Hosts: Anistoremus virginicus, Porkfish; Recorded by Melugin (1940) from Grand Balistes vetula, Queen triggerfish; Brachy­ Isle, La. genys chrysargyreus, Bronze grunt; Hae­ mulon jlavolineatum, French grunt; H. 83. Stephanostomum casum (Linton, 1910) plumieri, White grunt; H. sciurus, Yellow MacFarlane, 1936. grunt; H. macrostomum, Gray grunt. Hosts: Ocyurus chrysurus, Yellowtail; Lut­ janus analis, Muttonfish; Llltjanus griseus, 98. Diplangus parvus Manter, 1947. Gray snapper. Hosts: Haemulon jlavolineatum, French grunt; H. carbonarium, Caesar grunt. 84. Stephanostomum lineatum Manter, 1934. Hosts: Laemonema barbatulum; Phycis 99. Diplangus miolecithus Manter, 1947. cirratus,' Urophycis regius. Hosts: Haemulon album, Margate fish; H. parra, Sailor's choice. 85. Stephanostomum microstephanum Manter, 1934. 100. Brachyenteron parexocoeti Manter, 1947. Host: Epinephelus niveatus, Snowy grouper. Host: Parexocoetus mesogaster, Flying fish. 86. Stephanostomum megacephalum Manter, 1940. 101. Brachyenteron peristedioni Manter, 1934. Host: Caranx latus, Horse-eye jack. Host: Peristedion platycephalum. 87. Stephanostomum promicropsi Manter, 1947. Family FELLODISTOMATIDAE Host: Promicrops itaiara, Jewfish. 102. Pycnadena lata (Linton, 1910) Linton, 1911. 88. Stephanostomum sentum (Linton, 1910) Manter, Host: Calamus calamus, Saucer-eye porgy. 1947. 103. Pycnadenoides calami Manter, 1947. Hosts: Calamus bajonado, Jolt-head porgy; Host: Calamus bajonado, Jolt-head porgy. C. calamus, Saucer-eye porgy; Haemulon sciurus, Yellow grunt; H. plumieri, White 104. Bacciger harengulae Yamaguti, 1938. grunt. Host: Harengllla dupeola, Pilchard. 89. Stephanostomum coryphaenae Manter, 1947. 105. Antorchis urna (Linton, 1910) Linton, 1911. Host: Coryphaena hippurus, Dolphin. Hosts: Angelichthys ciliaris, Queen angelfish; A. isobelita, Angelfish; Pomacanthus arcua­ 90. Stephanostomum dentatum (Linton, 1900) Man­ tus, Black angelfish. ter, 1931. Hosts: Epinephelus adsencionis, Rock hind; 106. Megalomyzon robustus Manter, 1947. E. morio, Red grouper; Mycteroperca Host: Lachnolaimus maximus, Hogfish. venenosa, Yellow-fin grouper. 107. Proctoeces erythraeus Odhner, 1911. Hosts: Calamus bajonado, Jolt-head porgy; 91. Stephanostomum ditrematis (Yamaguti, 1939) C. calamus, Saucer-eye porgy. Manter, 1947. Hosts: Caranx latus, Horse-eye jack; C. 108. M esolecitha linearis Linton, 1910. ruber, Runner. Host: Acanthuru.~ caeruleus, Blue tang. Family ZOOGONIDAE 109. Tergestia laticollis (Rud., 1819) Odhner, 1911. Hosts: A uxis thazard, Frigate mackerel; 92. Steganoderma parexocoeti Manter, 1947. Host: Parexocoet1tS mesogaster, Flying fish. Euthynnus alletteratus, Little tuna. 110. 'l'ergestia pectinata (Linton, 1905) Manter, 93. Steganoderma hemiramphi Manter, 1947. 1940. Host: H emiramphlls brasiliensis, Halfbeak. Hosts: Trachurops crumenophthalmus, Gog­ 94. Steganoderma elongatum Manter, 1947. gle-eye; Priacanthus arenatus, Big-eye. Host: Strongylura timucu, Houndfish. 111. Tergestia acuta Manter, 1947. Host: Caranx bartholomaei, Yellow jack. 95. Diphterostomllm americanllm Manter, 1947. Host: Brachygenys chrysargyreus, Bronze 112. Benthotrema plenum Manter, 1943. grunt. Host: Unidentified Lizard fish. 113. Lissoloma brotlllae Manter, 1934. 96. Deretrema fusillum Linton, 1910. Host: Brotllla barbata. Hosts: Abudefduf saxatilis, Sergeant major; Decodon puellaris; Ocyurus chyrsurlls, Yel­ 114. Lomasoma gracilis (Manter, 1934) Manter, lowtail; Priacanthus arenatus, Big-eye; 1935. Upeneus parvus, Goatfish. Host: Peristedion miniatum. 25!l534 0-54--23 340 FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE

115. Lomasoma monolenei (Manter, 1934) Manter, 129. Hurleytrema eucinostomi Manter, 1942. 1935. Host: Eucinostomus lefroyi, Mojal'ra. Host: M onolene antillarum. 130. Monorchis latus Manter, 1942. 116. Lomasoma wardi (Manter, 1934) Manter, 1935. Hosts: A nisotremus virginicus, Porkfish i Hosts: Coelorhynchus carminatus, Grenadier; Haemulon plumieri, White grunt. Urophycis regius. 131. Paraproctotrema brevicaecum Manter, 1942. 117. Megenteron crassum Manter, 1934. Host: Caranx bartholomaei, Yellow jack. Host: Diplacanthopoma brachysoma. 132. Postmonorchis orthopristis Hopkins, 1941. 118. Steringophorus magnus Manter, 1934. Host: Haemulon flavolineatum, French grunt. Host: Unidentified eel. 133. Proctotrema longicaecum Manter, 1940. 119. Steringophorus profundus Manter, 1934. Host: Anisotremus virginicus, Porkfish. Host: Argentina striata. 134. Proctotrema truncatum (Linton, 1910) Manter, Family HAPLOSPLANCHNIDAE 1940. 120. Haplosplanchnus acutus (Linton, 1910) Man­ Hosts: Haemulon album, Margate fish; Jl. ter, 1937. flavolineatum, French grunt; H. plumieri, Hosts: Strongylura raphidoma, Houndfish; White grunt; H. sciurus, Yellow grunt. Strongylura timucu, Houndfish. 135. Proctotrema parvum Manter, 1942. 121. Haplosplanchnus adacutus Manter, 1937. Host: Haemulon flavolineatum, French grunt. Hosts: Abudefduf marginatus, Sergeant major; Halichoeres bivittatus, Slippery Family BIVESICULIDAE dick; H. maculipinna, Slippery dick. 136. Bivesicula hepsetiae Manter, 1947. Host: Hepsetia stipes, Hardhead. 122. Haplosplanchnus brachyurus Manter, 1937. Hosts: Cryptotomus auropunctatus, Parrot­ Family GORGODERIDAE fish; Pseudoscarus guacamaia, Rainbow 137. Phyllodistomum carangis Manter, 1947. parrotfish; P s e u d 0 s caru s coelestinus, Host: Caranx ruber, Runner. Parrotfish; Sparisoma aurofrenatum, Parrotfish; S. spinidens, Parrotfish; S. 138. Xystretrum solidum Linton, 1910. viride, Parrotfish. Hosts: Balistes capriscus, Triggerfish; Lacto­ phrys triqueter, Trunkfish. 123. Haplosplanchnus kyphosi Manter, 1947. Hosts: Kyphosus sectatrix, Bermuda chub; 139. X ystretrum pulchrum (Travassos, 1921) Man­ H. incisor, Yellow chub. ter, 1947. Host: Sphoeroides splengleri, Sout.hern puf­ 124. Haplosplanchnus obtU8US (Linton, 1910) Man­ fer. ter, 1937. Hosts: Acanthurus coeruleus, Blue tang; A. Family MEGAPERIDAE hepatu8, Doctorfish. 140. Megapera gyrina (Linton, 1907) Manter, 1934. 125. Haplosplanchnus pomacentri Manter, 1937. Host: Lactophrys tricornis, Trunkfish. Hosts: Pomacentrus leucostictus, Beau greg­ 141. M egapera orbicularis (Manter, 1933) Manter ory; P. xanthurus, Beau gregory. 1934. 126. Haplosplanchnus sparisomae Manter, 1937. Host: Lactophrys tricornis, Trunkfish. Hosts: Sparisoma pachycephalum, Parrot­ 142. Megapera ovalis (Manter, 1933) Manter, 1934. fish; P. viride, Parrotfish. Host: M onacanthus hispidus, Filefish. Family MONORCHIDAE 143. M egapera pseudura (Manter, 1933) Manter, 127. Genolopa ampullacea Linton, 1910. 1934. Hosts: Bathystoma striatum; B. rimator, Host: Lactophrys tricornis, Trunkfish. Tomtate; Brachygenys chrysargyreus, Bronze grunt; Haemulon album, Margate 144. Thysanopharynx elongatus Manter, 1933. fish; H. carbonarium, Caesar grunt; H. Host: Lactophrys tricornis, Trunkfish. flavolineatum, Franch grunt; H. macro­ Family CRYPTOGONIMIDAE stomum, Gray grunt; H. plumieri, White grunt; H. sciurus, Yellow grunt; Synodus 145. Siphodera vinaledwardsii (Linton, 1899) Lin­ foetens, Lizardfish. ton, 1910. Host: Ocyurus chrysurus, Yellowtail. 128. Hurleytrema chaetodoni Manter, 1942. Hosts: Chaetodon capistratus, Butterfly fish; J46. Metadena crassulata Linton, 1910. C. ocellatuB, Butterfly fish. Host: Lutjanus analis, Muttonfish. GULF OF MEXICO 341

147. Metadena globosa (Linton, 1910) Manter, 1947. 162. Lethadena profunda (Manter, 1934), Manter Host: Lutjanus griseus, Gray snapper. 1947. Hosts: Pronotogrammus aureorubens; Xeno­ 148. Metadena adglobosa Manter, 1947. dermichthys capei. Host: Lutjanus griseus, Gray snapper. 149. M etadena brotulae (Manter, 1934) Manter, 163. Adinsoma robustum (Manter, 1934) Manter, 1947. 1947. Hosts: Brotula barbata; Lophius piscatorius, Hosts: Chaunax pictus, Frogfish; Chloroph­ Angler fish. thalmus truculentu..~; M erluccius sp.; Para­ lichthy.~ oblongus, Flounder; Urophycis Family HEMIURIDAE chesteri; U. regius. 150. Hemiurus sp. of Manter, 1934. Host: Peristedion imberbe. 164. Ectenurus virgulus Linton, 1910. Hosts: Bothus ocellatus, Flounder; Haren­ 151. Parahemiurus merus (Linton, 1910) Woolcock, gula clupeola, Pilchard; Trachurops cru­ 1935. menophthalmus, Goggle-eye jack. Hosts: Abudefduf saxatilis, Sergeant major; Harengula clupeola, Pilchard; Ocyurus 165. Parectenurus americanus Manter, 1947. chrysurus, Yellowtail; Synodus foetens, Hosts: Caranx bartholomaei, Yellow jack; Lizardfish; Trachurops crumenophthalmus, Synodus foetens, Lizardfish. Goggle-eye jack. 166. Dinurus torna/us (Rud., 1819) Looss, 1907. 152. Anahemiurus microcercus Manter, 1947. Host: Coryphaena hippurus, Dolphin. Hosts: Calamus bajonado, Jolt-head porgy; C. calamus, Saucer-eye porgy; Eucinos­ 167. Dinurus breviductus Looss, 1907. tomus lefroyi, Mojarra. Host: Coryphaena hippurus, Dolphin.

153. Lecithochirium mecosaccum Manter, 1947. 168. Dinurus longisinus Looss, 1907. Hosts: Synodus foetens, Lizardfish; S. poeyi, Host: Coryphaena hippurus, Dolphin. Lizardfish. 154. Lecithochirium microstomum Chandler, 1935. 169. Dinurus barbatus (Cohn, 1903) Looss, 1907. Hosts: A ncylopsetta dilecta, Flounder; Prom­ Host: Coryphaena hippurus, Dolphin. icrops itaiara, Jewfish; Megalops atlanticus 170. Dinurus scombri Yamaguti, 1934. Tarpon. Host: Euthynnus alletteratus, Little tuna. Reported also by Chandler (1935) from Trichiurus lepturus, Cutlass fish, in 171. Stomachicola rubea (Linton, 1910) Manter, Galveston Bay and from the same host 1947. at Grand Isle, La., by Melugin (1940). Hosts: Gyrnnothorax funebris, Black moray; 155. Lecithochirium sp. of Manter, 1934. G. rnoringa, Moray. Host: Urophycis regius. Reported from Synodus foetens at Grand Isle, La., by Melugin (1940). It also 156. Lecithochirium texanum (Chandler, 1941) occurs at Tortugas (Linton, 1910). Manter, 1947. Host: Sarda sarda, Bonito. 172. Aponurus laguncula Looss, 1907. Reported by Chandler (1941) from Texas Host: Ocyurus chrysurus, Yellowtail. coast near Freeport. 173. Aponurus interrnedius Manter, 1934. 157. Sterrhurus floridensis Manter, 1934. Hosts: Chaunax pictus; unidentified eel; Hosts: At least 21 species of fishes at unidentified sole. Tortugas (see Manter, 1947). 174. Brachadena pyriformis Linton, 1910. 158. Sterrhurusfusiformis (Luhe, 1901) Looss, 1907. Hosts: Bathystoma striatum; B"'achygenys Hosts: Gymnothol"ax moringa, Spotted mo­ chrysargyreus, Bronze grunt; Calarnus ray; G. funebris, Black moray. bajonado, Jolt-head porgy; Chaetodon aya, 159. Sterrhurus microcercus Manter, 1947. Butterfly fish; C. sedentarius; Haemulon Host: Fistularia tubaccaria, Trumpet fish. album, Margate fish; H. macrostomum, Gray grunt; H. parra, Sailor's choice; 160. Sterrhurus praeclarus Manter, 1934. H. plumieri, White grunt; Ogcocephalus Host: Merluccius sp. cubifrons, Batfish. 161. Dissosaccuslaevis (Linton, 1898) Manter, 1947. Hosts: Helicolenus maderensis; Peristedion 175. Lecithaster acutus (Linton, 1910) Manter, 1947. longispathum; P. min i a tum; P. Hosts: Acanthurus hepatus, Doctor fish; A. platycephalum. caeruleus, Blue tang. 342 FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE

176. Leurodera decora Linton, 1910. 191. Sclerodistomum sphoeroidis Manter, 1947. Hosts: A nisotremus virginicus, Porkfish; Host: Sphoeroides spengleri, Southern puffer Brachygenys c'hrysargyreus, Bronze grunt; H aemulon carbonarium, Caesar grunt; 192. Tetrochetus coryphaenae Yamaguti, 1934. H. parra, Sailor's choice; H. plumieri, Host: Coryphaena hippurus, Dolphin. White grunt; H. sciurus, Yellow grunt; 193. Prosogonotrema bilabiatum Vigueras, 1940. H. flavolineatum, French grunt; Lutjanus Host: Ocyurus chrysurus, Yellowtail. griseus, Gray snapper; Acanthurus hepatus Reported from north coast of Cuba by (1), Doctor fish. Vigueras (1940b). 177. Theletrumfustiforme Linton, 1910. Vigueras considered it in a new family. Host: Pomacanthus arcuatus, Black angelfish. Family APOROCOTYLIDAE Reported also from P. arcuatus and P. paru, French angelfish, from the north 194. Deontacylix ovalis Linton, 1910. coast of Cuba by Vigueras (1940a). Hosts: Kyphosus sectatrix, Bermuda chub; K. incisor, Yellow chub. 178. Derogenes crassus Manter, 1949. Host: Callionymus agassizii. 195. Cardicola cardiocolum (Manter, 1947) Short, 1953. 179. Derogenes varicus (Mueller, 1784) Looss, 1901. 196. Cardicola laruei Short, 1953. Hosts: Helicole~us maderensis; M erluccius Host: Cynoscion arenarius, White trout; sp.; Scorpaena cristulata; Setarches par­ C. nebulosus, Speckled trout. matus; Urophycis regius. 180. Gonocerca crassa Manter, 1934. HOST SPECIFICITY OF TREMATODES OF Hosts: A ncylopsetta dilecta; Brotula barbata; MARINE FISHES OF THE GULF OF Coelorhynchus carminatus; Lophius pis­ MEXICO catorius; M erluccius sp.,; M. bilinearis; Paralichthys oblongus; Paralichthys sp.; Monogenea are rather highly specific in regard P. squamilentus; Phycis cirratus; Saurida to their hosts. Most species are known to infect normani; Setarches parmatus; Synodontid; Synodus intermedius; Urophycis regius. in nature only a single host species. If there are several hosts they are usually related. 181. Gonocerca phycidis Manter, 1925. COll~ Hosts: Coelorhynchus carminatus, Grenadier; Host specificity among the Digenea varies M erluccius sp.; Urophycis regius. siderably. It is not marked among a number of 182. Gonocercella atlantica Manter, 1940. species infecting birds and mammals. Judging Host: Monacanthus hispidus, Filefish. from the collection records at Tortugas, trema~ todes of fishes there have very considerable host 183. Parasterrhurus anurus Manter, 1934. Host: Argentina striata. specificity. Considered from the species level of 189 species collected (Manter 1947) 105 or 55.5 184. Hemiperina nicolli Manter, 1934. Hosts: Chaunax pictus, Frogfish; Dibranchus percent were found in only 1 species of host; atlanticus; Diplacanthopoma brachysoma. 43 from 2 hosts, 14 from 3 hosts, etc., although 1 species occurred in 14 hosts, 1 in 16 hosts, and 1 in 185. Hysterolecitha rosea Linton, 1910. non~ Hosts: Acanthurus bahianus, Ocean tang; 21 hosts. While it must be admitted that A. caeruleus, Blue tang; A. hepatus, occurrence of a trematode within a certain species Doctor fish. of host does not necessarily mean it cannot infect 186. M acradena perfecta Linton, 1910. that host, data derived from numerous examina~ Host: Acanthurus caeruleus, Blue tang. tions indicate rather clearly that such species 187. Macradenina acanthuri Manter, 1947. do not do so under natural conditions. The Host: Acanthurus caeruleus, Blue tang. fishes studied were mostly shallow-water species 188. Opisthodena dimidia Linton, 1910. living not far from one another. The fact that a Hosts: Kyphosus sectatrix, Bermuda chub; few species do in fact infect a wide variety of K. incisor, Yellow chub. hosts indicates that food habits are not necessarily 189. Dictysarca virens Linton, 1910. a barrier. The only experimental testing of host Host: Gymnothorax funebris, Black moray' specificity in this region was that by McCoy (1930) who found that Hamacreadium mutabile and 190. Hirudinella ventricosa (Pallas, 1774) Baird, y. 1853. gulella developed only in three species of the Host: Coryphaena hippurus, Dolphin. snapper family, Lutjanidae, and did not infect GULF OF MEXICO 343 any of five other speCIes of fishes belonging in a number of species found only at 50 fathoms or five other families. below at Tortugas but known from surface waters If a trematode species occurs in 2 or several of the North Atlantic. Studies since 1934 add 2 hosts, the hosts are usually related. For example, species which do extend into surface waters at 138 trematode species (73 percent of those col­ Tortugas and some 6 species found in distant lected) were limited to a single host genus. Only oceans. The following table shows the distribu­ rarely is a trematode species found in fishes be­ tion of these species. Numbers in parentheses longing in different families. Little is known indicate the number of host species involved. regarding the host specificity of larval stages for The table is based on a total of 41 adult species their intermediate hosts. It is sometimes greater, definitely identified. particularly for the molluscan host, than that of the adult parasite. TABLE 1.-Distribution of trematode fauna This tendency of trematodes to occur in a Species Depth Other localities limited number of hosts which are usually related [alhoms gives added interest to the geographical dis­ a. Species reported elsewhere tribution of these parasites. During my study of (1934) Helicometra [asciala (3) ..• _ 50-60 Great Britain (10). Tortugas trematodes, I was frequently impressed European (20). to discover that the hosts of the same or a related Mexican Pacitlc (2), Derogencs varicus (5). _ 12&-200 Maine coast (6). North Atlantic (many). trematode in distant oceans proved to be a fish North Pacltlc (many). related to the host at Tortugas. Galapagos Islands (1). Dissosaccuslaevis (4). _ 16&-200 Woods Hole (1), deepwater. Gonocerca phgcidis (3) ... _ 160-300 Maine coast (1). Lepidapedon elongalum (4) _ 160-300 Maine coast (6). GEOGRAPHICAL DISTRIBUTION OF British Isles (1). Woods Hole (6). TREMATODES OF FISHES AT TORTU­ Now known from Japan (1). Lepidapeaon rach/on (1) _ 200-325 Maine coast (1). GAS, FLORIDA British Isles (4). Woods Hole (4). TREMATODES FROM 50 FATHOMS OR BELOW Sterrhurus jloridensis (8) _ 50-197 Tortugas (shallow), (many). b. Species reported elsewhere Systematic dredgings south of Loggerhead Key since 1934 under the direction of Dr. W. H. Longley in the Deretrema [usillum (1) _ 50 Woods Hole (1). Tortugas, shallow (1). summers of 1930-32 presented an unusual op­ Derogenes crassus (1) _ 00 Japan (1), deepwater (1). Tasmania (1), shallow. portunity for collection of trematode parasites Gonocerca crassa (12) _. _ 4lHl00 Japan (1), deepwater (1). Lecithochir/um microstomum (1) __ 100 Tortugas (1), shallow. down to depths of some 500 fathoms (Manter, Lepidapedon n/coll/ (1)------00 Mexican Pacitlc (1), shallow. Mgzotus vitellosus (1) _ 50-60 Tortugas (1), shallow. 1934). It was found that fishes down to that Prosorhgnchus ozakii (1) _ 00 Mexican Paeitlc (I), Shallow. depth were as commonly infected as fishes of Pseudopecoeloides vulgaris (16) _'_ 50-315 Puget Sound (1), shallow. shallow (surface) waters, 49 species being col­ lected from about 90 species of fishes. These The above data suggest that a bathymetric trematodes are almost entirely different species barrier between the trematode faunas of surface from those occurring in the nearby surface water; waters and of depths 100 fathoms or more at only 3 species from below 100 fathoms occurred in Tortugas is more effective than any existing be­ lesser depths and 2 of these are atypical (Dis­ tween the deeper fduna and distant cool oceans. tomum jenestratum is an immature, very non­ The deep-sea fauna itself is evidently very wide­ specific species, while Sterrhurus floridensis has so spread, but a number of its trematode species little host specificity that precocious maturity in a occur in more shallow depths near or in the Arctic crustacean host is suggested). Even at 50 fathoms and Antarctic Seas. only 3 additional species (Deretrema fusillus, The above comparisons have been on the spe­ Myzoxenus vitellosus, and Helicometrina nimia) cies level. It is of some further interest to note Occur in surface waters at Tortugas. This marked that of 13 new genera named in 1934 from deep difference in the deeper water fauna might be water at Tortugas, 7 involving 15 species have expected since the fishes and invertebrates are since been reported elsewhere as follows: One (1 also different species. species) from Woods Hole; four (11 species) from A more interesting aspect of the deeper wlLter Japan; two (3 species) from Tasmania. The trematode fauna is the affinity it shows for distant, species from Tasmania were in shallow-water cool or cold water faunas. Manter (1934) noted fishes. 344 FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE

A remarkable incident occurred in connection as might not be expected. There is one striking with Lepidapedon nicolli and Pro8orhynchus ozakii. difference among these regions. Whereas at These two species, new at the time, both occurred Bermuda the species of host fishes are, in general, at Tortugas at 90 fathoms in the same individual identical with those involved at Tortugas,·in the fish, a snowy grouper, Epinephelus niveatus. In Pacific the hosts are, with very few exceptions, 1934, I collected these same two species again different but related species. As far as the from a single fish, an "unidentified, spotted, trematodes are concerned, the tropical American grouper-like fish" from the Pacific Ocean at Isabel Pacific occupies a position comparable to that Island, Mexico. It is unfortunate that this fish of Bermuda in relation to Tortugas, almost as could not be further identified. As far as I can though there were no land barrier between the two learn, Epinephelus niveatus does not occur in the oceans. It is well known that this land barrier has Pacific. not always existed. Have the trematode parasites retained their specific identity while their hosts, TREMATODES FROM SURFACE WATER FISHES perhaps both fish and molluscan, have evolved A comparison of 142 species of trematodes from into slightly different species? Trematode dis­ less than 50 fathoms at Tortugas with faunas tribution suggests that just as the Gulf Stream known elsewhere reveals, in general, (1) that the has made Bermuda practically an outpost of the Tortugas fauna is very unlike that of the coast of Gulf of Mexico and Caribbean faunas, the Maine or Massachusetts, the European Atlantic tropical and subtropical American Pacific is also or the Mediterranean, and (2) it is strikingly such an outpost persisting as evidence of the similar to the fauna at Bermuda and the tropical prehistoric continuity of the two oceans. Amercian Pacific. Further evidence of an ancient influence of Of the 142 species at Tortugas, 28 species (19.8 inter-oceanic continuity is the relative dissimilar­ percent) occur at Bermuda (Hanson 1950). ity of the Tortugas fauna to such adjacent regionS This similarity is to be expected since many of -as Beaufort, N. C. Here the trematodes have the same species of fishes occur in both regions. been as well sampled as in the Pacific, and only 15 In fact, at least 21 of the 28 species of trematodes of the Tortugas species are known there. This occur in the same host species in both regions. number is hardly more than half the number known The only other part of the world where such to occur in the Pacific. If the trematodes were trematodes have been collected sufficiently to accidentally distributed by migrants such as give adequate s~mpling and which compares with birds, this difference would not be expected. the similarity to Bermuda is the tropical American Fourteen Tortugas species, from less than 50 Pacific. Considering species from 50 fathoms or fathoms, occur at Woods Hole, Mass. Eleven less, 24 or 16.8 percent of 142 Tortugas species occur in Japanese waters. This extension of occur in the American Pacific. Also, there is species far into the Pacific is reflected in the reason to believe this similarity is even greater Bermuda fauna. Five of the Bermuda species than this figure indicates. Three additional occur in Japan and six in the tropical American species in the Pacific occur in Bermuda fishes which Pacific, as compared with seven at much nearer occur at Tortguas; still three more species are now Woods Hole, six at Beaufort, three in the European reported (Pearse 1949) from Beaufort, N. C., in Atlantic, and three in the Mediterranean (Hanson fishes which occur at Tortugas. Thus, the 1950). This greater affinity of the Bermuda probable total is at least 30 species, a figure trematodes for Japanese than for European waters practically identical with that derived from com­ is better understood in the light of the Pacific parison with Bermuda. Furthermore, if fishes affiinities of the Tortugas fauna. down to 100 fathoms were considered, the Pacific The distribution of these trematodes is, of percentage would be slightly higher. course, a result of numerous and complex factors. Thus, based on our present knowledge, one can For example, a few of the species considered h?re conclude that the Tortugas trematode fauna of are parasites of open sea fishes with a very Wide surface-water fishes is very similar to that of range. Their parasites would be expected to Bermuda, as would be expected, and equally as have a wide distribution. Omission of these similar to that of the tropical American Pacific, species, however, would not materially change the GULF OF MEXICO 345 proportional figures. The species from deeper chelys imbricata. It is an interesting fact that \Vater were considered separately because parasites none of these trematodes is known to occur as an from these depths have not been studied in most adult parasite of fishes.! Furthermore, despite regions. Presumably, these species have a wide the aquatic life of the turtles, these trematodes distribution at appropriate depths. If sufficient belong to reptilian parasitic groups (families and numbers of species are involved, the exceptional genera). Turtles have no representatives of such cases or errors in identification will be minimized. common trematode families of fishes as Hemi­ In most cases, such factors would probably more uridae, Opecoelidae, Fellodistomatidae, etc. A Or less balance one another in the various localities. similar situation exists in connection with the lIowever, it seems to me that the figures available trematodes of marine mammals which are quite are not suitable for statistical analysis because of different from species occurring in either turtles factors which cannot be given numerical. value. or fishes. For example, from purely geographical considera­ Only a few turtles from the Gulf have been tions one would expect the Pacific fauna to be examined for parasites. Additional records, par­ Very different from Tortugas, separated as it is ticularly from Ohelone and Eretmochelys, could by a land barrier and with practically no pos­ easily be made. Such records would make pos­ sibility of contact by way of the Antarctic or the sible comparisons between the parasites of this Partly fresh-water Panama Canal. More com­ region and other seas. plete knowledge of these parasites in all regions TREMATODES OF CARETTA CARETTA is needed. As this knowledge is gained it seems probable that a very interesting host-parasite­ Fifteen species of trematodes from Oaretta distribution picture will unfold, particularly at caretta at Tortugas, Fla., have been reported by the specific and generic levels. Linton (1910), Pratt (1914), Manter (1932), and Luhman (1935). These are as follows: TREMATODES OF TURTLES ASPIDOGASTREA Four species of marine turtles occur in the Gulf ASPIDOGASTRIDAE of Mexico. These are the loggerhead turtle, 1. Lophotaspis vallei (Stossich, 1899) Looss, Oaretta caretta, the green turtle, Ohelonia mydas, 1902. the hawk-bill turtle, Eretmochelys imbricata, and DIGENEA the leathery or leatherback turtle, Dermochelys PRONOCEPHALIDAE coriacea. All four of these turtles have a very 2. Cricocephalus albus (Kuhl & van Hasselt, \Vide distribution in warm seas. They are very 1822) Looss, 1899. 3. Diaschistorchis pandus (Braun, 1901) Johns­ favorable hosts for trematodes, some of which also ton, 1913. appear to be widely distributed, others being 4. Pleurogonius trigonocephalus (Rud., 1809) reported as yet only from certain localities. '['he Looss, 1901. actual distribution of these trematodes can be 5. Pyelosornurn longicaecurn Luhman, 1935. BRACHYCOELIIDAE known only after the examinations made in various 6. Cyrnatocarpus undulatus Looss, 1899. regions are more or less equivalent. Until then, 7. Orchidasma amphiorchis (Braun, 1899) the presence of a species is more significant than Braun, 1901. its apparent absence. With only two exceptions, PLAGIORCHIDAE the trematodes described from marine turtles are 8. Pachypsolus ovalis Linton, 1910. 9. Pachypsolus tertius Pratt, 1914. all Digenea. Since a molluscan host is also reo 10. Styphlotrerna solitarius (Loass, 1899) quired in their life cycle it is probable that a Odhner, 1910. number of these trematodes will have a distribu­ RHYTIDODIDAE tion limited by the distribution of the molluscan 11. Rhytidodcs sccundus Pratt, 1914. host. GORGODERIDAE Considering the wide individual range of these 12. Phyllodistornurn cyrnbijorrne, (Rud., 1819) Braun, 1899. turtles in the open sea, a surprising variety of trematodes succeed in parasitizing them. Fifty­ I The presence orasingle specimen or Orchlda.ma amph/orchi. in Corgphama hlppums reported by Manter (193!) is without doubt an accidental and one species have been reported from Chelone temporary condition probably due to recent Ingestion or a young turtle by rnYdas, 30 from Oaretta caretta, and 9 from Eretmo- the IIsh. 346 FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE

DIGENEA-Continued Japanese waters, two from Australia, one from SPIRORCHIDAE Bermuda, and one from Ireland. 13. Hapalotrema synorchis Luhman, 1935. 14. Neospirorchis pricei Manter & Larson, TREMATODES OF DERMOCHELYS CORIACEA, THE 1950. LEATHERY OR LEATHERBACK TURTLE 15. Carettacola bipora Manter & Larson, 1950. Apparently only one species of trematode, Five of these species (Pyelosomum longicaecum, Pyelo8omum renicapite (Leidy 1856) Ruiz 1946, Pachypsolus ovalis, P. tertius, Rhytidoeds secundus, has been recorded from this turtle and none from and Hapalotrema synorchis) are known as yet only the Gulf of Mexico. However, Dr. A. C. Chandler from the Gulf of Mexico. informs me (by correspondence) he has collected this species of trematode from a leather-back TREMATODES OF C~ELONIA MYDAS, THE GREEN TURTLE turtle washed up near Galveston, Texas, some years ago. Lists by Hughes, Higginbotham, and Clary (1941) and Ruiz (1946) show that 51 species of TREMATODES OF BIRDS OF THE GULF trematodes have been reported from Chelonia OF MEXICO mydas in various parts of the world. Of these, Almost no study of trematodes collected from 10 species or about one-fifth of the total, are also birds from the Gulf of Mexico has been made. known from Caretta caretta. I believe no other Obviously such trematodes must be numerous. species of host is known to harbor as many as 50 ItseemsJ>eyond the scope of this paper to assemble species of trematodes. Surely, no other reptile the scattered records of trematodes collected else­ approaches this number. where even from birds which occur in the Gulf. No definite records of species have been pub­ Pratt (1911), Linton (1928), and Manter (1930) lished from this host in the Gulf of Mexico. all mention Galactosomum cochleariforme (Rud.) Manter (1930) states that he collected three as present in Fregata magnificens, the man-of-war species of trematodes from Chelonia at Tortugas bird, at Tortugas, and Chandler (1951) recorded but had not then identified them. This collection Galactosomumjregatae and a new strigeid, Schwartz­ is still incompletely studied. Two of the species, itrema seamsteri, from a specimen of this bird however, can be reported as Desmogonius dP.s­ near Corpus Christi, Tex. mogonius Stephens, 1911, known from the same Records from oceanic birds from the Caribbean host from the coast of Nicaragua; and Polyangium region include: linguatula (Looss 1899) Looss 1900, known from the Mediterranean and from Brazil. More ex­ Galactosomurn johnsoni Price, 1934, from Sula leu­ aminations of Chelonia mydas will, without doubt, cogastra, brown booby, from Puerto Rico. reveal additional species in Gulf waters. Ruiz Galacto8omum fregatae Prudhoe, 1949, from Fregata magnijicens, man-of-war bird, from Trinidad. (1946) reports 10 species from this host from the Galactosomurn darbyi Price, 1934, from Pelecanus coast of Brazil. Price (1939a) reports Rhytido­ occidentalis, brown pelican, from Dominican doides intestinal-is Price, 1939, and R. simuis Republic. Price, 1939, from a Chelonia mydas which died in M esostephanus appendiculatoides (Price, 1934) Lutz, 1935, from Pelecanus occidentalis, brown pelican, the Washington Zoo but does not state the origin from Dominican Republic. of the turtle. Mesostephanu8 jajardensis (Price, 1934) Lutz, 1935, from Sula leucogaster, brown booby, from Puerto TREMATODES OF ERETMOCHELYS IMBRICATA, Rico. THE HAWK-BILL TURTLE TREMATODES OF MAMMALS Apparently only one trematode has been re­ ported from this turtle in the Gulf region. It is Marine mammals, particularly dolphins and Diaschistorchis pandus (Braun) reported from whales, are commonly infected with trematodes. Cuba by Vigueras (1935). This trematode has No collections from such hosts, however, appear an almost world-wide distribution and also infects to have been made in the Gulf of Mexico. Caretta caretta and Chelonia mydas. Five species Sokoloff and Caballero (1932) described a trem­ of trematodes are known from E. imbricata from atode, Schizamphistoma manati from a manatee, GULF OF MEXICO 347

Manatu8 latiro8tri8, from the delta of the Panu~o Pinna carnea River, near Tampico, Mexico. Cercaria N gasterostome. Thais deltoidea STUDIES ON LARVAL STAGES AND LIFE Cercaria 0______echinostome. CYCLES OF TREMATODES OF THE Except for the life cycles studied by McCoy GULF OF MEXICO (1929, 1930) little attempt has been made to Miller (1925, 1926, 1927, 1929) and Miller and associate these cercariae with adult trematodes. M-cCoy (1929) have studied cercariae collected The problem is usually difficult because many Chiefly from snails from coral reefs near the Biolog­ characters used in classification do not appear in ical Laboratory at Tortugl1s. Most of these stud­ the cercaria. The mollusks studied might be ies were on the behavior of the cercariae and only infected with trematodes from such birds as the brief preliminary descriptions of the cercariae were pelican, tern, or frigate bird, or from marine given. The cercariae were named alphabetically turtles, as well as from fishes. McCoy proved that as Cercaria A, Cercaria B, etc. In 1929, Cercaria Cercaria A developed into Hamacreadium muta­ P was referred to as Cercaria jloriden8is. As is bile, and Cercaria B into Hamacreadium gulella. usual among marine mollusks, only a small per­ He believed Cercaria P developed into some centage of individuals are infected. During the species of Acanthocha8mu8. Second intermediate first season (1925) Miller examined 4,341 mollusks hosts were Thala880ma biJa8ciatus, bluehead, or belonging to 33 genera and including 50 species Halichoere8 bivittatu8, slippery dick. Manter and varieties, and found only 45 individuals ill­ (1932, 1933a) found that Cercaria J from Col­ eluding 9 species infected. Miller's later work umbella mercatoria penetrated into and encysted indicates that the incidence of infection may vary in the shrimp, LY8mata intermedia. He found in greatly from year to year. Altogether, 22 species this same shrimp and in Crangon jormo8um the of cercariae were studied. The type of each cer­ metacercariae of Helicometrina nimia. He also caria is indicated. The following list includes the pointed out that Cercaria L greatly resembles a mollusks found infected and the cercariae found common juvenile trematode known as Di8tomum in each: j enestratum. Schechter (1943) reports cercariae of Parorchi8 Astrea americana Cercaria A cotylocercous. acanthus from the oyster-drilling snail, Thais Cercaria B cotylocercous. jloridana hay8ae from Barataria Bay, La. The Astrea longispina adult of this echinostomid trematode is a parasite Cercaria C immature sporocysts. of birds, e. g., the herring gull, Laru.<: argentatu8. Cerithium litteratum Cercaria D ______gymnocephalous. It has been reported from Cuba by Vigueras Cercaria E gymnocephalous. (1940a) from the flamingo, Phoenicopteru8 ruber, Cercaria F huge-tailed monostorne. and from Nycticorax nycticorax hoactli. Cercaria G Xiphidiocercaria. Cable and McLean (1943) described a "ratten­ Cercaria P binoculate, lophocercouH. konig" cercaria, C. clau8ii Monticelli, from the Cercaria Q trioculate monostome. Cercaria R______large furcocercous. gastropod, Lamellaria leucosphaera, from the west Cercaria S______distorne, gymnocepha!ous. coast of Florida. Miller (1929) described the same Cercaria T huge-tailed mOllostorne. colonial aggregation of individuals for his Cercaria Cercaria U huge-tailed rnOllostome. W from Cerithium litt(watum at Tortugas. Cercaria W______huge-tailed mOllostomc. Larval stages of gasterostomes are common Cerithiwn algicola Cercaria H ______gymIlocephalous. parasites of bivalves. Oysters both of the Atlantic Columbella mercatoria and Gulf coasts are commonly infected with such Cercaria 1______cotylocercous (?) larvae. Pearse and Wharton (1938) reported Cercaria J cotylocercous. "Bucephalus gracile8cens (Rudolphi)" from oysters Crepidula aculeata Cercaria K ______trichocercous. in Apalachicola Bay, Fla. A study of these Cercaria L cystophorous. gasterostome larvae in oysters of the Gulf of Glllphis lisleri Mexico is being made by Sewell H. Hopkins of Cercaria M cotylocercous. Texas A. and M. Research Foundation. 348 FISHERY BULLETIN OF THE FISH AND WILDLIFE SEdVICE

The miracidium of Lophotaspis vallei, an as­ (less than 50 fathoms) shows a marked similarity pidogastrid of the loggerhead turtle was studied to such faunas at Bermuda and in the tropical by Manter (1932). Wharton (1939) discovered American Pacific, a similarity approximately that a juvenile stage of this trematode occurred in twice as great as is shown to Beaufort, N. C., or the flag conch, Fasciolaria gigas, collected in Woods Hole, Mass. Further study of this phe~ Gulf County, Fla. As Wharton indicated, mira­ nomenon, at both the generic and specific level, cidia doubtless penetrate the conch and develop is suggested. without reproduction to the infective stage. LITERATURE CITED Turtles would thus become infected by eating infected conchs. CABLE, R. M., and McLEAN, RICHARD A. 1943. The occurrence of Cercaria clausii Monticelli, a SUMMARY marine rattenkonig larval trematode, on the west coast of Florida. Notulae Naturae, Acad. Nat. Sci. Over 200 species of Trematoda reported from Philadelphia, No. 129, pp. 1-7. fishes of the Gulf of Mexico include 10 species CHANDLER, A. C. of Monogenea, 2 species of Aspidogastrea, and 1935. Parasites of fishes in Galveston Bay. Proc. U. S. 196 species of Digenea. Of the Digenea, 18 species Nat. Mus. 83: 123-157,7 pis. a are gasterostomes, 178 species are prosostomes. 1941. Two new trematodes from the bonito, Sard Most of these trematodes have been studied sarda, in the Gulf of Mexico. Jour. Parasit. 27(2): 183-184. only from the eastern portion of the Gulf at 1951. Trematodes from the man-o-war bird, Fregata Tortugas, Fla. magnificens rothschildi, on the Texas coast, with th~ Trematodes of other vertebrates of the Gulf description of a new species, Schwartzitrema seamstert . have been studied very little. Fifteen species Tex. Jour. Sci. 3: 186-189. FAUST, ERNEST CARROLL, and TANG, CHUNG-CHANG. have been reported from Caretta caretta, the logger­ 1936. Notes on new aspidogastrid species, with a con­ head turtle, at Tortugas. Two species are re­ sideration of the phylogeny of the group. Parasit. ported in this paper from Chelonia mydas, the 28: 487-501. green turtle. Three species of Trematoda are FUJII, HAROI,D. known in the Gulf. At least five other species 1944. Three monogenetic trematodes from marine have been reported from oceanic birds in the fishes. Jour. Parasit. 30(3): 153-158, 2 pis. HANSON, MARY LOUISE. Caribbean, birds which are also common in the 1950. Some digenetic trematodes of marine fishes of Gulf. Bermuda. Proc. Helminth. Soc. Washington 17: Larval stages of trematodes in mollusks of the 74-89. Gulf include 22 species of cercariae at Tortugas. HUGHES, R. CHESTER; HIGGINBOTHAM, JOE W.; and Cercariae of Parorchis acanthus have been re­ CLARY, JASPER W. ported from Thais floridana from the Louisiana 1941. The trematodes of reptiles. Part II. Host cat­ coast. Here also oysters are commonly infected alog. Proc. Okla. Acad. Sci. 21: 35-43. with gasterostome larvae. LINTON, EDWIN. 1910. Helminth fauna of the Dry Tortuga9. II. Tre­ Only three life cycles are known. Hamacrea­ matodes. Carnegie lnst. Washington Pub. No. 133, dium mutabue and H. gulella, adults of which occur Pap. Tortugas Lab. 4: 11-98, 28 pis. in Lutjanidae (snappers), develop in a snail, 1928. Notes on trematode parasites of birds. Proc. Astrea americana, and then utilize Thalassoma U. S. Nat. Mus. 73 (1): 1-36, 11 pis. bijasciatus, bluehead, or Halichoeres bivittatus, LUHMAN, MARION slippery dick, as second intermediate host. Ju­ 1935. Two new trematodes from the loggerhead turtle (Caretta caretta). Jour. Parasit. 21 (4): 274-276. venile stages of Lophotaspis vallei from the logger­ MACCALLUM, G. A. head turtle infect the flag conch, Fasciolaria gigas. 1917. Some new forms of parasitic worms. ZoopathO­ A discussion of geographical distribution of logica 1 (2): 43-75. trematodes of fishes points out (1) that the trema­ MANTER, H. W. todes of fishes from 100 fathoms or deeper show 1930. Studies on the trematodes of Tortugas fisheS. practically no affinity to trematodes of surface Carnegie lnst. Year Book 29: 338-340. waters of the Gulf but do show considerable 1931. Further studies on trematodes of Tol'tugas fisheS. Carnagie lnst. Year Book 30: 386-387. affinity to species from fishes of distant but cold 1932. Continued studies on trematodes of TortUgaB• waters, (2) the trematode fauna of surface waters Carnegie lnst. Year Book 31: 287-288. GULF OF MEXICO 349

MANTER, H. W. PEARSE, A. S. 1933a. A new family of trematodes from marine fishes. 1949. Observations on flatworms and nemerteans col­ Trans. Am. Micros.. Soc. 52 (3): 233-242, 1 pI. lected at Beaufort, N. C. Proc. U. S. Nat. Mus. 1933b. The genus Helicometra and related trematodes 100: 25-38. from Tortugas, Florida. Carnegie lnst. Pub. No. 435, PRATT, HENRY S. Pap. Tortugas Lab. 28 (11): 167-182, 3 pis. 1910. Monocotyle floridana, a new monogenetic trema­ 1934. Some digenetic trematodes from deepwater fish tode. Pap. Mar. BioI. Lab. Tortugas, 4: 1-9. of Tortugas, Florida. Carnegie lnst. Pub. No. 435, 191!. On Galactosomum cochleariiorme Rudolphi. Zool. Pap. Tortugas Lab. 27: 257-345, 15 pis. Anzeig. 38: 143-148. 1935. The structure and taxonomic position of M eguso­ 1914. Trematodes of the loggerhead turtle (Caretta lena estrix Linton, 1910, (Trematoda) with notes on caretta) of the Gulf of Mexico. Arch. Parasit. 16: related trematodes. Parasit. 27 (3): 431-439. 411-427, 2 pIs. 1937. The status of the trematode genus Deradena 1916. The trematode genus Stephanochasmus Looss in Linton with a description of six species of Haplos­ the Gulf of Mexico. Parasit. 8 (3): 229-238, 1 pI. phanchnus Looss (Trematoda). Skrjabin Jubilee PRICE, EMMETT W. Volume, pp. 381-387. 1939a. North American monogenetic trematodes. III. 1940a. Gasterostomes (Trematoda) of Tortugas. Car­ The family Capsalidae (Capsaloidea). Jour. Wash­ negie lnst. Washington Pub. No. 524, Pap. Tortugas ington Acad. Sci. 29 (2): 63-92. Lab. 33: 1-19. 1939b. A new genus and two new species of digenetic 1940b. The geographical distribution of digenetic trematodes from a marine turtle. Proc. Helminth trematodes of marine fishes of the tropical American Soc. Washington 6 (1): 24-25. Pacific. Allan Hancock Pac. Exp. 2 (16): 531-547. 1942. Monorchidae (Trematoda) from fishes of Tortu­ READ, CI,ARK P. gas, Florida. Trans. Am. Micros. Soc. 61: 349-360, 1947. A new trematode, Opecoeloides polyfimbiatus n. 2 pis. sp., from the lizard fish, Synodus foeten.~. Jour. 1947. The digenetic trematodes of marine fishes of Tor­ Parasit. 33: 231-233. tugas, Florida. Am. Mid. Nat. 38 (2): 257-416,21 pIs. Rurz, JosE MANOEL. 1949. An additional trematode from Tortugas, Florida, 1946. Pronocephalidae (Trematoda). Estudo das and a new name for Opisthoporus Manter, 1947, pre­ especies brasileiras c reVlsao da familia. Mem. occupied. Am. Mid. Nat. 41 (2): 432-435. Inst.Butantan 19: 249-372. --- & LARSON, MARY I. SCHECHTER, VICTOR. 1950. Two new blood flukes from a marine turtle 1943. Two flatworms from the oyster-drilling snail, Caretta caretta. Jour. Parasit. 36 (6): 595-599. 'Thais floridana haysae Clench. Jour. Parasit. 29: 362. McCoY, OLIVER R. SHoR-r, ROBERT B. 1929. The life-history of a marine trematode, Hama­ 1953. A new blood fluke, Cardicola laruei n. g., n. sp. creadium mutabile Linton, 1910. Parasit. 21: 220-225. (Aporocotylidae) from marine fishes. Jour. Parasit. 1930. Experimental studies on two fish trematodes of 39 (2): 304-309. the genus Hamacreadium (Family Allocreadiidae). SOKOLOl'~" DEMETRIO, and CABALLEHO, EDUARDO. Jour. Parasit. 17: 1-13. 1932. Una nueva espccie de trematodo parasito del MELUGIN, JANE. intestino del manati. Anal. lnst. BioI. 3: 163-167. 1940. Studies on marine fish trematodes of Lou isiana. Abstr. Theses, Louisiana State Univ. (1938-39), VlGUERAS, 1. PEREZ. Univ. Bull. 32, n. s. 1, p. 89. 1935a. Sobre la presencia en Cuba de Diaschistorchis MILLER, H. M., JR. pandus (Braun) (Trematoda) parasito de Chelonia 1925. Preliminary report on the larval trematodes in­ imbricata. Rev. Parasit.,Clinical Lab., Hab. 1: 167-168. festing certain molluscs from Dry Tortugas. Carnegie 1935b. Tristomum poeyi n. sp. (Trematoda), parasito lnst. Year Book No. 24, pp. 232-238. de Makaira ampla Poey (Pisces). Mem. Soc. Hist., 1926. Behavior studies on Tortugas larval trematodes, Nat. Habana 9 (1): 43-44. with notes on the morphology of two additional 1940a. Macrorchitrema havanensis, n. gen., n. sp. species. Carnegie lnst. Year Book No. 25, pp. (Trematoda Paramphistomidae), parasito del intestino 243-247. de Holacanthus tricolor Bloch (Pisces). Anal. Hist. 1927. Further studies on the behavior of larval trema­ BioI. 11 (1): 197-207. todes from Dry Tortugas. Carnegie lnst. Year Book 1940b. Notas sobre algunas esp6cies nuevas de trema­ No. 26, pp. 224-226. todes y sobre otras poco concoidas. Pub. Revist. 1929. Continuation of study on behavior and reactions Univ. Habana No. 28 & 29, pp. 1-28, 16 pis. of marine cercariae from Tortugas. Carnegie lnst. 1940c. Prosogonotrematidae n. fam. y Prosogonotrema Year Book No. 28, pp. 292-294 bilabiatum n. gen., n. sp. (Trematoda, Distomata) MILLER, H. M., and McCoY, O. R. parasito de Ocyurus chrysurus (Bloch) (Pisces). 1929. An experimental study of the behavior of the Mem. Soc. Cub. Rist. Nat. 14 (3): 249-252, 1 pI. Cercaria floridensis in relation t,o its fish intermediate 1942. Notas helminthologicas. Univ. Habana Pub. host. Carnegie lnst. Year Book No. 28, pp. 295-297. bimestral, pp. 193-223. 350 FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE

VON WICKLEN, JANE HOGAN. WHARTON, G. W. 1946. The trematode genus Opecoeloides and related 1939. Studies on Lophotaspis vallei (Stossich, 1899) genera, with a description of Opecoeloide.~ polynemi, (Trematoda; Aspidogastridae). Jour. Parasit. 25: n. sp. Jour. Parasit. 32 (2): 156-163, 1 pI. 83-86. CESTODA

By ASA C. CHANDLER, Rice Institute The first work on cestodes of Gulf fishes was lesser extent other sciaenid fishes, very fre­ done by Linton on material collected by him at quently harbor the plerocerci of Poecilancistrium the Biological Laboratory of the Carn~gie In­ robustum or related species which are known to stitution at Dry Tortugas, Florida, in the sum­ fishermen as "spaghetti worms" because of their mers of 1906-1908. They are discussed in Linton great length. The adults of these worms are (l908b, 1909). No further work was done until probably parasitic in a shark or ray as are other Chandler (1935a, 1935b) reported some ob­ Trypanorhyncha. Some city health departments servations on helminths of fish in Galveston Bay, have considered banning drum from the markets Texas. Perez Vigueras (1936), in a report of because of their very frequent infestation, but the helminths in Cuba, reported three unidentified writer has counseled against this since these fish larvae of Trypanorhyncha from teleosts off the are an important cheap source of protein food. province of Havana. Potter (1937) described one additional new species of cestode from Dry TETRAPHYLLIDEA Tortugas, and Shuler (1938) reported on some Family DISCULICIPITIDAE cestodes collected from this same locality by Manter in 1930-1932, adding one more valid Disculicep,~ pileatus (Linton, 1890) Joyeux and Baer, 1935. new species. Chandler (1942) reported on some Host: CarcharimJ,s leucas, cub shark. cestodes from sharks taken near Englewood, Family CEPHALOBOTHRIIDAE Florida, adding two new species. Seamster HexacanaLis (?) marsupium (Linton, 1916) Dollfus, 1948. (1950, personal communication) reported a few Host: Stoasodon narinari, spotted sting ray; Dry Tor- cestode infections from fishes taken in or near tugas (Linton). . Corpus Christi Bay, Texas. Other than these Family PHYLLOBOTHRIIDAE few and incomplete investigations the cestodes of fishes in the Gulf of Mexico have not been Phyllobrothriwn foliaturn Linton, 1890. Host: Dasyat1tS sabina, southern sting ray; Dry Tor­ studied. Further studies will undoubtedly yield tugas (Linton). lhany new species and bring to light interesting Phyllobothriurn rnustelis (van Beneden, 1850) (~Orygrna­ geographical relations with the fauna of the tobothriurn angusturn Linton, 1890.) Caribbean Sea, the North Atlantic, and the Host: Carcharinus Leucas, cub shark; Dry Tortugas Pacific Coast of Mexico. No cestodes have yet (Linton) . PhyUobothriurn Lactuca van Beneden, 1850. been reported from marine reptiles or mammals in Host: N egaprion brevirostris, yellow shark; Dry Tor­ the Gulf. tugas (Shuler). The cestode fauna so far known consists almost Phyllobothriurn dasybati Yamaguti, 1934. entirely of Tetraphyllidea and Trypanorhyncha Host: N egaprion brevirostris, yellow shark; Dry Tor­ tugas (Shuler). Which, as adults, parasitize practically all elasmo­ Phyllobothrium turnidnm Linton, H)22. branchs in the Gulf and, as larvae, are found very Host: Scoliodon terrae-novae, sharp-nosed shark; Dry commonly in the flesh or viscera of teleosts, Tortllgas (Shuler). often in such food fishes as members of the Phyllobothrium centrururn Southwell, 1925 (=Anthocepha­ families Sciaenidae, Serranidae, and Lutjanidae. lurn gracile Linton, 1890). Although incapable of development in man they Host: Dasyatis sabina, southern sting ray; Dry Tor­ tugas (Linton). cause considerable economic loss because of Phyllobothrium sp. Popular antipathy to "wormy" fish. On the Host: Dasyatis sabina, southern sting ray; Corpus Texas coast the drum, Pogonias cromis, and to a Christi Bay (Seamster). 351 352 FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE

Rhinebothri1lm flexile Linton, 1890. Proteocephalus elongatus Chandler, 1935a. Host: Dasyatis sabina, southern sting ray; Dry Tortugas Host: Lepisosteus osseus, long-nosed gar; Galveston (Linton). Bay (Chandler). Echeneibothrium minimum van Beneden, 1850. Scolex pleuronectis Muller, 1788 (=S. polymorphus Rud., Host: Dasyatis sabina, southern sting ray; Corpus 1819, of many writers). Christi Bay (Seamster). A larval form in intestine, cystic duct, or gall bladder Anthobothrium laciniatum Linton, 1890. of many teleost fishes including Epinephelus striatus, Hosts: Hypoprion brevirostris; Dry Tortugas (Shuler); A uxis thazard (not a Gulf fish; probably Euthynnus allet­ Carcharinus leucas, cub shark; Dry Tortugas (Linton). teratus), M ycteroperca bonaci, Lutjanus griseus. and Ocyurus Anthobothrium variabile (Linton, 1890) Southwell, 1925. chrysurus at Dry Tortugas (Linton), and Galeichthys felis Host: Dasyatis sabina, southern sting ray; Dry Tortugas and Bagre marina from Galveston Bay (Chandler). (Linton). Anthobothrium sp. TRYPANORHYNCHA Host: Dasyatus sabina, southern sting ray; Corpus Christi Bay (Seamster) . Family TENTACULARIIDAE Nybelinia palliata (Linton, 1924) Joyeux and Baer, 1936. Family ONCHOBOTHRIIDAE Host: Sphyrna zygaena, hammer-head shark; Engle­ Acanthobothrium brev'/ssime Linton, 1909. wood, Fla. (Chandler). Host: Dasyatis sabina, southern sting ray; Dry Tortu­ Family EUTETRARHYNCHIDAE gas (Linton). Acanthobothrium coronatum (Rud., 1819) van Beneden, Eutetrarhynchus linea/us (Linton, 1909) Dollfus, 1942. 1850 (=Onchobothrium tortum Linton, 1916). Host: Ginglymostoma cirratum, nurse shark; Dry Tor­ Host: Stoasodon narinari, spotted sting ray; Dry Tor­ tugas (Linton, Shuler). tugas (Linton). Family OTOBOTHRIIDAE Acanthobothrium sp. Otobothrium penetrans Linton, 1907. Host: Dasyatis sabina, southern sting ray; Corpus Hosts: Carcharinus leucas, cub shark; Dry Tortugas Christi Bay (Seamster). (Linton); Carcharinus limbatus, spot-fin ground Thysanocephalum thysanocephalum (Linton, 1889) Braun, shark; Scoliodon terrae-novae, sharp-nosed shark; Dry 1900. Tortugas (Shuler). Host: Galeocerdo cuvier, tiger shark; Dry Tortugas Otobothrium curtum (Linton, 1909) Dollfus, 1942. (Linton). Hosts: Galeocerdo cuvier, tiger shark; immature, Thysanocephalum rugosum Chandler, 1942. M ycteroperca bonaci, black grouper; Epinephelus Host: Galeocerdo cuvier, tiger shark; Englewood, Fla. striatus, Nassau grouper; Dry Tortugas (Linton). (Chandler) . Otobothrium crenacolle Linton, 1890. Cylindrophorus lasius (Linton, 1890) Southwell, 1925. Host: Carcharinus leucas, cub shark; Dry Tortugas Host: Carcharinus leucas, cub shark; Dry Tortugas (Linton). (Linton). Poecilancistrium robustum (Chandler, Hl35b) Dollfus, 1942. Cylindrophorus exceptus (Linton, 1924) Southwell, 1925. Hosts: 1m. Cynoscion nebulosus, spottcd sca trout; Host: Carcharinus leucas, cub shark; Rockport, Tex. Galveston Bay (Chandler); also abundant in Pogonias (Seamster). cromis on Texas coast (Chandler, unpublished). Pedibothrium globicephalum Linton, 1909. Diplootobothrium springeri Chandler, 1942. Host: Ginglymostoma cirratum, nurse shark; Dry Tor­ Host: Sphyrna tudes, hammer-head shark; Englewood. tugas (Linton,. Fla. (Chandler). Pedibothrium longispine Linton, 1909. Family DASYRHYNCHIDAE Host: Ginglymostoma cirratum, nurse shark; Dry Tortu­ gas (Linton). Dasyrhynchus variouncinatus (Pintner, 1913) Pintner, 1928 Pedibothrium brevispine Linton, 1909. (= Tentacularia insignis (Linton, 1924) Shuler, 1938; Host: Ginglymostoma cirratum, nurse shark; Dry Tor­ Dasyrhynchus insigne Chandler, 1942). tugas (Linton). Host: Negaprion brevirostris, yellow shark; Dry Tor­ tugas (Linton, Shuler). Platybothrium hypoprioni Potter, 1937. Callitetrarhynchus gracilis (Rud., 1819) Dollfus, 1942 Host: Negaprion brevirostris, yellow shark; Dry Tortugas (= Rhynchobothrium speciosum Linton, 1897; 7'entacu­ (Potter). laria lepida Chandler, 1935a; Tentacularia pseudodera Onchobothrium sp. Linton, 1907. Shuler, 1938). Host: Stoasodon narinari, spotted sting ray; Dry Tortu­ Hosts: Negaprion brevirostris, yellow shark; Dr! gas (Linton). Tortugas (Shuler); Jm., Epinephel1ls straitus, Nassau Family PROTEOCEPHALIDAE grouper, Mycteroperca bonaci, black grouper, M· venenosa, yellow-fin grouper, Lutjanus griseus, gr~y Proteocephalus australis Chandler, 1935a. snapper; Dry Tortugas (Linton); Galeich/hys fel~8, Host: Lepisosteus osseus, long-nosed gar; Galveston gaff-topsail catfish, Bagre marina, sea catfish; Gal­ Bay (Chandler). veston Bay (Chandler). GULF OF MEXICO 353

Family LACISTORHYNCHIDAE Rhynchobothrium exile Linton, 1909 (probably a Cal­ litetrarhynchus according to Pintner, 1931; Dollfus Grillotia perelica (Shuler, 1938) Dollfus, 1942. disagrees) . Host: N egaprion brevirostris, yellow shark; Dry Tortugas Host: Galeocerdo cuvier, tiger shark; Dry Tortugas (Shuler) . , (Linton). In addition, various unidentified species Grillotia sp. (identified as G. heptanchi group by Dollfus). of Rhynchobothrium, both adults and larvae, were Host: 1m., Cynoscion nebulosus, Rockport, Tex. (un­ reported from various fishes from Dry Tortugas by published) . Linton (1909). Perez Vigueras reported three species of larval Trypanorhyncha from marine teleosts off Family PTEROBOTHRIIDAE Havana, Cuba, and Seamster (personal communication) reported a "Tetrarhynchus sp." from Carcharinus Pterobothrium heteracanthum Diesing, 1850 (=Syndes­ leucas off Rockport, Tex. mobothrium fillicolle Linton, 1890; Synbothrium fillicolle Linton (1897); Gymnorhynchus gigas Chandler, PSEUDOPHYLLIDEA 1935). Family PTYCHOBOTHRIIDAE Hosts: Dasyatis sabina, southern sting ray; Dry Tortugas (Linton); 1m., Calamus calamus, saucer-eye porgy; Plychobothrium belones (Duj., 1845) Lonnberg, 1869 Dry Tortugas (Linton); Galeichthy.~ felis, gaff-topsail (= Dibothrium resliforme Linton, 1891). catfish, Micropogon undulatu,~, croaker; Galveston Hosts: Strongylura notata, needlefish, S. rhaphidoma, Bay (Chandler). houndfish; Dry Tortugas (Shuler). Clestobolhrium crassiceps (Rud., 1819) Liihe, 1899. Pterobothrium lintoni (MacCallum, 1916) Dollfus, 1942 Hosts: M erluccius bilinearis, silver hake, Chlorophthalmus (= Tetrarhynchus erinaceus Linton, 1897; Gymnorhyn­ truculentus, green-eye, M erluccius sp.; Dry Tortugas chus malleus (Linton, 1924; Chandler, 1935). (Shuler). Host: 1m., Galeichthysfelis, gaff-topsail catfish; Galveston Bothriocephalus sp. (Linton, 1908) (= Dibolhrium sp., Bay (Chandler). Linton, 1908). Host: 1m., Hepsetia stipes, silverside; Dry Tortugas Incertae Sedis (Linton). Rhynchobothrium simile Linton, 1909 (possibly a Lacis­ CYCLOPHYLLIDEA torhynchus according to Dollfus, 1942). Incertae Sedis Host: Ginglymostoma cirratum, nurse shark; Dry Tortugas (Linton). Glossocercus cyprinodontis Chandler, 1935 (probably Dilepididae). Rhynchobothrium tenuispine Linton, 1890. Host: 1m., Cyprinodon variegatus; Galveston Bay Host: Ginglymostoma cirratum, nurse shark; Dry (Chandler). Tortugas (Liilton). Cysticercoides menidiae Chandler, 1935. Rhynchobothrium binuncum Linton, 1909. Host: 1m., Menidia berrylina peninsulae, silversides; Host: Dasyatis sabina, southern sting ray; Dry Tortugas Galveston Bay (Chandler).

(Linton). NOTE.-lllbllogmphy follows chapter OD Nematoda, p. 368. Blank page retained for pagination ACANTHOCEPHALA

By ASA C. CHANDLER, Rice Institute

In contrast to the trematodes and cestodes, very low bays throughout the Gulf where small Crus­ few species of Acanthocephala have been reported tacea, which probably serve as intermediate hosts, from Gulf animals. To some extent, certainly, abound; but they will probably not be found this is due to the scantiness of investigations, par­ abundantly in oceanic or reef-dwelling fishes. ticularly on parasites of fishes of bays and estu­ aries, but enough work has been done to make it EOACANTHOCEPHALA evident that the Acanthocephala are not as abun­ Family NEOECHINORHYNCHIDAE dant in the Gulf as they are in more northern waters, e. g., Woods Hole, Massachusetts. Linton Atactorhynchu8 verecundu8 Chandler, 1935a (1907) called attention to this relative scarcity of Host: Cyprinodon variegatu8; Galveston Bay (Chandler) Acanthocephala, both of species and of individuals, PALAEACANTHOCEPHALA in southern seas, for he observed it not only at Dry Tortugas in the Gulf but also at Beaufort, Family GORGORHYNCHIDAE North Carolina, and at Bermuda. In a later col­ Gorgorhynchu8 gibber Chandler, 1934 lection at Dry Tortugas (Linton 1909) he re­ Host: GaieichthY8 feli8, gaff-topsail catfish; Galveston rnarked that Acanthocephala were found in only Bay (Chandler) 7 of the 32 species of fish examined, and in every Nippo8trongylu8 ornatu8 (Van Cleave, 1918) Van Cleave case few or even only 1 was found, all belonging and Lincicome, 1940 (= Rhadinorhynchus pristi8 (Rud., 1802) of Linton, 1891-1909) to a single species which he called Echinorhynchus Hosts: A uxis thazard (Euthynnus alletteratus?), frigate pristis, now known as Nipporhynchus ornatus. In mackerel, Haemulon sciurus, yellow grunt, Haemulon a personal communication Seamster (1950) rc­ plumieri, white grunt, Lutjanus griscus, gray snapper, ported only 1 specimen of 30 fish examined in and four other species, unnamed; Dry Tortugas Corpus Christi Bay, Texas, representing 16 species, (Linton) Filisoma fidum Van Cleave and Manter, 1947 to harbor an acanthocephalan. Chandler (1935a), Host: K yph08US sectatrix, rudder fish; Dry Tortugas on the other hand, in an examination of 23 species (Van Cleave and Manter) of teleost fish from Galveston Bay represented by Family RHADINORHYNCHIDAE from 1 to over 100 specimens of each found a predominance of Acanthocephala as common adult Illiosentis furcatus Van Cleave and Lincicome, 1939 Parasites. However, no elasmobranchs were ex­ Host: M enticirrhus americanu8, southern kingfish; amined so the varied tetraphyllidean and trypa­ Grand Isle, La. (Van Cleave and Lincicome), Corpus Christi Bay, Tex. (Seamster) norhynchan cestodes found practically universally 'Telo8enti8 tenuicorni8 (Linton, 1891) Van Cleave, 1947 in the spiral valves of these hosts did not come Hosts: llIicropogon undulatu8, croaker, Leio8tomu.~ into the picture. The three adult species of xanthuru8, spot; Galveston Bay (Chandler) Acanthocephala found were all fairly common in Family CENTRORHYNCHIDAE their respecHve hosts and were the only adult Parasites that one could depend on finding in Arhythmorhynchu8 duocinctus Chandler, 1935a repeated examinations of particular hosts. Acan­ Host: 1m., ParalichthY8 lethostigmu8, southern flounder; thocephala will probably be found to be fa,irly Galveston Bay (Chandler) cornmon in fishes frequenting the shores and shal- NOTE.-Blbliogrnphy follows chapter on Nematoda, p. 358.

259534 0-54--24 355 Blank page retained for pagination NEMATODA

By ASA C. CHANDLER, Rice Institute

Remarkably few nematodes, either adults or Most of the nematodes found by Linton have larvae, have been identified in fishes or other been inadequately described and have not been animals of the Gulf. The only work done on them given specific names. The larval forms found by known to the writer is Linton's work at Dry Tor­ Chandler were all described as fully as the material tugas (1907-1909) and my own work on fishes of available would permit and were all tentatively Galveston Bay (1935a). In addition, a single designated new species until the adults become species of oxyurid, Laurotrava8soxyuris trava8sosi, known. Was described by Perez Viguel'as (1938) from a ASCARIDATA teleost on the north coast of Cuba. Family OXYURIDAE Linton called attention to the fact that nema­ todes were sparingly represented, as were Acantho­ Laurotravassoxyuris travassosi Perez Vigueras, 1938. cephala, in the fishes he examined at Dry Tortugas Host: Holacanthus tricolor, rock beauty; Havana (Perez Vigueras). as compared with the large numbers, particularly of encysted immature forms, in fishes at Woods Family ASCARIDIDAE lIole, Massachusetts. In the Gulf fishes immature Acanthocheilus sp. Linton, 1909. encysted forms were found in only 7 of 32 species Host: Ginglymostoma cirratum, nurse shark; Dry Tor­ tugas (Linton). e:x:amined and in small numbers in all except the Amphicaecum parvum Chandler, 1935a. Yellow-fin grouper, Mycteroperca venenosa, whereas Host: 1m., Dorosoma cepedianum, gizzard shad; Galves­ in the north such nematodes were found in the ton Bay (Chandler). \1iscera of a large number of species of fishes, often Contracaecum chaunaxi Olsen, 1952. in great numbers. Linton found adult nematodes Host: Chaunax sp.; Dry Tortugas (Olsen). Contracaecum collieri Chandler, 1935a. in only 6 species of fish, "(l elasmobranch and 5 Hosts: 1m., Cyprinodon variegatus, broad killifish, Para­ teleosts)" but except for the species found in the lichthys lethostigmus, southern flounder; Galveston stomach of a nurse shark only 1 or 2 specimens Bay (Chandler). Were found in a host. Contracaecum histiophori Yamaguti, 1935. Host: Istiophorus americanus; Florida coast (Olsen). Chandler found nematodes, like Acanthoce­ Contmcaecum ogcocephali Olsen, 1952. Phala, to be relatively more frequent in the inshore Host: Ogcocephalus radt'utus; Dry Tortugas (Olsen). fishes of Galveston Bay than in the oceanic and Contracaecum robustum Chandler, 1935a. reef fishes examined by Linton, but they could Hosts: 1m., Mugil cephalus, mullet, Fundulus sp. (probably grandis) , common killifish; Galveston Bay certainly not be considered abundant, either in (Chandler). species or individuals. In 23 species of teleosts Goezia minuta Chandler, 1935a. stUdied 3 adult nematodes were found, each Host: Bagre marina, sea catfish; Galveston Bay (Chan­ dler). represented by only 1 or 2 specimens in single Heterotyphlum eurycheilum Olsen, 1952. host individuais. Nine species of immature nema­ Host: Promicrops itaiara; Dry Tortugas (Olsen). todes were found, but only two, both belonging Rhaphidascaris anchoviellae Chandler, 1935a. to the genus Contracaecum, occurred frequently in Hosts: 1m., Anchoa epsetus, striped anchovy, Menidia berrylina peninsulae, silversides; Galveston Bay their particular hosts and then in only moderate (Chandler). Immature ascarids in small numbers numbers. Seamster (1950), in a personal com­ recorded by Linton (1907) from the following hosts lll.unication, reported finding 1 nematode infesta­ at Dry Tortugas: Sphyraena barracuda, Haemulon sciurus, H. flavolineatum, H. macrostomum, Mycter­ tion in 30 fish b{jlonging to 16 genera from Corpus operca bonaci, M. venenosa, Ocyuru8 chrysurus,' Christi BaYi this nematode was not identified. Epinephel,us striatus, and Hepsetia stipes. 357 358 FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE

Rhaphidasearis lutiani Olsen,_ 1952. LINTON, E. Host: Lutianus analis; Dry Tortugas (Olsen). 1907. Preliminary report on animal parasites collected Terranova ginglymostomae Olsen, 1952. at Tortugas, Florida, June 30 to July 18, 1906. Host: Gingylostoma eirratum; Dry Tortu~as (Olsen). Carnegie lnst. Washington Yearbook No.5, 1906, Terranova secundum (Chandler, 1935) Olsen, 1952. pp.112-117. Host: 1m., Triehiurus lepturus, cutlass fish; Galveston 190811.. Preliminary report on animal parasites. Car­ Bay (Chandler). negie Inst. Washington Yearbook No.6, 1907, pp. Terranova triehiuri (Chandler, 1935) Olsen, 1952. 106-123. Host: 1m., Triehiurus lepturus, cutlass fish; Galveston 1908b. Helminth fauna of the Dry Tortugas. I. Bay (Chandler). Cestodes. Carnegie Inst. Washington Pub. 102, pp. 157-190. Family CUCULLANIDAE 1909. Preliminary report on helminths. Carnegie Inst. Washington Yearbook No.7, 1908, pp. 124-127. Diehelyne lastigatus Chandler, 193511.. 1910. Notes on the flesh parasites of marine food fishes. Host: Seiaenops oeellatus, redfish; Galveston Bay Bull. Bur. Fish. 28, 1908, Pt. 2, pp. 1195-1209: (Chandler). 1916. Notes on two cestodes from the spotted sting ray· Diehelyne diploeaeeum Chandler, 1931511.. Jour. Parasit. 3: 34-38, 1 pI. Host: I etalurus lureatus, channel cat, usually found in OLSEN, L. S. fresh water, Galveston Bay (Chandler). 1952. Some nematodes parasitic in marine fishes. Pub. Inst. Marine Sci. 2: 175-215. Family PHILOMETRIDAE PEARSE, A. S., and WHARTON, G. W. Philornetra sp. (Linton, 1907) (=Iehthyonema sp. Linton, 1938. The oyster "leech," Styloehus inimieus Palombi, 1907) . associated with oysters on the coasts of Florida. Hosts: Lntjanus griseus, gray snapper, Strongylura ma­ Ecol. Monogr. 8: 605-655. rina, billfish; Dry Tortugas (Linton). PEREZ, VIGUERAS, I. 1936. Notas sobre Ill. fauna parasitologica de Cuba Incertae Sedis (Parte 1: Vermes). Mem. Soc. Cub. de Hist. Nat. 9 (1): 45-49 (2): 59-66; 10 (2): 53-86. Aganwnema immanis Chandler, 193511. (possibly a Phil­ POTTER, C. C. ome/ra) . 1937. A new cestode from a shark (Hypoprion bre­ Hosts: Fundulus sp. (grandis?) , Cyprinodon variega/us, virostris Poey). Proe. Helminth Soc. Washington killifishes; Galveston Bay (Chandler). 4: 70-86. Agamonema vomi/or Chandler, 193511.. SCHECHTER, VI CTOR. Host: Ictalurus lureatns, channel cat (usually found in 1943. Two flatworms from the oyster-drilling snail, fresh water); Galveston Bay (Chandler). Thais floridana haysae Clench. Jour. Parasit. 29: :362. BIBLIOGRAPHY SHULER, R. H. 11l38. Some cestodes of fish from Tortugas, Florida. CHANDLER, ASA C. Jour. Parasit. 24 (1): 57-63. 1934. A revision of the genus Rhadinorhynchus (Acan­ VAN CU;AVE, H ..J. thocephala) with descriptions of new genera and ]\)47. On the occurrence of the acanthocephalan genUS species. Parasit. 26 (3): 352-:~58, pI. 14. Telosentis in North America. Jour. Parasit. 33: 193511.. Parasites of fishes in Galveston Bay. Proc. 126-133. U. S. Nat. Mus. 83 (21l77): 123-157, pIs. 6-12. --- and LINCICOME, D. H. 11l35b. A new Tetrarhynchid larva frolll Galveston Bay. 11l31l. On a new genus and species of Rhadinorhynchidac Jour. Parasit. 21 (3): 214-215. (Acanthocephala). Parasit. 31: 413-416, pI. 12. 1942. Some cestodes from Florida sharks. Proc. U. S. 1940. A reconsideration of the acanthocephalan familY Nat. Mus. 92 (3135): 25-31. Hhadinorhynchidae. Jour. Parasit. 26 (1): 75-81. DOLLFUS, R. P. --- and MANTER, H. W. 11l42. Etudes critiques sur les TMrarhynques. du 1\)47. A new species of the acanthocephalan gennS MuseulIl de Paris. Arch. Mus. National d'Hist. Nat., Filisoma from the Dry Tortugas, Florida. Jonr. Ser. 6, 19, 466 pp. Parasit. 33: 487-490.