Zoogeography of Digenetic Trematodes from West African Marine Fishes1

Home , Caproidae, Coitocaecum, Cynoglossus canariensis, Ephippion, Haploporidae, Helicometra

192 PROCEEDINGS OF THE HELMINTHOLOGICAL SOCIETY

JACOB H. FISCHTHAL Department of Biological Sciences, State University of New York at Binghamton, Binghamton, New York 13901.

ABSTRACT: Of the 107 species of trematodes found in West African (Mauritania to Gabon) marine fishes, 100 are allocated to 64 genera in 24 families while seven are immature didymozoids. Many of these genera are located in most of the world's seas with the exception of the polar seas; only five are endemic to West Africa. The data for the 41 species known from West Africa and elsewhere, and those morphologically closest to the 55 endemic species, indicate that they are very widely distributed, particularly in the Western and North Atlantic, and Mediterranean. Historical and present- day events concerning physical and biological environmental factors and their effects on actual and potential hosts as well as on life cycle stages of the trematodes have resulted in the geographical distribution reported. The distribution of marine fishes has been emphasized to explain in part the trematode distribution.

Studies on the geographical distribution of (Gulf of Guinea from 5° S to 15° N) and digenetic trematodes of marine fishes in various warm temperate Mauritania have been pre- seas have been presented by Manter (1955, sented by Ekman (1953), Buchanan (1958), 1963, 1967), Szidat (1961), and Lebedev Longhurst (1962), and Ingham (1970). (1969), but West African waters were not included as sufficient data were not available Zoogeographical Distribution until more recently. The digenetic trematodes Of the 107 species of trematodes found in of West African marine fishes (mainly shore West African fishes, 100 are allocated to 64 and shelf inhabitants) have been reported by genera in 24 families while seven are immature Dollfus (1929, 1937a, b, 1946, 1951, 1960), didymozoids of unknown generic status (Ap- Dollfus and Capron (1958), Thomas (1959, pendix I). A list of the host species and the 1960), Nikolaeva (1965), Fischthal and trematodes found in each is given in Appendix Thomas (1968a, b, c, 1969, 1970a, b, 1971, II. The data on the geographical distribution 1972a, b, c), and Fischthal and Williams of the genera (Table 1) indicate that many (1971). are located in most of the world's sea? with According to Ekman (1953) the West Afri- the exception of the Arctic and Antarctic, can tropical marine fauna in general is poorer and that only five (7.8%) are endemic to than any other tropical coastal fauna (Western West Africa (Table 2). The latter figure is Atlantic, Indo-West Pacific, Eastern Pacific) comparable to that for the North Atlantic because of lower water temperatures, the con- (7.5%). The data for the species (Tables 1, siderably shallower layer of tropical water, the 3) indicate that the 41 species elsewhere known sandy sea floor which is unsuitable for coral and those morphologically closest to the 55 reef formation, and the open coast which pro- endemic species are very widely distributed, vides little or no shelter from the surf. He but with much larger numbers occurring in further noted that the region contains only an the Western Atlantic, North Atlantic, and Med- insignificant number of endemic genera, al- iterranean, and mostly in tropical and warm though endemic species are much more nu- temperate waters. Intermediate numbers occur merous and represent a fairly high percentage in the tropical Eastern Pacific, Japanese region, of the total number in several groups. Data and Red Sea. on the oceanography of tropical West Africa Discussion

1 Contribution from the Department of Biological Explanations for the geographical distribu- Sciences, State University of New York at Binghamton, Binghamton, New York 13901. tion of the trematodes noted above relate in

Table 1. Summary of geographical distribution of 58 genera1 and 96 species of digenetic trematodes from West African marine fishes. Under No. spp. the first column of numbers indicates how many of 41 West African species are also known from the locality stated; the second column indicates how many species morphologically most like the 55 endemic species are known from the locality stated.

Localitv No. genera No. spp. Locality No. genera No. spp.

N. Europe 21 9— 3 Tropical E. Pacific 31 12—10 British Isles 8— 2 Mexico 8—10 Belgium, Norway 3 — 0 Panama 6— 3 Baltic Sea 5— 1 Ecuador 4— 1 Eastern N. Atlantic 15 11— 2 Galapagos Isl. 11 4— 7 Europe 10— 2 Central Pacific 31 3 — 4 Morocco 1 — 0 Hawaii 3 — 4 Madeira Isl. 1— 0 Line Isl. 1 — 0 1— 0 Iceland Japan 39 13 — 21 Western N. Atlantic 38 20—11 Japan 9—16 Canada 2 — 1 Korea 0 — 2 U. S. 14 — 9 Yellow Sea 2— 0 Bimini 7— 3 E. China Sea 1 — 3 Bermuda 7— 2 Taiwan 1 — 2 2— 0 Bahama East Indies 31 5— 8 Azores 3 2— 0 S. China Sea 4— 0 Eastern S. Atlantic 18 8— 2 Philippines 3— 7 South-West Africa 8— 2 Mariana Isl. 1— 0 Western S. Atlantic 17 6— 1 Celebes 2— 3 Brazil 3— 1 Borneo 0— 1 Argentina 3— 0 Indian Ocean 26 9— 7 Mediterranean— Black Sea 31 17 — 6 India 4 — 6 Mediterranean 17— 6 Ceylon 0— 1 Black Sea 7— 3 Madagascar 1— 1 Gulf of Aden 5 — 0 Caribbean 35 19 — 10 Red Sea 24 12 — 4 Gulf of Mexico 42 20—17 New Zealand— Australia 30 8 — 8 Arctic 9 1 — 1 Australia J g Barents Sea .1 — 1 Tasmania 1— 0 N. Pacific 25 6— 8 New Zealand 5— 0 Bering Sea 2— 0 New Caledonia 2 — 2 Okhotsk Sea 3— 1 Fiji Isl. 0— 3 Alaska 1— 0 Antarctic 4 0 — 0 Canada 1— 1 U. S. 5 — 6

1 Endemic genera excluded are: Diplomonorcheides Thomas, 1959; Elopsium Fischthal and Thomas, 1972; Neochoano- dera Fischthal and Thomas, 1970; Ncolcpocrcadium Thomas, 1960; Pedunculotrema Fischthal and Thomas, 1970. Pal- aeorchis Szidat, 1943, has only freshwater fishes from Japan and Europe as hosts. great part to historical and present-day events the Tethys fauna, and was essentially tropical concerning physical and biological environ- and more homogeneous than the present-day mental factors (geological occurrences, cli- tropical-subtropical shelf fauna. This probably mates, temperatures, currents, salinity, bottom applied to the trematodes as well. Valentine conditions, shore configurations, animal and (1967) noted that fluctuating climates act plant populations, food chain, etc.) and their as a sort of diversity pump, enhancing species effects on actual and potential hosts as well diversity during climatic deterioration and re- as on stages in the life cycles of the trematodes. taining some fraction of the new lineages Migrations have been accomplished by adult during climatic improvement. He indicated fishes and mollusks and their pelagic larvae; that the Tethyan fauna was enriched by this mollusks probably have moved long distances process acting within the Tethys and also by via currents while attached to floating vegeta- recruitment of lineages from the extra-Tethyan tion or debris. provincial margins, especially during improving Ekmart (1953) noted that an immense climatic phases. Trematodes probably were Tethys Sea, already existing in the Lower similarly affected. Cambrian and continuing into the late Tertiary, During the course of geological time various included the Eastern Pacific, tropical Atlantic, barriers arose to divide the Tethys Sea into Mediterranean, and Indo-West Pacific. The more or less separate seas. Other seas were fauna of this sea was part of one major unit, also affected by the appearance or removal of

Table 2. Number of genera and per cent endemic Table 3. Summary of geographical distribution in of digenetic trematodes of marine fishes from var- littoral provinces (Hedgpeth, 1957) of digenetic ious geographical regions.1 trematodes of West African marine fishes.

Locality No. genera % genera endemic No. of spp. No. of 41 spp. morphologically N. Atlantic 58 7.5 elsewhere closest to 55 Tropical W. Atlantic 145 25 Province known endemic spp. Mediterranean— Atlantic 89 23 Red Sea-Gulf of Aden 38 25 Arctic 3 2 India 55; 24 Boreal 8 :.2 East Indies 84 31 Antiboreal 4 i Japan 168 32 Boreal- Antiboreal 6 i New Zealand— Australia 76 17 Warm temperate 33 27 Hawaii 169 40 Tropic 32 36 Tropical E. Pacific 78 11 W. Africa 64 7.8

1 Figures for Hawaii based on data by Yamaguti (1970) others, except W. Africa, from Lebedev (1969). in part. Much of the displaced Mediterranean fauna (with their trematodes) migrated to West Africa which still supports some of them. On barriers. Ruggieri (1967) noted that the sep- the American side of the Atlantic (Florida, aration of the Mediterranean from the Indian Georgia, neighboring states) colder climates Ocean occurred during the Miocene, but Tor- of the Miocene also affected the tropical fauna tonese (1964) indicated the Pliocene. The but not as catastrophically as in the North and former author further noted that towards the Eastern Atlantic and the Mediterranean. The end of the Miocene the two straits connecting major difference was that during the Pliocene the Mediterranean with the Atlantic disap- and Quaternary the tropical climate and fauna peared and the Mediterranean was transformed reappeared on the American side, whereas only into a series of lagoons which either dried an insignificant part reappeared in tropical up or became gradually desalinified, thus deci- West Africa and especially the Mediterranean. mating its marine fauna. During the Pliocene Tortonese (1964) noted that the largest group the Gibraltar straits formed and the Atlantic comprising the present-day Mediterranean fish flowed anew into the Mediterranean, reestab- fauna (about 550 species) is Atlantic (boreal, lishing truly marine conditions. Those elements West African, or amphi-Atlantic); other spe- of the Miocene fauna that had persisted in the cies are endemic (about 70 or 13%), cosmo- Atlantic outside Gibraltar were reintroduced politan, or Indo-Pacific. In great part it is (with their trematodes) to the Mediterranean because of the historical events related above together with new species previously absent that significantly low percentages of endemic from the area; some of the latter may have genera generally occur in the fauna of West been from West Africa. Ruggieri indicated Africa, the Mediterranean, and the North At- that the Atlantic facing the Gibraltar straits lantic, and also why many identical as well probably was the true asylum for the Indo- as closely related species of trematodes and Pacific relicts (with their trematodes) during other groups are found in these regions. the Miocene salinity crisis in the Mediterra- Ben-Tuvia (1966) indicated that at least nean; some of these relicts along with other 24 of about 800 species of fishes have migrated Mediterranean fishes may have migrated south- from the Red Sea to the Mediterranean since ward to West Africa during this time. the opening of the Suez Canal in 1869, some Ekman (1953), Briggs (1966, 1970), and as far west as Lampedusa near Sicily; no re- Ruggieri (1967) reported that during the late liable records exist of a reverse migration al- Tertiary and early Quaternary there was a though a few are found in the Suez Canal. considerable change in the tropical nature of Some 18 of the Red Sea species are now among the North Atlantic and Mediterranean due the more common forms along the Mediter- primarily to the advent of colder climates. ranean coast of Israel and nine are sufficiently The northern fauna (with their trematodes) abundant to be commercially exploited. Some migrated southward when possible and si- of the trematodes from Red Sea fishes possibly multaneously the tropical fauna disappeared have become established in closely related

Mediterranean fishes or in other fishes which that probably occur on both sides of Central feed on the same intermediate hosts. Wide- America, only about 12 are identical (ex- ranging Mediterranean fishes then may have cluding 13 circumtropical and a few euryhaline moved into the Atlantic and to West Africa. species); this barrier is about 99% effective. Briggs (1970) reported that during the Rosenblatt reported that the shore fish fauna Oligocene some boreal species (including of both sides of the Americas show great fishes and their trematodes) north of the similarity at the family, subfamily, and generic Bering Land Bridge moved out of the Arctic levels, but not at the species level; however, Basin to the North Atlantic. During the there are a large number of pairs of sibling Miocene and again in the Pliocene the Bering species. He further noted that a fair number Land Bridge disappeared, opening a seaway of genera in the Western Atlantic are found between the North Pacific and Arctic. The also in the Indo-West Pacific. About 10 genera movement of species (including trematodes) (2%) of the total for the Western Atlantic was (and still is) predominantly northward are common to and limited to the tropical from the Pacific to the Arctic Ocean to the Atlantic. The historical events relating to the North Atlantic. These events explain in great continuous Tethys Sea and subsequent forma- part Manter's (1963) observation that of 54 tion of the South and Central American land species of digenetic trematodes of marine fishes bridges during the Pliocene to separate the from the British Columbian region of Canada, Atlantic and Pacific account in great part for 21 (40%) also occur in the North Atlantic. the present-day distribution of fishes in this Briggs also noted that present-day species region. This also explains in part Manter's in the North Atlantic have broad latitudinal (1963) observation that of 226 species of ranges. Some, with their trematodes, occur digenetic trematodes from Caribbean fishes, in West Africa. about 18% also occur in the tropical Eastern Briggs (1967) observed that the Mid- Pacific. It also explains in part why many Atlantic Barrier is a broad, deepwater expanse trematodes from West African fishes occur in of ocean separating the West African tropics the Eastern Pacific and Indo-West Pacific from those of the Western Atlantic. About since, as noted above, about 31% of the 380 species of tropical shore fishes are found West African shore fish species also occur in in the Eastern Atlantic and 900 species in the the tropical Western Atlantic and a fair number Western Atlantic, giving a total of 1,280. of genera from the latter locality also are About 118 (9.2%) of this total are trans- found in the Indo-Pacific. Atlantic species, making the barrier about 91% The Old World Land Barrier has been 98% effective. The migration has been and still effective in separating the shore fishes of is from west to east. The 118 trans-Atlantic the Indo-West Pacific from the Atlantic. Briggs species comprise 31% of the tropical West (1967) indicated that the fish fauna of tropical African shore fish fauna; many genera are Southeast Africa in the area between Mozam- also in common. Briggs noted further that bique and Algoa Bay numbers about 1,000 West African invertebrate groups also show species and in tropical West Africa about 380, appreciable percentages of the species as being a total of 1,380. Migration has been from trans-Atlantic (6% of the prosobranch mol- east to west around the Cape of Good Hope, lusks). These facts serve greatly to explain involving only 31 species (8 to both sides why such a large number of trematodes species of the Atlantic; 4 confined to West Africa; from West African fishes are identical with 3 to St. Helena Island; 16 circumtropical); or closely related to those in the tropical however, many genera are in common. This Western Atlantic and its neighboring warm distribution of fishes has to some extent af- temperate zones. fected the distribution of their trematodes. Briggs (1961, 1967) and Bosenblatt (1967) Briggs (1961, 1964, 1967) noted that the have discussed the New World Land Barrier. East Pacific Barrier is a vast stretch of about The former author noted that of the total 3,000 miles of deepwater lying between Pol- number of shore fish species (about 1,000) ynesia and America. He reported 62 trans-

Pacific species of shore fishes (13 of which terranean. Successful reciprocal migrations are, are circumtropical) out of a total of about at least, very rare and may be completely 1,037 species for both regions (about 387 lacking. Furthermore, judging from the general species assumed to be in the Line Islands indications of relationship among the four lying at easternmost Polynesia in the Equa- great tropical faunas, this process has been torial Countercurrent; about 650 species going on for many millions of years. Some between southern Mexico and Peru); many of the dominant species are so successful that common genera are in the two regions. This they have been able to establish and maintain barrier is 94% effective. The successful mi- circumtropical distributions (indicating more grations were recent or current invasions, and or less regular migrations across the East were almost exclusively from west to east Pacific, Old World land, and mid-Atlantic via the countercurrent. Briggs (1967) related barriers). The Western Atlantic tropics may that 18 species of mollusks are trans-Pacific. be considered a secondary center of evolu- Some of the trematodes probably moved across tionary radiation. Many species produced in this barrier with their vertebrate and molluscan this area have proved capable of migrating hosts. eastward to colonize the Eastern Atlantic Briggs (1960) reported that at least 107 region. However, species originating in the species of marine fishes form homogeneous Eastern Atlantic are apparently incapable of worldwide circumtropical species populations. successfully invading the western side. Again, Additional species are worldwide but form the advantage seems to lie with the arsa that subspecific populations in parts of their range. possesses the richer fauna and higher level of Other species are nearly circumtropical but competition." The distributions exhibited by have not been found in the Eastern Pacific. the digenetic trematodes of marine fishes from A majority of the 107 fish species can be con- West Africa and from other regions are in sidered to be typical of the open seas (90 agreement with Briggs' observations and con- pelagic, 14 littoral, 3 benthic). In West Africa clusions. five of these 107 species were found infected with at least one species of trematode which Literature Cited occurs in the same host species elsewhere. Ben-Tuvia, A. 1966. Red Sea fishes recently The circumtropical (or nearly so) distribu- found in the Mediterranean. Copeia (2): 254- tion of many genera and species of fishes has 275. aided considerably the wide distribution of Briggs, J. C. 1960. Fishes of worldwide (cir- their trematodes; additionally, new and more cumtropical) distribution. Copeia (3): 171- localized hosts have been acquired by some. 180. Briggs (1967) summarized the information . 1961. The East Pacific Barrier and the concerning tropical marine fishes and inverte- distribution of marine shore fishes. Evolu- brates as follows: "There is no doubt that tion 15: 545-554. ———-. 1964. Additional transpacific shore the Indo-West Pacific Region has served as fishes. Copeia (4): 706-708. the evolutionary and distributional center for . 1966. Oceanic islands, endemism, and the entire marine tropics. Its fauna is almost marine paleotemperatures. Syst. Zool. 15: unbelievably rich with, for example, more 153-163. than 3,000 species of shore fishes. It seems . 1967. Relationship of the tropical shelf clear that the unusually stable ecosystems and regions. Stud. Trop. Oceanogr. Miami 5: high level of competition provide the proper 569-578. 1970. A faunal history of the North environment for the evolution of dominant Atlantic Ocean. Syst. Zool. 19: 19-34. species that can successfully invade the other Buchanan, J. R. 1958. The bottom fauna regions. From the Indo-West Pacific, dominant communities across the continental shelf off species migrate eastward across the open Accra, Ghana (Gold Coast). Proc. Zool. Soc. ocean to America, westward around the Cape London 130: 1-56. Dollfus, R. P. 1929. Helmintha I. Trematoda of Good Hope into the Atlantic, and north- et Acanthocephala. In T. Monod (ed.), Con- ward through the Suez Canal into the Medi- tribution a 1'etude de la faune du Cameroun.

2 partie. Faune Colon. Frang. (Gruvel) 3: 73- Bull. Inst. Fond. Afr. Noire, Ser. A, Sci. Nat. 114. 34. In press. . 1937a. Les trematodes Digenea des , and . 1972b. Digenetic trem- selaciens (plagiostomes). Catalogue par hotes. atodes of marine fishes from Senegal. Bull. Distribution geographique. Ann. Parasit. 15: Inst. Fond. Afr. Noire, Ser. A, Sci. Nat. 34. 57-73, 164-176, 259-281, 557. In press. —. 1937b. Parasitologia Mauritanica. Hel- —, and . 1972c. Digenetic trem- mintha (III). Trematodes de selaciens et de atodes of fish from the Volta River drainage cheloniens. Bull. Comite Etudes Hist, et system in Ghana prior to the construction of Sclent. Afr. Occid. Franc. 19: 397-519. 1936. the Volta dam at Akosombo in May 1964. J. . 1946. Sur trois especes de distomes, Helm. 46: 91-105. dont une a 17 ventouses (Enentemm (Jean- -, and M. O. Williams. 1971. Some di- cadenatia) brumpti n. sp.) parasites du genetic trematodes of marine fishes from poisson marin Kyphosus sectatrix (L.). Ann. Sierra Leone. J. Helm. 45: 41-50. Parasit. 21: 119-128. Hedgpeth, J. W. 1957. Marine biogeography. . 1951. Metacercaire de trematode (Gas- Chapter 13: Treatise on marine ecology and terostomata) enkystee chez des Sparisoma, paleoecology. Geol. Soc. Amer. Mem. 67 1: Rupiscartes et Blennius de Goree (Senegal). 359-382. Bull. Inst. Frang. Afr. Noire, Sen A, Sci. Ingham, M. C. 1970. Coastal upwelling in the Nat. 13: 762-770. northwestern Gulf of Guinea. Bull. Mar. Sci. . 1960. Sur trois helminthes (distomien, Miami 20: 1-34. tetrarhynque, palaeacanthocephale) d'un Me- Lebedev, B. I. 1969. [Basic regularities in dis- lanoglaea ventralis K. Barnard 1941, poisson tribution of monogeneans and trematodes of teleosteen. Bull. Inst. Frang. Afr. Noire, Ser. marine fishes in the world oceans.] Zool. Zh. A, Sci. Nat. 22: 385-394. 48:41-50. (Bussian text.) -, and A. Capron. 1958. Uii Aephnidio- Longhurst, A. R. 1962. A review of the ocean- genes (Trematoda Digenea) chez un Labrax ography of the Gulf of Guinea. Bull. Inst. Frang. Afr. Noire, Ser. A, Sci. Nat. 24: 633- (Serranidae) du Senegal. Bull. Inst. Frang. 663. Afr. Noire, Ser. A, Sci. Nat. 20: 311-319. Maiiter, H. W. 1955. The zoogeography of trem- Ekman, S. 1953. Zoogeography of the sea. 417 atodes of marine fishes. Exp. Parasit. 4: 62— p. Sidgwick and Jackson, London. 86. Fischthal, J. H., and J. D. Thomas. 1968a. . 1963. The zoogeographical affinities of Digenetic trematodes of some freshwater and trematodes of South American freshwater marine fishes from Ghana. Proc. Helm. Soc. fishes. Syst. Zool. 12: 45-70. Wash. 35: 126-140. -. 1967. Some aspects of the geograph- , and . 1968b. Digenetic trem- ical distribution of parasites. J- Parasit. 53: atodes of marine fishes from Ghana: Families 1-9. Acanthocolpidae, Bucephalidae, Didymozo- Nikolaeva, V. M. 1965. [On the develop- idae. Proc. Helm. Soc. Wash. 35: 237-247. mental cycle of trematodes belonging to the , and . 1968c. Siphodera ghanen- family Didymozoidae (Monticelli, 1888) sis sp. n. (Cryptogonimidae), a digenetic Poche, 1907.] Zool. Zh. 44: 1317-1327. trematode from an estuarine fish from (Russian text.) Ghana. J. Parasit. 54: 765-766. Rosenblatt, R. H. 1967. The zoogeographic re- , and . 1969. Digenetic trem- lationships of the marine shore fishes of trop- atodes of marine fishes from Ghana: Family ical America. Stud. Trop. Oceanogr. Miami Monorchiidae. J. Helm. 43: 11-30. 5: 579-592. , and . 1970a. Digenetic trem- Ruggieri, G. 1967. The Miocene and later atodes of marine fishes from Ghana: Family evolution of the Mediterranean Sea. In C. G. Opecoelidae. Proc. Helm. Soc. Wash. 37: Adams and D. V. Ager (eds.), Aspects of 129-141. Tethyan Biogeography. Syst. Assoc. Publ. 7: , and . 1970b. Digenetic trem- 283-290. atodes of marine fishes from Ghana: Family Szidat, L. 1961. Versuch einer Zoogeographie Lepocreadiidae. J. Helm. 44: 365-386. des Stid-Atlantik mit Hilfe von Leitparasiten , and . 1971. Some hemiurid trem- der Meeresfische. Parasit. Schriftenreihe 13: atodes of marine fishes from Ghana. Proc. 1-98. Helm. Soc. Wash. 38: 181-189. Thomas, J. D. 1959. Trematodes of Ghanaian , and . 1972a. Additional hemiurid sub-littoral fishes. I. The family Monorchi- and other trematodes of fishes from Ghana. idae. J. Parasit. 45: 95-113.

. 1960. Trematodes of Ghanaian sub- Japan; Galapagos. P. caudovatus Manter, 1940 littoral fishes. II. The family Lepocreadiidae —also Suez. Rhipidocotyle ghanensis Fischthal Nicoll, 1935, sensu Cable and Hunninen, and Thomas, 1968. R. senegalensis Fischthal 1942. Lib. Horn. Dr. Edu. Caballero y C., p. 321-328. and Thomas, 1972. Cryptogonimidae.1 Para- Tortonese, E. 1964. The main biogeographical cryptogonimns ghanensis Fischthal and Thomas, features and problems of the Mediterranean 1968. Didymozoidae. Allonematobothrium fish fauna. Copeia (1): 98-107. ghanense Fischthal and Thomas, 1968. Di- Valentine, J. W. 1967. The influence of cli- dymozoidae (Monilicaecum) larvae I Niko- matic fluctuations on species diversity within laeva, 1965—also South-West Africa. Di- the Tethyan provincial system. In C. G. Adams and D. V. Ager (eds.), Aspects of dymozoidae gen. sp. larvae V Nikolaeva, 1962 Tethyan Biogeography. Syst. Assoc. Publ. 7: —also Red Sea. Immature Didymozoid B 153-166. Fischthal and Kuntz, 1964—also Philippines. Yamaguti, S. 1970. Digenetic trematodes of Immature Didymozoid D Fischthal and Hawaiian fishes. Keigaku Publ., Tokyo, 436 p. Thomas, 1968. Immature Didymozoid E Fischthal and Thomas, 1968. Immature Di- dymozoid F Fischthal and Thomas, 1968. Appendix I Immature Didymozoid G Fischthal and Thomas, List of digenetic trematodes from West 1968. Enenteridae. Cadenatellabrumpti (Doll- African marine fishes, and other fus, 1946) Nahhas and Cable, 1964—also localities from which known Bimini. C. cadenati (Dollfus, 1946) Nagaty, 1948. Enenterum pimelepteri Nagaty, 1942— Acanthocolpidae. Stephanostomum afri- also Red Sea. Fellodistomatidae. Elopsium canum Fischthal and Williams, 1971. S. ghanense Fischthal and Thomas, 1972. Marke- bicoronatum (Stossich, 1883) Manter, 1940— vitschiella sp. Fischthal and Thomas, 1968. also Mediterranean, Tyrrhenian, Adriatic, Black, Monascus typicus (Odhner, 1911) Looss, 1912 Yellow, E. China Seas. S. casum (Linton, —also Mediterranean, Adriatic, Black, Red 1910) McFarlane, 1934—also Bermuda; Ba- Seas; Gulf of Aden; India. Steringotrema hama; Caribbean; Gulf of Mexico; U. S. At- divergent (Rudolphi, 1809) Odhner, 1911— lantic; Mexican, Canadian Pacific; Galapagos; also Mediterranean, Tyrrhenian, Baltic: Seas; Philippines; New Caledonia; Red Sea. S. British Isles. Gorgoderidae. Nagmia africana coryphaenae Manter, 1947—also Bimini; Ca- Fischthal and Thomas, 1972. IV. senegalensis ribbean; Gulf of Mexico. S. ghanense Fischthal Fischthal and Thomas, 1972. Halipegidae. and Thomas, 1968. S. megacephalum Manter, Gonocercella trachinoti (MacCallum, 1913) 1940—also Caribbean; Gulf of Mexico; Mexi- Yamaguti, 1954—also U. S. Atlantic; Gulf can, Panama, Ecuador Pacific; Red Sea. S. of Mexico. Haploporidae. Megasolena hys- sierraleonense Fischthal and Williams, 1971. terospina (Manter, 1931) Overstreet, 1969— S. trachinoti Fischthal and Thomas, 1968. also U. S. Atlantic; Caribbean; Gulf of Mexico. Azygiidae. Otodistomum veliporum (Creplin, Hemiuridae. Adinosoma robusta (Manter, 1837) Stafford, 1904—also Mediterranean, 1934) Manter, 1947—also Gulf of Mexico. Black Seas; European, Moroccan, U. S., Cana- Aponurus lagunculus Looss, 1907—also Med- dian Atlantic; Iceland; British Isles; North, iterranean, Tyrrhenian, Adriatic, Black Seas; Baltic Seas; Argentina; New Zealand; Sea of South-West Africa; Gulf of Mexico; U. S., Japan; Alaska; U. S., Mexican Pacific. Bu- Mexican Pacific; Celebes; S. China Sea; Gulf cephalidae. Bucephaloides ghanensis Fischthal of Aden; Red Sea. Dinurus barbatus (Cohn, and Thomas, 1968. B. gracilescens (Rudolphi, 1819) Hopkins, 1954—also Mediterranean, 1903) Looss, 1907—also European Atlantic; Adriatic Seas; British Isles; South-West Africa; Caribbean; Gulf of Mexico; Mexican, Panama Barents Sea; Far Eastern Seas of Siberia. B. Pacific. D. breviductus Looss, 1907—also ovatus (Linton, 1900) Hopkins, 1954—also 1 Siphodera ghanensis described by Fischthal and Thomas U. S. Atlantic; Panama Pacific. Prosorhynchus (1968c) is from freshwater fishes from Ghana and Gabon, aculeatus Odhner, 1905, metacercaria—also but the two other species in the genus, S. vmaladwardsn (Linton, 1901) Linton, 1910, and S. cirrhiti Yamaguti, Mediterranean, North, Baltic Seas; British Isles; 1970, are from marine fishes.

European, U. S. Atlantic; Caribbean; Gulf Dollfus and Capron, 1958 also India. Diplo- of Mexico; Argentina; Red Sea. D. tornatus proctodaeum ghanense (Fischthal and Thomas, (Rudolphi, 1819) Looss, 1907—also European, 1970) Fischthal and Thomas, 1972. D. hau- U. S. Atlantic; Azores; Caribbean; Gulf of strum (MacCallum, 1918) LaRue, 1926—also Mexico; Gulf of Aden; Red Sea. Ectenurus Bimini; U. S. Atlantic; Caribbean; Hawaii. lepidus Looss, 1907—also Mediterranean, Tyr- Holorchis legendrei Dollfus, 1946—also Med- rhenian, Adriatic, Black Seas; South-West Af- iterranean; European Atlantic. Homalometron rica; Caribbean; Brazil; Hawaii; New Zealand; senegalense Fischthal and Thomas, 1972. Lep- Gulf of Aden. E. virgulus Linton, 1910—also idapedon ghanense Fischthal and Thomas, Bimini; Bermuda; Bahama; U. S. Atlantic; 1970. Lepocreadioides cynoglossi Fischthal Caribbean; Gulf of Mexico; Argentina; Gulf and Thomas, 1970. Lepocreadium ghanense of Aden; Red Sea. Lecithaster africanus Fischthal and Thomas, 1970. Neochoanodera Fischthal and Thomas, 1971. L. ghanensis ghanensis Fischthal and Thomas, 1970. Ne- Fischthal and Thomas, 1971. Lecithochirium olepocreadium caballeroi Thomas, 1960. Opech- ghanense Fischthal and Thomas, 1972. L. ona bacillaris (Molin, 1859) Looss, 1907—also microstomum Chandler, 1935—also U. S. At- Adriatic, Black, Baltic, Red Seas; European lantic; Caribbean; Gulf of Mexico; Brazil; Atlantic; South-West Africa; British Isles India; Mexican, Panama Pacific; Galapagos; Taiwan; New Zealand-Australia region. O. ghanensis Madagascar. Lecithocladium augustiovum Fischthal and Thomas, 1970. O. pseudobacil- Yamaguti, 1953—also Philippines; Celebes; laris Fischthal and Thomas, 1970. Pseudo- Red Sea. L. excisum (Rudolphi, 1819) Liihe, creadium ghanense Fischthal and Thomas, 1901—also Mediterranean, Adriatic, Black, 1970. Monodhelminthidae. Monodhelmis tor- Irish, North, Baltic Seas; European, U. S. pedinis Dollfus, 1937. Monorchiidae. Diplo- Atlantic; South-West Africa; Gulf of Mexico; monorcheides magnacetabulum Thomas, 1959. Japan; S. China Sea; New Zealand. L. mecode- Lasiotocus accraensis Fischthal and Thomas, rum Fischthal and Thomas, 1971. L. unibul- 1969. L. attenuatus Fischthal and Thomas, bolabrum Fiscllthal and Thomas, 1971. Para- 1969. L. chaetodipteri Thomas, 1959. L. hemiurus merus (Linton, 1910) Woolcock, cynoglossi Thomas, 1959. L. ghanensis 1935—also South-West Africa; U. S. Atlantic; Fischthal and Thomas, 1969. L. synapturae Bimini; Bermuda; Caribbean; Gulf of Mexico; Fischthal and Thomas, 1969. Palaeorchis Brazil; U. S., Ecuador Pacific; Bering, Okhotsk, senegalensis Fischthal and Thomas, 1972. S. China Seas; Japan. Prosorchis ghanensis Parahurleytrema trachinoti (Thomas, 1959) Fischthal and Thomas, 1972. Sterrhurus gha- Nahhas and Powell, 1965. Proctotrema am- nensis Fischthal and Thomas, 1972. S. fusi- phitmncatum Fischthal and Thomas, 1969. formis (Liihe, 1901) Looss, 1907—also Med- Proctotrematoides ophichthi Fischthal and iterranean, Adriatic Seas; British Isles; Azores; Thomas, 1969. Opecoelidae. Coitocaecum European, U. S. Atlantic; Bermuda; Caribbean; cadenati Dollfus, 1960. Helicometra fasciata Gulf of Mexico; Ecuador Pacific; Japan. S. (Rudolphi, 1819) Odhner, 1902—also Medi- musculus Looss, 1907—also Mediterranean, terranean, Black Seas; N. Atlantic; South-West Adriatic, Black Seas; Bimini; Bermuda; U. S. Africa; Caribbean; Mexican Pacific; Japan; Atlantic; Caribbean; Gulf of Mexico; Japan. New Caledonia; Tasmania; Red Sea. Pedun- Tubulovesicula lindbergi (Layman, 1930) cidotrema capecoastense Fischthal and Thomas, Yamaguti, 1934—also Madeira Isl.; Canadian, 1970. P. ghanense Fischthal and Thomas, 1970. U. S., Panama Pacific; Bering, Okhotsk, S. Plagioporus gerridis Fischthal and Thomas, China, Red Seas; Japan. Hirudinellidae. 1970. Podocotyle temensis Fiscllthal and Hirudinella ventricosa (Pallas, 1774) Baird, Thomas, 1970. Podocotyloides chloroscombri 1853—also Bimini; Bermuda; Caribbean; Gulf Fischthal and Thomas, 1970. Poracanthium of Mexico; Mexican, Panama, Ecuador Pacific; ghanense Fischthal and Thomas, 1970. Pseudo- Galapagos; Hawaii; Line Isl.; Mariana Isl. pecoelus ghanensis Fischthal and Thomas, 1970. Lepocreadiidae. Aephnidiogenes africanus P. tortugae von Wicklen, 1946—also Caribbean; Fischthal and Thomas, 1972. A. senegalensis Gulf of Mexico. P. vulgaris (Manter, 1934)

Copyright © 2011, The Helminthological Society of Washington 200 PROCEEDINGS OF THE HELMINTHOLOGICAL SOCIETY von Wicklen, 1946—also Gulf of Mexico; New Batrachoides liberiensis (Steind.), Zealand. Opistholebetidae. Pycnadena afri- Batrachoididae cana Fischthal and Williams, 1971. Pyc- Sterrhurus ghanensis (G) nadenoides ghanensis Fischthal and Thomas, Blennius sanguinolentus Pallas, Blenniidae 1968. P. senegalensis Fischthal and Thomas, Prosorhynchus aculeatus, metacercana (S) 1972. Pelorohelminthidae. Pelorohelmins gha- Brachydeuterus auritus (Cuv. and Val.), nensis Fischthal and Thomas, 1968. Pleor- Pomadasyidae chiidae. Pleorchis ghanensis Fischthal and Didymozoidae (Monilicaecum) larvae I (G) Thomas, 1968. Proctoecidae. Mesolecitha Immature Didymozoid B (G) ghanensis Fischthal and Thomas, 1968. Pty- Immature Didymozoid E (G) chogonimidae. Ptychogonimus megastomus Zoogonus mirus (G) (Rudolphi, 1819) Liihe, 1900—also Medi- Caesiomorus glaucus (L.), Carangidae terranean, Tyrrhenian, Adriatic Seas; British Lecithocladium excisum (G) Isles; European Atlantic; North Sea; Bermuda; Opechona ghanensis (G) Caribbean; Japan; New Zealand. Sclerodisto- Opechona pseudobacillaris (G) matidae. Sclerodistomum italicum (Stossich, Capros aper (L.), Caproidae 1893) Looss, 1912—also Adriatic Sea. Syn- Pycnadenoides senegalensis (S) coeliidae. Syncoelium katuwo Yamaguti, 1936 Caranx africanus Steind., Carangidae —also U. S. Pacific; Japan. Zoogonidae. Di- Ectenurus virgulus (G) phtherostomum anisotremi Nahhas and Cable, Immature Didymozoid E (G) 1964—also Caribbean; Gulf of Mexico. Zo- Caranx crysos (Mitchill) ogonus mirus Looss, 1901—also Adriatic Sea; Ectenurus virgulus (G) Gulf of Marseille. Caranx hippos (L.) Parahemiurus merus (G) Appendix II Poracanthium ghanense (G) Stephanostomum megacephalum (G) Alphabetical list of host species with Cephalacanthus volitans (L.), Dactylopteridae trematodes found in each1 Lecithocladium unibulbolabrum (G) (S) Acanthocybium solandri (Cuv. and Val.), Sterrhurus ghanensis (G) Scombridae Sterrhurus musculus (G) Hirudinella ventricosa (G) Chaetodipterus lippei Steind., Ephippidae Acanthurus monroviae (Steind.), Acanthuridae Lasiotocus chaetodipteri (G) Mesolecitha ghanensis (G) Megasolena hysterospina (SL) Prosorchis ghanensis (G) Neochoanodera ghanensis (G) Alutera punctata Cuv., Monacanthidae Chloroscombrus chrysurus (L.), Carangidae Diploproctodaeum haustrum (S) Ectenurus lepidus (G) Antennarius commersonii (Lac.), Antennariidae Monascus typicus (G) Immature Didymozoid E (S) Podocotyloides chloroscombri (G) Rhipidocotyle senegalensis (S) Clupisudis niloticus (Ehrenberg),2 Anus heudeloti Cuv. and Val., Ariidae Osteoglossidae Monodhelmis torpedinis (G) Sterrhurus musculus (G) Arius latiscutatus Gunther Corvina nigra Val., Sciaenidae Pelorohelmins ghanensis (G) Stephanostomum bicoronatum (S) Arnoglossus imperialis (Raf.), Bothidae Coryphaena hippurus L., Coryphaenidae Lecithocladium excisum (S) Dinurus barbatus (G) Bathygobius separator (Cuv. and Val.), Dinurus breviductus (G) (S) Gobiidae Dinurus tornatus (G) Helicometra fasciata (G) Stephanostomum coryphaenae (G) Cynoglossus canariensis Steind., Cynoglossidae 1 Trematcxles indicated by (C) are from Cameroon, (G) Ghana, (M) Mauritania, (S) Senegal, and (SL) Sierra Leone. - Freshwater fish.

Diplomonorcheides magnacetabulum (G) Fodiator acutus (Val.), Exocoetidae Cynoglossus goreensis Steind. Neolepocreadium caballeroi (G) Diplomonorcheides magnacetabulum (G) Galeoides decadactylus (Bloch), Polynemidae Immature Didymozoid E (G) Ectenurus lepidus (G) Lasiotocus cynoglossi (G) Lecithaster africanus (G) Lepocreadioides ghanensis (G) Lecithochirium ghanense (G) Parahemiurus merus (G) Lecithocladium excisum (G) Cynoglossus senegalensis (Kaup) Lecithocladium mecoderum (G) Diplomonorcheides magnacetabulum (G) Poracanthium ghanense (G) (SL) Immature Didymozoid E (G) Stephanostomum sierraleonense (SL) Lasiotocus cynoglossi (G) (SL) Sterrhurus musculus (G) Lepocreadioides ghanensis (SL) Galeorhinus mustelus (L.), Carcharhinidae Cynoscion macrognathus (Bleeker), Sciaenidae Ptychogonimus megastornus (M) Immature Didymozoid E (G) Gephyroberyx darwini (Johnson), Pleorchis ghanensis (G) Trachichthyidae Pseudocreadium ghanense (G) Adinosoma robusta (S) Pseudopecoelus ghanensis (G) Genes melanopterus Bleeker, Liognathidae Cypselurus heterurus (Ra£.), Exocoetidae Pedunculotrema ghanense (G) Immature Didymozoid F (G) Genes nigri Giinther Cypselurus lutkeni Jordan and Evermann Plagioporus gerridis (G) Lecithaster ghanensis (G) Ghjphisodon saxatilis (L.), Pomacentridae Decapterus rhonchus (Geoff.), Carangidae Helicometra fasciata (G) Ectenurus lepidus (G) Gymnothorax vicinus (Castelnau), Muraenidae Monascus typicus (G) Sterrhurus fusiformis (G) Dentex canariensis Steind., Sparidae Hydrocyon brevis Giinther,2 Characidae Sclerodistomum italicum (S) Tubulovesicula lindbergi (G) Diagramma mediterraneum Guichenot, Hyporhamphus calabaricus (Giinther), Pomadasyidae Hemirhamphidae Holorchis legendrei (S) Lecithaster ghanensis (G) Drepane punctata (L.), Drepanidae Ilisha melanota Derscheid, Clupeidae Pseudocreadium ghanense (G) Helicometra fasciata (G) Pycnadenoides ghanensis (G) Lecithocladium excisum (G) Elops lacerta Cuv. and Val., Elopidae Kyphosus sectatrix (L.), Kyphosidae Elopsium ghanense (G) Cadenatella brumpti (S) Engraulis encrasicholus (L.), Engraulidae Cadenatella cadenati (S) Parahemiurus merus (G) Enenterum pimelepteri (S) Ephippion guttifer (Bennett), Tetraodontidae Labrax punctatus (Bloch), Serranidae Diploproctodaeum ghanense (G) Aephnidiogenes senegalensis (S) Labrisomus nuchipinnis (Quoy and Gaimard), Epinephelus aeneus (Geoff.), Serranidae Clinidae Allonematobothrium ghanense (G) Helicometra fasciata (G) Lepidapedon ghanense (G) Sterrhurus musculus (G) Lagocephalus laevigatus L., Tetraodontidae Diploproctodaeum ghanense (G) Epinephelus goreensis (Cuv. and Val.) Immature Didymozoid D (G) Podocotyle temensis (G) Lecithochirium ghanense (G) Prosorhynchus caudovatus (G) Parahemiurus merus (G) Ethmalosa dorsalis (Cuv. and Val.), Clupeidae Larimus peli Bleeker, Sciaenidae Parahemiurus merus (G) Helicometra fasciata (G) Euthynnus alleteratus (Raf.), Scombridae Immature Didymozoid G (G) Immature Didymozoid B (G) Pseudopecoelus tortugae (G) Lecithochirium microstomum (G)

Syncoelium katuwo (S) 2 Freshwater fish.

Lethrinus atlanticus Guv. and Val., Lethrinidae Paracryptogonimus ghanensis (G) Lasiotocus accraensis (G) Pedunculotrema capecoastense (G) Lobotes surinamensis (Bloch), Lobotidae Pleorchis ghanensis (G) Bucephaloides ovatus (C) Proctotrema amphitruncatum (G) Lophius sp., Lophiidae Pycnadenoides ghanensis (G) Bucephaloides gracilescens (S) Pomadasys suillus (Val.) Lutjanus guineensis Bleeker, Lutjaiiidae Palaeorchis senegalensis (S) Paracryptogonimus ghanensis (G) Psettodes belcheri Bennett, Psettodidae Lutjanus maltzani (Steind.) Immature Didymozoid E (G) Prosorhynchus caudovatus (G) Parahemiurus merus (G) Lutjanus modestus Bleeker Rhipidocotijle ghanensis (G) Helicometra fasciata (G) Sterrhurus ghanensis (G) Pycnadena africana (SL) Pseudotolithus senegalensis (Giinther), Stephanostomum casum (G) Sciaenidae Melanoglaea ventralis Barnard, Ateleopidae Stephanostomum africanum (SL) Coitocaecum cadenati (S) Pseudupeneus cyclostomus (Lac.), Mullidae Mustelus canis (Mitchill), Carcharhinidae Didymozoidae gen. sp. larvae V (G) Ptychogonimus megastomus (S) Pteroplatea micrura (Schneider), Trygonidae Mijxus curvidens (Val.), Mugilidae Immature Didymozoid E (G) Stephanostomum megacephalum (G) Rhinobatus albomaculatus Norman, Narcacion torpedo Klein, Torpedinidae Rhinobatidae Monodhelmis torpedinis (M) Immature Didymozoid E (G) Ophichthus semicinctus (Richardson), Poracanthium ghanense (G) Ophichthyidae Rhinoptera marginata (Geoff.), Myliobatidae Proctotrematoides ophichthi (G) Nagmia africana (S) Otolithus brachygnathus (Bleeker), Sciaenidae Rupiscartes atlanticus (Val.), Blenniidtie Stephanostomum africanum (S) Prosorhynchus aculeatus, metacercaria (S) Pagellus bogaraveo (Briinnich), Sparidae Sardinella cameronensis Regan, Clupeidae Parahemiurus merus (S) Parahemiurus merus (G) Pycnadenoides senegalensis (S) Sargus cervinus Val., Sparidae Steringotrema diver gens (S) Aephnidio genes africanus (S) Pagrus ehrenbergi Guv. and Val., Sparidae Sciaena sp., Sciaenidae Markevitschiella sp. (G) Sterrhurus ghanensis (G) Paraconger notialis Kantor, Congridae Scomber colias Gmelin, Carangidae Sterrhurus ghanensis (G) Lecithocladium excisum (G) Parakuhlia boulengeri Pellegrin, Kuhliidae Lepocreadium ghanense (G) Holorchis legendrei (S) Opechona bacillaris (G) Pentanemus quinquarius (L.), Polynemidae Scomberomorus tritor (Cuv. and Val.), Poracanthium ghanense (G) Scombridae Periophthalmus koelreuteri (Pallas), Gobiidae Bucephaloides ghanensis (G) Lecithaster ghanensis (G) Didymozoidae (Monilicaecum) larvae I (G) Phyllogramma regani Pellegrin, Muraenesocidae Immature Didymozoid E (G) Sterrhurus ghanensis (G) Lecithochirium ghanense (G) Tubulovesicula lindbergi (G) Lecithocladium excisum (G) Pomadasys jubelini (Cuv. and Val.), Scorpaena scrofa L., Scorpaenidae Pomadasyidae Helicometra fasciata (S) Aephnidio genes senegalensis (G) (SL) Pseudopecoelus vulgaris (S) Diphtherostomum anisotremi (G) Scorpaena senegalensis Steind. Immature Didymozoid E (G) Sterrhurus musculus (G) Lasiotocus attenuatus (G) Scyris alexandrinus (Geoff.), Carangidae Lasiotocus chaetodipteri (G) Immature Didymozoid E (G)

Selar crumenophthalmus (Bloch), Carangidae Lecithochirium ghanense (G) Ectenurus virgulus (S) Lecithocladium augustiovum (G) Lecithochirium ghanense (S) Neolepocreadium caballeroi (G) Monascus typicus (G) (S) Parahemiurus merus (G) Parahemiurus merus (G) Stephanostomum trachinoti (G) Sterrhurus musculus (G) Sterrhurus ghanensis (G) Seriola dumerili (Risso), Carangidae Trachinotus goreensis (Cuv. and Val.) Sclerodistomum italicum (S) Gonocercella trachinoti (G) Smarts melanurus Val., Maenidae Lecithochirium ghanense (G) Holorchis legendrei (S) Lecithocladium augustiovum (G) Solea hexophthalma Bennett, Soleidae Neolepocreadium caballeroi (G) Homalometron senegalense (S) Parahemiurus merus (G) Sparisoma cretense (L.), Sparisomidae Parahurleytrema trachinoti (G) Prosorhynchus aculeatus, metacercaria (S) Stephanostomum ghanense (G) Syacium micrurum Ranzini, Bothidae Trichiurus lepturus L., Trichiuridae Sterrhurus ghanensis (G) Lecithochirium ghanense (G) Synaptura lusitanica Capello, Soleidae Lecithochirium microstomum (G) Lasiotocus cynoglossi (G) Pseudopecoelus tortugae (G) Lasiotocus ghanensis (G) Trygon marmorata (Steind.), Dasyatidae Lasiotocus synapturae (G) Nagmia senegalensis (S) Temnodon saltator Val., Pomacentridae Umbrina canariensis Val., Sciaenidae Sclerodistomum italicum (S) Stephanostomum bicoronatum (S) Tetraodon pustulus Murray, Tetraodontidae Umbrina cirrosa (L.) Diploproctodaeum ghanense (G) Pycnadenoides ghanensis (G) Torpedo narce Risso, Torpedinidae Umbrina ronchus Val. Otodistomum veliporum (M) Pleorchis ghanensis (S) Trachinocephalus myops (Schneider), Pycnadenoides ghanensis (G) Synodidae Stephanostomum bicoronatum (S) Lecithochirium ghanense (G) Umbrina steindachneri Cadenat Sterrhurus ghanensis (G) Stephanostomum bicoronatum (S) Trachinotus glaucus (L.), Carangidae Upeneus prayensis Cuv. and Val., Mullidae Aponurus lagunculus (G) Lecithocladium augustiovum (S) Ectenurus virgulus (G) Vomer setapinnis (Mitchill), Carangidae Lecithaster ghanensis (G) Didymozoidae (Monilicaecurn) larvae I (G)

Redescription of Trichuris fossor Hall, 1916 (Nematoda: Trichuridae) from the Northern Pocket Gopher, Thomomys talpoides1 KENNETH S. TODD, JR., AND DONALD L. LEPP College of Veterinary Medicine, University of Illinois, Urbana, Illinois

Trichuris fossor Hall, 1916, was described basis of measurements of two males and one from the northern pocket gopher, Thomomys female specimen. Chandler (1945) briefly talpoides (syn. fossor), from Colorado on the described Trichuris fossor from Thomomys bottae from California. During a survey of 1 Supported in part by General Research Support Grant FR-05460 from the NIH. the parasites of Thomomys talpoides from