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Checklist of the helminth parasites of vertebrates in Costa Rica.

Abstract: Helminth parasites of vertebrates have been studied in Costa Rica for more than 50 years. Survey work on this group of parasites is far from complete. We assembled a database with all the records of helminth parasites of wild and domestic vertebrates in Costa Rica. Information was obtained from different sources such as literature search (all published accounts) and parasite collections. Here we present a checklist with a parasite-host list as well as a host-parasite list. Up to now, 303 species have been recorded, including 81 species of digeneans, 23 monogeneans, 63 cestodes, 12 acanthocephalans, and 124 nematodes. In total, 108 species of vertebrates have been studied for helminths in Costa Rica (31 species of fishes, 7 amphibians, 14 reptiles, 20 birds, and 36 mammals). This represents only 38% of the vertebrate fauna of Costa Rica since about 2,855 species of vertebrates occur in the country. Interestingly, 58 species (19.1%) were recorded as new species from Costa Rica and most of them are endemic to particular regions. Considering the valuable information that parasites provide because it is synergistic with all the information about the natural history of the hosts, helminth parasites of vertebrates in Costa Rica should be considered within any initiatives to accomplish the national inventory of biological resources. Starting with this compilation work, the Coleccion de Helmintos de Costa Rica (CHCR), hosted at the Facultad de Microbiologia, Universidad de Costa Rica, has re-emerged and it is our hope that it will have the standards of quality to assure that it will become the national depository of helminths in the country.

Key Words: Helminths, Costa Rica, Biodiversity, Biological inventories, Parasites, Vertebrates.

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Costa Rica possesses a strong tradition in the study of parasites of domestic and wildlife vertebrates. For more than 50 years the helminth fauna of these hosts has been studied by national parasitologists and microbiologists (see for example Jimenez-Quiros and Brenes-Madrigal 1957, Brenes-Madrigal and Jimenez-Quiros 1959, Brenes-Madrigal et al. 1959a, Brenes-Madrigal et al. 1960a, Brenes-Madrigal and Arroyo-Sancho 1962a y b, Brenes-Madrigal et al. 1966). Since the very beginning in his career, Dr. Rodrigo Brenes-Madrigal, from the Facultad de Microbiologia, Universidad de Costa Rica, established a close collaboration with Dr. Eduardo Caballero and his research group from the Instituto de Biologia, Universidad Nacional Autonoma de Mexico (UNAM) to contribute to the understanding of the Costa Rican helminth parasite biodiversity and as a result, several papers were published between the late 50's and early 80's (see Caballero et al. 1957, Caballero and Brenes-Madrigal 1958, Brenes-Madrigal and Bravo-Hollis 1959, Lamothe-Argumedo and Brenes-Madrigal 1983). A first attempt to compile the information of the helminth parasites of Costa Rica was made by Brenes-Madrigal (1961) who presented up to that year a checklist of about 150 species described as helminth parasites of wild and domestic vertebrates as well as human beings.

However, the extraordinary biological richness of Costa Rica has attracted the attention of several foreign parasitologists who have contributed, more recently, to increase the amount of information about the helminths of vertebrates in Costa Rica and in fact, a group of parasitologists leaded by Dr. Daniel Brooks have developed a research program lo inventory the parasites of particular regions of the country, i.e. the Area de Conservacion de Guanacaste (ACG), in the province of Guanacaste (further information about the data base of parasites of vertebrates in the ACG can be obtained at: http://brooksweb.zoo.utoronto.ca) (see for example Marques et al. 1996, Perez-Ponce de Leon et al. 1998, Brooks et al. 1999, Platt 2000, Zelmer and Brooks 2000, Carreno et al. 2001, Leon-Regagnon et al. 2001, Choudhury el al. 2002).

Of all the helminth specimens collected from Costa Rican hosts, only a few were saved in the collection of the Facultad de Microbiologia, University of Costa Rica and most of them have been deposited in foreign parasite collections, i.e. the Coleccion Nacional de Helmintos, Mexico City, Mexico, the U.S. National Parasite Collection, Beltsville, Maryland, U.S.A., and in the Harold W. Manter Laboratory of Parasitology, Lincoln, Nebraska, U.S.A., since there was not a national parasite collection in Costa Rica. However, this is changing. Recently, the University of Costa Rica has issued an initiative to support biological collections, among other kind of collections. This interest is completely congruent with the initiative developed by Costa Rica to establish an inventory of the biological diversity within its territory, and to facilitate their efforts to preserve biodiversity. The starting point for planning the conservation and use of biodiversity is the compilation of information through biological surveys and inventories. Biological collections are one of the most important components of this inventory work, considering both the specimens preserved with high standards of quality and the information associated with each specimen (database) (Reaka-Kudla et al. 1997). So the objective of this paper is two-fold, to update all the available information about helminth parasites of vertebrates in Costa Rica, and to set the basis for the re-establishment of the Coleccion Helmintologica de la Facultad de Microbiologia as the Coleccion de Helmintos de Costa Rica which should have the status of National Collection, and will become the depository of the helminth parasites of vertebrates and invertebrates in the near future, calling the attention to all national and foreign parasitologists, that some of the specimens collected in the country, and their published accounts should be deposited in this re-emerged collection.

MATERIALS AND METHODS

To accomplish the goals of this paper, we conducted a search in two kinds of sources: Parasite collections and bibliographic references. Information from the Coleccion Nacional de Helmintos in Mexico City, from the U.S. National Parasite Collection, and the Parasitological Collection of the Harold W. Manter Laboratory of Parasitology, University of Nebraska-Lincoln, both in the U.S.A., was obtained with permission of their curators. All records from Costa Rican hosts were then extracted and entered in a database built on Microsoft Access 2000. At the same time, a bibliographical search was conducted in sources such as Helminthological Abstracts and Biological Abstracts. We did put special attention to all published accounts done in Costa Rican journals where some records have also been published. These records were also entered in the database. The database will be available upon request to B.R.O., curator of the Coleccion de Helmintos de Costa Rica.

The checklist is divided in two parts. First we present a parasite-host list. It is presented in phylogenetic order starting with Platyhelminthes (Digenea, Monogenea and Cestoda), and following with Acanthocephala and Nematoda. Within each phylum and class, families are presented alphabetically and within family, species ate also presented in alphabetical order. To allocate species in families we followed taxonomical treatments such as Yamaguti (1963, 1971), Amin (1985, 1992), Schmidt (1986), Khalil et al. (1994), and Anderson et al. (1973-1984). Each record contains the information about the species name, authority and year, habitat, host(s) with the group to which it belongs in parentheses (see symbology section), locality (particular locality when possible and province), and bibliographical reference. In those cases where specimens were deposited in some parasite collection, the collection acronym is mentioned. Finally, in most cases we present notes to point out possible taxonomical changes, synonyms, and new combinations of names. In all cases, we tried to consider the inclusion of the most well accepted taxonomical name following authorities. Most records are made from adults, but when larval forms were recorded, it is pointed out after the parasite name in parentheses.

The second part is the host-parasite list. This list is ordered following a traditional classification of vertebrates (Pisces, Amphibia, Reptilia, Aves, and Mammalia). Within each vertebrate group, host species are presented in alphabetical order, with the list of helminths parasitizing each of them ordered in the same way, indicating the group to which each helminth parasite belongs (see symbology). We used the name of the host as described in the original papers, however we recognize that some of them may have changed after taxonomical revisions or more accurate determinations.

RESULTS

In this checklist we present records for 303 species included in 206 genera of helminths. Eighty-one of the 303 species are digeneans, 23 are monogeneans, 63 are cestodes, 12 are acanthocephalans and finally, 124 are nematodes. Next we present the information for each species of helminth.

Parasite-Host List

Symbology: A= amphibian; B= bird; F= fish; M= mammal; R= reptile; (NP)= not published; (NA)= information not available; CMNPA refers to Canadian Museum of Nature, Invertebrate Zoology Collection, Ottawa, Canada; CHCR to the Coleccion de Helmintos de Costa Rica, San Jose, Costa Rica; CNHE to the Coleccion Nacional de Helmintos, Mexico City, Mexico; HWML to the Harold W. Manter Laboratory, University of Nebraska-Lincoln, Nebraska, U.S.A.; IOC to the Instituto Oswaldo Cruz, Rio de Janeiro, Brazil; USNPC to the U.S. National Parasite Collection, Beltsville, Maryland, U.S.A., and MNHG to the Museum de Histoire Naturelle, Geneva, Switzerland.

PLATYHELMINTHES Gegenbaur, 1859

Digenea (Van Beneden, 1858)

Acanthocolpidae Luhe, 1909

1. Manteria brachydera (Manter, 1940) Caballero, 1950.

Stomach, Caranx caballus, Oligoplites altus, Oligoplites refulgens (F), Golfo de Nicoya, Puntarenas (Ponciano-Rodriguez 1986). Specimens: CNHE.

2. Stephanostomum casum (Linton, 1910) McFarlane, 1934.

Intestine, Caranx caballus (F), Playa Ocotal, Guanacaste (Perez-Ponce de Leon et al. 1998). Specimens: CNHE.

Acanthostomidae Poche, 1926

3. Proctocaecum acuti (Caballero and Brenes-Madrigal, 1958) Lamothe-Argumedo and Ponciano-Rodriguez, 1985. Intestine, Crocodylus acutus (R), Los Chiles de Grecia, Alajuela (Caballero and Brenes-Madrigal 1958). Specimens: CHCR, CNHE.

Notes: This species was described as Acanthostomum acuti by Caballero and Brenes-Madrigal (1958) and transferred to Atrophecaecum by Groschaft and Barus (1970). Synonym of Acanthostomum scyphocephalum (Nasir 1974), Caimanicola marajoara (Brooks 1980), and A. americanum (Moravec 2001).

4. Stunkardiella minima (Stunkard, 1938) Lamothe-Argumedo and Ponciano-Rodriguez, 1985.

Intestine, Rhamdia rogersi (F), Rio Tiribi, Paso Ancho, San Jose (Caballero and Brenes-Madrigal 1957). Specimens: CHCR, CNHE, HWML.

Notes: Specimens originally recorded as Acanthostomum gnerii Szidat, 1954 by Caballero and Brenes-Madrigal (1957). Lamothe-Argumedo and Ponciano (1985) transferred this record to S. minima, however this criterion was not accepted by Brooks and Holcman (1993).

Brachycoeliidae (Looss, 1899) Johnston, 1912

5. Mesocoelium travassosi Pereira and Cuocolo, 1940.

Intestine, Bufo marinus (A), Tilaran, Guanacaste (Caballero and Brenes-Madrigal 1958). Specimens: CHCR, CNHE, HWML.

Bucephalidae Poche, 1907

6. Bucephalopsis sp.

Intestine, Caranx caballus (F), "Fish Oceanographic Region No. 77" (NP). Specimens: HWML.

7. Bucephalus introversus Manter, 1940.

Stomach, Caranx hippos and Seriola sp. (F), Puerto Culebra, Guanacaste (Manter 1940).

Cephalogonimidae (Looss, 1899) Nicoll, 1914

8. Cephalogonimus americanus Stafford, 1902.

Intestine, Rana pipiens (A), Coris, Cartago (Brenes-Madrigal et al. 1959b). Specimens: CHCR.

Cyclocoelidae Kossack, 1911

9. Haematotrephus facioi (Brenes-Madrigal and Arroyo-Sancho, 1962).

Aerial sacs, Jacana spinosa (B), Aranjuez, Puntarenas (Brenes-Madrigal and Arroyo-Sancho 1962). Specimens: CHCR.

Notes: This species was described as Cyclocoelum facioi Brenes-Madrigal and Arroyo-Sancho, 1962, however it is listed within the genus Haematotrephus Stossich, 1902 by Yamaguti (1971).

10. Hyptiasmus costaricense (Brenes-Madrigal and Arroyo-Sancho, 1962)

Aerial sac and thoracic cavity, Momotus momotus (B), Quepos, Puntarenas (Brenes-Madrigal and Arroyo-Sancho 1962). Specimens: CHCR.

Notes: Originally recorded as Cyclocoelum costaricense Brenes-Madrigal and Arroyo-Sancho, 1962 and listed as a member of the genus Hyptiasmus Kossack, 1911 by Yamaguti (1971).

Dicrocoeliidae Odhner, 1910

11. Controrchis biliophilus Price, 1928

Bile ducts, Alouatta palliata palliata (M), Tilaran, Guanacaste (Jimenez-Quiros and Brenes-Madrigal 1957), La Pacifica, Guanacaste (Stuart et al. 1990). Specimens: CHCR, CNHE, HWML.

Notes: Jimenez-Quiros and Brenes-Madrigal (1957) determined his specimens as Controrchis caballeroi Jimenez-Quiros and Brenes-Madrigal, 1957, a species that was later synonymized with C. biliophilus by Jimenez-Quiros and Brenes Madrigal (1958).

12. Lubens lubens (Braun, 1901) Strom, 1940 Gall bladder, Calocitta formosa (B), Tilaran, Guanacaste (Brenes-Madrigal et al. 1966). Specimens: CHCR.

Notes: Species described as Lubens centroamericanus Brenes-Madrigal, Arroyo-Sancho and Munoz-Montoya, 1966 and synonymized with L. lubens by Lamothe-Argumedo (1979).

13. Lutztrema obliquum (Travassos, 1917) Travassos, 1941

Bile ducts, Gymnostinops montezuma (B), Chitaria, Cartago (Jimenez-Quiros and Arroyo-Sancho 1960b). Specimens: CHCR.

14. Platynosomum costaricense Brenes-Madrigal, Arroyo-Sancho and Munoz, 1966.

Gall bladder, Buteo nitidus (B), Tilaran, Guanacaste (Brenes-Madrigal et al. 1966). Specimens: CHCR.

15. P. illiciens (Braun, 1901) Travassos, 1916. Gall bladder, Leucopternis semiplumbea (B), Sarapiqui, Heredia (Brenes-Madrigal and Arroyo-Sancho 1962). Specimens: CHCR.

16. Robertdigenea dollfusi Caballero and Caballero-Rodriguez, 1970.

Bile ducts and liver, Procyon lotor crassidens (M), Balsa de Mora, San Jose (Caballero and Caballero-Rodriguez 1970). Specimens: CNHE.

17. Zonorchis costaricensis Brenes-Madrigal and Jimenez-Quiros, 1959.

Gall bladder, Gymnostinops montezuma (B), Peralta, Cartago (Brenes-Madrigal and Jimenez-Quiros 1959). Specimens: CHCR.

18. Z. macroovaricus Jimenez-Quiros and Arroyo-Sancho, 1960.

Bile ducts, Pteroglossus torquatus (B), Tilaran, Guanacaste (Jimenez-Quiros and Arroyo-Sancho 1960b). Specimens: CHCR.

Didymozoidae Poche, 1907

19. Didymozoidae gen. sp. (metacercariae). Intestine, Haemulon maculicauda (F), Bahia Culebra, Guanacaste (Perez-Ponce de Leon et al. 1998). Specimens: CNHE.

Diplostomidae Poirier, 1886

20. Alaria nasuae La Rue and Townsend, 1927 Intestine, Canis latrans (M), San Jose, San Jose (NP). Specimens: USNPC.

Eucotylidae Skrjabin, 1924

21. Tanaisia freitasi Brenes-Madrigal and Arroyo-Sancho, 1962

Kidneys, Gymnostinops montezuma (B), Orosi, Cartago (Brenes-Madrigal and Arroyo-Sancho 1962). Specimens: CHCR.

22. T. winteri Brenes-Madrigal and Arroyo-Sancho, 1962.

Kidneys, Jacana spinosa (B), Aranjuez, Puntarenas (Brenes-Madrigal and Arroyo-Sancho 1962). Specimens: CHCR.

Eumegacetidae Travassos, 1922.

23. Eumegacetes macroorchis Brenes-Madrigal and Arroyo-Sancho, 1962.

Cloaca, Crotophaga sulcirostris (B), Aranjuez, Puntarenas (Brenes-Madrigal and Arroyo-Sancho 1962). Specimens: CHCR.

Fasciolidae Railliet, 1895

24. Fasciola hepatica Linneo, 1758.

Liver, Bos taurus (M), Cerrillos, Camago (Brenes-Madrigal 1961), Santa Cruz de Turrialba (Chang-Diaz and Cartin-Gonzalez 1983), San Pedro Montes de Oca, San Jose (NP). Specimens. CHCR, USNPC.

Fellodistomidae (Nicoll, 1909) Nicoll, 1913

25. Tergestia laticollis (Rudolphi, 1819) Stossich, 1899.

Intestine, Caranx caballus (F), Playa Ocotal, Guanacaste (Perez-Ponce de Leon et al. 1998), Puerto Culebra, Guanacaste (Manter 1940). Specimens: CNHE, HWML.

Gorgoderidae Looss, 1901

26. Gorgoderina megalorchis Bravo-Hollis, 1948.

Urinary bladder, Bufo marinus (A), Piedad de Santa Ana, San Jose (Caballero et al. 1957). Specimens: CHCR, CNHE.

Haematoloechidae Odening, 1964

27. Haematoloechus meridionalis Leon-Regagnon, Brooks and Zelmer, 2001.

Lungs, Rana vaillanti (A), Rio Pizote, Guanacaste (Leon-Regagnon et al. 2001). Specimens: CNHE, USNPC.

28. Neohaematoloechus helibravoae Caballero-Rodriguez, 1989.

Lungs, Rana palmipes (A), NA (NP). Specimens: CNHE.

Notes: This species is considered nomina nuda.

Hemiuridae Luhe, 1901

29. Ectenurus virgulus Linton, 1910.

Intestine, Fistularia commersonii (F), Playa Ocotal, Guanacaste (Perez-Ponce de Leon et al. 1998). Specimens: CNHE.

30. Halipegus eschi Zelmer and Brooks, 2000.

Esophagus, Rana vaillanti (A), Rio Pizote, Guanacaste (Zelmer and Brooks 2000). Specimens: USNPC.

31. Lecithochirium microstomum Chandler, 1935.

Intestine, Fistularia commersonii (F), Playa Ocotal, Guanacaste (Perez-Ponce de Leon et al. 1998). Specimens: CNHE.

32. L. monticelli (Linton, 1898) Crowcroft, 1946.

Stomach, Synodus sp. (F), Puntarenas (Bravo-Hollis and Arroyo-Sancho 1962a). Specimens: CNHE.

Notes: According to Yamaguti (1971) this material may belong to a different species.

33. Mecoderus oligoplites Manter, 1940.

Stomach, Oligoplites sp. (F), Puntarenas (Bravo-Hollis and Arroyo-Sancho 1962a), Oligoplites altus and Oligoplites refulgens (F), Golfo de Nicoya, Puntarenas (Ponciano-Rodriguez 1986). Specimens: CNHE.

34. Pseudolecithaster sp.

Intestine, Echidna nocturna (F), Playa Cuajiniquil, Guanacaste (Perez-Ponce de Leon et al. 1998). Specimens: CNHE.

35. Theletrum lamothei Perez-Ponce de Leon, Leon-Regagnon and Monks, 1998.

Intestine, Echidna nocturna (F), Playa Cuajiniquil, Guanacaste (Perez-Ponce de Leon et al. 1998). Specimens: CNHE, USNPC.

Heronimidae Ward, 1917

36. Heronimus mollis (Leidy, 1856) Stunkard, 1964.

Lungs, Kinosternon cruentatum (R), Santa Ana, San Jose (Caballero et al. 1957). Specimens: CNHE.

Notes: Species originally recorded as Heronimus chelydrae MacCallum, 1902.

Heterophyidae Odhner, 1914

37. Euryhelmis costaricensis Brenes-Madrigal, Arroyo-Sancho and Jimenez-Quiros, 1960

Intestine, Mustela frenata (M), Tres Rios, Cartago (Brenes-Madrigal et al. 1960a). Specimens: CHCR.

Homalometridae (Cable and Hunnien, 1942)

38. Crassicutis cichlasomae Manter, 1936.

Intestine, Cichlasoma sp. (F), Liberia, Guanacaste (Bravo-Hollis and Arroyo-Sancho 1962b). Specimens: CNHE.

39. C. opisthoseminis Bravo-Hollis and Arroyo-Sancho, 1962.

Intestine, Cichlasoma sp. (F), Liberia, Guanacaste (Bravo-Hollis and Arroyo-Sancho 1962b). Specimens: CNHE.

Lecithodendriidae (Luhe, 1931) Odhner, 1910

40. Castroia silvai Travassos, 1928.

Intestine, Micronycteris hirsutus (M), Rio Grande de Atenas, Alajuela (Caballero and Montero-Gei 1963). Specimens: CNHE.

Notes: According to Martin (1969), morphology of the specimens differs from the original description.

41. Langeronia provitellaria Sacks, 1952.

Intestine, Rana warchewitschii and Bufo marinus (N), Coris, Cartago (Brenes-Madrigal et al. 1959a). Specimens: CHCR.

Notes: Specimens originally determined as Langeronia macrocirra Caballero and Bravo-Hollis, 1949 and later transferred to L. provitellaria by Christian (1970).

42. Limatulum costarricensis (Caballero, y Brenes-Madrigal, 1957) Caballero, 1964.

Intestine, Eptesicus propinquus (M), Finca Monte de Heredia (Caballero and Brenes-Madrigal 1957). Specimens: CHCR, CNHE.

Note: Originally recorded as Ochoterenatrema costarricensis Caballero and Brenes-Madrigal, 1957 and later transferred to the genus Limatulum Travassos, 1921 by Caballero (1964).

43. Prosthodendrium cordiforme (Braun, 1900) Macy, 1936.

Intestine, Eptesicus propinquus (M), Finca Monte de Heredia (Caballero and Brenes-Madrigal 1957). Specimens: CHCR, CNHE.

44. Pseudosonsinotrema chabaudi (Caballero and Caballero-Rodriguez, 1969).

Intestine, Rana pipiens (A), Sarapiqui, Heredia (Caballero and Caballero-Rodriguez 1969), Turrialba, Cartago (Sullivan 1971). Specimens: CNHE, HWML, USNPC.

Notes: Specimens collected by Caballero and Caballero-Rodriguez (1969) was included in the genus Brenesia Caballero and Caballero-Rodriguez, 1969, which was later synonymized with Pseudosonsinotrema by Sullivan (1974). Additionally, species described by Sullivan (1971) as Pseudosonsinotrema echinophallus Sullivan, 1970 was synonymized with P. tabai (Sullivan, 1974).

Lepocreadiidae (Odhner, 1905) Nicoll, 1935

45. Bianium plicitum (Linton, 1928) Stunkard, 1930.

Intestine, Sphoeroides sp. (F), Mata de Limon, Puntarenas (Caballero and Brenes-Madrigal 1957). Specimens: CHCR, CNHE.

46. B. simonei (Travassos, Teixeira de Freitas and Buhrnheim, 1965) Yamaguti, 1971

Intestine, Arothron hispidus (F), Bahia Prieta, Guanacaste (Perez-Ponce de Leon et al. 1998). Specimens: CNHE.

47. Hypocreadium myohelicatum Bravo-Hollis and Manter, 1957.

Intestine, Epinephelus itajara (F), Golfo de Santa Clara, Guanacaste (Perez-Ponce de Leon et al. 1998). Specimens: CNHE.

Leucochloridiidae (Poche, 1907) Dollfus, 1934

48. Leucochloridium costaricense Brenes-Madrigal and Arroyo-Sancho, 1962.

Intestine, Gymnostinops montezuma (B), Cachi, Cartago (Brenes-Madrigal and Arroyo-Sancho 1962). Specimens: CHCR.

Macroderoididae McMullen, 1937

49. Glypthelmins facioi Brenes-Madrigal, Arroyo-Sancho, Jimenez-Quiros and Delgado-Flores, 1959.

Intestine, Rana pipiens (A), Coris, Cartago (Brenes-Madrigal et al. 1959b), Turrialba, Cartago (Sullivan 1976). Specimens: CHCR, USNPC.

50. G. hepaticus (Lutz, 1928).

Liver, Bufo marinus (A), Piedades de Santa Ana, San Jose (Caballero et al. 1957). Specimens: CHCR, CNHE.

Notes: Originally recorded as Choledocystus intermedius Caballero, Bravo-Hollis and Zerecero, 1944, but synonymized with G. hepaticus by Sullivan (1977).

51. G. palmipedis (Lutz, 1928) Travassos, 1930.

Intestine, Bufo marinus (A), Chapernal, Puntarenas (Brenes-Madrigal and Arroyo-Sancho 1960). Specimens: CHCR.

52. Parallopharynx gonzalezi Brenes-Madrigal, Arroyo-Sancho and Montero-Gei, 1960

Intestine, Ctenosaura similis and Basiliscus sp., (R), Rio Grande de Atenas, Alajuela (Brenes-Madrigal et al. 1960b). Specimens: CHCR, HWML.

53. Wallinia chavarriai Choudhury, Daverdin and Brooks, 2002.

Intestine, Astyanax aenaeus (F), Rio Sapoa, Guanacaste, and Bryconamericus scleroparius (F), Rio Sapoa and Quebrada Limonal, Guanacaste (Choudhury et al. 2002). Specimens: HWML, USNPC.

Microphallidae Travassos, 1920

54. Maritremopsis longibursatus (Caballero and Brenes-Madrigal, 1964) Deblock, 1973.

Intestine, Chamaeptes unicolor (A), Caserio de Tiquiares, San Jose (Caballero and Brenes-Madrigal 1964). Specimens: CHCR, CNHE.

Notes: Originally described as Ornithotrema longibursatum Caballero and Brenes-Madrigal, 1964 and transferred to the genus Maritremopsis Deblock, 1963 by Deblock (1973).

55. M. prolixum (Caballero and Montero-Gei, 1961) Deblock, 1973.

Bile ducts, Philander opossum (M), Rio Grande de Atenas, Alajuela (Caballero and Montero-Gei 1961). Specimens: CNHE.

Notes: Originally described as Maritrema prolixum Caballero and Montero-Gei, 1961, and later transferred to the genus Pseudomaritrema Belopolskaya, 1952 by Caballero (1964), and to the genus Metamaritrema by Yamaguti (1971).

56. Ornithotrema momoti Caballero, Brenes-Madrigal and Arroyo-Sancho, 1963.

Intestine, Momotus momotus (B), Sarapiqui, Heredia (Caballero et al. 1963a). Specimens: CHCR, CNHE.

57. Spelotrema skrjabini (Caballero, 1958) Yamaguti, 1971.

Intestine, Didelphis marsupialis (M), Bello Horizonte, Escazu, San Jose (Caballero 1958). Specimens: CHCR, CNHE.

Notes: Originally described as Carneophallus skrjabini Caballero, 1958.

Monorchiidae Odhner, 1911

58. Lasiotocus costaricae (Manter, 1940) Yamaguti, 1954.

Intestine and caeca, "Medialuna" (F), Puerto Culebra, Guanacaste (Manter 1940). Specimens: HWML, USNPC.

Notes: Originally described as Proctotrema costaricae Manter, 1940.

Opecoelidae Ozaki, 1925

59. Helicometra sinuata (Rudolphi, 1819) Odhner, 1902.

Intestine, "Moray" (F), Puerto Culebra, Guanacaste (Manter 1940). Specimens: HWML.

Opisthorchiidae (Looss, 1899) Braun, 1901

60. Amphimerus costarricenis Lamothe-Argumedo and Brenes-Madrigal, 1983.

Bile ducts, Felis catus (M), Boruca, San Jose (Lamothe-Argumedo and Brenes-Madrigal 1983). Specimens: CNHE.

61. A. neotropicalis Caballero, Montero-Gei and Caballero-Rodriguez, 1963.

Pancreatic ducts, Philander opossum (M), Rio Grande de Atenas, Alajuela (Caballero et al. 1963b). Specimens: CNHE.

Note: Nasir and Diaz (1972) synonymized this species with A. speciosus (Stiles and Hassall, 1896) and Correa-Gomes (1979) with A. pseudofelineus (Ward, 1911).

Paragonimidae Dollfus, 1939

62. Paragonimus caliensis Little, 1968.

Lungs, Chironecter minima and Procyon lotor (M), Gamalotillo de Puriscal, San Jose and Didelphis marsupialis (M), Parrita, Puntarenas (Brenes-Madrigal et al. 1984).

63. P. mexicanus Miyazaki and Ishii, 1968.

Lungs, Canis familiaris and Conepatus semistriatus (M), NA (Brenes-Madrigal et al. 1984), Didelphis marsupialis (M), Tabarcia de Mora, San Jose (Brenes-Madrigal et al. 1980), Felis catus (M), Balsa de Atenas, Alajuela, Rincon de Osa, Puntarenas, Tabarcia de Mora, San Isidro de Balsa and Cangrejal de Acosta , San Jose (Brenes-Madrigal et al. 1980), Felis pardalis (M), Bagaces, Guanacaste (Brenes-Madrigal et al. 1980), Philander opossum (M), Rio Grande de Atenas, Alajuela (Caballero and Montero-Gei 1961), Balsa de Atenas and Tornos de Atenas, Alajuela, Rincon de Osa, Puntarenas, and San Isidro de Balsa, San Jose (Brenes-Madrigal et al. 1980), Procyon lotor (M), San Isidro de Balsa, San Jose (Brenes-Madrigal et al. 1980), Urocyon cinereoargenteus (M), Balsa de Atenas and Turrucares, Alajuela (Brenes-Madrigal et al. 1980), Carrizal de Alajuela, Alajuela (Caballero 1956). Specimens: CNHE.

Notes: Brenes-Madrigal et al. (1980), synonymized all records of Paragonimus rudis (Diesing, 1850), by Caballero (1956) and by Caballero and Montero-Gei (1961) with P. mexicanus.

Paramphistomidae Fischoeder, 1901

64. Cotylophororon sp.

Stomach, Bos taurus (M), San Carlos, Alajuela (NP). Specimens: USNPC.

65. Megalodiscus temperatus (Stafford, 1905) Harwood, 1932.

Intestine, Rana pipiens (A), Coris, Cartago (Brenes-Madrigal et al. 1959b). Specimens: CHCR.

66. Paramphistomum cervi (Schrank, 1790) Stomach, Bos taurus (M), San Jose, San Jose (NP). Specimens: CHCR.

67. P. liorchis Fischoeder, 1901.

Rumin, Odocoileus virginianus (M), Parque Nacional Santa Rosa, Guanacaste (Carreno et al. 2001). Specimens: USNPC.

Plagiorchiidae (Luhe, 1901) Ward, 1917

68. Ochetosoma bravoae Brenes-Madrigal and Arroyo-Sancho, 1960.

Esophagus, "Colubridae" (R), Bello Horizonte, Escazu, San Jose (Brenes-Madrigal and Arroyo-Sancho 1960). Specimens: CHCR.

69. O. miladelarocai Caballero y Vogelsang, 1947.

Esophagus, Drymobius margaritiferus (R), Coris, Cartago (Brenes-Madrigal et al. 1959a). Specimens: CHCR.

70. Plagiorchis sp.

Intestine, amphibians, birds and turtles, NA (Brenes-Madrigal 1961).

Proterodiplostomidae Dubois, 1936

71. Prolecithodiplostomum constrictum Dubois, 1936.

Intestine, Caiman crocodylus (R), La Selva, Heredia and Palma Sur, Puntarenas (Brooks et al. 1977). Specimens: HWML.

72. Proterodiplostomum medusae (Dubois, 1937) Caballero, Hidalgo-Escalante and Grocott, 1957.

Intestine, Crocodylus acutus (R), Los Chiles de Grecia, Alajuela (Caballero and Brenes-Madrigal, 1958), Caiman crocodylus (R), La Selva, Heredia (Brooks et al. 1977). Specimens: CHCR, CNHE, HWML.

Psilostomidae Looss, 1900

73. Mesaulus grandis (Rudolphi, 1819) Braun, 1902.

Intestine, Ajaia ajaja (B), Mata de Limon, Puntarenas (Caballero and Brenes-Madrigal 1958). Specimens: CHCR, CNHE.

Notes: Originally recorded as Cotylotretus grandis (Rudolphi, 1819) Odhner, 1910 and transferred to Mesaulus Braun, 1902.

Rhopaliidae (Looss, 1899) Viana, 1924

74. Rhopalias baculifer Braun, 1900.

Intestine, Philander opossum (M), Rio Grande de Atenas, Alajuela (Caballero and Montero-Gei 1961). Specimens: CNHE, HWML.

75. R. coronatus (Rudolphi, 1819) Stiles and Hassall, 1898.

Intestine, Didelphis marsupialis (M), Bello Horizonte, Escazu, San Jose (Caballero et al. 1957), Philander opossum (M), Rio Grande de Atenas, Alajuela (Caballero and Montero-Gei 1961), Cariari, Limon (Siebert 1970). Specimens: CHCR, CNHE, HWML.

76. R. horridus (Diesing, 1850) Stiles and Hassall, 1898.

Intestine, Didelphis marsupialis (M), Bello Horizonte, Escazu, San Jose (Caballero et al. 1957), Philander opossum (M), Rio Grande de Atenas, Alajuela (Caballero and Montero-Gei 1961). Specimens: CHCR, CNHE.

77. R. macracanthus Chandler, 1932. Intestine, Philander opossum (M), Cariari, Limon (Siebert 1970).

Stomylotrematidae (Travassos, 1922) Poche, 1926

78. Stomylotrema ucremium Brenes-Madrigal, Arroyo-Sancho and Munoz-Montoya, 1966.

Intestine, Icterus galbula (B), Alajuela, Alajuela (Brenes-Madrigal et al. 1966). Specimens: CHCR.

Telorchiidae Dollfus, 1925

79. Telorchis sp.

Intestine, turtle (R), NA (Brenes-Madrigal 1961), Geomyda manni (R), NA (NP). Specimens: USNPC.

80. T. pseudoaculeatus Dollfus, 1929

Intestine, Kinosternon cruentatum (R), Santa Ana, San Jose (Caballero and Brenes-Madrigal 1957). Specimens: CHCR, CNHE.

Notes: According to Nasir (1974), this species is a synonym of T. aculeatus (Von Linstow, 1879).

Urotrematidae Poche, 1926

81. Urotrema scabridum Braun, 1900.

Intestine, Eptesicus propinquus (M), Monte Heredia, Heredia (Caballero et al. 1957). Specimens: CHCR, CNHE.

Monogenoidea

Bychowsky, 1937

Allodiscocotylidae Tripathi, 1959

1. Hargicola oligoplites (Hargis, 1957) Lebedev, 1970.

Gills, Oligoplites altus (F), Golfo de Nicoya, Puntarenas (NP). Specimens: CNHE.

Cemocotylidae Yamaguti, 1963

2. Cemocotylella elongata (Meserve, 1938) Price, 1962.

Gills, Caranx caballus (F), Golfo de Nicoya, Puntarenas (NP). Specimens: CNHE, HWML.

Chauhaneidae Lebedev, 1972

3. Ahpua piscicola Caballero and Bravo-Hollis, 1973.

Gills, Caranx caballus, Oligoplites altus (F), Golfo de Nicoya, Puntarenas (NP). Specimens: CNHE, HWML.

Dactylogyridae Bychowsky, 1933

4. Ameloblastella chavarriai (Price, 1938) Kristky, Mendoza and Scholz, 2000. Gills, Rhamdia rogersi (F), San Pedro Montes de Oca, San Jose (Price 1938). Notes: This species was described as Cleidodiscus chavarriai Price, 1938, transferred to the genus Urocleidoides Mizelle and Price, 1964 by Molnar et al. (1974) and established as the type species of Ameloblastella Kritsky, Mendoza and Scholz, 2000 by Kritsky et al. (2000).

5. Aphanoblastella travassosi (Price, 1938) Kristky, Mendoza and Scholz, 2000. Gills, Rhamdia rogersi (F), San Pedro Montes de Oca, San Jose (Price 1938). Notes: This species was described as Cleidodiscus travassosi Price, 1938, transferred to the genus Urocleidoides by Molnar et al. (1974) and established as the type species of Aphanoblastella Kritsky, Mendoza and Scholz, 2000 by Kritsky et al. (2000). Specimens: HWML.

6. Jainus hexops Kritsky and Leiby, 1972. Gills, Astyanax fasciatus (F), 3 km. SW of Rincon, Puntarenas (Kritsky and Leiby 1972). Specimens: USNPC.

7. Palombitrema heteroancistrum Price and Bussing, 1968.

Gills, Astyanax fasciatus (F), Rio Montenegro, 23 Km NW of Canas, Guanacaste (Price and Bussing, 1968), 3 km. SW of Rincon, Puntarenas (Kritsky and Leiby 1972). Specimens: USNPC.

Notes: Price and Bussing (1968) created the genus Palombitrema Price and Bussing, 1968 to include this species; Kritsky and Leiby (1972) synonymized it with Urocleidoides and Sunano (1997) re-established its validity.

8. Urocleidoides costaricensis (Price and Bussing, 1967) Kritsky and Leiby, 1972. Gills, Astyanax fasciatus (F), Canas, Guanacaste (Price and Bussing 1967). Specimens: USNPC.

Notes: Species described as Cleidodiscus costaricensis Price and Bussing, 1967, transferred to Urocleidoides by Kritsky and Leiby (1972) and considered as incerme sedis by Kritsky et al. (1986).

9. Urocleidoides strombicus (Price and Bussing, 1967) Kritsky and Thatcher, 1974.

Gills, Astyanax fasciatus (F), Canas, Guanacaste (Price and Bussing 1967). Specimens: USNPC.

Notes: Described as Cleidodiscus strombicus Price and Bussing, 1967 later included in Urocleidoides by Kritsky and Thatcher (1974) and considered as incertae sedis by Kritsky et al. (1986).

Diclidophoridae Cerfontaine, 1895

10. Hargicotyle louisianensis (Hargis, 1955) Mamaev, 1972.

Gills, Menticirrhus sp., (F), Puntarenas (Bravo-Hollis and Arroyo-Sancho 1962a). Specimens: CNHE.

Notes: Originally recorded as Choricotyle louisiana Hargis, 1955, the type species of Hargicotyle Mamaev, 1972.

Discocotylidae Price, 1936

11. Heterobothrium ecuadori (Meserve, 1938).

Gills, Sphoeroides sp. (F), Mata de Limon, Puntarenas (Caballero and Brenes-Madrigal 1957). Specimens: CNHE.

Notes: Specimens originally described as Tagia ecuadori Sproston, 1946. However, Euzet and Birgi (1975), propose Tagia as a synonym of Heterobothrium Cerfontaine, 1895.

12. Pseudomazocraes monsivaisae Caballero and Bravo-Hollis, 1955.

Gills, Vomer declivifrons (F), Puntarenas (NP). Specimens: CNHE, HWML.

Gyrodactylidae Cobbold, 1864

13. Anacanthocotyle anacanthocotyle Kritsky and Fritts, 1970.

External surface, Astyanax fasciatus (F), 3 km., SW of Rincon, Puntarenas (Kritsky and Fritts 1970). Specimens: USNPC.

14. Gyrodactylus bullatarudis Turnbull, 1956. External surface, Poecilia sphenops (F), 3 km., SW of Rincon, Puntarenas (Kritsky and Fritts 1970). Specimens: USNPC.

15. G. costaricensis Kritsky and Fritts, 1970. External surface, Astyanax fasciatus, Poecilia sphenops (F), 3 km., SW of Rincon, Puntarenas (Kritsky and Fritts 1970). Specimens: USNPC.

16. G. neotropicalis Kritsky and Fritts, 1970. External surface, Astyanax fasciatus (F), 3 km., SW of Rincon, Puntarenas (Kritsky and Fritts 1970). Specimens: USNPC.

Hexabothriidae Price, 1942

17. Erpocotyle carcharhini Watson and Thorson, 1976.

Gills, Carcharhinus leucas (F), Barra del Colorado, Rio Colorado, Limon (Watson and Thorson 1976). Specimens: USNPC.

18. Heteroncocotyle leucas Hargis, 1955. Gills, Carcharhinus leucas (F), Barra del Colorado, Rio Colorado, Limon (Watson and Thorson 1976). Specimens: USNPC.

Microbothriidae Price, 1936

19. Dermophthirius maccallumi Watson and Thorson, 1976.

Skin, Carcharhinus leucas (F), Barra del Colorado, Rio Colorado, Limon (Watson and Thorson 1976). Specimens: USNPC.

Microcotylidae Taschenberg, 1879

20. Jaliscia caballeroi (Bravo-Hollis, 1960) Mamaev and Egorova, 1977.

Gills, "tunna" (F), Golfo de Nicoya, Puntarenas, (NP). Specimens: CNHE.

21. Polymicrocotyle manteri Lamothe-Argumedo, 1967.

Gills, Lutjanus colorado (F), Golfo de Nicoya, Puntarenas, (NP). Specimens: CNHE.

Polystomatidae Gamble, 1896

22. Neopolystoma fentoni Platt, 2000. Conjunctival sac, Kinosternon leucostomum (R), Quebrada Costa Rica, Guanacaste, and Rhinoclemmys pulcherrima (R), Quebrada El Duende and Quebrada San Gerardo, Guanacaste (Platt 2000). Specimens CHIOC, USNPC.

Protomicrocotylidae Poche, 1926

23. Neomicrocotyle pacifica (Meserve, 1938) Yamaguti, 1968.

Gills, Xurel marginatus (F), Puerto Culebra, Guanacaste (Meserve 1938). Specimens: USNPC.

Notes: Species originally described as Protomicrocotyle pacifica Meserve, 1938.

Cestoidea Rudolphi, 1808

Anoplocephalidae Cholodkovsky, 1902

1. Anoplocephala magna (Abildgaard, 1789) Sprengel, 1905.

Intestine, Equus caballus (M), NA (Brenes-Madrigal 1961).

2. A. perfoliata (Goeze, 1782) Blanchard, 1848.

Intestine, Equus caballus (M), NA, Limon (Brenes-Madrigal, 1961), San Pedro Montes de Oca, San Jose (NP). Specimens: USNPC.

3. Anoplocephaloides floresbarroetae Rausch, 1976.

Bile duct, Sylvilagus brasiliensis (M), Aserri, San Jose (Flores-Barroeta et al. 1961).

Notes: The species was misidentified by Flores-Barroeta et al. (1961) as Paranoplocephala infrequens (Douthitt, 1915). Later, based on this record, Rausch (1976) erected Anoplocephaloides flores-barroetae Rausch, 1976. In 1990, this species was transferred to the genus Leporidotaenia Genov, Murai, Georgiev and Harris, 1990 by Genov et al. (1990), however this genus was considered a synonym of Anoplocephaloides by Beveridge (1994).

4. A. mamillana (Mehlis, 1831) Rausch, 1976.

Intestine, Equus caballus (M), NA (Brenes-Madrigal 1961).

Notes: Originally described as Paranoplocephala mamillana (Mehlis, 1831).

5. Avitellina centripunctata (Rivolta, 1874) Gough, 1911.

Intestine, Canis familiaris (M), NA (Brenes-Madrigal 1961).

Notes: This species is a typical parasite of ruminants.

6. Mathevotaeinia bivittata (Janicki, 1904) Yamaguti, 1959.

Intestine, Marmosa sp. (M), Rio Grande de Atenas, Alajuela (Flores-Barroeta et al. 1961).

Notes: Originally recorded as Oochoristica bivittata Janicki, 1904.

7. M. didelphidis (Rudolphi, 1819) Spasskii, 1951.

Intestine, Marmosa sp. (M), Rio Grande de Atenas, Alajuela (Flores-Barroeta et al. 1961).

Notes: Originally recorded as Oochoristica didelphidis (Rudolphi, 1819).

8. M. pennsylvanica (Chandler and Melvin, 1951) Yamaguti, 1959.

Intestine, Didelphis marsupialis etensis (M), Bello Horizonte, Escazu, San Jose (Flores-Barroeta et al. 1958a). Specimens: CNHE.

Notes: Originally recorded as Oochoristica pennsylvanica Chandler and Melvin, 1951.

9. Moniezia sp. Intestine, Capra hircus (M), NA (Alvarado et al. 1990).

10. Moniezia benedeni (Moniez, 1879) Blanchard, 1891.

Intestine, Bradypus griseus griseus (M), Turrialba, Cartago (Flores-Barroeta et al. 1958b), Bos taurus (M), San Jose, San Jose (Brenes-Madrigal, 1961). Specimens: USNPC.

11. M. expansa (Rudolphi, 1805) Blanchard, 1891.

Intestine, Bos taurus (M), NA (Brenes-Madrigal 1961).

12. Oochoristica guanacastensis Brooks, Perez Ponce de Leon, and Garcia, 1999. Intestine, Ctenosaura similis (R), Parque Nacional Santa Rosa, Guanacaste (Brooks et al. 1999). Specimens: CNHE, USNPC.

Cathetocephalidae Dailey and Overstrett, 1973

13. Cathetocephalus thatcheri Dailey and Ovestreet, 1973

Spiral valve, Carcharhinus leucas (F), Barra del Colorado, Rio Colorado, Limon (Watson and Thorson 1976).

Dasyrhynchiade Dollfus, 1935

14. Dasyrhynchus variouncinatus (Pintner, 1913) Pintner, 1928.

Spiral valve, Carcharhinus leucas (F), Barra del Colorado, Rio Colorado, Limon (Watson and Thorson 1976).

Davaineidae Fuhrmann, 1907

15. Davainea proglottina (Davaine, 1860) Blanchard, 1891.

Intestine, Gallus gallus (B), NA (Brenes-Madrigal 1961).

16. Raillietina sp.

Intestine, Gallus gallus (B), NA; Sigmodon hispidus (M), Canas, Guanacaste Rodriguez-Ortiz et al. 2000). Specimens: USNPC.

17. R. echinobothrida (Megnin, 1880) Fuhrmann, 1924.

Intestine, Gallus gallus (B), NA (Brenes-Madrigal 1961).

18. R. tetragona (Molin, 1858) Fuhrmann, 1924.

Intestine, Gallus gallus (B), San Jose, San Jose (Flores-Barroeta and HidalgoEscalante 1960), NA (Brenes-Madrigal, 1961). Specimens: CNHE.

Notes: Originally recorded as Inermicapsifer tetragona (Molin, 1858).

Dilepididae Railliet and Henry, 1909

19. Amoebotaenia sp.

Intestine, Gallus gallus (B), NA (Brenes-Madrigal 1961).

20. A. cuneata (Linstow, 1872).

Intestine, Gallus gallus (B), NA (Brenes-Madrigal 1961).

21. Anomotaenia sp.

Intestine, Turdus grayi (B), NA (Brenes-Madrigal 1961).

22. Choanotaenia sp.

Intestine, Gallus gallus (B), NA (Brenes-Madrigal 1961).

23. C. infundibulum (Bloch, 1799) Railliet, 1896.

Intestine, Gallus gallus (B), NA (Brenes-Madrigal 1961).

24. Paricterotaenia sp.

Intestine, Sturnaella magna (B), Canton Cartago, Cartago (NP), Oriolus oriolus (B), San Jose San Jose (NP). Specimens: USNPC.

Dipylidiidae Stiles, 1896

25. Dipylidium caninum (Linneo, 1758) Leuckart, 1863.

Intestine, Canis familiaris, Felis catus (M), NA (Brenes-Madrigal 1961); C. familiaris, San Jose, San Jose (Vargas and Contreras 1998).

Hymenolepididae Railliet and Henry, 1909

26. Aploparaksis sp.

Intestine, Turdus grayi (B), NA (Brenes-Madrigal 1961).

Notes: Recorded as Haploparaxis sp.

27. Diorchis acuminatus (Clerc, 1902) Clerc, 1903.

Intestine, Undetermined host (B), NA (Brenes-Madrigal 1961).

Notes: The original distribution range of this species is in Europe (Schmidt 1986).

28. Hymenolepis sp.

Intestine, Gallus gallus, Gymnostinops montezuma, Ostinops quatemozinus (B), San Jose San Jose, (NP). Specimens: USNPC.

29. H. cantaniana Polonio, 1860.

Notes: This species is listed in Brenes-Madrigal (1961), but no information is provided.

30. H. diminuta (Rudolphi, 1819) Weinland, 1858.

Intestine, Rattus norvegicus (M), San Jose, San Jose (Vives and Zeledon 1957); Sigmodon hispidus (M), Canas, Guanacaste (Rodriguez-Ortiz et al. 2000).

31. Vampirolepis nana (Von Siebold, 1852) Spasskii, 1954.

Intestine, Rattus norvegicus (M), San Jose, San Jose (Vives and Zeledon 1957).

Lacistorhynchidae Guiart, 1927

32. Callitetrarhynchus gracilis (Rudolphi, 1819) Pintner, 1931.

Spiral valve, Carcharhinus leucas (F), Barra del Colorado, Rio Colorado, Limon (Watson and Thorson 1976). Specimens: USNPC.

Onchobothriidae Braun, 1900

33. Acanthobothrium campbelli Marques, Brooks and Monks, 1995.

Spiral valve, Urotrygon chilensis (F), Costa de Pajaros, Puntarenas (Marques et al. 1995). Specimens: HWML, MNHG.

34. A. cimari Marques, Brooks and Monks, 1995.

Spiral valve, Dasyatis longus (F), Punta Morales, Golfo de Nicoya, Puntarenas (Marques et al. 1995). Specimens: HWML, MNHG.

35. A. costarricense Marques, Brooks and Monks, 1995.

Spiral valve, Dasyatis longus (F), Punta Morales, Puntarenas (Marques et al. 1995). Specimens: HWML, MNHG.

36. A. franus Marques, Centritto and Stewart, 1997.

Spiral valve, Narcine entemedor (F), Playa Cuajiniquil, Golfo de Santa Elena, Guanacaste (Marques et al. 1997). Specimens: CHIOC, CNHE, USNPC.

37. A. inbiorium Marques, Centritto and Stewart, 1997.

Spiral valve, Narcine entemedor (F), Playa Cuajiniquil, Golfo de Santa Elena, Guanacaste (Marques et al. 1997).

Specimens: CHIOC, CNHE, USNPC.

38. A. nicoyaense Brooks and McCorquodale, 1995.

Spiral valve, Aetobatis narinari (F), Punta Morales, Golfo de Nicoya, Puntarenas (Brooks and McCorquodale 1995).

Specimens: HWML, MNHG.

39. A. puntarenasense Marques, Brooks and Monks, 1995.

Spiral valve, Dasyatis longus (F), Punta Morales, Golfo de Nicoya, Puntarenas (Marques et al. 1995). Specimens: HWML, MNHG.

40. A. vargasi Marques, Brooks and Monks, 1995.

Spiral valve, Dasyatis longus (F), Punta Morales, Golfo de Nicoya, Puntarenas (Marques et al. 1995). Specimens: HWML, MNHG.

41. Acanthobothroides pacificus Marques, Brooks and Molina-Urena, 1996.

Spiral valve, Himantura pacifica (F), Playa Panama, Guanacaste (Marques et al. 1996). Specimens: HWML, MNHG.

42. Phoreiobothrium triloculatum Linton, 1901.

Spiral valve, Carcharhinus leucas (F), Barra del Colorado, Rio Colorado, Limon (Watson and Thorson 1976). Specimens: USNPC.

43. Platybothrium hypoprioni Potter, 1927. Spiral valve, Carcharhinus leucas (F), Barra del Colorado, Rio Colorado, Limon (Watson and Thorson 1976).

Otobothriidae Dollfus, 1942

44. Othobothrium penetrans Linton, 1907. Spiral valve, Carcharhinus leucas (F), Barra del Colorado, Rio Colorado, Limon (Watson and Thorson 1976). Specimens: USNPC.

Phyllobothriidae Braun, 1900

45. Anthobothrium laciniatum Linton, 1890. Spiral valve, Carcharhinus leucas (F), Barra del Colorado, Rio Colorado, Limon (Watson and Thorson 1976).

46. A. pristis Woodland, 1934.

Spiral valve, Pristis perotteti (F), Barra del Colorado, Rio Colorado, Limon (Watson and Thorson 1976).

47. Phyllobothrium leuci Watson and Thorson, 1976.

Spiral valve, Carcharhinus leucas (F), Barra del Colorado, Rio Colorado, Limon (Watson and Thorson 1976).

48. P. nicaraguensis Watson and Thorson, 1976.

Spiral valve, Carcharhinus leucas (F), Barra del Colorado, Rio Colorado, Limon (Watson and Thorson 1976).

49. P. pristis Watson and Thorson, 1976. Spiral valve, Carcharhinus leucas (F), Barra del Colorado, Rio Colorado, Limon (Watson and Thorson 1976).

50. Rhinebothrium geminum Marques, Brooks and Molina-Urena, 1996. Spiral valve, Himantura pacifica (F), Playa Panama, Guanacaste (Marques et al. 1996). Specimens: HWML, MNHG.

Proteocephalidae La Rue, 1911

51. Ophiotaenia sp. Intestine, Constrictor constrictor (R), San Jose, San Jose (Brenes-Madrigal 1961). Specimens: USNPC.

52. O. perspicua La Rue, 1911. Intestine, Bothrops sp., (R), Muelle San Carlos, Alajuela (Flores-Barroeta et al. 1961).

Pterobothriidae Pintner, 1931

53. Pterobothrioides carvajali Campbell and Beveridge, 1997.

Spiral valve, Dasyatis longus (F), Playa Cuajiniquil, Golfo de Santa Elena, Guanacaste (Campbell and Beveridge 1997). Specimens: USNPC.

Taeniidae Ludwig, 1886

54. Echinococcus granulosus (Batsch, 1786) Rudolphi, 1810 (hydatidid cyst).

Lungs, Bos taurus (M), Los Cartagos, Alajuela (Velazquez and Morales 1983).

55. E. oligarthrus Diesing, 1863. Intestine, Felis concolor costaricensis (M), Talamanca, San Jose (Brenes-Madrigal et al. 1973).

56. Taenia sp.

Intestine, Agouti paca (M), NA (Matamoros et al. 1991).

57. T. omissa Luhe, 1910 (cysticerci). Lungs, Odocoileus virginianus (M), Parque Nacional Santa Rosa, Guanacaste (Carreno et al. 2001). Specimens: USNPC.

58. T. pisiformis (Bloch, 1780) Gmelin, 1790. Intestine, Urocyon cinereoargenteus (M), Carrizal, Alajuela (Flores-Barroeta et al. 1958b).

Notes: This species was originally recorded as Taenia laticollis Rudolphi, 1819.

59. T. saginata Goeze, 1782 (cysticerci). Muscle, Bos taurus (M), Cartago and Guanacaste (Brenes-Madrigal 1961).

60. T. solium Linneo, 1758 (Cysticerci). Muscle, Sus scrofa (M), NA (Brenes-Madrigal 1961).

61. T. taeniaeformis Batsch, 1796 (adult and cysticerci).

Liver, Rattus norvegicus, R. rattus (M), San Jose, San Jose (Vives and Zeledon 1957; Intestine, Felis wiedii nicaraguense (M), Mata de Limon, Puntarenas (Flores-Barroeta et al. 1958b), F. catus (M), San Jose, San Jose (Brenes-Madrigal 1961).

Tentaculariidae Poche, 1926

62. Nybelinia bisulcata (Linton, 1889) Poche, 1926.

Spiral valve, Carcharhinus leucas (F), Barra del Colorado, Rio Colorado, Limon (Watson and Thorson 1976).

Trilocularidae Yamaguti, 1959

63. Escherbothrium molinae Berman and Brooks, 1994.

Spiral valve, Urotrygon chilensis (F), Costa de Pajaros and Punta Morales, Golfo de Nicoya, Puntarenas (Berman and Brooks 1994).

ACANTHOCEPHALA

(Rudolphi, 1808)

Centrorhynchidae Van Cleave, 1916

1. Centrorhynchus sp.

NA, Mustela noveboracensis (M), San Jose, San Jose (N-P). Specimens: USNPC.

2. C. tumidulus (Rudolphi, 1819) Travassos, 1923 (cysthacanth).

Intestine, Rana sp. (A), NA (Brenes-Madrigal 1961).

Echinorhynchidae Cobbold, 1876

3. Acanthocephalus lutzi (Von Linstow, 1896) Meyer, 1931 (cysthacanth).

NA, Leptodactylus gracilis (A), San Jose, San Jose (NP). Specimens: USNPC.

Illiosentidae Golvan, 1960

4. Koronacantha pectinaria (Van Cleave, 1940).

Intestine, "Medialuna" (F), Bahia Culebra, Guanacaste (Van Cleave 1940), Microlepidotus brevipinnis (F), Playas Panama and Hermosa, and Golfo de Santa Elena, Playa Cuajiniquil, Guanacaste (Monks et al. 1997). Specimens: CNHE, HWML, USNPC.

Notes: Van Cleave (1940) described this species as Tegorhynchus Van Cleave, 1921, but was later transferred to Koronacantha Monks and Perez-Ponce de Leon, 1996 by Monks et al. (1997).

Moniliformidae Van Cleave, 1924

5. Moniliformis moniliformis (Bremser, 1811) Travassos, 1915.

Intestine, Rattus norvegicus, R. rattus (M), San Jose, San Jose (Vives and Zeledon 1957). Specimens: CNHE, USNPC.

Oligacanthorhynchidae

Southwell and MacFie, 1925

6. Oligacanthorhynchidae gen. sp.

Subcutaneous tissue, Tonatia silvicola (M), Peninsula de Osa, Puntarenas (Gibson and McCarthy 1987).

7. Oncicola oncicola (Ihering, 1892) Travassos, 1916.

Muscle, Gallus gallus (B), Parrita, Puntarenas (Zeledon and Arroyo-Sancho 1960).

8. Prosthenorchis sp.

Intestine, Saimiri oerstedi (M), Parque Nacional Corcovado (Appleton and Boinski 1991).

9. P. elegans (Diesing, 1851) Travassos, 1915

Intestine, Cebus capucinus (M), NA (NP). Specimens: USNPC.

Plagiorhynchidae Golvan, 1960

10. Plagiorhynchus angrense (Travassos, 1926) Schmidt and Kuntz, 1966.

Intestine, Mustela frenata (M), NA (Brenes-Madrigal 1961).

Notes: In accordance with Brenes-Madrigal (1961), this record is doubtful because this species has been only recorded in birds. In Costa Rica it was recorded as Prosthorhynchus angrensis Travassos, 1926.

Polymorphidae Meyer, 1931

11. Polymorphus corynosoma Travassos, 1915

Intestine, Mustela frenata (M), NA (Brenes-Madrigal 1961).

Notes: Recorded as Hexaglandula corynosoma (Travassos, 1915). According to Amin (1992), it is a typical parasite of herons.

12. P. minutus (Goeze, 1782) Luhe, 1911. Intestine, "birds", NA (Brenes-Madrigal 1961).

NEMATODA

(Rudolphi, 1808)

Ancylostomatidae (Looss, 1905)

1. Ancylostoma braziliense Gomes de Faria, 1910.

Intestine, Potos flavus arborensis (M), NA (Brenes-Madrigal 1961).

2. A. caninum (Ercolani, 1859).

Intestine, Canis familiaris (M), San Jose, San Jose (Brenes-Madrigal 1961, Vargas and Contreras 1998), Canis latrans (M), and Felis catus (M), San Jose, San Jose (Brenes-Madrigal 1961). Specimens: USNPC.

3. Bunostomum phlebotomum (Railliet in Rizzo, 1900).

Intestine, Bos taurus (M), San Isidro de Coronado, San Jose (Brenes-Madrigal 1961). Specimens: USNPC.

4. B. trigonocephalum (Rudolphi, 1808). Intestine, Capra hircus (M), Sabanilla de Montes de Oca, San Jose (Brenes-Madrigal 1961).

5. Gaigeria pachyscelis Railliet and Henry, 1910.

Intestine, Bos taurus (M), Turrialba, Cartago and Capra hircus (M), Sabanilla de Montes de Oca, San Jose (Brenes-Madrigal 1961).

6. Globocephalus urosubulatus (Alessandrini, 1909).

Intestine, Sus scrofa (M), Puriscal and Santa Ana, San Jose (Brenes-Madrigal 1961).

7. Monodontus sp.

Intestine, Sigmodon hispidus (M), Canas, Guanacaste (Rodriguez-Ortiz et al. 2000).

Angiostrongylidae

(Boehm and Gebauer, 1934)

8. Angiostrongylus costaricensis Morera and Cespedes, 1971

Mesenteric arteries, Ateles geoffroyi (M), NA (Berrocal et al. 1997), Nasua narica (M), NA (Monge et al. 1978), Sigmodon hispidus (M), San Rafael, Alajuela (Rodriguez-Ortiz et al. 1999, 2000)

Notes: Chabaud (1972) included this species in the genus Parastrongylus Baylis, 1928, which is considered a subgenus of Angiostrongylus Kamensky, 1905 by Anderson (1978).

Ascarididae Baird, 1853

9. Ascaris lumbricoides Linneo, 1758. Intestine, Alouatta palliata palliata (M), La Pacifica, Guanacaste (Stuart et al. 1990), Canis familiaris (M), NA (Brenes-Madrigal 1961).

10. A. suum Goeze, 1782.

Intestine, Sus scrofa (M), Matadero Municipal, San Jose (Brenes-Madrigal 1961).

11. Lagochilascaris sp.

Larynx, Felis pardalis mearnsi (M), San Vito de Java, Puntarenas (Brenes-Madrigal et al. 1972).

12. Neoascaris vitulorum (Goeze, 1782).

Intestine, Bos taurus (M), Turrialba, Cartago (Brenes-Madrigal 1961).

Notes: According to Hartwich (1974), Neoascaris Travassos, 1927 is a synonym of Toxocara Stiles, 1905.

13. Ophidascaris sp.

NA, Constrictor constrictor (R), NA (Brenes-Madrigal 1961).

14. Parascaris equorum (Goeze, 1782).

Intestine, Equus caballus (M), NA, Limon (Brenes-Madrigal 1961).

15. Porrocaecum sp. (adult and larvae). NA, "goose" (B), and "cobra imperator" (R), San Jose, San Jose (NP). Specimens: USNPC.

16. P. angusticolle (Molin, 1860).

Intestine, "Carraco" (B), NA (Brenes-Madrigal 1961).

17. Toxocara canis (Werner, 1782).

Intestine, Canis familiaris (M), San Jose, San Jose (Brenes-Madrigal 1961, Vargas and Contreras 1998).

18. T. mystax (Zeder, 1800)

Intestine, Felis catus (M), NA (Brenes-Madrigal 1961).

Ascaridiidae Travassos, 1919

19. Ascaridia sp.

Intestine, Columbidae (B), San Jose (NP). Specimens: USNPC.

20. A. columbae (Gmelin, 1790).

Intestine, Columba fasciata (B), NA (Brenes-Madrigal 1961).

21. A. galli (Schrank, 1788) Freeborn, 1923.

Intestine, "duck", Gallus gallus, Meleagris gallopavo (B), NA (Brenes-Madrigal 1961).

Atractidae (Railliet, 1917) Travassos, 1919

22. Atractis caballeroi Brenes-Madrigal and Bravo-Hollis, 1960.

Intestine, Kinosternon cruentatum (R), Santa Ana, San Jose (Brenes-Madrigal and Bravo-Hollis 1960). Specimens: CHCR, CNHE.

Notes: Bursey and Flanagan (2002) transferred this species tothe genus Klossinemella Costa, 1961.

23. A. impura Caballero, 1944.

Intestine, Kinosternon cruentatum (R), Santa Ana, San Jose (Brenes-Madrigal and Bravo-Hollis 1960). Specimens: CHCR, CNHE.

Notes: Bursey and Flanagan (2002) recognized this species as valid.

24. A. longicaudatum (Brenes-Madrigal and Bravo-Hollis, 1960)Baker, 1987.

Intestine, Ctenosaura similis (R), Liberia, Guanacaste and Mata de Limon, Puntarenas (Brenes-Madrigal and Bravo-Hollis 1960). Specimens: CHCR, CNHE.

Notes: Baker (1987) synonymized the genus Cyrtosomum Gedoelst, 1919 with Atractis Dujardin, 1985, transferring to this the species described by Brenes-Madrigal and Bravo-Hollis (1960). Bursey and Flanagan (2002) re-established the genus Cyrtosomum as valid, including in this genus Cyrtosomum longicaudatum Brenes-Madrigal and Bravo-Hollis, 1960.

Chabertiidae (Popova, 1952)

Lichtenfels, 1980

25. Oesophagostomum radiatum (Rudolphi, 1803).

Colon and intestine, Bos taurus (M), San Pedro Montes de Oca, San Jose and Turrialba, Cartago (Brenes-Madrigal 1961). Specimens: USNPC.

26. O. venulosum (Rudolphi, 1809). Intestine, Capra hircus, "sheep" and "rats" (M), NA (Brenes-Madrigal 1961). Note: This species is not a common parasite of rodents.

Cosmocercidae (Railliet, 1916) Travassos, 1925

27. Aplectana itzocanensis Bravo-Hollis, 1943.

Intestine, Bufo marinus (A), San Jose, San Jose (Brenes-Madrigal and Bravo-Hollis 1959). Specimens: CHCR, CNHE.

Notes: The original record was made as Oxysomatium itzocanensis (Bravo-Hollis, 1943). According with Baker (1985) and Goldberg et al. (1998), the specimens may belong to Aplectana incerta Caballero, 1949.

28. Oxysomatium sp.

Intestine, Constrictor constrictor and "culebra coral" (R), NA (Brenes-Madrigal 1961).

Diaphanocephalidae Travassos, 1920

29. Kalicephalus sp.

Intestine, "cobra imperator" (R), San Jose, San Jose (NP). Specimens: USNPC.

30. K. inermis macrovulvus (Caballero, 1954)

Intestine, Bothrops atrox (R), Hospital San Juan de Dios, San Jose (Brenes-Madrigal and Bravo-Hollis 1960). Specimens: CHCR, CNHE.

Notes: Species described as Kalicephalus macrovulvus Caballero, 1954 and considered as a subspecies of Kalicephalus inermis Molin, 1861 by Schad (1962).

31. K. subulatus Molin, 1861

Intestine, Constrictor constrictor (R), San Juan, San Jose (NP). Specimens: USNPC.

Dictyocaulidae (Skrjabin, 1933)

32. Dictyocaulus sp.

Lungs, Capra hircus and Bos taurus (M), NA (Brenes-Madrigal 1961). Specimens: USNPC.

33. D. arnfieldi (Cobbold, 1884)

Lungs, Equus caballus (M), NA, Limon (Brenes-Madrigal 1961).

34. D. eckerti Skrjabin, 1931.

Lungs, Bos taurus (M), Turrialba, Cartago (Brenes-Madrigal 1961).

35. D. viviparus (Bloch, 1782).

Lungs, Bos taurus (M), Irazu and Paraiso-Orosi, Cartago (Echandi-Quiros 1964).

Diplotriaenidae Skrjabin, 1916

36. Diplotriaena muscisaxicolae

Schuurmans-Stekhoven, 1952.

Aerial sacs, Nesotriccus ridgwayi (B), Isla Cocos, Puntarenas (Anderson et al. 1980). Specimens: USNPC.

Gnathostomatidae Lane, 1923

37. Echinocephalus jenzeni Hoberg, Brooks, Molina-Urena and Erbe, 1998.

Spiral valve, Himantura pacifica (F), Playa Panama, Guanacaste (Hoberg et al. 1998). Specimens: USNPC.

38. Spiroxys sp.

Intestine, Geomyda manni (R), San Juan, San Jose (Brenes-Madrigal 1961). Specimens: USNPC.

Gongylonematidae (Hall, 1916)

39. Gongylonema neoplasticum (Fibiger and Ditlevsen, 1914)

Stomach, Rattus novergicus (M), San Jose, San Jose (Vives and Zeledon 1957).

40. G. pulchrum Molin, 1857

Esophagus, Odocoileus virginianus (M), Parque Nacional Santa Rosa, Guanacaste (Carreno et al. 2001). Specimens: USNPC.

Habronematidae (Chitwood and Wehr, 1932)

41. Habronema megastoma (Rudolphi, 1819) Seurat, 1914.

Stomach, Equus caballus (M), NA (Brenes-Madrigal 1961).

42. H. microstoma ((Schneider, 1866).

Stomach, Equus caballus (M), San Pedro Montes de Oca, San Jose (Brenes-Madrigal 1961). Specimens: USNPC.

43. Parabronema sp.

Intestine, Alouatta palliata (M), Reserva Biologica La Selva (Stoner 1996).

Heligmonellidae

(Skrjabin and Schikhobalova, 1952)

44. Hassalstrongylus aduncus (Chandler, 1933) Durette-Desset, 1972.

Intestine, Sigmodon hispidus (M), San Rafael Ojo de Agua, Alajuela (Rodriguez-Ortiz et al. 1999, 2000), Canas, Guanacaste (Rodriguez-Ortiz et al. 2000).

45. Nippostrongylus brasiliensis (Travassos, 1914).

Intestine, Rattus novergicus (M), San Jose, San Jose (Vives and Zeledon 1957).

Heteroxynematidae

(Skrjabin and Schikhobalova, 1948)

46. Aspiculuris tetraptera (Nitzsch, 1821).

Intestine, Rattus rattus (M), Escazu, San Jose (Vives and Zeledon 1957).

Kathlaniidae Lane, 1914)

Travassos, 1918

47. Cruzta sp.

NA, "cobra imperator" (R), San Jose, San Jose (NP). Specimens: USNPC.

Metastrongylidae Leiper, 1908

48. Metastrongylus elongatus (Dujardin, 1845).

Trachea, Bronchial tube, Sus scrofa (M), NA (Brenes-Madrigal 1961).

Molineidae (Skrjabin and Schulz, 1937)

49. Nematodirus sp.

NA, Equus caballus (M), NA (Marin 1955).

Notes: According to Brenes-Madrigal (1961) the record is doubtful because the genus is not parasite of equines.

50. Oswaldocruzia subauricularis (Rudolphi, 1819) Travassos, 1917.

Intestine, Bufo marinus (M), Tilaran, Guanacaste and Paso Ancho, San Jose (Brenes-Madrigal and Bravo-Hollis 1959). Specimens: CHCR, CNHE.

Onchocercidae (Leiper, 1911)

51. Dipetalonema gracile (Rudolphi, 1809) Diesing, 1861.

Mesentery, Cebus capucinus (M), NA (NP). Specimens: USNPC.

52. Dirofilaria immitis (Leidy, 1856) Railliet and Henry, 1911 (adults and larvae).

Intestine and heart, Canis familiaris (M), NA, Heredia (Sancho et al. 1989); San Jose, San Jose (Leuterer and Gothe 1993, Vargas and Contreras 1998).

53. Foleyella sp. Blood, Rana forreri (A), Area de cosnservacion de Guanacaste, Guanacste (Desser 2001). Specimens: CMNPA.

54. Litomosoides carinii (Travassos, 1911). Body cavity, Sigmodon hispidus (M), Canas, Guanacaste (Rodriguez-Ortiz et al. 2000).

55. L. penai Jimenez-Quiros and Arroyo-Sancho, 1960. Body cavity, Hemiderma perspicillata (M), Santa Ana, San Jose (Jimenez-Quiros and Arroyo-Sancho 1960a). Specimens: CHCR.

56. Ochoterenella sp. Blood, Bufo marinus (A), Area de cosnservacion de Guanacaste, Guanacste (Desser 2001). Specimens: CMNPA.

57. O. caballeroi Esslinger, 1987. Mesentery, Bufo marinus (A), San Jose, San Jose (Esslinger 1987). Specimens: CNHE. Notes: Original identification: Ochoterenella digiticauda Caballero, 1944.

58. O. digiticauda Caballero, 1944 (adults and larvae). Mesentery, Bufo marinus (A), Santa Ana, San Jose (Brenes-Madrigal and Bravo-Hollis 1959). Specimens: CHCR, CNHE. Notes: Some of these specimens, identified as Ochoterenella digiticauda, were used by Esslinger (1987) to describe O. caballeroi.

59. Onchocerca cervicalis Railliet and Henry, 1910. Cervical ligament, Equus caballus (M), Esparza, Puntarenas (Velazquez et al. 1983).

60. O. cervipedis Wehr and Dikmans, 1935. Dermis, Odocoileus virginianus (M), Parque Nacional Santa Rosa, Guanacaste (Carreno et al. 2001). Specimens: USNPC.

61. Oswaldofilaria brevicaudata (Rodhain and Vuylsteke, 1937). Dermis and blood, Ctenosaura similis (R), Reserva Natural de Guanacaste, Guanacaste (Desser 1997).

62. Setaria cervi (Rudolphi, 1819). Body cavity, Bos taurus (M), NA (Brenes-Madrigal 1961).

63. S. equina (Abildgaard, 1789). Body cavity, eyes, scrotum, mesentery and pleura, Equus caballus (M), NA (Brenes-Madrigal 1961).

64. S. labiatopapillosa (Aless., 1838). Body cavity, Bos taurus (M), San Jose, San Jose (NP). Specimens: USNPC.

65. S. yehi (Durette-Desset, 1966). Rectum, Odocoileus virginianus (M), Parque Nacional Santa Rosa, Guanacaste (Carreno et al. 2001). Specimens: USNPC.

Oxyuridae Cobbold, 1864

66. Oxyuris sp. Intestine, Ateles geoffroyi (M), NA (Berrocal et al. 1997).

67. Oxuyris equi (Schrank, 1788). Intestine, Equus caballus (M), San Pedro Montes de Oca, San Jose (NP), NA, Limon(Brenes-Madrigal 1961). Specimens: USNPC. Notes: Specimens deposited at the USNPC were determined as Oxyuris curvula Rudolphi, 1803.

68. Skrjabinema sp. Intestine, Capra hircus (M), NA (Alvarado et al. 1990).

69. S. ovis (Skrjabin, 1915) Wereschtchagin, 1926. Intestine, Capra hircus (M), NA (Velazquez and Solis 1983). Notes: This species was found in a goat, which had been imported into Costa Rica from the U.S.A.

70. Trypanoxyuris sp. Intestine, Ateles geoffroyi (M), Parque Nacional Santa Rosa, Guanacaste (Stuart et al. 1990).

71. T. minutus (Schneider, 1866) Intestine, Alouatta palliata palliata (M), La Pacifica, Guanacaste (Stuart et al. 1990).

Pharyngodonidae Travassos, 1919

72. Ozolaimus cirratus (von Linstow, 1906) Railliet and Henry, 1912. Intestine, Ctenosaura similis (R), Liberia, Guanacaste and Mata de Limon, Puntarenas (Bravo-Hollis and Brenes-Madrigal 1959). Specimens: CHCR, CNHE.

73. O. ctenosauri Caballero, 1938. Intestine, Ctenosaura similis (R), Liberia, Guanacaste and Mata de Limon, Puntarenas (Bravo-Hollis and Brenes-Madrigal 1959). Specimens: CHCR, CNHE. Notes: Originally described as Ozolaimus monhystera (Von Linstow, 1902) and redetermined as O. ctenosauri by Moravec et al. (1996).

74. O. megatyphlon (Rudolphi, 1819) Dujardin, 1845. Intestine, Ctenosaura similis (R), Liberia, Guanacaste and Mata de Limon, Puntarenas (Bravo-Hollis and Brenes-Madrigal 1959). Specimens: CHCR, CNHE.

75. Spauligodon oxkutcabiensis (Chitwood, 1938). Intestine, Sceloporus malachiticus (R), San Jose, San Jose (Goldberg and Bursey 1992). Specimens: USNPC.

Physalopteridae (Railliet, 1893) Leiper, 1908

76. Physalopteridae gen. sp. Intestine, Alouatta palliata (M), Reserva Biologica La Selva (Stoner 1996).

77. Physaloptera sp. (larvae). Stomach, Bothrops athrox (R), NA (Brenes-Madrigal 1961), Sceloporus malachitichus (R), Volcan Irazu, Cartago (Goldberg and Bursey 1992). Specimens: USNPC.

Protostrongylidae Leiper, 1926

78. Parelaphostrongylus tenuis (Dougherty, 1945). Brain and craneal sinus, Odocoileus virginianus (M), Parque Nacional Santa Rosa, Guanacaste (Carreno et al. 2001). Specimens: USNPC.

Rhabdiasidae Railliet, 1915

79. Rhabdias bufonis (Schrank, 1788) Stiles and Hassal, 1905. Lungs, Bufo marinus (A), San Jose, San Jose (NP). Specimens: USNPC. Notes: According to Baker (1987), this species is only found in Europe.

80. R. sphaerocephala Goodey, 1924. Lungs, Bufo marinus (A), Rio Tiribi, San Jose and La Pena, Alajuela (Brenes-Madrigal and Bravo-Hollis 1959), NA (Colam, 1971). Specimens: CHCR, CNHE. Notes: According to Baker (1987) records from the America's needs confirmation.

Spirocercidae (Chitwood and Wehr, 1932)

81. Ascarops sp. (larvae). Stomach, Sceloporus malachiticus (R), San Jose, San Jose, San Ramon de Tres Rios and Volcan Irazu, Cartago (Goldberg and Bursey 1992). Specimens: USNPC.

82. Leiurus leptocephalus (Rudolphi, 1819) Leuckart, 1850. Intestine, Bradypus griseus (M), Turrialba, Cartago (Jimenez-Quiros and Brenes-Madrigal 1956). Specimens: CHCR, CNHE.

83. Mastophorus muris (Gmelin, 1790) Chitwood, 1938. Intestine, Rattus novergicus (M), San Jose, San Jose (Campos and Vargas 1978).

84. Physocephalus sexalatus (Molin, 1860) Diesing, 1861. Intestine, Sus scrofa (M), NA (Brenes-Madrigal 1961).

85. Spirocerca lupi (Rudolphi, 1809) Esophagus, Canis familiaris (M), NA, Heredia (Sancho et al. 1989).

Spiruridae Oerley, 1885

86. Protospirura sp. Intestine, Sigmodon hispidus (M), Canas, Guanacaste (Rodriguez-Ortiz et al. 2000).

87. P. muris (Gmelin, 1790). Intestine, Rattus novergicus (M), San Jose, San Jose (Vives and Zeledon 1957, Campos and Vargas 1978).

Strongylidae Baird, 1853

88. Cylicocyclus sp. Intestine, Bos taurus (M), San Pedro Montes de Oca, San Jose (NP). Specimens: USNPC.

89. C. insigne Boulenger, 1917. Colon and intestinal caeca, Equus caballus (M), San Pedro Montes de Oca, San Jose (NP). Specimens: USNPC.

90. Eucyathostomum webbi Pursglove, 1976 Intestine, Odocoileus virginianus (M), Parque Nacional Santa Rosa, Guanacaste (Carreno et al. 2001). Specimens: USNPC.

91. Strongylus edentatus (Looss, 1900) Colon and intestinal caeca, Equus caballus (M), NA (Brenes-Madrigal 1961). Specimens: USNPC. Notes: Originally recorded as Alfortia edentatus (Looss, 1900) Skrjabin, 1933.

92. S. equinus Mueller, 1780 Colon and intestinal caeca, Equus caballus (M), NA, Limon (Brenes-Madrigal 1961).

93. S. vulgaris (Looss, 1900). Colon and intestinal caeca, Equus caballus (M), NA (Brenes-Madrigal 1961). Notes: Recorded as Delafondia vulgaris (Looss, 1900) Skrjabin, 1933.

94. Triodontophorus tenuicollis Boulenger, 1916. Intestinal caeca, Equus caballu (M), San Pedro Montes de Oca, San Jose (NP). Specimens: USNPC.

Strongyloididae Chitwood and McIntosh, 1934

95. Strongyloides sp. Intestine, Agouti paca (M), Cantones Atenas, Central, Poas, San Carlos and Vega, Alajuela, Cantones Cartago and El Guarco, Cartago, Canton Central, Heredia, Canton Matina, Limon, Canton Osa, Puntarenas, Cantones Central, Santa Ana and Terrazu, San Jose (Matamoros et al. 1991); Capra hircus (M), NA (Alvarado et al. 1990).

96. S. cebus Darling, 1911 Intestine, Alouatta palliata, Ateles geoffroyi and Cebus capucinus (M), Curu, Puntarenas, Los Inocentes, La Pacifica and Parque Nacional Santa Rosa, Guanacaste (Stuart et al. 1998).

97. S. ratti Sandground, 1925 Intestine, Rattus novergicus (M), San Jose, San Jose (Vives and Zeledon 1957).

98. S. sigmodontis Melvin and Chandler, 1950 Intestine, Sigmodon hispidus (M), Canas, Guanacaste (Rodriguez-Ortiz et al. 2000).

Syngamidae Leiper, 1912

99. Mammomonogamus laryngeus (Railliet, 1899) Ryzhikov, 1948 Larynx, Bos taurus (M), NA (Cordero et al. 1981).

100. Stephanurus dentatus Diesing, 1839 Liver and kidney, Bos taurus, Sus scrofa (M), (Brenes-Madrigal 1961).

101. Syngamus sp. Larynx, Felis pardalis mearnsi (M), San Vito de Java, Puntarenas (Brenes-Madrigal et al. 1972).

102. S. trachea (Montagu, 1811) Trachea, Gallus gallus and Meleagris gallopavo (B), NA (Brenes-Madrigal 1961).

Tetrameridae Travassos, 1914

103. Tetrameres fissipina (Diesing, 1861) Intestine, Gallus gallus (B), Tres Rios, Cartago (Brenes-Madrigal et al. 1962). Specimens: CHCR.

Thelaziidae Skrjabin, 1915

104. Oxyspirura mansoni (Cobbold, 1879) Intestine, Gallus gallus (B), Turrialba, Cartago (Brenes-Madrigal et al. 1962). Specimens: USNPC.

105. Thelazia sp. Stomach, Constrictor constrictor (R), NA (Brenes-Madrigal 1961). Notes: Members of Thelazia are common parasites in eyes of mammals or birds so the record is doubtful in accordance with Brenes-Madrigal (1961).

Trichostrongylidae (Leiper, 1908) Leiper, 1912

106. Ashworthius patriciapillitae Hoberg, Abrams, Carreno and Lichtenfels, 2002 Abomasum, Odocoileus virginianus (M), Parque Nacional Santa Rosa, Guanacaste (Carreno et al. 2001, Hoberg et al. 2002). Specimens: USNPC. Notes: Describes as Ashworthius sp. by Carreno et al. (2001).

107. Cooperia sp. Intestine, Bos taurus (M), Turrialba, Cartago (Brenes-Madrigal 1961), Capra hircus (M), NA (Alvarado et al. 1990).

108. Haemonchus sp. Abomasum, Capra hircus (M), NA (Alvarado et al. 1990).

109. H. contortus (Rudolphi, 1803) Abomasum, Bos taurus (M), Sabanilla de Montes de Oca, San Jose and Capra hircus (M), Turrialba, Cartago (Brenes-Madrigal 1961).

110. Mecistocirrus sp. Bos taurus (M), San Pedro Montes de Oca, San Jose (NP). Specimens: USNPC.

111. M. digitatus (Von Linstow, 1906) Abomasum, Bos taurus (M), Turrialba, Cartago (Brenes-Madrigal 1961), Heredia (Porras 1987), San Jose and San Pedro Montes de Oca, San Jose (NP). Specimens: USNPC.

112. Ostertagia sp. Abomasum and intestine, Bos taurus (M), Turrialba, Cartago and Capra hircus (M), Sabanilla de Montes de Oca, San Jose (Brenes-Madrigal 1961).

113. Mazamastrongylus sp. Abomasum, Odocoileus virginianus (M), parque Nacional Santa Rosa, Guanacaste (Carreno et al. 2001). Specimens: USNPC. Notes: According to Durette-Desset (1983), the genus Mazamostrongylus Cameron, 1935 is synonym of Spiculopteragia (Orloff, 1933).

114. Trichostrongylus sp. Intestine, Bos taurus (M), Turrialba, Cartago (Brenes-Madrigal 1961), Sigmodon hispidus (M), San Rafael Ojo de Agua, Alajuela (Rodriguez-Ortiz et al. 1999).

115. T. sigmodontis Baylis, 1945 Intestine, Sigmodon hispidus (M), Canas, Guanacaste (Rodriguez-Ortiz et al. 2000).

Trichuridae (Ransom, 1911) Railliet, 1915

116. Anatrichosoma buccalis Pence and Little, 1972. Mouth, Didelphis marsupialis (M), Tres Rios, Cartago (Pence and Little 1972). Specimens: USNPC.

117. Capillaria sp. Intestine, Agouti paca (M), Canton Central, Heredia (Matamoros et al. 1991), Bos taurus (M), Santa Ana, San Jose (Brenes-Madrigal 1961), Kinosternon cruentatum (R), Santa Ana, San Jose (Brenes-Madrigal and Bravo-Hollis 1960). Specimens: CHCR, CNHE.

118. C. annulosa (Dujardin, 1845). Crop and oesophagus, NA (B), NA (Brenes-Madrigal 1961). Notes: Validity of this records needs confirmation since the species is parasite of rodents in Europe.

119. C. obsignata Madsen, 1945. Intestine, Columba livia (B), NA (Brenes-Madrigal 1961), Gallus gallus (B), Guacimo, Limon (Brenes-Madrigal et al. 1962). Specimens: CHCR. Notes: The species was transferred to the genus Baruscapillaria Moravec, 1982 by Moravec (1982).

120. Trichosomoides crassicauda (Bellingham, 1840). Urinary bladder, Rattus novergicus (M), San Jose, San Jose (Vives and Zeledon 1957).

121. Trichuris sp. Intestine, Agouti paca (M), Canton Matina, Limon, Carnton Cartago, Cartago and Canton San Carlos, Alajuela (Matamoros et al. 1991).

122. T. ovis (Abildgaard, 1795) Intestine, Bos taurus (M), Sabanilla de Montes de Oca, San Jose and Turrialba, Cartago (Brenes-Madrigal 1961). Notes: Originally recorded as Trichocephalus ovis Abildgaard, 1795.

123. T. suis (Schrank, 1788) Colon and intestinal ceca, Sus scrofa (M), NA (Brenes-Madrigal 1961). Notes: Originally recorded as Trichocephalus suis Schrank, 1788.

124. T. vulpis (Froelich, 1789) Intestine, Canis familiaris (M), San Jose, San Jose (Brenes-Madrigal 1961, Vargas and Contreras 1998). Notes: Recorded as Trichocephalus vulpis Froelich, 1789 and Trichuris canis.
Parasite-Host List

Symbology: * new species. A = acanthocephalan; C = cestode;
D = digenean; M = monogenean; N = nematode.

                            Pisces

Host                          Helminth Species

Aetobatis narinari            Acanthobothrium nicoyaense * (C)
Arothron hispidus             Bianium simonei (D)
Astyanax aenaeus              Wallinia chavariiae (D)
Astyanax fasciatus            Anacanthocotyle anacanthocotyle * (M)
                              Gyrodactylus costaricensis (M)
                              Gyrodactylus neotropicalis * (M)
                              Jainus hexops * (M)
                              Palombitrema heteroancistrium * (M)
                              Urocleidoides costaricensis * (M)
                              Urocleidoides strombicirrus * (M)
Bryconamericus scleloparius   Wallinia chavarriae * (D)
Caranx caballus               Ahpua piscicola (M)
                              Bucephalopsis sp. (D)
                              Cemocotylella elongata (M)
                              Manteria brachydera (D)
                              Stephanostomum casum (D)
                              Tergestia laticollis (D)
Caranx hippos                 Bucephalus introversus (D)
Carcharhinus leucas           Anthobothrium laciniatum (C)
                              Callitetrarhynchus gracilis (C)
                              Cathetocephalus thatcheri (C)
                              Dasyrhynchus variouncinatus (C)
                              Dermoplithirius maccallumi (M)
                              Erpocotyle carcharhini (M)
                              Heteroncocotyle leucas (M)
                              Nybelinia bisulcata (C)
                              Otobothrium penetrans (C)
                              Phoreiobothrium triloculatum (C)
                              Phyllobothrium leuci (C)
                              Phyllobothrium nicaraguensis (C)
                              Phyllobothrium pristis (C)
                              Platybothrium hypoprioni (C)
Cichlasoma sp.                Crassicutis cichlasomae (D)
                              Crassicutis opisthoseminis * (D)
Dasyatis longus               Acanthobothrium cimari * (D)
                              Acanthobothrium costarricense * (C)
                              Acanthobothrium puntarenasense * (C)
                              Acanthobothrium vargasi * (C)
                              Pterobothrioides carvajali (C)
Echidna nocturna              Pseudolecithaster sp. (D)
                              Theletrum lamothei * (D)
Epinephelus itajara           Hypocreadium myohelicatum (D)
Fistularia commersonii        Ectenurus virgulus (D)
                              Lecithochirium microstomum (D
Haemulon maculicauda          Didymozoidae gen. sp. (D)
Himantura pacifica            Acanthobothroides pacificus * (C)
                              Echinocephalus jenzeni * (N)
                              Rhinebothrium geminum * (C)
Lutjanus colorado             Polymicrocotyle manteri (M)
Menticirrhus sp.              Hargicotyle louisianensis (M)
Microlepidotus brevipinnis    Koronacantha pectinaria (A)
Narcine entemedor             Acanthobothrium franus * (C)
                              Acanthobothrium inbiorium * (C)
Oligoplites altus             Ahpua piscicola (M)
                              Hargicola oligoplites (M)
                              Manteria brachydera (D)
                              Mecoderus oligoplites (D)
Oligoplites refulgens         Manteria brachydera (D)
                              Mecoderus oligoplitis (D)
Oligoplites sp.               Mecoderus oligoplitis (D)
Poecilia sphenops             Gyrodactylus bullatarudis (M)
                              Gyrodactylus costaricensis * (M)
Pristis perotteti             Anthobothrium pristis (C)
Rhamdia rogersi               Ameloblastella chavarriai * (M)
                              Aphanoblastella travassosi * (M)
                              Stunkardiella minima (D)
Seriola sp.                   Bucephalus introvecsus * (D)
Sphoeroides sp.               Bianium plicitum (D)
                              Heterobothrium ecuadori (M)
Synodus sp.                   Lecithochirium monticellii (D)
Urotrygon chilensis           Acanthobothrium campbelli * (C)
                              Escherbothrium molinae * (C)
Vomer declivifrons            Pseudomazocraes monsivaisae (M)
Xurel marginatus              Neomicrocotyle pacifica * (M)

                          Amphibians

Host                          Helminth Species

Bufo marinus                  Aplectana itzocanensis (N)
                              Glypthelmins hepaticus (D)
                              Glypthelmins palmipedis (D)
                              Gorgoderina megalorchis (D)
                              Langeronia provitellaria (D)
                              Mesocoelim travassosi (D)
                              Ochoterenella caballeroi (D)
                              Ochoterenella digiticauda (N)
                              Ochoterenella sp. (N)
                              Oswaldocruzia subauricularis (N)
                              Rhabdias bufonis (N)
                              Rhabdias sphaerocephala (N)
Leptodactylus gracilis        Acanthocephalus lutzi (A)
Rana forreri                  Foleyella sp. (N)
Rana palmipes                 Neohaematoloechus helibravoae * (D)
Rana pipiens                  Cephalogonimus americanus (D)
                              Glypthelmins facioi * (D)
                              Megalodiscus temperatus (D)
                              Pseudosonsinotrema chabaudi * (D)
Rana vaillanti                Haematoloechus meridionalis * (D)
                              Halipegus eschi * (D)
Rana warchewitschii           Langeronia provitellaria (D)
Rana sp.                      Centrorhynchus tumidulus (A)

                            Reptiles

Host                          Helminth Species

Basiliscus sp.                Parallopharynx gonzalezi * (D)
Bothrops atrox                Kalicephalus inermis (N)
                              Physaloptera sp. (N)
Bothrops sp.                  Ophiotaenia perspicua (C)
Caiman crocodilus             Prolecithodiplostomum constrictum (D)
                              Proterodiplostomum medusae (D)
Constrictor constrictor       Kalicephalus subulatus (N)
                              Ophidascaris sp. (N)
                              Ophiotaenia sp. (C)
                              Oxysomatium sp. (N)
                              Thelazia sp. (N)
Crocodylus acutus             Proctocaecum acuti * (D)
                              Proterodiplostomum medusae (D)
Ctenosaura similis            Atractis longicaudatum * (N)
                              Oochoristica guanacastense * (C)
                              Oswaldofilaria brevicaudata (N)
                              Ozolaimus cirratus (N)
                              Ozolaimus ctenosauri (N)
                              Ozolaimus megatyphlon (N)
                              Parallopharynx gonzalezi (D)
Drymobius margaritiferus      Ochetosoma miladelarocai (D)
Geomyda manni                 Telorchis sp. (D)
                              Spiroxys sp. (N)
Kinosternon cruentatum        Atractis caballeroi * (N)
                              Atractis impura (N)
                              Capillaria sp. (N)
                              Heronimus mollis (D)
                              Telorchis pseudoaculeatus (D)
Kinosternon leucostomum       Neopolystoma fentoni * (M)
Rhinoclemmys pulcherrima      Neopolystoma fentoni (M)
Sceloporus malachiticus       Ascarops sp. (N)
                              Physaloptera sp. (N)
                              Spauligodon oxkutzcabiensis (N)

                             Aves

Host                          Helminth Species

Ajaia ajaja                   Mesaulus grandis (D)
Buteo nitidus                 Platynosomum costarricense * (D)
Calocitta formosa             Lubens lubens (D)
Chamaeptes unicolor           Maritremopsis longibursatus * (D)
Columba fasciata              Ascaridia columbae (N)
Columba livia                 Capillaria obsignata (N)
Crotophaga sulcirostris       Eumegacetes macroorchis * (D)
Gallus gallus                 Amoebotaenia cuneata (C)
                              Amoebotaenia sp. (C)
                              Ascaridia galli (N)
                              Capillaria obsignata (N)
                              Choanotaenia infundibulum (C)
                              Choanotaenia sp. (C)
                              Davainea proglottina (C)
                              Hymenolepis sp. (C)
                              Oncicola oncicola (A)
                              Oxyspirura mansoni (N)
                              Raillietina echinobothrida (C)
                              Raillietina sp. (C)
                              Raillietina tetragona (C)
                              Syngamus trachea (N)
                              Tetrameres fissispina (N)
Gymnostinops montezuma        Hymenolepis sp. (C)
                              Leucochloridium costarricense * (D)
                              Lutztrema obliquum (D)
                              Tanaisia freitasi * (D)
                              Zonorchis costarricensis * (D)
Icterus galbula               Stomylotrema ucremium * (D)
Jacana spinosa                Haematotrephus facioi * (D)
                              Tanaisia winteri * (D)
Leucoplernis semiplumbea      Platynosomum illiciens (D)
Melaegris gallopavo           Syngamus trachea (N)
                              Ascaridia galli (N)
Momotus momotus               Hyptiasmus costarricense * (D)
                              Ornithotrema momoti * (D)
Nesotriccus ridgwayi          Diplotriaena muscisaxicolae (N)
Oriolus oriolus               Paricterotaenia sp. (C)
Ostinops quatemozinus         Hymenolepis sp. (C)
Pteroglossus torquatus        Zonorchis macroovaricus (D)
Sturnaella magna              Paricterotaenia sp. (C)
Turdus grayi                  Anomotaena sp. (C)
                              Aploparaksis sp. (C)

                            Mammalia

Host                          Helminth Species

Agouti paca                   Capillaria sp. (N)
                              Strongyloides sp. (N)
                              Taenia sp. (C)
                              Trichuris sp. (N)
Alouatta palliata             Ascaris lumbricoides (N)
                              Controrchis biliophilus (D)
                              Parabronema sp. (N)
                              Physalopteridae gen. sp. (N)
                              Strongyloides cebus (N)
                              Trypanoxyuris minutus (N)
Ateles geoffroyi              Angiostrongylus costaricensis (N)
                              Oxyuris sp. (N)
                              Strongyloides cebus (N)
                              Trypanoxyuris sp. (N)
Bos taurus                    Bunostomum phlebotomum (N)
                              Capillaria sp. (N)
                              Cooperia sp. (N)
                              Cotylophoron sp. (N)
                              Cylicocyclus sp. (N)
                              Dictyocaulus eckerti (N)
                              Dictyocaulus sp. (N)
                              Dictyocaulus viviparus (N)
                              Echinococcus granulosus (C)
                              Fasciola hepatica (D)
                              Gaigeria pachyscelis (N)
                              Haemonchus contortus (N)
                              Mammomonogamus laryngeus (N)
                              Mecistocirrus digitatus (N)
                              Mecistocirrus sp.
                              Moniezia benedeni (C)
                              Moniezia expansa (C)
                              Neoascaris vitulorum (N)
                              Oesophagostomum radiatum (N)
                              Ostertagia sp. (N)
                              Paramphistomum cervi (D)
                              Setaria cervi (N)
                              Setaria labiatopapillosa (N)
                              Stephanurus dentatus (N)
                              Taenia saginata (C)
                              Trichostrongylus sp. (N)
                              Trichuris ovis (N)
Bradypus griseus              Leiurus leptocephalus (N)
                              Moniezia benedeni (C)
Canis familiaris              Ancylostoma caninum (N)
                              Ascaris lumbricoides (N)
                              Avitellina centripunctata (C)
                              Dipilydium caninum (C)
                              Dirofilaria immitis (N)
                              Paragonimus mexicanus (D)
                              Spirocerca lupi (N)
                              Toxocara canis (N)
                              Trichuris vulpis (N)
Canis latrans                 Alaria nasuae (D)
                              Ancylostoma caninum (N)
Capra hircus                  Bunostomum trigonocephalum (N)
                              Cooperia sp. (N)
                              Dictyocaulus sp. (N)
                              Gaigeria pachyscelis (N)
                              Haemonchus contortus (N)
                              Haemonchus sp. (N)
                              Moniezia sp. (C)
                              Oesophagostomum venulosum (N)
                              Ostertagia sp. (N)
                              Skrjabinema ovis (N)
                              Skrjabinema sp. (N)
                              Strongyloides sp. (N)
Cebus capucinus               Dipetalonema gracile (N)
                              Prostenorchis elegans (A)
                              Strongyloides cebus (N)
Chironecter minima            Paragonimus caliensis (D)
Conepatus semistriatus        Paragonimus mexicanus (D)
Didelphis marsupialis         Anatrichosoma buccalis (N)
                              Mathevotaenia pennsylvanica (C)
                              Paragonimus caliensis (D)
                              Paragonimus mexicanus (D)
                              Rhopalias coronatus (D)
                              Rhopalias horridus (D)
                              Spelotrema skrjabini * (D)
Eptesicus propinquus          Limatulum costarricensis * (D)
                              Prosthodendrium cordiforme (D)
                              Urotrema scabridum (D)
Equus caballus                Anoplocephala magna (C)
                              Anoplocephala perfoliata (C)
                              Anoplocephaloides mamillana (C)
                              Cylicocyclus insigne (N)
                              Dictyocaulus arnfieldi (N)
                              Habronema megastoma (N)
                              Habronema microstoma (N)
                              Nematodirus sp. (N)
                              Onchocerca cervicalis (N)
                              Oxyuris equi (N)
                              Parascaris equorum (N)
                              Setaria equina (N)
                              Strongylus edentatus (N)
                              Strongylus equinus (N)
                              Strongylus vulgaris (N)
                              Triodontophorus tenuicollis (N)
Felis catus                   Amphimerus costarricensis * (D)
                              Ancylostoma caninum (N)
                              Dipylidium caninum (N)
                              Paragonimus mexicanus (D)
                              Taenia taeniaeformis (C)
                              Toxocara mystax (N)
Felis concolor                Echinococcus oligarthrus (C)
Felis pardalis                Lagochilascaris sp. (N)
                              Paragonimus mexicanus (D)
                              Syngamus sp. (N)
Felis wiedii                  Taenia taeniaeformis (C)
Hemiderma perspicillatum      Litomosoides penai * (N)
Marmosa sp.                   Mathevotaenia bivittata (C)
                              Mathevotaenia didelphidis (C)
Micronycteris hirsutus        Castroia silvai (D)
Mustela frenata               Euryhelmis costaricensis * (D)
                              Plagiorhynchus angrense (A)
                              Polymorphus corynosonta (A)
Mustela noveboracensis        Centrorhynchus sp. (A)
Nasua narica                  Angiostrongylus costaricensis (N)
Odocoileus virginianus        Ashworthius patriciapillitae * (N)
                              Eucyathostomum webbi (N)
                              Gongylonema pulchrum (N)
                              Mazamastrongylus sp. (N)
                              Onchocerca cervipedis (N)
                              Paramphistomum liorchis (D)
                              Parelaphostrongylus tenuis (N)
                              Setaria yehi (N)
                              Taenia omissa (C)
Philander opossum             Amphimerus neotropicalis * (D)
                              Maritremopsis prolixum * (D)
                              Paragonimus mexicanus (D)
                              Rhopalias baculifer (D)
                              Rhopalias coronatus (D)
                              Rhopalias horridus (D)
                              Rhopalias macracanthus (D)
Potos flavus                  Ancylostoma braziliense (N)
Procyon lotor                 Paragonimus caliensis (D)
                              Paragonimus mexicanus (D)
                              Robertdigenea dollfusi * (D)
Rattus novergicus             Gongylonema neoplasticum (N)
                              Hymenolepis diminuta (C)
                              Mastophorus muris (N)
                              Moniliformis moniliformis (A)
                              Nippostrongylus brasiliensis (N)
                              Protospirura muris (N)
                              Strongyloides ratti (N)
                              Taenia taeniaeformis (C)
                              Trichosomoides crassicauda (N)
                              Vampirolepis nana (C)
Rattus rattus                 Aspiculuris tetraptera (N)
                              Moniliformis moniliformis (A)
                              Taenia taeniaeformis (C)
Saimiri oerstedi              Prosthenorchis sp. (A)
Sigmodon hispidus             Angiostrongylus costaricensis (N)
                              Hassalstrongylus aduncus (N)
                              Hymenolepis diminuta (C)
                              Litomosoides carinii (N)
                              Monodontus sp. (N)
                              Protospirura sp. (N)
                              Raillietina sp. (C)
                              Strongyloides sigmodontis (N)
                              Trichostrongylus sigmodontis (N)
                              Trichostrongylus sp. (N)
Sus scrofa                    Ascaris suum (N)
                              Globocephalus urosubulatus (N)
                              Metastrongylus elongatus (N)
                              Physocephalus sexalatus (N)
                              Stephanurus dentatus (N)
                              Taenia solium (C)
                              Trichuris suis (N)
Sylvilagus brasiliensis       Anoplocephaloides floresbarroetae * (C)
Tonatia silvicola             Oligacanthorhynchidae gen. sp. (A)
Urocyon cinereoargenteus      Paragonimus mexicanus (D)
                              Taenia pisiformis (C)

                       Unidentified hosts

Host                          Helminth Species

"Amphibian"                   Plagiorchis sp. (D)
"Atun"                        Jaliscia caballeroi (M)
"Aves"                        Diorchis acuminatus (C)
"Aves"                        Polymorphus minutus (A)
"Aves"                        Capillaria annulosa (N)
"Aves" (?)                    Plagiorchis sp. (D)
"Carraco"                     Porrocaecum angusticolle (N)
"Cobra imperator"             Porrocaecum (N), Kalicephalus sp. (N) y
                                Cruzia sp. (N)
"Colubridae"                  Ochetosoma bravoiae * (D)
"Columbidae"                  Ascaridia sp. (N)
"Culebra coral"               Oxysomatium sp. (N)
"Ganzo"                       Porrocaecum sp. (N)
"Medialuna" sp.               Lasiotocus costaricae * (D)
"Medialuna" sp.               Koronacantha pectinaria (A)
"Moray"                       Helicometra sinuata (D)
"Ovejas"                      Oesophagostomum venulosum (N)
"Patos"                       Ascaridia galli (N)
"Rats"                        Oesophagostomum venosolum (N)
"Tortuga"                     Telorchis sp. (D)
"Tortuga"                     Plagiorchis sp. (D)


DISCUSSION

In this checklist we present information of 303 species of helminths parasitizing 108 species of vertebrates in Costa Rica. Helminth species are represented by 81 species of digeneans, 23 of monogeneans, 63 of cestodes, 12 of acanthocephalans, and 124 of nematodes. Strikingly, almost all the records we present herein correspond to adult forms infecting mostly the intestine of their hosts, or habitats such as the liver or lungs. Vertebrates infected with helminths are included in all major groups, such as fishes (including elasmobranchs and teleosts, with 31 species), amphibians (7 species), reptiles (14 species), birds (20 species), and mammals (36 species), including both wildlife and domestic. The largest number of helminth species has been described from Mammals, however this might be ah artifact of the sampling since this is the group on which more species have been analyzed for helminths in Costa Rica. Characteristically, most records of helminths are presented as isolated reports describing one species in a particular host, and only a few surveys of the helminth fauna of a particular host have been conducted (see for example Carreno et al. 2001). However, in this checklist we present a comprehensive list of the helminth fauna of each host species when it is available, considering all the published accounts, e.g. the toad Bufo marinus possess a helminthological record of 12 species of helminths, recorded in 12 reports (papers) from 10 localities along its distributional range in Costa Rica. In contrast, the shark Carcharhinus leucas is parasitized with 14 species of helminths in one single locality (Watson and Thorson 1976).

Clearly, the inventory of the helminth parasites of vertebrates in Costa Rica is in its first steps and a large number of species will be described in the next years. So far, only 108 species of vertebrates have been studied for helminths (most of them representing isolated reports and not complete surveys of each host species along its distributional range), and from them, 303 helminth species were recorded. However, only 3.8% of the vertebrate fauna of the country has been studied, considering that there are about 2,855 species of vertebrates in Costa Rica (Janzen 1983, Stiles and Skutch 1989, Bussing 1998). Interestingly, 58 species were recorded as new species from Costa Rica and most of them are endemic to particular regions. This represents approximately 19.1% of endemic species of helminths in vertebrates of Costa Rica. In comparison, after more than 70 years of survey work of helminth parasites of vertebrates in Mexico, approximately 1,632 species of helminths hosted in 947 species of vertebrates have been recorded. Approximately, 20% of the helminth fauna of vertebrates from Mexico has been described so far, according with estimates presented by Perez-Ponce de Leon and Garcia-Prieto (2001b).

Considering that every species of vertebrate supports a diverse biota of symbiotic organisms, and that parasites are integral components of biodiversity, we have learned that documenting the diversity of helminth parasites inhabiting vertebrate hosts in Costa Rica (and in any other part of the world) is very important by several reasons. They can act as agents to control and regulate host populations and they can also act as agents of disease representing a threat for domestic and wildlife vertebrates and also a threat for the conservation programs, especially those directed to preserve endangered species and ecosystems. The ecology and evolutionary history of these parasites is tied to that of their hosts, so they can be used as key indicators of ecosystem health, and also as indicators to monitor global change. At the same time, they provide valuable information to the understanding of the biogeographical and coevolutionary history of the association (Gardner and Campbell 1992, Hoberg 1997a, 1997b, Marcogliese and Cone 1997, Overstreet 1997, Brooks and Hoberg 2000, Brooks et al. 2001, Perez-Ponce de Leon and Garcia-Prieto 2001a).

There is no doubt about the importance of the helminth parasites as part of the inventory of natural resources in Costa Rica and we would like also to emphasize the important role that these parasites may playas agent of diseases in human beings. Several of the helminths mentioned in the checklist are the cause of some disease that directly causes health problems, or at least the potential exists for some of them to become emerging pathogens (see Morera and Cespedes 1971, Brenes-Madrigal et al. 1974, and Brenes-Madrigal et al. 1983) for cases of human infections with Angiostrongylus costaricensis, Wuchereria bancrofti, and Raillietina sp., respectively).

To summarize, helminth parasites of vertebrates in Costa Rica need to be considered within any initiatives to accomplish the national inventory of biological resources. The information provided by them is synergistic with all the information about the natural history of the hosts that is being assembled from field data and, in addition to that, the risk of emerging diseases will be assessed and probably all the information will be used when decisions to handle and protect biodiversity are taken. In this sense, to build up and preserve a national depository of the specimens (and the information associated to each of them) is a necessary step. Universities and government agencies have to be convinced to support now, and in the future, the establishment of biological collections where all specimens will be saved, so infrastructure and personnel have to be provided to guarantee the quality of the specimens within collections, as well as to provide a service to the society and the scientific community to make sure that the knowledge and the information contained in collections is available for all of them. At this point, the Coleccion de Helmintos de Costa Rica (CNCR) has re-emerged and we will try to hold the promise that it will have the standards of quality to assure that it will become the national depository of helminths in the country.

ACKNOWLEDGMENTS

We would like to thank Scott Gardner and Skip Sterner, Harold W. Manter Laboratory of Parasitology, University of Nebraska-Lincoln, and Eric Hoberg and Patt Pillit, U.S. National Parasite Collection, Beltsville, Maryland, for providing information form their databases and bibliography. Agustin Jimenez provided useful bibliographic references. Federico Villalobos Brenes provided some information about localities in Costa Rica. Funds to complete this paper were partially provided by the agreement between the Universidad de Costa Rica and the Instituto Mexicano de Cooperacion Internacional through the project: "Support to catalog and maintain the Helminthological collection of the University of Costa Rica". LGP thanks finantial support provided by Fernando Alvarez, Department of Zoology, University of Mexico.

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Received 12-VI 2002.

Corrected 10-VII-2002.

Accepted 16-VII-2002.

Beatriz Rodriguez-Ortiz (1), Luis Garcia-Prieto (2), and Gerardo Perez-Ponce de Leon (2)

(1) Laboratorio de Helmintologia, Facultad de Microbiologia, Universidad de Costa Rica, San Jose, Costa Rica, e-mail: brodrigu@cariari.ac.ucr.cr

(2) Laboratorio de Helmintologia, Instituto de Biologia, UNAM. Ap. Postal 70-153. C.P. 04510, Mexico D.F., e-mail: ppdleon@servidor.unam.mx; gprieto@ibiologia.unam.mx

RESUMEN

Los helmintos de vertebrados silvestres y domesticos han sido estudiados por mas de 50 anos en Costa Rica; sin embargo, la informacion sobre los mismos se encuentra dispersa en diversas fuentes. Con el fin de recopilar dicha informacion, construimos una base de datos que contiene gran parte de los registros sobre helmintos de vertebrados silvestres y domesticos descritos para Costa Rica; estos fueron obtenidos a partir de busquedas bibliograficas electronicas y de consultas a colecciones parasitologicas, conformando un listado que contiene 303 especies, incluyendo 81 especies de digeneos, 23 de monogeneos, 63 de cestodos, 12 de acantocefalos y 124 de nematodos. Tales registros proceden del estudio de 108 especies de vertebrados (31 especies de peces, 7 anfibios, 14 reptiles, 20 aves y 36 mamiferos), lo cual representa unicamente el 3.8% de la fauna de vertebrados pues en el pais se han descrito aproximadamente 2,855 especies. De manera interesante, 58 de las 303 especies (19.1%) que han sido registradas representan especies nuevas y, en su mayoria endemicas. Dado que los parasitos aportan informacion relevante sobretodo cuando se conjunta de manera sinergistica con aquella obtenida de la historia natural de los hospederos a los que parasitan, consideramos que los helmintos parasitos deben ser incorporados en las iniciativas para completar el inventario biologico nacional. Este trabajo de compilacion representa el primer paso para la re-constitucion de la Coleccion de Helmintos de Costa Rica (CHCR), albergada por la escuela de Microbiologia de la Universidad de Costa Rica, que de alcanzar y mantener los estandares internacionales de calidad necesarios, debera convertirse en el depositario nacional de los helmintos de este pais.
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Author:Rodriguez-Ortiz, Beatriz; Garcia-Prieto, Luis; Perez-Ponce de Leon, Gerardo
Publication:Revista de Biologia Tropical
Date:Jun 1, 2004
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