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Revista de Biología Tropical

On-line version ISSN 0034-7744

Rev. biol. trop vol.52 n.2 San José Jun. 2004

 

Checklist of the helminth parasites of vertebrates
in Costa Rica

Beatriz Rodríguez-Ortíz 1 , Luis García-Prieto 2 , and Gerardo Pérez-Ponce de León 2

1 Laboratorio de Helmintología, Facultad de Microbiología, Universidad de Costa Rica, San José, Costa Rica, e-mail: brodrigu@cariari.ac.ucr.cr

2 Laboratorio de Helmintología, Instituto de Biología, UNAM. Ap. Postal 70-153. C.P. 04510, México D.F., e-mail: ppdleon@servidor.unam.mx; gprieto@ibiologia.unam.mx

Received 12-VI-2002. Corrected 10-VII-2002. Accepted 16-VII-2002.

 

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 3.8% 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 Colección de Helmintos de Costa Rica (CHCR), hosted at the Facultad de Microbiología, Universidad de Costa Rica, has reemerged 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.

 

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 Jiménez-Quirós and Brenes-Madrigal 1957, Brenes-Madrigal and Jiménez-Quirós 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 Microbiología, Universidad de Costa Rica, established a close collaboration with Dr. Eduardo Caballero and his research group from the Instituto de Biología, Universidad Nacional Autónoma de México (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 to inventory the parasites of particular regions of the country, i.e. the Área de Conservación 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, Pérez-Ponce de León et al. 1998, Brooks et al. 1999, Platt 2000, Zelmer and Brooks 2000, Carreño et al. 2001, León-Régagnon et al. 2001, Choudhury et al. 2002).

Of all the helminth specimens collected from Costa Rican hosts, only a few were saved in the collection of the Facultad de Microbiología, University of Costa Rica and most of them have been deposited in foreign parasite collections, i.e. the Colección 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 reestablishment of the Colección Helmintológica de la Facultad de Microbiología as the Colección 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 reemerged 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 Colección 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 Colección 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 are 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 Colección de Helmintos de Costa Rica, San José, Costa Rica; CNHE to the Colección 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, Río de Janeiro, Brazil; USNPC to the U.S. National Parasite Collection, Beltsville, Maryland, U.S.A., and MNHG to the Muséum de Histoire Naturelle, Geneva, Switzerland.

Platyhelminthes

Gegenbaur, 1859

Digenea

(Van Beneden, 1858)

Acanthocolpidae Lühe, 1909

1. Manteria brachydera (Manter, 1940) Caballero, 1950. Stomach, Caranx caballus, Oligoplites altus, Oligoplites refulgens (F), Golfo de Nicoya, Puntarenas (Ponciano-Rodríguez 1986). Specimens: CNHE.

2. Stephanostomum casum (Linton, 1910) McFarlane, 1934. Intestine, Caranx caballus (F), Playa Ocotal, Guanacaste (Pérez-Ponce de León et al. 1998). Specimens: CNHE.

Acanthostomidae Poche, 1926

3. Proctocaecum acuti (Caballero and Brenes-Madrigal, 1958) Lamothe- Argumedo and Ponciano-Rodríguez, 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- Rodríguez, 1985. Intestine, Rhamdia rogersi (F), Río Tiribí, Paso Ancho, San José (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), Tilarán, 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), Corís, 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), Tilarán, Guanacaste (Jiménez-Quirós and Brenes-Madrigal 1957), La Pacífica, Guanacaste (Stuart et al. 1990). Specimens: CHCR, CNHE, HWML.

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

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

Notes: Species described as Lubens centroamericanus Brenes-Madrigal, Arroyo-Sancho and Muñóz-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 (Jiménez-Quirós and Arroyo-Sancho 1960b). Specimens: CHCR.

14. Platynosomum costaricense Brenes-Madrigal, Arroyo-Sancho and Muñóz, 1966. Gall bladder, Buteo nitidus (B), Tilarán, Guanacaste (Brenes- Madrigal et al. 1966). Specimens: CHCR.

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

16. Robertdigenea dollfusi Caballero and Caballero-Rodríguez, 1970. Bile ducts and liver, Procyon lotor crassidens (M), Balsa de Mora, San José (Caballero and Caballero-Rodríguez 1970). Specimens: CNHE.

17. Zonorchis costaricensis Brenes-Madrigal and Jiménez-Quirós, 1959. Gall bladder, Gymnostinops montezuma (B), Peralta, Cartago (Brenes- Madrigal and Jiménez-Quirós 1959). Specimens: CHCR.

18. Z. macroovaricus Jiménez-Quirós and Arroyo-Sancho, 1960. Bile ducts, Pteroglossus torquatus (B), Tilarán, Guanacaste (Jiménez-Quirós and Arroyo-Sancho 1960b). Specimens: CHCR.

Didymozoidae Poche, 1907

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

Diplostomidae Poirier, 1886

20. Alaria nasuae La Rue and Townsend, 1927 Intestine, Canis latrans (M), San José, San José (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, Cartago (Brenes-Madrigal 1961), Santa Cruz de Turrialba (Chang-Díaz and Cartin- González 1983), San Pedro Montes de Oca, San José (NP). Specimens. CHCR, USNPC.

Fellodistomidae (Nicoll, 1909)

Nicoll, 1913

25. Tergestia laticollis (Rudolphi, 1819) Stossich, 1899. Intestine, Caranx caballus (F), Playa Ocotal, Guanacaste (Pérez-Ponce de León 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 José (Caballero et al.1957). Specimens: CHCR, CNHE.

Haematoloechidae Odening, 1964.

27. Haematoloechus meridionalis León-Règagnon, Brooks and Zelmer, 2001. Lungs, Rana vaillanti (A), Río Pizote, Guanacaste (León-Règagnon et al. 2001). Specimens: CNHE, USNPC.

28. Neohaematoloechus helibravoae Caballero-Rodríguez, 1989. Lungs, Rana palmipes (A), NA (NP). Specimens: CNHE.

Notes: This species is considered nomina nuda.

Hemiuridae Lühe, 1901

29. Ectenurus virgulus Linton, 1910. Intestine, Fistularia commersonii (F), Playa Ocotal, Guanacaste (Pérez-Ponce de León et al. 1998). Specimens: CNHE.

30. Halipegus eschi Zelmer and Brooks, 2000. Esophagus, Rana vaillanti (A), Río Pizote, Guanacaste (Zelmer and Brooks 2000). Specimens: USNPC.

31. Lecithochirium microstomum Chandler, 1935. Intestine, Fistularia commersonii (F), Playa Ocotal, Guanacaste (Pérez-Ponce de León 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-Rodríguez 1986). Specimens: CNHE.

34. Pseudolecithaster sp. Intestine, Echidna nocturna (F), Playa Cuajiniquil, Guanacaste (Pérez-Ponce de León et al. 1998). Specimens: CNHE.

35. Theletrum lamothei Pérez-Ponce de León, León-Règagnon and Monks, 1998. Intestine, Echidna nocturna (F), Playa Cuajiniquil, Guanacaste (Pérez- Ponce de León et al. 1998). Specimens: CNHE, USNPC.

Heronimidae Ward, 1917

36. Heronimus mollis (Leidy, 1856) Stunkard, 1964. Lungs, Kinosternon cruentatum (R), Santa Ana, San José (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 Jiménez- Quirós, 1960 Intestine, Mustela frenata (M), Tres Ríos, 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 (Lühe, 1931) Odhner, 1910

40. Castroia silvai Travassos, 1928. Intestine, Micronycteris hirsutus (M), Río 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), Corís, 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 Ochoterena-trema 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-Rodríguez, 1969). Intestine, Rana pipiens (A), Sarapiqui, Heredia (Caballero and Caballero-Rodríguez 1969), Turrialba, Cartago (Sullivan 1971). Specimens: CNHE, HWML, USNPC.

Notes: Specimens collected by Caballero and Caballero-Rodríguez (1969) was included in the genus Brenesia Caballero and Caballero-Rodríguez, 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 Limón, Puntarenas (Caballero and Brenes-Madrigal 1957). Specimens: CHCR, CNHE.

46. B. simonei (Travassos, Teixeira de Freitas and Bührnheim, 1965) Yamaguti, 1971 Intestine, Arothron hispidus (F), Bahía Prieta, Guanacaste (Pérez-Ponce de León et al. 1998). Specimens: CNHE.

47. Hypocreadium myohelicatum Bravo-Hollis and Manter, 1957. Intestine, Epinephelus itajara (F), Golfo de Santa Clara, Guanacaste (Pérez-Ponce de León et al. 1998). Specimens: CNHE.

Leucochloridiidae (Poche, 1907)

Dollfus, 1934

48. Leucochloridium costaricense Brenes-Madrigal and Arroyo-Sancho, 1962. Intestine, Gymnostinops montezuma (B), Cachí, Cartago (Brenes- Madrigal and Arroyo-Sancho 1962). Specimens: CHCR.

Macroderoididae McMullen, 1937

49. Glypthelmins facioi Brenes-Madrigal, Arroyo-Sancho, Jiménez-Quirós and Delgado-Flores, 1959. Intestine, Rana pipiens (A), Corís, 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 José (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), Río Grande de Atenas, Alajuela (Brenes-Madrigal et al. 1960b). Specimens: CHCR, HWML.

53. Wallinia chavarriai Choudhury, Daverdin and Brooks, 2002. Intestine, Astyanax aenaeus (F), Río Sapoa, Guanacaste, and Bryconamericus scleroparius (F), Río 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), Caserío de Tiquiares, San José (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), Río 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), Sarapiquí, Heredia (Caballero et al. 1963a). Specimens: CHCR, CNHE.

57. Spelotrema skrjabini (Caballero, 1958) Yamaguti, 1971. Intestine, Didelphis marsupialis (M), Bello Horizonte, Escazú, San José (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 José (Lamothe-Argumedo and Brenes-Madrigal 1983). Specimens: CNHE.

61. A. neotropicalis Caballero, Montero-Gei and Caballero-Rodríguez, 1963. Pancreatic ducts, Philander opossum (M), Río Grande de Atenas, Alajuela (Caballero et al. 1963b). Specimens: CNHE.

Note: Nasir and Díaz (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 José 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 José (Brenes-Madrigal et al. 1980), Felis catus (M), Balsa de Atenas, Alajuela, Rincón de Osa, Puntarenas, Tabarcia de Mora, San Isidro de Balsa and Cangrejal de Acosta , San José (Brenes-Madrigal et al. 1980), Felis pardalis (M), Bagaces, Guanacaste (Brenes-Madrigal et al. 1980), Philander opossum (M), Río Grande de Atenas, Alajuela (Caballero and Montero-Gei 1961), Balsa de Atenas and Tornos de Atenas, Alajuela, Rincón de Osa, Puntarenas, and San Isidro de Balsa, San José (Brenes-Madrigal et al. 1980), Procyon lotor (M), San Isidro de Balsa, San José (Brenes-Madrigal et al. 1980), Urocyon cinereoargenteus (M), Balsa de Atenas and Turrúcares, 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), Corís, Cartago (Brenes-Madrigal et al. 1959b). Specimens: CHCR.

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

67. P. liorchis Fischoeder, 1901. Rumin, Odocoileus virginianus (M), Parque Nacional Santa Rosa, Guanacaste (Carreño et al. 2001). Specimens: USNPC.

Plagiorchiidae (Lühe, 1901) Ward, 1917

68. Ochetosoma bravoae Brenes-Madrigal and Arroyo-Sancho, 1960. Esophagus, "Colubridae" (R), Bello Horizonte, Escazú, San José (Brenes- Madrigal and Arroyo-Sancho 1960). Specimens: CHCR.

69. O. miladelarocai Caballero y Vogelsang, 1947. Esophagus, Drymobius margaritiferus (R), Corís, 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 Limón, 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), Río 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, Escazú, San José (Caballero et al. 1957), Philander opossum (M), Río Grande de Atenas, Alajuela (Caballero and Montero-Gei 1961), Cariari, Limón (Siebert 1970). Specimens: CHCR, CNHE, HWML.

76. R. horridus (Diesing, 1850) Stiles and Hassall, 1898. Intestine, Didelphis marsupialis (M), Bello Horizonte, Escazú, San José (Caballero et al. 1957), Philander opossum (M), Río Grande de Atenas, Alajuela (Caballero and Montero-Gei 1961). Specimens: CHCR, CNHE.

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

Stomylotrematidae (Travassos, 1922) Poche, 1926

78. Stomylotrema ucremium Brenes-Madrigal, Arroyo-Sancho and Muñóz- 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 José (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 José (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 José (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 Rincón, Puntarenas (Kritsky and Leiby 1972). Specimens: USNPC.

7. Palombitrema heteroancistrum Price and Bussing, 1968. Gills, Astyanax fasciatus (F), Río Montenegro, 23 Km NW of Cañas, Guanacaste (Price and Bussing, 1968), 3 km. SW of Rincón, 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 Suriano (1997) re-established its validity.

8. Urocleidoides costaricensis (Price and Bussing, 1967) Kritsky and Leiby, 1972. Gills, Astyanax fasciatus (F), Cañas, 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 incertae sedis by Kritsky et al. (1986).

9. Urocleidoides strombicus (Price and Bussing, 1967) Kritsky and Thatcher, 1974. Gills, Astyanax fasciatus (F), Cañas, 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 Limón, 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 Rincón, Puntarenas (Kritsky and Fritts 1970). Specimens: USNPC.

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

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

Hexabothriidae Price, 1942

17. Erpocotyle carcharhini Watson and Thorson, 1976. Gills, Carcharhinus leucas (F), Barra del Colorado, Río Colorado, Limón (Watson and Thorson 1976). Specimens: USNPC.

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

Microbothriidae Price, 1936

19. Dermophthirius maccallumi Watson and Thorson, 1976. Skin, Carcharhinus leucas (F), Barra del Colorado, Río Colorado, Limón (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, Limón (Brenes-Madrigal, 1961), San Pedro Montes de Oca, San José (NP). Specimens: USNPC.

3. Anoplocephaloides floresbarroetae Rausch, 1976. Bile duct, Sylvilagus brasiliensis (M), Aserrí, San José (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 floresbarroetae 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), Río 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), Río 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, Escazú, San José (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 José, San José (Brenes-Madrigal, 1961). Specimens: USNPC.

11. M. expansa (Rudolphi, 1805) Blanchard, 1891. Intestine, Bos taurus (M), NA (Brenes-Madrigal 1961).

12. Oochoristica guanacastensis Brooks, Pérez Ponce de León, and García, 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, Río Colorado, Limón (Watson and Thorson 1976).

Dasyrhynchiade Dollfus, 1935

14. Dasyrhynchus variouncinatus (Pintner, 1913) Pintner, 1928. Spiral valve, Carcharhinus leucas (F), Barra del Colorado, Río Colorado, Limón (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), Cañas, Guanacaste Rodríguez-Ortíz 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 José, San José (Flores-Barroeta and Hidalgo-Escalante 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), Cantón Cartago, Cartago (NP), Oriolus oriolus (B), San José San José (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 José, San José (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 José San José, (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 José, San José (Vives and Zeledón 1957); Sigmodon hispidus (M), Cañas, Guanacaste (Rodríguez-Ortíz et al. 2000).

31. Vampirolepis nana (Von Siebold, 1852) Spasskii, 1954. Intestine, Rattus norvegicus (M), San José, San José (Vives and Zeledón 1957).

Lacistorhynchidae Guiart, 1927

32. Callitetrarhynchus gracilis (Rudolphi, 1819) Pintner, 1931. Spiral valve, Carcharhinus leucas (F), Barra del Colorado, Río Colorado, Limón (Watson and Thorson 1976). Specimens: USNPC.

Onchobothriidae Braun, 1900

33. Acanthobothrium campbelli Marques, Brooks and Monks, 1995. Spiral valve, Urotrygon chilensis (F), Costa de Pájaros, 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-Ureña, 1996. Spiral valve, Himantura pacifica (F), Playa Panamá, Guanacaste (Marques et al. 1996). Specimens: HWML, MNHG.

42. Phoreiobothrium triloculatum Linton, 1901. Spiral valve, Carcharhinus leucas (F), Barra del Colorado, Río Colorado, Limón (Watson and Thorson 1976). Specimens: USNPC.

43. Platybothrium hypoprioni Potter, 1927. Spiral valve, Carcharhinus leucas (F), Barra del Colorado, Río Colorado, Limón (Watson and Thorson 1976). Otobothriidae Dollfus, 1942

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

Phyllobothriidae Braun, 1900

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

46. A. pristis Woodland, 1934. Spiral valve, Pristis perotteti (F), Barra del Colorado, Río Colorado, Limón (Watson and Thorson 1976).

47. Phyllobothrium leuci Watson and Thorson, 1976. Spiral valve, Carcharhinus leucas (F), Barra del Colorado, Río Colorado, Limón (Watson and Thorson 1976).

48. P. nicaraguensis Watson and Thorson, 1976. Spiral valve, Carcharhinus leucas (F), Barra del Colorado, Río Colorado, Limón (Watson and Thorson 1976).

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

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

Proteocephalidae La Rue, 1911

51. Ophiotaenia sp. Intestine, Constrictor constrictor (R), San José, San José (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 (Velázquez and Morales 1983).

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

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

57. T. omissa Lühe, 1910 (cysticerci). Lungs, Odocoileus virginianus (M), Parque Nacional Santa Rosa, Guanacaste (Carreño 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 José, San José (Vives and Zeledón 1957; Intestine, Felis wiedii nicaraguense (M), Mata de Limón, Puntarenas (Flores- Barroeta et al. 1958b), F. catus (M), San José, San José (Brenes-Madrigal 1961).

Tentaculariidae Poche, 1926

62. Nybelinia bisulcata (Linton, 1889) Poche, 1926. Spiral valve, Carcharhinus leucas (F), Barra del Colorado, Río Colorado, Limón (Watson and Thorson 1976).

Trilocularidae Yamaguti, 1959

63. Escherbothrium molinae Berman and Brooks, 1994. Spiral valve, Urotrygon chilensis (F), Costa de Pájaros 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 José, San José (NP). 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 José, San José (NP). Specimens: USNPC. Illiosentidae Golvan, 1960

4. Koronacantha pectinaria (Van Cleave, 1940). Intestine, "Medialuna" (F), Bahía Culebra, Guanacaste (Van Cleave 1940), Microlepidotus brevipinnis (F), Playas Panamá 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 Pérez- Ponce de León, 1996 by Monks et al. (1997).

Moniliformidae Van Cleave, 1924

5. Moniliformis moniliformis (Bremser, 1811) Travassos, 1915. Intestine, Rattus norvegicus, R. rattus (M), San José, San José (Vives and Zeledón 1957). Specimens: CNHE, USNPC. Oligacanthorhynchidae

Southwell and MacFie, 1925

6. Oligacanthorhynchidae gen. sp. Subcutaneous tissue, Tonatia silvicola (M), Península de Osa, Puntarenas (Gibson and McCarthy 1987).

7. Oncicola oncicola (Ihering, 1892) Travassos, 1916. Muscle, Gallus gallus (B), Parrita, Puntarenas (Zeledón 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. m inutus (Goeze, 1782) Lühe, 1911. Intestine, "birds", NA (Brenes- Madrigal 1961).

Nematoda

(Rudolphi, 1808)

Ancylostomatidae (Looss, 1905)

1. Ancylostoma braziliense Gómes de Faria, 1910. Intestine, Potos flavus arborensis (M), NA (Brenes-Madrigal 1961).

2. A. caninum (Ercolani, 1859). Intestine, Canis familiaris (M), San José, San José (Brenes-Madrigal 1961, Vargas and Contreras 1998), Canis latrans (M), and Felis catus (M), San José, San José (Brenes-Madrigal 1961). Specimens: USNPC.

3. Bunostomum phlebotomum (Railliet in Rizzo, 1900). Intestine, Bos taurus (M), San Isidro de Coronado, San José (Brenes-Madrigal 1961). Specimens: USNPC.

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

6. Globocephalus urosubulatus (Alessandrini, 1909). Intestine, Sus scrofa (M), Puriscal and Santa Ana, San José (Brenes-Madrigal 1961).

7. Monodontus sp. Intestine, Sigmodon hispidus (M), Cañas, Guanacaste (Rodríguez-Ortíz et al. 2000). Angiostrongylidae

(Boehm and Gebauer, 1934)

8. Angiostrongylus costaricensis Morera and Céspedes, 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 (Rodríguez-Ortíz 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 Pacífica, Guanacaste (Stuart et al. 1990), Canis familiaris (M), NA (Brenes-Madrigal 1961).

10. A. suum Goeze, 1782. Intestine, Sus scrofa (M), Matadero Municipal, San José (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, Limón (Brenes-Madrigal 1961).

15. Porrocaecum sp. (adult and larvae). NA, "goose" (B), and "cobra imperator" (R), San José, San José (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 José, San José (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 José (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 José (Brenes-Madrigal and Bravo-Hollis 1960). Specimens: CHCR, CNHE.

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

23. A. impura Caballero, 1944. Intestine, Kinosternon cruentatum (R), Santa Ana, San José (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 Limón, 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 José 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 José, San José (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 José, San José (NP). Specimens: USNPC.

30. K. inermis macrovulvus (Caballero, 1954) Intestine, Bothrops atrox (R), Hospital San Juan de Dios, San José (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 José (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, Limón (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), Irazú 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-Ureña and Erbe, 1998. Spiral valve, Himantura pacifica (F), Playa Panamá, Guanacaste (Hoberg et al. 1998). Specimens: USNPC.

38. Spiroxys sp. Intestine, Geomyda manni (R), San Juan, San José (Brenes-Madrigal 1961). Specimens: USNPC.

Gongylonematidae (Hall, 1916)

39. Gongylonema neoplasticum (Fibiger and Ditlevsen, 1914) Stomach, Rattus novergicus (M), San José, San José (Vives and Zeledon 1957).

40. G. pulchrum Molin, 1857 Esophagus, Odocoileus virginianus (M), Parque Nacional Santa Rosa, Guanacaste (Carreño 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 José (Brenes-Madrigal 1961). Specimens: USNPC.

43. Parabronema sp. Intestine, Alouatta palliata (M), Reserva Biológica 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 (Rodríguez-Ortíz et al. 1999, 2000), Cañas,

Guanacaste (Rodríguez-Ortíz et al. 2000).

45. Nippostrongylus brasiliensis (Travassos, 1914). Intestine, Rattus novergicus (M), San José, San José (Vives and Zeledón 1957). Heteroxynematidae (Skrjabin and Schikhobalova, 1948)

46. Aspiculuris tetraptera (Nitzsch, 1821). Intestine, Rattus rattus (M), Escazú, San José (Vives and Zeledón 1957).

Kathlaniidae Lane, 1914)

Travassos, 1918

47. Cruzia sp. NA, "cobra imperator" (R), San José, San José (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 (Marín 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), Tilarán, Guanacaste and Paso Ancho, San José (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 José, San José (Leuterer and Gothe 1993, Vargas and Contreras 1998).

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

54. Litomosoides carinii (Travassos, 1911). Body cavity, Sigmodon hispidus (M), Cañas, Guanacaste (Rodríguez-Ortíz et al. 2000).

55. L. peñai Jiménez-Quirós and Arroyo-Sancho, 1960. Body cavity, Hemiderma perspicillata (M), Santa Ana, San José (Jiménez-Quirós and Arroyo-Sancho 1960a). Specimens: CHCR.

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

57. O. caballeroi Esslinger, 1987. Mesentery, Bufo marinus (A), San José, San José (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 José (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 (Velázquez et al. 1983).

60. O. cervipedis Wehr and Dikmans, 1935. Dermis, Odocoileus virginianus (M), Parque Nacional Santa Rosa, Guanacaste (Carreño 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 José, San José (NP). Specimens: USNPC.

65. S. yehi (Durette-Desset, 1966). Rectum, Odocoileus virginianus (M), Parque Nacional Santa Rosa, Guanacaste (Carreño 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 José (NP), NA, Limón (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 (Velázquez and Solís 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 Pacífica, 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 Limón, Puntarenas (Bravo-Hollis and Brenes-Madrigal 1959). Specimens: CHCR, CNHE.

73. O. ctenosauri Caballero, 1938. Intestine, Ctenosaura similis (R), Liberia, Guanacaste and Mata de Limón, Puntarenas (Bravo-Hollis and Brenes- Madrigal 1959). Specimens: CHCR, CNHE.

Notes: Originally described as Ozolaimus monhystera (Von Linstow, 1902) and re-determined as O. ctenosauri by Moravec et al. (1996).

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

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

Physalopteridae (Railliet, 1893)

Leiper, 1908

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

77. Physaloptera sp. (larvae). Stomach, Bothrops athrox (R), NA (Brenes- Madrigal 1961), Sceloporus malachitichus (R), Volcán Irazú, 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 (Carreño et al. 2001). Specimens: USNPC.

Rhabdiasidae Railliet, 1915

79. Rhabdias bufonis (Schrank, 1788) Stiles and Hassal, 1905. Lungs, Bufo marinus (A), San José, San José (NP). Specimens: USNPC.

Notes: According to Baker (1987), this species is only found in Europe.

80. R. sphaerocephala Goodey, 1924. Lungs, Bufo marinus (A), Río Tiribi, San José and La Peña, 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 José, San José, San Ramón de Tres Ríos and Volcán Irazú, Cartago (Goldberg and Bursey 1992). Specimens: USNPC.

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

83. Mastophorus muris (Gmelin, 1790) Chitwood, 1938. Intestine, Rattus novergicus (M), San José, San José (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), Cañas, Guanacaste (Rodríguez-Ortíz et al. 2000).

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

Strongylidae Baird, 1853

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

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

90. Eucyathostomum webbi Pursglove, 1976 Intestine, Odocoileus virginianus (M), Parque Nacional Santa Rosa, Guanacaste (Carreño 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, Limón (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 José (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, Cantón Central, Heredia, Cantón Matina, Limón, Cantón Osa, Puntarenas, Cantones Central, Santa Ana and Terrazú, San José (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 Pacífica and Parque Nacional Santa Rosa, Guanacaste (Stuart et al. 1998).

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

98. S. sigmodontis Melvin and Chandler, 1950 Intestine, Sigmodon hispidus (M), Cañas, Guanacaste (Rodríguez-Ortíz 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 Ríos, 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, Carreño and Lichtenfels, 2002 Abomasum, Odocoileus virginianus (M), Parque Nacional Santa Rosa, Guanacaste (Carreño et al. 2001, Hoberg et al. 2002). Specimens: USNPC.

Notes: Describes as Ashworthius sp. by Carreño 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 José and Capra hircus (M), Turrialba, Cartago (Brenes- Madrigal 1961).

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

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

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

113. Mazamastrongylus sp. Abomasum, Odocoileus virginianus (M), parque Nacional Santa Rosa, Guanacaste (Carreño 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 (Rodríguez-Ortíz et al. 1999).

115. T. sigmodontis Baylis, 1945 Intestine, Sigmodon hispidus (M), Cañas, Guanacaste (Rodríguez-Ortíz et al. 2000).

Trichuridae (Ransom, 1911)

Railliet, 1915

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

117. Capillaria sp. Intestine, Agouti paca (M), Cantón Central, Heredia (Matamoros et al. 1991), Bos taurus (M), Santa Ana, San José (Brenes- Madrigal 1961), Kinosternon cruentatum (R), Santa Ana, San José (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), Guácimo, Limón (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 José, San José (Vives and Zeledón 1957).

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

122. T. ovis (Abildgaard, 1795) Intestine, Bos taurus (M), Sabanilla de Montes de Oca, San José 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 José, San José (Brenes-Madrigal 1961, Vargas and Contreras 1998).

Notes: Recorded as Trichocephalus vulpis Froelich, 1789 and Trichuris canis.










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 an 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 Carreño 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 Pérez-Ponce de León and García-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, Pérez-Ponce de León and García-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 play as 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 Céspedes 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 Colección 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. Agustín Jiménez 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 Cooperación 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.

Resumen

Los helmintos de vertebrados silvestres y domésticos han sido estudiados por más de 50 años en Costa Rica; sin embargo, la información sobre los mismos se encuentra dispersa en diversas fuentes. Con el fin de recopilar dicha información, construimos una base de datos que contiene gran parte de los registros sobre helmintos de vertebrados silvestres y domésticos descritos para Costa Rica; éstos fueron obtenidos a partir de búsquedas bibliográficas electrónicas y de consultas a colecciones parasitológicas, conformando un listado que contiene 303 especies, incluyendo 81 especies de digéneos, 23 de monogéneos, 63 de céstodos, 12 de acantocéfalos 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 mamíferos), lo cual representa únicamente el 3.8% de la fauna de vertebrados pues en el país 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 mayoría endémicas. Dado que los parásitos aportan información relevante sobretodo cuando se conjunta de manera sinergística con aquella obtenida de la historia natural de los hospederos a los que parasitan, consideramos que los helmintos parásitos deben ser incorporados en las iniciativas para completar el inventario biológico nacional. Este trabajo de compilación representa el primer paso para la re-constitución de la Colección de Helmintos de Costa Rica (CHCR), albergada por la escuela de Microbiología de la Universidad de Costa Rica, que de alcanzar y mantener los estándares internacionales de calidad necesarios, deberá convertirse en el depositario nacional de los helmintos de este país.

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