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Raúl Pineda, et al. •Helminth Parasites of Viviparous Fishes in Mexico 437

Helminth Parasitesof Viviparous

Fishes in Mexico

Raúl Pineda-López1,Guillermo Salgado-Maldonado2,

Eduardo Soto-Galera3,Norma Hernández-Camacho1,

Anita Orozco-Zamorano1,Salvador Contreras-Robledo1,

Guillermina Cabañas-Carranza2,and Rogelio Aguilar-Aguilar2

1Universidad Autónoma de Querétaro, Facultad de CienciasNaturales-Biología Laboratorio de Parasitología,

Querétaro, México.2Universidad Nacional Autónoma de México, Instituto de

Biología, Departamento de Zoología, México, DF, México.3Laboratorio de Ictiología y Limnología, Escuela Nacional de

Ciencias Biológicas, Instituto Politécnico Nacional,México, DF, México.

• Viviparous Fishes Genetics, Ecology, and Conservation438

Helminth Parasitesof Viviparous

Fishes in Mexico

Raúl Pineda-López,Guillermo Salgado-

Maldonado,Eduardo Soto-Galera,

Norma Hernández-Camacho, Anita

Orozco-Zamorano,Salvador Contreras-

Robledo, GuillerminaCabañas-Carranza,

and Rogelio Aguilar-Aguilar

Abstract

This study is aimed at collecting the extant publisheddata on the helminth parasites of the MexicanGoodeidae and Poeciliidae, as well as contributingoriginal data from the author’s research, with the endof describing the taxonomic composition of thisfauna and exploring their evolutionary andzoogeographic origins. The parasitological data in thisstudy cover 9 goodeid species from 7 genera, and 20poeciliid species from 6 genera, collected from theLerma, Santiago, Pánuco, Balsas, Ayuquila, andGrijalva-Usumacinta basins as well as from theYucatán Peninsula, Mexico. A total of 32 helminthspecies were recorded in the collected goodeid fishspecies. The data demonstrate that Posthodiplostomumminimum (MacCallum, 1921) and Clinostomumcomplanatum (Rudolphi, 1814) metacercariae arefrequent in the goodeids. The most abundanthelminth record in the collected goodeid host specieswas found in Goodea atripinnis, which had 16helminth species, and in Girardinichthys multiradiatusand Alloophorus robustus, which each had 13 helminthspecies. A total of 46 helminth species were recordedin the 20 collected poeciliid fish. The metacercariae ofPosthodiplostomum minimum (MacCallum, 1921)and Centrocestus formosanus (Nishigori, 1924) are veryfrequent in Mexican poeciliids. The host speciesPoecilia mexicana was found to have the mostabundant helminth community, with 24 species. Thedata indicate 4 components that determine helminthparasite community composition in these fishfamilies, which are described below in order of thenumber of helminth species in each component.1) Generalist, allogenic, mostly Nearctic helminthspecies. They have colonized goodeids and poeciliidsas secondary intermediate hosts or paratenic hostsand conclude their lifecycle in birds. 2) Helminthspecies commonly associated with other fish species.Goodeid and poeciliid species have acquired these viasympatry with other fish species. 3) Anthropogenicallyintroduced helminth species. 4) Specialist helminthspecies. These appear to restrict their parasiticrelationship to goodeid or poeciliid fish. They havedeveloped, or evolved, together with their hosts andare probably endemic or ancient species. The datapresented here also indicate the numericalpredominance of trematode and nematode species,the low representation of monogeneans and cestodes,and only a few species records for theacanthocephalan group. A large allogenic componentis also characteristic of helminth communities inMexican freshwater fish. The helminth communitiesrecorded in Poeciliidae were notably richer than thoseobserved in the Goodeidae.

Resumen

El presente estudio recaba datos bibliográficos y otrosoriginados de nuestras propias investigaciones, paradescribir la composición taxonómica de lascomunidades de helmintos parásitos de peces de lasfamilias Goodeidae y Poeciliidae de México, yexplorar sus orígenes evolutivos y zoogeográficos. Losdatos parasitológicos que se presentan incluyen 9especies de goodeidos de 7 géneros y 20 especies depoecílidos de 6 géneros, recolectados de las cuencas delos ríos Lerma, Santiago, Pánuco, Balsas, Ayuquila yGrijalva-Usumacinta, así como de los cuerpos de aguade la Península de Yucatán. Se registran un total de 32especies de helmintos parásitos de goodeidos. Losdatos demuestran que las metacercarias dePosthodiplostomum minimum (MacCallum, 1921) ylas de Clinostomum complanatum (Rudolphi, 1814)son frecuentes entre los peces de esta familia. Elregistro helmintológico más abundante entre lasespecies de goodeidos examinadas lo presentó Goodeaatripinnis con 16 especies de helmintos, en tanto queGirardinichthys multiradiatus y Alloophorus robustus,presentaron cada una 13 especies de helmintos. En las20 especies de poecílidos se registraron 46 especies dehelmintos las metacercarias de Posthodiplostomumminimum (MacCallum, 1921) y de Centrocestusformosanus (Nishigori, 1924) son los helmintos másfrecuentes entre los poecílidos de México. Poeciliamexicana es la especie de hospedero con mayornúmero de especies de helmintos, con 24 especies.Pueden distinguirse 4 componentes de estascomunidades, por orden del número de especies queincluyen, éstos son: 1) componente de especiesgeneralistas, alogénicas, en su mayoría neárticas, quehan colonizado a los goodeidos y poecílidos comohospederos intermediarios secundarios o bien, comohospederos paraténicos, y que concluyen su ciclo devida en aves. 2) Especies de helmintos adquiridas porsimpatría con otras especies de peces; estas especiesson comunes en otros grupos de peces. 3) Especiesintroducidas antropogénicamente. 4) Especiesespecialistas de helmintos. Éstas se han desarrolladocon sus hospederos y probablemente son endémicas ymuy antiguas. Los datos también muestran lapredominancia numérica de las especies detremátodos y nemátodos; el número reducido deespecies de monogéneos y céstodos, en tanto que sólose registró una especie de acantocéfalo. Uncomponente numeroso de especies alogénicas estambién una característica común entre los helmintosparásitos de peces de agua dulce de México.Finalmente, es notable que las comunidades dehelmintos de los Poeciliidae son mucho más ricas quelas de los Goodeidae.

• Viviparous FishesHarry J. Grier and Mari Carmen Uribe, book editors.New Life Publications, Homestead, Florida, 2005. p 437-456.


Introduction

Viviparous fish in Mexico belong to thefamilies Goodeidae and Poeciliidae,both in the order Cyprinodontiformes.

The Goodeinae is endemic to the Mexican pla-teau and is currently composed of 16 genera, in-cluding: Allodontichthys, Alloophorus, Allotoca,Ameca, Ataenobius, Chapalichthys, Characodon,Girardinichthys, Goodea, Hubbsina, Ilyodon,Skiffia, Xenophorus, Xenotaenia, Xenotoca andZoogoneticus. The Poeciliidae are neotropical inorigin and are widely distributed, ranging fromthe southern United States to Argentina. Severalgenera of Poeciliids have been recorded in Me-xico: Belonesox, Brachyrhaphis, Carlhubbsia,Gambusia, Heterandria, Phallichthys, Poecilia,Poeciliopsis, Priapella, Xenodexia, and Xipho-phorus (Espinosa-Pérez et al., 1993).

Data on the helminth parasites of some Mexi-can goodeids and poeciliids have been publishedas part of the general inventories of helminthsof freshwater fish (Moravec et al., 1995a,b;Scholz et al., 1995a,b, 1996; Salgado-Maldo-nado et al., 2001a,b, 2004a,b; Aguilar-Aguilaret al., 2004; Martínez-Aquino et al., 2004).Taxonomic treatments of a particular helminthspecies have also been published (Pérez, 2001).The parasitology of these fish families, however,has not been addressed holistically, and the avail-able data have not been analyzed as a whole. Inresponse, this study is aimed at collecting theextant published data on the helminth parasitesof the Mexican Goodeidae and Poeciliidae, aswell as contributing original data from theauthor’s research, with the end of describing the

taxonomic composition of this fauna and ex-ploring their evolutionary and zoogeographicorigins.

Materials and Methods

The data included in the results section comesfrom the freshwater fish helminth parasite data-base for Mexico assembled by the second authorof this study and financed by the National Com-mission for Study and Exploitation of Biotic Re-sources (Comisión Nacional para el Estudio yAprovechamiento de los Recursos Bióticos–Conabio). This database will be available on theConabio webpage (www//http.conabio.xolo.mx),and portions of these data have been publishedpreviously as inventories (Scholz et al., 1995a,b,1996; Moravec et al., 1995a,b; Salgado-Mal-donado et al., 2001a,b, 2004a,b; Aguilar-Aguilaret al., 2004). Original data from the authors’ ownresearch is also included in the present study.

Data were collected by catching goodeid andpoeciliid fish in chosen locations in a numberof the hydrological basins of Mexico, includingthe Lerma, Santiago, Pánuco, Balsas, Ayuquilain the Sierra de Manantlán, Grijalva-Usuma-cinta and surface waters of the Yucatán Penin-sula. The collected fish were kept alive untilhelminthological examination was done 8 hoursafter capture. The examination included all or-gan systems, except the blood and bones. Dataon the number of helminth species and numberof helminth individuals per species were recov-ered during the examinations. Every helminth


found in every fish host was counted andsamples were taken for morphological study andtaxonomic determination (see Salgado-Mal-donado, 1979). Details of the methodology, andcollection dates and locations can be found inSalgado-Maldonado et al., 2001a,b, 2004a,b;Aguilar-Aguilar et al., 2004. Taxonomic deter-minations for monogeneans, trematodes, ces-todes, nematodes and acanthocephalans werevalidated with the help of the taxonomic spe-cialists E. Mendoza Franco (Cinvestav-IPNUnidad Mérida [monogeneans]), F. Moravec(nematodes), and T. Scholz (Academy of Sci-ences of the Czech Republic [cestodes and meta-cercariae]). Vouchers of all collected helminthspecies were deposited in the National HelminthCollection of the Institute of Biology, NationalAutonomous University of Mexico; the ParasiteCollection of the Autonomous University ofQuerétaro; the Parasitological Collection of theAutonomous University Juárez of Tabasco; andthe Parasitological Collection of the Academyof Sciences of the Czech Republic (IPCAS).

Results

The parasitological data in this study cover 9goodeid species from 7 genera, and 20 poeciliidspecies from 6 genera, collected from the Lerma,Santiago, Pánuco, Balsas, Ayuquila, and Grijal-va-Usumacinta basins as well as from the Yuca-tán Peninsula (Tables 1 and 2).

Goodeidae

A total of 32 helminth species were recorded inthe collected goodeid fish species. These in-cluded 5 adult trematodes and 8 metacercariae;1 monogenean; 2 adult cestodes and 6 metaces-todes; 4 adult nematodes and 5 larvae, and 1cystacanth (Table 1). Of these 32 species, 26(81%) were found in the Lerma and Santiago,11 (34%) in the Ayuquila, 4 (12%) in the Bal-sas, and only 1 (3%) in the Pánuco. All the hel-minths collected from goodeids in the Ayuquila,Balsas and Pánuco rivers were also present in thefish from the waters of the Lerma and Santiagobasins. The only exception were five species re-corded from the Ayuquila and not found in theLerma-Santiago: the trematodes Dendrorchis sp.,Saccocoelioides cf. sogandaresi Lumsden, 1961,Magnivitellinum simplex Kloss, 1966, Ascocotyle(Ascocotyle) tenuicollis Price, 1935, and thenematode Capillaria cyprinodonticola Huffmanand Bullock, 1973.

The data demonstrate that Posthodiplos-tomum minimum MacCallum, 1921, andClinostomum complanatum Rudolphi, 1814,metacercariae are frequent in the goodeids, withthe former parasitizing 7 fish species and thelatter 6 fish species. The cestode Bothriocephalusacheilognathi Yamaguti, 1934, was recorded in5 goodeid fish species, while the adult trema-tode Margotrema bravoae Lamothe-Argumedo,1970, the nematode Rhabdochona lichtenfelsiSánchez-Álvarez, García-Prieto and Pérez, 1998,and the Spiroxys sp. nematode larvae all parasitize4 goodeid species. The nematode R. lichthenfelsiwas recorded in goodeids from 4 of the studiedhydrological basins.

The most abundant helminth record in thecollected goodeid host species was found inGoodea atripinnis, which had 16 helminth spe-cies, and in Girardinichthys multiradiatus andAlloophorus robustus, which each had 13 hel-minth species. The remaining goodeid specieshad from 2 to 10 helminth species (Table 1).

Of the 32 helminth species, 50% are autoge-nic, they complete their lifecycle in the samebody of water, and the other 50% are allogenic,mainly maturing in birds. Almost all the hel-minths recorded in goodeids are widely distrib-uted generalists and have been recorded in otherhost species. Two nominal helminth species areexclusive to goodeids: Margotrema bravoae andRhabdochona lichthenfelsi. There is also a clearlydistinguishable anthropogenically introducedcomponent in the goodeids of the Mexicanhighland plateau, which includes C. formosanusmetacercariae, the monogenean Gyrodactylus cf.elegans von Nordmann, 1832, the cestode B.acheilognathi, and the nematode Psudocapillariatomentosa Dujardin, 1843.

Poeciliidae

A total of 46 helminth species were recorded inthe 20 collected poeciliid fish species, including2 adult trematodes, 21 metacercaria, 5 monoge-neans, 1 adult cestode, 6 metacestodes, 4 adultand 5 larval nematodes, 1 acanthocephalan, and1 cystacanth (Table 2). Of the 46 helminth spe-cies, 23 (50%) were found in the Grijalva-Usumacinta, 20 (43%) in the Pánuco, 17 (37%)from the Yucatán Peninsula, 13 (28%) in theBalsas, 14 (30%) in the Papaloapan, 11 (24%) inthe Ayuquila, and 10 (22%) in the Lerma-San-tiago. Some of the helminth species collectedfrom poeciliids from the Yucatán Peninsula arenot present in other hydrological basins, includ-


ing A. ? astorquii, A. (Leighia) megacephala, A.(Phagicola) diminuta, D. papillifera and Fal-caustra sp. The same is true of poeciliids from theGrijalva-Usumacinta basin, the only basin inwhich P. bychowskyi, R. ondatrae, D. (Austro-diplostomum) compactum, E. leopoldinae, A.(Leighia) chandleri, A. (L.) mcintoshi, A. (Phagi-cola) macrostoma, A. (P.) mollienisicola, Protero-diplostomidae, S. seculus, U. reticulatus andGnathostoma sp. were recorded. Overall, 20 hel-minth species were only found in the poeciliidsof southeast Mexico, i.e. those from the YucatánPeninsula or Grijalva-Usumacinta basin (par-ticularly in Tabasco), including the 17 listedabove plus 3 species recorded in both regions (G.tropica, S. minima and A. (Phagicola) nana.There were also 2 helminth species found only inthe poeciliids of the Papaloapan basin, includingMexiconema cichlasomae Moravec, Vidal-Martí-nez and Salgado-Maldonado, 1992; Spinitectusmexicanus Caspeta-Mandujano, Moravec andSalgado-Maldonado, 2000; and Rhabdochonasp. (a species distinct from others of the samegenus and which are recorded in this study). Themetacercariae of Posthodiplostomum minimumMacCallum, 1921, and Centrocestus formosanusNishigori, 1924, are very frequent in Mexicanpoeciliids, the former parasitizing 12 poeciliidspecies and the latter 11. The cestode Bothrio-cephalus acheilognathi Yamaguti, 1934, was re-corded in 10 host species as were Uvulifer sp.metacercaria. Other abundant species includedthe trematode S. cf. sogandaresi and Contra-caecum sp. larvae, both of which were recordedin 9 poeciliid host species.

The cestode B. acheilognathi and Contra-caecum sp. larvae were the most widely distrib-uted helminths in poeciliids, as they wererecovered in 6 of the sampled hydrological ba-sins. The trematode S. cf. sogandaresi, and P.minimum and C. formosanus metacercaria werealso widely distributed, all of them being re-corded in 5 of the sampled basins.

The host species Poecilia mexicana was foundto have the most abundant helminth commu-nity, with 24 species. This fish species was fol-lowed by P. petenensis, which had 20 helminthspecies, and Gambusia yucatana, with 17 spe-cies. The remaining poeciliid species hostedfrom 2 to 15 helminth species (Table 2).

Of the 46 helminth species recorded inpoeciliids, 20 (43%) are autogenic, completingtheir lifecycle in the same body of water, and 26(57%) are allogenic, mainly maturing in birds.Almost all the helminths recorded in the

Poeciliids are widely distributed generalists andhave been recorded in other host taxa. Three ofthe helminth species, however, appear to be ex-clusive to poeciliids: Urocleidoides reticulatus,Spinitectus mexicanus and Rhabdochona xipho-phori. There is also a clearly distinguishableanthropogenically introduced component inMexican poeciliids, including C. formosanusmetacercaria and the cestode B. acheilognathi.

Discussion

The results highlight the need for more samplingand field work aimed at completing the hel-minth database for goodeids and poeciliids inMexico. To date, only 7 goodeid genera and 9goodeid species of the 19 known genera and 51known species in Mexico have had any studydone of their helminth fauna. The situation issimilar for poeciliids, for which helminth stud-ies have been done for 6 genera and 20 species ofthe 11 known genera and 95 known species inMexico. Only the most common and accessiblegoodeid and poeciliid species have been exam-ined, leaving a data gap for species with morelimited distributions or from rarer populations.

The currently available data is from generalfish samples and samples that only circumstan-tially report goodeid and poeciliid data. The datafrom the present study, in contrast, are based onthe determinations of taxonomic specialists andthe reference material has been deposited in stableinstitutional helminth collections to ensure itsconservation and accessibility for future reference.To bring the level of knowledge on the helminthfauna of goodeids and poeciliids in Mexico to acomparable level of that currently available forcichlid (Cichlidae) parasite fauna (one of the bestunderstood as far as helminths are concerned[Salgado-Maldonado et al., 1997; Vidal-Martínezet al., 2001]), sampling programs are needed thatare directed at chosen goodeid and poeciliid spe-cies. This is particularly difficult since many ofthese species have very limited geographic distri-butions and consist of scarce populations. Manyof the species in these families are also threatenedand require special permits for research collec-tion. Of the two families, the poeciliids are morewidely distributed, though many of the speciesare small and live in microhabitats within bodiesof water that are hard to access and sample, andthus require specialized collection tools and tech-niques.

The data indicate 4 components that deter-mine helminth parasite community composi-


tion in these fish families, which are describedbelow in order of the number of helminth spe-cies in each component. 1) Generalist, allogenichelminth species. These are transported by birdsand have a wide geographic distribution, withmost being Nearctic though some have world-wide distribution. They have colonized goodeidsand poeciliids as secondary intermediate hostsor paratenic hosts and conclude their lifecyclein birds. 2) Helminth species commonly associ-ated with other fish species. Goodeid andpoeciliid species have acquired these by livingin the same locations as other infected fish, thatis, these are helminths acquired via sympatrywith other fish species. 3) Anthropogenicallyintroduced helminth species. 4) Specialist hel-minth species. These appear to restrict theirparasitic relationship to goodeid or poeciliidfish. They have developed, or evolved, togetherwith their hosts and are probably endemic orancient species.

A total of 38% of the recorded helminths ingoodeids, and 57% in poeciliids are allogenic,that is, widely distributed generalist species thatreach sexual maturity in birds and are trans-ported and dispersed by them. This componentin the goodeids includes C. complanatum,Diplostomum sp., P. minimum, Ligula intestinalisLinnaeus, 1758, Cyclustera cf. ralli Underwoodand Dronen, 1986, Valipora campilancristrotaWedl, 1855, Contracaecum sp., Eustrongylidessp., Spiroxys sp., Falcaustra sp., and Polymorphusbrevis Van Cleave, 1916. In poeciliids it includesC. complanatum, Diplostomum (A.) compatum,Diplostomum sp., P. minimum, Uvulifer sp.,Contracaecum sp., Eustrongylides sp., andPolymorphus sp., as well as 9 heterophyd and 5dilepidid species. It also includes the more geo-graphically restricted allogenic species R.ondatrae, Apharingostrigea sp., E. leopoldinae,and Gnathostoma sp. These allogenic helminthspecies have low host specificity and are widelydistributed among freshwater fish in Mexico(Scholz et al., 1995 a,b, 1996; Moravec, 1998;Scholz and Salgado-Maldonado, 2001b;Salgado-Maldonado et al., 2001a,b, 2004a,b;Vidal Martínez et al., 2001). Some are distrib-uted throughout North America, and still othersare found world-wide (Yamaguti, 1975; Gibson,1996; Moravec, 1998; Hoffman, 1999). Meta-cercaria of the heterophyd Ascocotyle spp. arewidely distributed in the fish of southeastMexico and on the Gulf of Mexico slope (Scholzet al., 1997, 2001). Metacestodes of theDilepididae family are widely distributed in the

freshwater fish of the Mexican plateau (Sholzand Salgado-Maldonado, 2001b; Salgado-Maldonado et al., 2001b).

The second component consists of helminthspecies that are infrequent in the parasite com-munities of goodeids and poeciliids and haveinvaded them from the parasite’s primary pre-ferred hosts, i.e. other fish species with whichthey live in sympatry. These include M. simplex,which is frequent in characids (mostly Astyanax)(see Jiménez Guzmán, 1973; Scholz et al.,1995b; Salgado-Maldonado et al., 2001a,2004a,b, 2005); Ascocotyle (Ascocotyle) tenuicollis,a metacercaria frequently associated withcichlids (Scholz et al., 2001; Vidal-Martínez etal., 2001); Proteocephalus pusillus Ward, 1910,which parasitizes salmonids (Schmidt, 1986;Hoffman, 1999); Capillaria cyprinodonticola,which parasitizes cyprinodontids and poeciliids(Moravec, 1998), and Ochetosoma brevicaecum(Caballero y Caballero, 1941) metacercariae,which parasitize amphibians (Jiménez-Ruiz etal., 2002). This second helminth component inpoeciliids includes Genarchella tropica, a para-site in characids; A. ? astorquii, P. bychowskyi, S.minima, R. ondatrae, which mainly parasitizevarious cichlid species in southeast Mexico (Sal-gado-Maldonado et al, 1997; Vidal-Martínez etal., 2001); R. kidderi, a parasite of cichlids; andR. lichtenfelsi, which parasitizes goodeids.

The third, anthropogenically introduced hel-minth component includes Centrocestusformosanus metacercariae, the monogeneanGyrodactylus cf. elegans, the cestode Bothrioceph-alus acheilognathi, and the nematode Pseudo-capillaria tomentosa. All four are found ingoodeids, but only the metacercariae and thecestode were recorded in poeciliids in this study.

Human-introduced helminths are a para-digm. They are the most successful helminthparasites of freshwater fish in Mexico based onthe number of host species they infect and theirwide distribution. The success of Bothriocephalusacheilognathi, for example, is directly favored byaquaculture policies carried out in centralMexico, especially in the highland plateau (seeSalgado-Maldonado and Pineda-López, 2003).“Controlled” production and constant “sowing”of Asian carps is common aquaculture practicein the Mexican altiplano. These fish species arere-introduced year after year into natural bod-ies of water as well as into artificial ones such asreservoirs and “bordos” (artificial, temporaryponds used for storing water during the dry sea-son) (Juárez-Palacios and Palomo-Martínez,


1985; Arredondo-Figueroa and Juárez-Palacios,1986). This Asian cestode’s current wide distri-bution and presence records clearly reflect thispractice. The parasite’s life cycle coincides nicelywith the carp cultivation cycle. The helmintheggs are initially deposited with the host feceson the pond or reservoir bottom. These pondsare then fertilized to aid and increase the pro-duction of the plankton needed to feed the smallcarp raised there. This is generally done with-out disinfecting or applying any treatment toeliminate the cestode eggs in the muddy bot-toms. This cycle ensures the conjunction in timeand space of the parasite eggs, the zooplanktoniccopepods that serve as intermediate hosts, andthe fish, which is the definitive host. The intro-duction of infected carp into reservoirs and otherlarge bodies of water in the Mexican altiplanofavors the infection of native fish with this ces-tode. This parasite has a notable lack of hostspecificity, which complements the already pro-pitious conditions for transmission betweencultivated and wild fish populations. Bothrio-cephalus acheilognathi has been recorded in 49freshwater fish species from 26 genera 7 fami-lies and 5 orders in Mexico (Salgado-Maldonadoand Pineda-López, 2003).

These practices of repeated introduction anddistribution of non-native carp have also favoredother parasites such as P. tomentosa and G. cf.elegans (Moravec et al., 2001; Salgado-Maldo-nado and Pineda-López, 2003).

Dispersion of C. formosanus metacercaria inMexico appears to be related to distribution ofits first intermediate host, the snail Thiaratuberculata. This species was initially introducedinto the Caribbean, the southern United States,and South America to help in controlling othersnail species that transmit schistosomiasis (seeScholz and Salgado-Maldonado, 2001a). T.tuberculata disperses actively and can establishpopulations in new environments with relativeease. It grows quickly and can reach sizablepopulation densities in a short time. These in-termediate host characteristics, coupled with thelack of host specificity in C. formosanus meta-cercariae, favor this trematode’s transmission rateand it is spread to a wide variety of fish species(Scholz and Salgado-Maldonado, 2001a).

Introduced helminth species’ aggressivenessand pathology in native fish species has beenthoroughly documented (Hoffman, 1999;Scholz and Salgado-Maldonado, 2001a;Salgado-Maldonado and Pineda-López, 2003).However, the effect of these parasite species on

wild fish populations has not yet been reliablytested. Goodeids, and especially poeciliids, playa role in this dispersion because they are excel-lent distributors of introduced parasite speciesto other native fish species. Poeciliids are knownto play this role in other parts of the world, suchas Hawaii (Font, 1997), Australia (Dove, 2000),and in the sinkholes of the Yucatán Peninsula(Scholz et al., 1996).

The fourth component in the helminth com-munities of Mexican goodeids and poeciliidsconsists of host-specific species, which have de-veloped with these host families. Nematodes ofthe family Rhabdochonidae, in particular, havewidely diversified in Mexican freshwater fish(Salgado-Maldonado et al., 2004b). Rhabdo-chona lichthenfelsi appears to have developed inassociation with goodeids in the Mexican high-land plateau, as indicated by its host records inthe present study. Trematodes of the genusMargotrema have also been collected only ingoodeids from the Mexican highland plateau.Although a second species has recently beendescribed for this genus (Pérez, 2001), the mor-phological variability of the M. bravoaeLamothe-Argumedo, 1970, type species is stillnot well studied. The differences between thesetwo species are very tenuous and the argumentsin favor of the second quite weak. None the less,the genus and type species validity are undis-puted, meaning that in the Mexican altiplanosome helminth parasites have speciated alongwith goodeids and are specific to these hosts.

To date, the two helminth species apparentlyhost-specific to poeciliids are Rhabdochonaxiphophori Caspeta-Mandujano, Moravec andSalgado-Maldonado 2000, and Spinitectusmexicanus Caspeta-Mandujano, Moravec andSalgado-Maldonado 2000, both of which haveonly been collected from poeciliids. It is also pos-sible that the trematodes indicated as Sacco-coelioides cf. sogandaresi, which includes acomplex of species that have not yet been mor-phologically differentiated, is closely associatedwith poeciliids. This is supported by their pres-ence in the records and their abundance data inthe different basins (Salgado-Maldonado et al.,2001a,b, 2004a,b, 2005).

The relative poorness of the endemic hel-minths of the Mexican highland plateau is likelydue to their young geological age. Salgado-Maldonado et al. (2001b) suggest that helminthparasite communities of the Mexican altiplano,especially those in goodeids, are no more than 5million years old. This, consequently, has con-


ditioned their poorness in terms of host speciesspecificity, which makes them invadable by in-troduced species. The presence of only threenominal helminth species specific to this fishfamily is congruent with this proposal, especiallyif the geological age of the fresh water bodies inthe altiplano and the evolutionary differentia-tion of the Goodeidae are taken into account(Barbour, 1973; Echelle and Echelle, 1984).This relative youth and consequent species poor-ness make these communities imminentlyinvadeable by allogenic, generalist and anthro-pogenically introduced helminth species.

Based on this, it is probable that the ances-tral marine group from which the goodeidsoriginated lost most of its parasites after beingisolated in the freshwater bodies of the Mexicanaltiplano during the Pliocene, 5 million yearsago (Barbour, 1973; Echelle and Echelle, 1984).However, some of these parasites persisted, suchas the ancestors of the current genus Margo-trema, which later differentiated in isolationwith their hosts. Other parasites such as theRhabdochona nematodes experienced an adap-tive radiation process, dispersing themselvesamong the fish species of central Mexico, withvarious helminth species developing in differ-ent host groups (Caspeta-Mandujano et al.,2000a, 2001; Salgado Maldonado et al., 2004b,2005). In the case of R. lichtenfelsi, the currentdata appear to indicate that this species differ-entiated in conjunction with the goodeids. Dur-ing later evolutionary development, thehelminth community of the goodeids, initiallyquite poor, was enriched by the invasion of al-logenic generalist species and by the set of spe-cies acquired through sympatry with other fishgroups. The current helminth community com-position of goodeids now exhibits the effect ofinvasion by anthropogenically introduced hel-minth species.

The presence of allogenic generalist helminthspecies with wide geographic distribution, andtransported by birds, are a constant in the Mexi-can freshwater fish studied to date (Salgado-Maldonado et al., 2001b). In some hydrologicbasins, the Lerma and the Santiago, for example,the allogenic species component is more signifi-cant in terms of number of species than in other

basins. This phenomenon is related to the smallsize of freshwater fish like the goodeids, and theirhabitat in shallow waters along the migrationroutes of Nearctic birds (Salgado-Maldonado etal., 2001b). Like the allogenic species, the in-troduced helminth species are excellent coloniz-ers and tend to occupy the “empty niches” (sensuKennedy, 1990) they find in these species-poorhelminth communities.

The geographic origins of the poeciliids aredifferent than those of the goodeids, but thecurrent conformation of helminth communitiesindicates that their development has followed asimilar path. In other words, these young com-munities with few specialists are invadable byallogenic helminth species and species fromother fish groups. For viviparous fish, popula-tions can be started with a single founding fe-male, suggesting a limited infection source inrecently established, dynamic populations, andpartially explaining these helminth communi-ties’ species poorness.

The data presented here also indicate that byanalyzing the taxonomic composition of hel-minth communities in a single host family, thesame described patterns are reflected in the dif-ferent studied hydrological basins (Salgado-Maldonado et al., 2001a,b, 2004a,b, 2005).These patterns include the numerical predomi-nance of trematode and nematode species, thelow representation of monogeneans and ces-todes, and only a few species records for theacanthocephalan group. A large allogenic com-ponent is also characteristic of helminth com-munities in Mexican freshwater fish.

The helminth communities in poeciliidsshown in the inventory exhibit variable richness(from 2 to 23 species). The host species withthe fewest parasites, however, appear to showthis trend because of insufficient sampling orgeographically restricted distribution (e.g., P.latipunctata from the Yucatán Peninsula, P.reticulata from the Balsas and P. butteri from theAyuquila). Despite this, the helminth commu-nities recorded in Poeciliidae were notably richerthan those observed in the Goodeidae. This mayresult from the Poeciliidae being neotropical,their wider geographic distribution, and thedensity and contiguity of their populations.


Table 1.Parasite host list of helminths collected from viviparous fishes of Mexico:Goodeidae: A. = Allophorus, G. = Girardinichthys, G. = Goodea atripinnis, I. = Ilyodon, X. = Xenotoca;Basin, BR = Río Balsas, L = Río Lerma, PS = Río Pánuco, SM = Río Ayuquila, Sierra de Manantlán.

Parasite Host Site of infection Basin Reference

Adult trematodaFamily Gorgoderidae

Dendrorchis sp. I. furcidens Intestine SM Salgado-Maldonado et al., 2004aFamily Haploporidae

Saccocoelioides cf.sogandaresi A. zonistius Intestine SM Salgado-Maldonado et al., 2004a

I. furcidens Intestine SM Salgado-Maldonado et al., 2004aI. whitei Intestine BR Salgado-Maldonado et al., 2001a

Family MacroderoididaeMagnivitellinum simplex I. furcidens Intestine SM Salgado-Maldonado et al., 2004aMargotrema bravoae A. robustus Intestine L Pérez, 2001

A. diazi Intestine L Pérez, 2001A. zonistius Intestine SM Salgado-Maldonado et al., 2004aG. multiradiatus Intestine L Sánchez-Nava et al., 2004

Margotrema guillerminae A. robustus Intestine L Pérez, 2001Metacercariae

Family ClinostomidaeClinostomum complanatum A. robustus Liver, mesentery L Salgado-Maldonado et al., 2001b

A. diazi Liver, mesentery L Salgado-Maldonado et al., 2001bA. zonistius Body cavity SM Salgado-Maldonado et al., 2004aG. atripinnis Liver L Salgado-Maldonado et al., 2001bI. furcidens Body cavity SM Salgado-Maldonado et al., 2004aX. variatus ? L Salgado-Maldonado et al., 2001b

Family DiplostomidaeDiplostomum sp. G. atripinnis ? L Salgado-Maldonado et al., 2001bPosthodiplostomumminimum A. robustus Liver, mesentery L Salgado-Maldonado et al., 2001b

muscle, eyesA. diazi Liver, muscle, L Pérez et al., 2000

MesenteryA. zonistius Mesentery SM Salgado-Maldonado et al., 2004aG. multiradiatus Mesentery, liver L Sánchez-Nava et al., 2004G. atripinnis Liver, muscle, L Salgado-Maldonado et al., 2001b

mesentery, eyesMuscle, liver, BR Salgado-Maldonado et al., 2001aeyes, mesentery,body cavity

I. furcidens Mesentery SM Salgado-Maldonado et al., 2004aX. variatus ? L Salgado-Maldonado et al., 2001b

Tylodelphys sp. G. multiradiatus Body cavity, L Sánchez-Nava et al., 2004mesentery, eyes

G. atripinnis Body cavity L Salgado-Maldonado et al., 2001b



Family HeterophyidaeAscocotyle (Ascocotyle)tenuicollis A. zonistius Heart SM Salgado-Maldonado et al., 2004aCentrocestus formosanus I. furcidens Gills SM Salgado-Maldonado et al., 2004a

I. whitei Gills BR Salgado-Maldonado et al., 2001aG. atripinnis Gills L Salgado-Maldonado et al., 2001b

Family PlagiorchiidaeOchetosoma brevicaecum G. multiradiatus Mesentery L Sánchez-Nava et al., 2004

A. diazi Intestine L Salgado-Maldonado et al., 2001bG. atripinnis Intestine L Pérez et al., 2000

MonogeneaFamily Gyrodactylidae

Gyrodactylus cf. elegans G. multiradiatus Fins L Sánchez-Nava et al., 2004Adult cestoda

Family BothriocephalidaeBothriocephalus acheilognathi

A. robustus Intestine L Salgado-Maldonado et al., 2001bA. diazi Intestine L Salgado-Maldonado et al., 2001bG. multiradiatus Intestine L Sánchez-Nava et al., 2004G. atripinnis Intestine L Salgado-Maldonado et al., 2001bX. variatus Intestine L Salgado-Maldonado et al., 2001b

Family ProteocephalidaeProteocephalus pusillus G. atripinnis Intestine L Salgado-Maldonado et al., 2001b

MetacestodesFamily Cyclophyllidea

Cyclophyllidea gen. sp. A. diazi Gall bladder L Pérez et al., 2000Family Diphyllobothriidae

Ligula intestinalis G. atripinnis Body cavity L Salgado-Maldonado et al., 2001bG. multiradiatus Body cavity L Sánchez-Nava et al., 2004

Family DilepididaeCyclustera cf. ralli G. multiradiatus Mesentery L Sánchez-Nava et al., 2004

X. variatus Mesentery L Salgado-Maldonado et al., 2001bValipora campylancristrota G. multiradiatus Gall bladder L Sánchez-Nava et al., 2004Dilepididae gen. sp. A. zonistius Mesentery SM Salgado-Maldonado et al., 2004a

I. furcidens Mesentery SM Salgado-Maldonado et al., 2004aFamily Proteocephalidae

Proteocephalidae gen. sp. A. robustus Liver, intestine L Salgado-Maldonado et al., 2001bmesentery

A. diazi Liver, intestine, L Salgado-Maldonado et al., 2001bmesentery

G. atripinnis Liver, intestine, L Salgado-Maldonado et al., 2001bmesentery



Adult nematodesFamily Capillaridae

Capillaria cyprinodonticola I. furcidens Intestine SM Salgado-Maldonado et al., 2004aPseudocapillaria tomentosa A. robustus Intestine L Salgado-Maldonado et al., 2001b

G. atripinnis Intestine L Salgado-Maldonado et al., 2001bFamily Rhabdochonidae

Rhabdochona lichthenfelsi A. robustus Intestine L Salgado-Maldonado et al., 2001bA. diazi Intestine L Salgado-Maldonado et al., 2001bG. atripinnis Intestine L Salgado-Maldonado et al., 2001b

Intestine BR Salgado-Maldonado et al., 2001aIntestine PR Salgado-Maldonado et al., 2004b

I. furcidens Intestine SM Salgado-Maldonado et al., 2004aLarval nematodes

Family AnisakidaeContracaecum sp. A. robustus Mesentery L Pérez et al., 2000

G. multiradiatus Mesentery L Sánchez-Nava et al., 2004G. atripinnis Mesentery, L Salgado-Maldonado et al., 2001b

body cavityX. variatus Mesentery L Salgado-Maldonado et al., 2001b

Family DioctophymatidaeEustrongylides sp. A. robustus Mesentery L Pérez et al., 2000

G. atripinnis Intestine L Salgado-Maldonado et al., 2001bFamily Kathlaniidae

Falcaustra sp. G. multiradiatus Mesentery, L Sánchez-Nava et al., 2004intestine

Family GnathostomatidaeGnathostoma sp. A. robustus Liver L Salgado-Maldonado et al., 2001bSpiroxys sp. A. robustus Mesentery, L Salgado-Maldonado et al., 2001b

intestineA. diazi Mesentery, L Salgado-Maldonado et al., 2001b

intestineG. multiradiatus Mesentery, L Sánchez-Nava et al., 2004

intestineX. variatus Intestine L Salgado-Maldonado et al., 2001b

Acanthocephala larvaeFamily Polymorphidae

Polymorphus brevis A. robustus Mesentery, L Salgado-Maldonado et al., 2001bmuscle

A. diazi Mesentery L Salgado-Maldonado et al., 2001bG. multiradiatus Mesentery L Sánchez-Nava et al., 2004G. atripinnis Intestine L Salgado-Maldonado et al., 2001bX. variatus Intestine L Salgado-Maldonado et al., 2001b


Table 2.Parasite host list of helminths collected from viviparous fishes of Mexico: PoeciliidaeBasin: BR = Río Balsas, CY = Yucatán Peninsula, GU = Grijalva-Usumacinta basin, L = Río Lerma, PP = Río Papaloapan,PS = Río Pánuco, SM = Río Ayuquila, Sierra de Manantlán, S = Río Santiago, MO = Sierra Madre Oriental.


Adult trematodaFamily Derogenidae

Genarchella tropica G. yucatana Intestine CY Scholz et al., 1995aP. petenensis Intestine GU Salgado-Maldonado et al., 2005

Family HaploporidaeSaccocoelioides cf.sogandaresi P. latipunctata Intestine CY Scholz et al., 1995a

P. mexicana Intestine PR Salgado-Maldonado et al., 2004bIntestine GU Salgado-Maldonado et al., 2005Intestine PP Present work

P. petenensis Intestine GU Salgado-Maldonado et al., 2005P. sphenops Intestine BR Salgado-Maldonado et al., 2001aP. gracilis Intestine BR Salgado-Maldonado et al., 2001a

Intestine PR Salgado-Maldonado et al., 2004bP. velifera CY Scholz et al., 1995bPoecilia sp. Intestine CY Scholz et al., 1995aX. helleri Intestine SM Salgado-Maldonado et al., 2004a

Intestine PP Present workXiphophorus sp. Intestine PR Salgado-Maldonado et al., 2004b

MetacercariaeFamily Acanthostomidae

Atrophecaecum ? astorquii G. yucatana Fins CY Scholz et al., 1995bP. latipunctata Fins CY Scholz et al., 1995bP. petenensis Fins CY Scholz et al., 1995bP. velifera CY Scholz et al., 1995b

Perezitrema bychowskyi G. yucatana GU Salgado-Maldonado et al., 2005Stunkardiella minima G. yucatana Fins CY Scholz et al., 1995b

P. petenensis GU Salgado-Maldonado et al., 2005Family Cathaemasiidae

Ribeiroia ondatrae P. petenensis GU Salgado-Maldonado et al., 2005Family Clinostomidae

Clinostomum complanatum H. bimaculata Mesentery PR Present workP. mexicana Muscle PR Salgado-Maldonado et al., 2004b

Muscle L Present workBody cavity PP Present workeyes

P. gracilis Mesentery PR Present workP. infans Liver L Present workX. helleri Gills PP Present work



Family DiplostomidaeDiplostomum (Austrodiplostomum)compactum P. mexicana GU Salgado-Maldonado et al., 2005

P. petenensis GU Salgado-Maldonado et al., 2005Diplostomum sp. P. mexicana Eyes PR Salgado-Maldonado et al., 2004b

P. sphenops Eyes, brain, BR Salgado-Maldonado et al., 2001abody cavity L Salgado-Maldonado et al., 2001b

P. reticulata Eyes, brain, BR Salgado-Maldonado et al., 2001abody cavity

P. gracilis Eyes PR Salgado-Maldonado et al., 2004bPosthodiplostomumminimum B. belizanus GU Salgado-Maldonado et al., 2005

G. yucatana GU Salgado-Maldonado et al., 2005G. vittata Mesentery, fat PR Salgado-Maldonado et al., 2004b

liver, muscleH. bimaculata Muscle, eyes BR Salgado-Maldonado et al., 2001a

liverP. butleri Mesentery SM Salgado-Maldonado et al., 2004aP. mexicana Liver, PR Salgado-Maldonado et al., 2004b

mesentery, fatbody cavitymuscle, kidney,eyes, intestineMesentery L Present work

P. mexicana GU Salgado-Maldonado et al., 2005P. petenensis GU Salgado-Maldonado et al., 2005P. sphenops Mesentery, BR Salgado-Maldonado et al., 2001a

body cavityMesentery L Present workMesentery L Salgado-Maldonado et al., 2001b

P. gracilis Fat PR Salgado-Maldonado et al., 2004bP. infans Muscle, liver BR Salgado-Maldonado et al., 2001a

Liver, mesentery L Salgado-Maldonado et al., 2001bX. helleri Mesentery SM Salgado-Maldonado et al., 2004aXiphophorus sp. Muscle, eyes PR Salgado-Maldonado et al., 2004b

fatUvulifer sp. G. yucatana CY Scholz et al., 1995b

P. butleri Skin SM Salgado-Maldonado et al., 2004aP. mexicana Skin PR Salgado-Maldonado et al., 2004bP. sphenops Skin, fins BR Salgado-Maldonado et al., 2001aP. petenensis Muscle CY Scholz et al., 1995bP. velifera CY Scholz et al., 1995bP. baenschi Skin SM Salgado-Maldonado et al., 2004aP. gracilis Skin, fins BR Salgado-Maldonado et al., 2001a

Fins PP Present workX. helleri Skin SM Salgado-Maldonado et al., 2004aXiphophorus sp. Skin PR Salgado-Maldonado et al., 2004b



Family EchinostomatidaeEchinochasmus leopoldinae G. yucatana GU Salgado-Maldonado et al., 2005

P. mexicana GU Salgado-Maldonado et al., 2005P. petenensis GU Salgado-Maldonado et al., 2005

Family HeterophyidaeAscocotyle(Ascocotyle) tenuicollis B. belizanus GU Salgado-Maldonado et al., 2005

G. yucatana GU Salgado-Maldonado et al., 2005G. vittata Mesentery PR Salgado-Maldonado et al., 2004bH. bimaculata Heart PP Present workP. butleri Heart, gills SM Salgado-Maldonado et al., 2004aP. latipunctata Heart, mesentery CY Scholz et al., 1995bP. mexicana GU Salgado-Maldonado et al., 2005

Heart PP Present workP. petenensis Heart, mesentery CY Scholz et al., 1995b

GU Salgado-Maldonado et al., 2005P. velifera CY Scholz et al., 1995bX. helleri Fat PP Present work

Ascocotyle(Leighia) chandleri G. yucatana GU Salgado-Maldonado et al., 2005Ascocotyle(Leighia) megacephala

P. velifera Intestinal wall CY Scholz et al., 1995bGU Salgado-Maldonado et al., 2005

Ascocotyle(Phagicola) diminuta P. petenensis Gills CY Scholz et al., 1995b

P. velifera CY Scholz et al., 1995bAscocotyle(Phagicola) macrostoma

B. belizanus GU Salgado-Maldonado et al., 2005Ascocotyle(Phagicola) mollienisicola


Ascocotyle(Phagicola) nana B. belizanus GU Salgado-Maldonado et al., 2005

G. yucatana Mesentery CY Scholz et al., 1995bH. bimaculata Intestinal wall PP Present workP. petenensis Mesentery CY Scholz et al., 1995b

GU Salgado-Maldonado et al., 2005P. velifera CY Scholz et al., 1995bX. helleri Mesentery SM Salgado-Maldonado et al., 2004a

Intestinal wall PP Present work



Centrocestus formosanus G. yucatana Gills GU Salgado-Maldonado et al., 2005Heterandria sp. Gills PP Scholz and Salgado-Maldonado, 2001aP. gracilis Gills BR Salgado-Maldonado et al., 2001a

Gills PR Salgado-Maldonado et al., 2004bP. mexicana Gills PR Salgado-Maldonado et al., 2004b

GU Salgado-Maldonado et al., 2005Gills L Present workGills PP Present work

P. petenensis GU Salgado-Maldonado et al., 2005P. reticulata Gills PP Present workP. sphenops Gills BR Salgado-Maldonado et al., 2001a

Gills L Salgado-Maldonado et al., 2001bGills PP Scholz and Salgado-Maldonado, 2001a

P. baenschi Gills SM Salgado-Maldonado et al., 2004aP. butleri Gills SM Salgado-Maldonado et al., 2004aP. infans Gills L Salgado-Maldonado et al., 2001bPoeciliopsis sp. Gills L Salgado-Maldonado et al., 2001bX. helleri Gills SM Salgado-Maldonado et al., 2004a

Gills PP Scholz and Salgado-Maldonado, 2001aXiphophorus sp. Gills PR Salgado-Maldonado et al., 2004b

Family ProterodiplostomidaeProterodiplostomidae gen. sp.

P. petenensis GU Salgado-Maldonado et al., 2005Family Strigeidae

Apharingostrigea sp. P. mexicana Body cavity PR Salgado-Maldonado et al., 2004bP. gracilis Body cavity PR Salgado-Maldonado et al., 2004b

MonogeneaFamily Dactylogyridae

Salsuginus neotropicalis B. belizanus Gills CY Mendoza-Franco and Vidal Martínez, 2001Salsuginus seculus G. yucatana Gills GU Salgado-Maldonado et al., 2005Urocleidoides reticulatus B. belizanus GU Salgado-Maldonado et al., 2005


Urocleidoides sp. G. yucatana Gills CY Scholz et al., 1995aP. mexicana Gills PR Present work

Dactylogyridae gen. sp. Xiphophorus sp. Gills PR Salgado-Maldonado et al., 2004bFamily Gyrodactylidae

Gyrodactylus sp. G. yucatana Gills CY Scholz et al., 1995aP. mexicana Fins PR, L Present work

Fins PP Present workP. sphenops Gills L Salgado-Maldonado et al., 2001bP. gracilis Gills BR Salgado-Maldonado et al., 2001a

Fins PP Present workP. infans Gills BR Salgado-Maldonado et al., 2001a



Adult CestodaFamily Bothriocephalidae

Bothriocephalusacheilognathi G. vittata Intestine PR Salgado-Maldonado et al., 2004b

G. yucatana Intestine CY Scholz et al., 1996Intestine GU Salgado-Maldonado et al., 2005

H. bimaculata Intestine BR Salgado-Maldonado et al., 2001aIntestine L Present work

P. butleri Intestine SM Salgado-Maldonado et al., 2004aP. mexicana Intestine PR Salgado-Maldonado et al., 2004bP. reticulata Intestine BR Salgado-Maldonado et al., 2001aP. sphenops Intestine BR Salgado-Maldonado et al., 2001aP. baenschi Intestine SM Salgado-Maldonado et al., 2004aP. gracilis Intestine BR Salgado-Maldonado et al., 2001aPoecilia sp. Intestine MO Aguilar-Aguilar et al., 2004

MetacestodaFamily Dilepididae

Dendrouterina papillifera P. velifera CY Scholz et al., 1996Glossocercus auritus P. catemaconis Mesentery PP Scholz and Salgado-Maldonado, 2001

P. mexicana Fat PR Salgado-Maldonado et al., 2004bMesentery PP Present work

P. sphenops Body cavity, BR Salgado-Maldonado et al., 2001amesentery, liver

P. gracilis Body cavity, BR Salgado-Maldonado et al., 2001amesentery, liver

Glossocercus caribaensis P. petenensis GU Salgado-Maldonado et al., 2005Parvitaenia cochlearii P. gracilis Liver BR Salgado-Maldonado et al., 2001aValipora minuta P. sphenops Liver,

gall bladder BR Salgado-Maldonado et al., 2001aP. gracilis Liver,

gall bladder BR Salgado-Maldonado et al., 2001aFamily Protecocephalidae

Proteocephalidae gen. sp. P. infans Mesentery L Salgado-Maldonado et al., 2001bAdult nematoda

Family CapillaridaeCapillaria cyprinodonticola P. mexicana Intestine, liver PR Salgado-Maldonado et al., 2004b

P. sphenops Liver BR Salgado-Maldonado et al., 2001aFamily Cystidicolidae

Spinitectus mexicanus H. bimaculata Intestine PP Caspeta-Mandujano et al., 2000bP. mexicana Intestine PP Present work

Family DaniconematidaeMexiconema cichlasomae X. helleri Abdominal cavity PP Moravec, 1998

Family RhabdochonidaeRhabdochona kidderi G. yucatana Intestine CY Moravec et al., 1995a

P. mexicana Intestine PR, L Present workP. infans Intestine PR, L Present work



Rhabdochona lichtenfelsi P. mexicana Intestine PR Salgado-Maldonado et al., 2004bRhabdochona xiphophori Xiphophorus sp. Intestine PR Salgado-Maldonado et al., 2004b

Nematoda larvaeFamily Anisakidae

Contracaecum sp. G. yucatana GU Salgado-Maldonado et al., 2005H. bimaculata Mesentery PP Present workP. mexicana Mesentery, liver PR Salgado-Maldonado et al., 2004b

body cavity,Intestine L Present work

P. petenensis Mesentery CY Moravec et al., 1995bP. sphenops Liver, mesentery BR Salgado-Maldonado et al., 2001a

MuscleP. velifera Mesentery CY Moravec et al., 1995bP. baenschi Body cavity SM Salgado-Maldonado et al., 2004aP. gracilis Fat PR Salgado-Maldonado et al., 2004bP. infans Mesentery L Salgado-Maldonado et al., 2001bX. montezumae Mesentery, liver PR Salgado-Maldonado et al., 2004b

Family DioctophymatidaeEustrongylides sp. H. bimaculata Mesentery PR, L Present work

P. butleri Body cavity, SM Salgado-Maldonado et al., 2004aMesentery

P. mexicana Body cavity PR Salgado-Maldonado et al., 2004bMesentery L Present work

P. sphenops Muscle BR Salgado-Maldonado et al., 2001aP. gracilis Muscle BR Salgado-Maldonado et al., 2001aX. helleri Mesentery SM Salgado-Maldonado et al., 2004a

Family GnathostomidaeGnathostoma sp. B. belizanus GU Salgado-Maldonado et al., 2005Spiroxys sp. G. yucatana Body cavity, CY Moravec et al., 1995b

mesenteryP. butleri Mesentery SM Salgado-Maldonado et al., 2004aP. mexicana Body cavity, fat PR Salgado-Maldonado et al., 2004b

Mesentery PP Present workP. petenensis Mesentery CY Moravec et al., 1995bP. gracilis Intestine PR Salgado-Maldonado et al., 2004bX. helleri Mesentery PP Present work

Family KathlanidaeFalcaustra sp. P. velifera CY Moravec et al., 1995b

Adult acanthocephalaFamily Neoechinorhynchidae

Octospiniferoides chandleri H. bimaculata Intestine PP Salgado-Maldonado et al., 1992P. petenensis GU Salgado-Maldonado et al., 2005

Acanthocephala (Cystacanth)Family Polymorphidae

Polymorphus sp. X. helleri Intestine SM Salgado-Maldonado et al., 2004a


Acknowledgment

We acknowledge the Consejo Nacional de Cien-cia y Tecnología (CONACyT) for funding thisresearch through grant 31599-B: “Evaluación dela transferencia de parásitos exóticos entre pecesnativos e introducidos”.

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