A Comprehensive Update on Helminth Parasite Biodiversity and Richness in Peruvian Amphibians

Abstract

This study aimed to comprehensively update and expand the knowledge on the diversity and richness of helminth parasites found in Peruvian amphibians. A systematic search was conducted across primary databases, encompassing both indexed and non-indexed articles, to compile the most recent data. As of the present study, a total of 83 distinct helminth taxa have been documented in association with 78 anuran species of the order Anura, marking a 176.7% increase from previously recorded figures. Nematodes exhibited the highest species richness, totalling 52 taxa (62.65%), followed by trematodes (21 taxa, 25.3%), acanthocephalans and cestodes (4 taxa each, 4.8%), and monogeneans (2 taxa, 2.4%). The overwhelming majority (85.5%) of the collected parasites (71 taxa) were identified as mature helminths, with the remaining 14.5% (12 taxa) in their larval stages. Notably, Cosmocerca brasiliense Travassos, 1925 (Cosmocercidae), and Physaloptera sp. (Physalopteridae) were the most prevalent nematodes, having infected the broadest range of host species. Rhinella marina Linnaeus, 1758 (Bufonidae), emerged as the anuran host with the highest diversity, harboring 17 distinct helminth species. These findings underscore the crucial role of helminth parasites in shaping amphibian ecosystems and their significance as bioindicators of environmental health. Protecting both amphibian hosts and their associated helminth parasites is paramount, as it is intrinsically linked to the preservation of ecological equilibrium within these ecosystems.

1. Introduction

South America represents 33.8% (n = 2937) of the total 8689 species of global amphibian diversity. Among South American countries, Peru ranks third in terms of amphibian diversity, with 670 recorded amphibian species (22.81%), following Brazil (n = 1174; 39.97%) and Colombia (n = 832; 28.33%) [1].

Campião et al. [2] published a list of helminth parasites found in South American amphibians, reporting a total of 30 helminth species in Peru, which positioned Peru fourth in terms of helminth biodiversity. However, this review did not consider data published in national journals, many of which are difficult to access and have since discontinued publication or exist only in printed form in Peruvian libraries.

In recent years, there has been an increase in studies focusing on helminth parasites affecting amphibians in Peru [2,3,4,5,6,7,8,9]. Despite this increasing attention, the available information remains limited. It is necessary to continue these studies to document the diversity of helminths, which may face extinction if their hosts become threatened. Such research also contributes to the understanding of parasitic infection dynamics, aids in the use of these parasites as bioindicators of contamination in a specific area, and informs the development of conservation strategies [10,11,12,13].

Given the dispersed nature of information related to Peruvian amphibian parasites, this study aims to consolidate these data to facilitate future research. The objective of this research is to compile a comprehensive checklist of helminth parasites infecting amphibians in Peru and analyze their diversity and richness.

2. Materials and Methods

The database was compiled through an extensive bibliographical review of helminth parasites reported from amphibians in Peru. Searches were conducted using several keywords on various platforms, including Zoological Records (Web of Knowledge), Biological Abstracts, Helminthological Abstracts, Google Scholar, Aquatic Sciences and Fisheries Abstracts, Pubmed, ScienceDirect, and Scopus. In each database, search words included “anuran parasites Peru”, “amphibian parasites Peru”, and amphibian helminths Peru”. Additionally, articles published in Spanish that were not indexed in electronic databases were included. Dissertation, theses, and abstracts of scientific meetings do not constitute formal publications, and thus were not considered. The helminth parasite richness was quantified as the total number of parasite species observed in each host species.

This paper consists of two sections: The first section provides a list of helminth parasites, including information about their hosts, locality, site of infection, stage, and the corresponding references. The list of parasite–host associations is presented in phylogenetic order, beginning with the phylum Platyhelminthes (Monogenea, Trematoda, and Cestoda), followed by the phyla Nematoda and Rotifera. Helminth taxonomy follows specific reference for each group: Amin [14] for Acanthocephala, Anderson et al. [15] for Nematoda, Gibson et al. [16], Jones et al. [17], Bray et al. [18] for Trematoda, and Campião et al. [2], Cohen et al. [19] for monogenean, and Khalin et al. [20] for Cestoda. The second section presents a list of amphibian hosts, categorizing them based on their conservation status according to the IUCN (International Union for Conservation of Nature) and their associated parasites, organized by families and amphibian species in alphabetical order. Taxonomic information regarding amphibians is updated based on the AW database [1].

New records for Peru were also included in cases where helminth species were found in amphibian hosts at the Helminthological and Minor Invertebrates Collection at the Museum of Natural History, San Marcos University (MUSM), Peru, even if they had not been previously published in a scientific journal. The collection’s acronym is provided alongside the record. Parasite and amphibian names were updated in accordance with recent publications.

3. Results

The results obtained in this research reveal that in Peru, there is available information on parasites associated with 78 anuran hosts distributed across 10 families: Bufonidae (10 spp.), Dendrobatidae (4 spp.), Hemiphractidae (6 sp.), Hylidae (27 spp.), Leptodactylidae (11 spp.), Microhylidae (3 spp.), Pipidae (1 sp.), Ranidae (1 sp.), Strabomantidae (8 spp.), and Telmatobiidae (7 spp.). The Bufonidae and Telmatobidae families exhibited the highest number of helminth parasites, with 49 and 29 taxa, respectively (Figure 1). Meanwhile, only one species of helminth parasite has been reported to infect Peruvian pipids and ranids so far.

To date, a total of 83 taxa of helminth parasites from 28 families infecting amphibians have been recorded in Peru. Among the different types of parasites, nematodes exhibited the highest species richness (n = 52, 62,65%), followed by trematodes (n = 21; 25.3%), acanthocephalans (n = 4; 4.8%), cestodes (n = 4; 4.8%), and monogeneans (n = 2, 2.4%). These helminth parasites were associated with 60 nominal species and 23 undetermined species (

Table 1. Total of nominal and undetermined parasite species of amphibia from Peru according to the taxonomic group.

Taxonomic Group	Nominal sp.	Undetermined sp.	Total
Monogenea	2	0	2
Trematoda	17	4	21
Cestoda	1	3	4
Nematoda	39	13	52
Acanthocephala	1	3	4

) within the context of 78 species of anuran host. The overwhelming majority (85.5%) of the registered parasites (71 taxa) were identified as mature helminths, with the remaining 14.5% (12 taxa) in their larval stages. Sixty-eight percent (n= 53) of the amphibian hosts were found to harbor two or more parasites per host (Figure 2).

Table 2. Number of anuran hosts of Peru, richness of helminths average, maximum richness, number and percentage of anuran families with larvae, and the percentage found in conservation lists of IUCN.

	Hosts	Average Richness	Maximum Richness	Number of Anuran Families with Larvae	Percentage Larvae	Percentage Conservation IUCN
Bufonidae	10	7.80	17	5	50	40
Dendrobatidae	4	1.25	2	0	0	0
Hempractidae	6	2.7	6	0	0	33.3
Hylidae	27	2.44	5	17	62.96	0
Leptodactylidae	11	3.45	6	6	54.55	9
Microhylidae	3	2.33	3	2	66.67	0
Pipidae	1	1	1	1	1	0

displays information on the Anura hosts in Peru, including the mean richness of helminths, the maximum richness, the number and percentage of Anura families with larval parasites, and the percentage of amphibians listed in the IUCN conservation database. Of all the collected parasites, 85.5% (n = 71) were identified as mature forms of helminths, while 14.5% (n = 12) were in larval stages. Nematodes such as Cosmocerca brasiliense Travassos, 1925; Physaloptera sp. Rud, 1819; and Aplectana hylambatis (Baylis, 1927) exhibited a broad range of host species. Anurans such as Rhinella marina Linnaeus, 1758 (17 spp.); Rhinella margaritifera (Laurenti, 1768) (16 spp.); and Telmatobius jelskii (Peters, 1863) (16 spp.) were found to host the highest diversity of helminth parasites.

We compiled a total of 47 articles published over a 60-year period (1963 to 2023) on helminth parasites in amphibian hosts from Peru. The highest number of articles published on amphibian hosts falls within the period from 2000 to 2023, with 22 articles, representing 46.81% of the total. The second-highest period for article publication spans from 1980 to 1999, with 15 articles, accounting for 31.9% of the total. The earliest period, from 1963 to 1979, saw the publication of 10 papers, representing 21.27% of the total number of articles. Figure 3 displays the curve representing the total accumulated records of helminth taxa between 1963 and 2023 in Peruvian amphibians, while Figure 4 illustrates the curve depicting the accumulated number of new helminth taxa during the same period in Peruvian amphibians.

The regions in Peru where amphibian parasites were most frequently recorded included Cuzco (39.75%), Lima (32.53%), Junín (16.87%), Arequipa (15.66%), and Pasco (13.25%) (Figure 5 and Figure 6). The most commonly infected organs were the intestine, small intestine, and urinary bladder, accounting for 28%, 19%, and 16% of recorded infections, respectively, across 11 different sites within the host amphibians.

Regarding the conservation status of the host amphibians, as per the IUCN, 63 anuran species are classified as ‘Least Concern’ (LC), two as ‘Near Threatened’ (NT), three are ‘Vulnerable’ (VU), six are ‘Endangered’ (EN), three are ‘Critically Endangered’ (CR), and the status of two species remains undetermined.

• PARASITE–HOST LIST

• Phylum Platyhelminthes Gegenbaur, 1959
• Class Monogenea Van Beneden, 1858
• Family Polystomatidae (Carus, 1863) Gamble, 1896
• Mesopolystoma samiriense Vaucher, 1981
• Host: Osteocephalus taurinus Steindachner, 1862
• Locality: Amazonas
• Site of infection: urinary bladder
• Stage: adult
• References: [19,21]

• Wetapolystoma almae Gray, 1993
• Host: Rhinella margaritifera (Laurenti, 1768)
• Locality: Cuzco, Madre de Dios
• Site of infection: urinary bladder
• Stage: adult
• References: [2,19,22]

• Class Trematoda Rudolphi, 1808
• Subclass Digenea Carus, 1863
• Superfamily Gorgoderoidea Looss, 1899
• Family Brachycoellidae Johnston, 1912
• Iquitos ceii Mañe-Garzon & Gil, 1963
• Host: Lithobates palmipes Spix, 1824
• Locality: Loreto
• Site of infection: intestine
• Stage: adult
• References: [7,23]

• Family Gorgoderidae Looss, 1899
• Gorgoderina alobata Lees & Mitchell, 1965
• Host: Telmatobius sp. Wiegmann, 1834
• Locality: not indicated
• Site of infection: urinary bladder
• Stage: adult
• Reference: Present study (MUSM 2185)

• Gorgoderina attenuata (Stafford, 1902)
• Host: Telmatobius jelskii (Peters, 1863)
• Locality: Lima
• Site of infection: urinary bladder
• Stage: adult
• Reference: [24]

• Gorgoderina carli Baer, 1930
• Host: Telmatobius brachydactylus (Peters, 1873)
• Locality: Pasco
• Site of infection: urinary bladder
• Stage: adult
• Reference: Present study (MUSM 1420a)

• Gorgoderina chilensis Dioni, 1947
• Hosts: Telmatobius brachydactylus, Telmatobius macrostomus Peters, 1873, Rhinella spinulosa Wiegmann, 1834
• Locality: Lima, Pasco
• Site of infection: urinary bladder
• Stage: adult
• References: [6,7,25], Present study (MUSM 1420b)

• Gorgoderina darwini Mane-Garzon & Gonzalez, 1978
• Host: Atelopus aff. bomolochus Peters 1973
• Locality: Piura
• Site of infection: urinary bladder
• Stage: adult
• Reference: Present study (MUSM 1764a,b)

• Gorgoderina gracilis Wongsawad, Sey, Rojanapaibul, Wongsawad, Marayong, Rojtinnakorn, Suwattanacoupt & Pachananwan, 2000
• Host: Telmatobius jelskii
• Locality: Lima
• Site of infection: urinary bladder
• Stage: adult
• Reference: present study: MUSM 1762.

• Gorgoderina parvicava Travassos, 1922
• Hosts: Atelopus aff. bomolochos, Telmatobius macrostomus, Leptodactylus rhodonotus (Gunther, 1868), Rhinella arequipensis (Vellard, 1959), Rhinella limensis (Werner, 1901), Rhinella spinulosa Wiegmann, 1834, Telmatobius culeus (Garman, 1876), Telmatobius jelskii, Telmatobius peruvianus Wiegmann, 1834
• Locality: Arequipa, Cuzco, Ica, Junín, Lima, Piura, Puno
• Site of infection: urinary bladder
• Stage: adult
• References: [2,3,5,7,25,26,27,28,29,30], present study (MUSM 114)

• Gorgoderina parvicava minuta Tantaleán & García, 1993
• Hosts: Telmatobius macrostomus
• Locality: Pasco
• Site of infection: urinary bladder
• Stage: adult
• References: [7,25,31]

• Gorgoderina megacysta Mane-Garzon & Gonzalez, 1978
• Hosts: Telmatobius brachydactylus, Telmatobius culeus, Leptodactylus rhodonotus
• Locality: Junín, Pasco
• Site of infection: urinary bladder
• Stage: adult
• Reference: present study (MUSM 110, 261 and 1315)

• Gorgoderina sp.
• Host: Telmatobius macrostomus, Telmatobius jelskii
• Locality: Lima, Pasco
• Site of infection: urinary bladder
• Stage: adult
• References: [7,24,25]

• Phyllostomum (Microlecithus) sp.
• Host: Rhinella spinulosa
• Locality: Huánuco
• Site of infection: urinary bladder
• Stage: adult
• Reference: [27]

• Family Mesocoeliidae Dollfus, 1933
• Mesocoelium monas (Rudolphi, 1819) Freitas, 1958
• Host: Rhinella marina Linnaeus, 1758
• Locality: Cuzco
• Site of infection: small intestine
• Stage: adult
• References: [7,27]

• Mesocoelium waltoni Brunetti, 1912
• Host: Rhinella marina
• Locality: Cuzco
• Site of infection: small intestine
• Stage: adult
• References: [7,27]

• Family lecithodendriidae Odhner, 1911
• Mosesia sp.
• Host: Rhinella limensis
• Locality: Lima
• Site of infection: small intestine
• Stage: adult
• References: [27,32]

• Superfamily Plagiorchioidae Lühe, 1901
• Family Plagiorchiidae Lühe, 1901
• Rudolphitrema rudolphi (Travassos, 1924) Travassos, 1926
• Host: Atelopus ignescens (Cornalia, 1849)
• Locality: Cajamarca
• Site of infection: small intestine
• Stage: adult
• References: [7,27,28]

• Family Glypthelminthidae Cheng, 1959
• Glypthelmins parva Travassos, 1924
• Hosts: Dendropsophus sarayacuensis (Shreve, 1935), Trachycephalus coriaceus (Peters, 1867), Scinax pedromedinae (Henle, 1991)
• Locality: Cuzco
• Site of infection: small intestine
• Stage: adult
• References: [2,7,33]

• Family Haematoloechidae Freitas & Lent, 1939
• Haematoloechus arequipensis Ibañez & Cordova, 1979
• Host: Telmatobius peruvianus
• Locality: Arequipa
• Site of infection: lungs
• Stage: adult
• References: [7,26,27,28]

• Haematoloechus medioplexus Looss, 1899
• Host: Rhinella limensis
• Locality: Lima
• Site of infection: lungs
• Stage: adult
• Reference: present study: MUSM 246

• Haematoloechus pukinensis Ibañez & Cordova, 1979
• Hosts: Telmatobius peruvianus, Telmatobius jelskii
• Locality: Arequipa, Lima
• Site of infection: urinary bladder
• Stage: adult
• References: [3,7,26,27,28]

• Haematoloechus sp. Looss, 1899
• Host: Rhinella spinulosa
• Locality: Ica
• Site of infection: urinary bladder
• Stage: adult
• References: [7,27]

• Class Cestoda Rudolphi, 1808
• Order Cyclophyllidea Van Deneden in Braum, 1990
• Family Nematotaeniidae Lühe, 1910
• Cylindrotaenia americana Jewell, 1916
• Hosts: Allobates marchesianus (Melin, 1941), Atelopus aff. bomolochos, Lithodytes lineatus (Schneider, 1799), Rhinella marina, Rhinella margaritifera, Rhinella poeppigii (Tschudi, 1845), Rhinella spinulosa, Scinax pedromedinae, Telmatobius jelskii, Telmatobius marmoratus (Duméril &. Bibron, 1841)
• Locality: Amazonas, Arequipa, Cuzco, Lima, Piura, San Martin, Ucayali
• Site of infection: small intestine
• Stage: adult
• References: [2,3,4,6,9,25,26,27,28,29,33,34]

• Family Proteocephalidae La Rue, 1911
• Ophiotaenia sp1.
• Host: Telmatobius jelskii
• Locality: Lima
• Site of infection: small intestine
• Stage: adult
• Reference: [3]

• Ophiotaenia sp2.
• Host: Telmatobius macrostomus
• Locality: Pasco
• Site of infection: small intestine
• Stage: adult
• Reference: [35]

• Order Diphyllobothriidea Lühe, 1910
• Family Diphyllobothriidae Lühe, 1910
• Spirometra sp. Faust, Campbell et Kellogg, 1929
• Hosts: Pristimantis nephophilus Duellman & Pramuk, 1999, Pristimantis rhodostichus (Duellman & Pramuk, 1999)
• Locality: San Martín
• Site of infection: subcutaneous level
• Stage: larvae
• References: [36,37]

• Phylum Nematoda Rudolphi, 1808
• Order Ascaridida Skrjabin & Shulz, 1940
• Superfamily Ascaridoidea Chabaud, 1965
• Family Ascarididae Baird, 1853
• Brevimulticaecum sp. Mozgovoi, 1951
• Hosts: Boana fasciata (Günther, 1858), Boana cinerascens (Spix, 1824), Leptodactylus bolivianus Boulenger, 1898, Leptodactylus pentadactylus (Laurenti, 1768), Scarthyla goinorum (Bokermann, 1962), Sphaenorhynchus lacteus (Daudin, 1800), Trachycephalus coriaceus
• Locality: Cuzco
• Site of infection: cysts in gastrointestinal wall
• Stage: larvae
• References: [2,33]

• Ophidascaris sp. Baylis, 1921
• Host: Callimedusa tomopterna (Cope, 1868)
• Locality: Cuzco
• Site of infection: mesentery and serous stomach
• Stage: larvae
• References: [2,33]

• Porrocaecum sp. Railliet & Henry, 1912
• Host: Pipa pipa (Linnaeus, 1758)
• Locality: Cuzco
• Site of infection: mesentery and serous stomach
• Stage: larvae
• References: [2,33]

• Superfamily Cosmocercoidea Skrjabin & Schikhobalova, 1951
• Family Atractidae (Railliet, 1917) Travassos, 1919
• Schrankiana inconspicata Freitas, 1959
• Host: Leptodactylus rhodonotus
• Locality: Cuzco
• Site of infection: intestine
• Stage: adult
• References: [2,33]

• Schrankiana larvata (Vaz, 1933) Fahel, 1952
• Hosts: Leptodactylus mystaceus (Spix, 1824), Leptodactylus pentadactylus
• Locality: Cuzco
• Site of infection: intestine
• Stage: adult
• References: [2,33]

• Schrankiana schrankai (Travassos, 1925)
• Host: Leptodactylus mystaceus
• Locality: Cuzco
• Site of infection: intestine
• Stage: adult
• References: [2,33]

• Schrankianella brasili (Travassos, 1927)
• Host: Leptodactylus pentadactylus
• Locality: Cuzco
• Site of infection: intestine
• Stage: adult
• References: [2,33]

• Family Cosmocercidae Travassos, 1925
• Aplectana hylambatis (Baylis, 1927)
• Hosts: Atelopus aff. bomolochos, Elachistocleis ovalis (Schneider, 1799), Gastrotheca marsupiata (Duméril & Bibron, 1841), Gastrotheca griswoldi, Gastrotheca peruana, Pleurodema marmoratum, Leptodactylus bolivianus, Leptodactylus leptodactyloides (Andersson, 1945), Leptodactylus pentadactylus, Leptodactylus rhodonotus, Lithodytes lineatus, Rhinella arequipensis, Rhinella spinulosa, Rhinella margaritifera, Rhinella marina, Rhinella poeppigii, Telmatobius jelskii, Telmatobius marmoratus
• Locality: Amazonas, Ancash, Apurimac, Arequipa, Cajamarca, Cuzco, Huánuco, Junín, La Libertad, Lima, Pasco, Piura, San Martin, Ucayali
• Site of infection: large intestine, small intestine
• Stage: adult
• References: [2,3,4,5,6,8,9,29,30,33,34,38,39,40]

• Aplectana membranosa Schneider, 1866
• Hosts: Gastrotheca marsupiata
• Locality: Abancay, Cajamarca, Huánuco, Junín, Piura
• Site of infection: intestine
• Stage: adult
• References: [8]

• Aplectana vellardi Travassos, 1926
• Hosts: Gastrotheca peruana, Rhinella arequipensis, Rhinella marina, Rhinella spinulosa
• Locality: Ancash, Arequipa, La Libertad, Lima, San Martin
• Site of infection: intestine, large intestine
• Stage: adult
• References: [6,8,9,38,40]

• Cosmocerca brasiliense Travassos, 1925
• Hosts: Adenomera andreae (Müller, 1923), Adenomera hylaedactyla (Cope, 1868), Dendropsophus koechlini (Duellman & Trueb, 1989), Dendropsophus leali (Bokermann, 1964), Dendropsophus leucophyllatus (Beireis, 1783), Dendropsophus marmoratus (Laurenti, 1768), Dendropsophus parviceps (Boulenger, 1882), Dendropsophus rhodopeplus (Günther, 1858), Dendropsophus schubarti (Bokermann, 1963), Edalorhina perezi Jiménez de la Espada, 1871, Gastrotheca stictopleura (Duellman, Lehr, and Aguilar, 2001), Boana boans (Linnaeus, 1758), Boana calcarata (Troschel, 1848), Boana cinerascens, Boana fasciata (Günther, 1858), Leptodactylus bolivianus, Leptodactylus pentadactylus, Leptodactylus petersii (Steindachner, 1864), Osteocephalus taurinus, Callimedusa atelopoides Duellman, Cadle & Cannatella, 1988, Pithecopus palliatus Peters, 1873, Callimedusa tomopterna, Phyllomedusa vaillanti Boulenger, 1882, Pristimantis fenestratus (Steindachner, 1864), Pristimantis peruvianus (Melin, 1941), Pristimantis toftae (Duellman, 1978), Scarthyla goinorum, Scinax garbei (Miranda-Ribeiro, 1926), Scinax ictericus Duellman & Wiens, 1993, Scinax pedromedinae, Sphaenorhynchus lacteus, Oreobates cruralis (Boulenger, 1902), Rhinella marina, Trachycephalus coriaceus
• Locality: Cuzco, Pasco
• Site of infection: intestine
• Stage: adult
• References: [2,8,33]

• Cosmocerca cajamarquensis Ibañez, 1982
• Host: Atelopus peruensis Grey & Cannatella, 1985
• Locality: Cajamarca
• Site of infection: intestine
• Stage: adult
• References: [40,41]

• Cosmocerca cruzi Rodrigues and Fabio, 1970
• Host: Gastrotheca excubitor
• Locality: Cusco
• Site of infection: intestine
• Stage: adult
• References: [8]

• Cosmocerca parva Travassos, 1925
• Hosts: Ameerega picta (Tschudi, 1838), Ameerega trivittata (Spix, 1824), Edalorhina perezi, Elachistocleis ovalis, Hamptophryne boliviana (Parker, 1927), Boana fasciata, Leptodactylus leptodactyloides, Leptodactylus mystaceus, Oreobates quixensis (Jiménez de la Espada, 1872), Callimedusa atelopoides, Pristimantis fenestratus, Pristimantis peruvianus, Pristimantis toftae, Rhaebo glaberrimus (Günther, 1869), Rhinella marina, Rhinella margaritifera, Scarthyla goinorum, Scinax garbei, Scinax ictericus
• Locality: Cuzco, Ucayali
• Site of infection: intestine
• Stage: adult
• References: [2,33,34,40]

• Cosmocerca podicipinus Baker & Vaucher, 1984
• Hosts: Allobates marchesianus, Allobates femoralis (Boulenger, 1884), Pristimantis imitatrix (Duellman, 1978), Leptodactylus leptodactyloides, Rhinella margaritifera
• Locality: Cuzco
• Site of infection: intestine
• Stage: adult
• References: [2,33]

• Cosmocercella phyllomedusae Bacher & Vaucher, 1986
• Hosts: Pithecopus palliatus, Callimedusa tomopterna, Phyllomedusa vaillanti
• Locality: Cuzco
• Site of infection: intestine
• Stage: adult
• References: [2,33]

• Cosmocercoides dukae (Holl, 1928)
• Hosts: Rhinella arequipensis, Rhinella limensis
• Locality: Lima
• Site of infection: intestine
• Stage: adult
• References: [38,40]

• Cosmocercoides lilloi Ramallo, Bursey y Goldberg, 2007
• Hosts: Gastrotheca excubitor, Gastrotheca griswoldi, Gastrotheca monticola, Gastrotheca peruana
• Locality: Ancash, Cajamarca, Cusco, Junín, Huánuco, La Libertad, Piura
• Site of infection: intestine
• Stage: adult
• References: [8]

• Raillietnema gubernaculatum Freitas & Ibañez, 1965
• Hosts: Elachistocleis ovalis, Rhinella limensis
• Locality: Cuzco, La Libertad
• Site of infection: intestine
• Stage: adult
• References: [2,33,40,42]

• Raillietnema ibañezi Cordova, 1998
• Host: Rhinella spinulosa
• Locality: Arequipa
• Site of infection: intestine
• Stage: adult
• References: [40,43]

• Cosmocercidae gen. sp. Railliet, 1916
• Host: Telmatobius jelskii
• Locality: Ayacucho, Huancavelica, Junín
• Site of infection: intestine
• Stage: larvae
• References: [3]

• Family Kathlaniidae (Lane, 1914) Travassos, 1918
• Falcaustra mascula (Rudolphi, 1819) Freitas & Lent, 1941
• Hosts: Rhinella limensis, Rhinella spinulosa, Telmatobius jelskii, Telmatobius marmoratus
• Locality: Arequipa, Cuzco, Ica, Lima
• Site of infection: small intestine
• Stage: adult
• References: [3,38,39,40,44]

• Falcaustra condorcanquii Ibañez & Cordova, 1976
• Hosts: Telmatobius macrostomus, Rhinella marina, Telmatobius jelskii, Telmatobius peruvianus
• Locality: Arequipa, Junín, Lima, San Martin
• Site of infection: small intestine
• Stage: adult
• References: [2,9,38,40,45]

• Falcaustra sp. Lane, 1915
• Hosts: Rhinella limensis, Rhinella marina
• Locality: Huánuco, Lima
• Site of infection: small intestine, stomach
• Stage: adult
• References: [5,38,40]

• Oxyascaris sp.Travassos, 1920
• Host: Gastrotheca peruana, Pleurodema marmoratum
• Locality: Ancash, La Libertad, Junín
• Site of infection: large intestine, small intestine
• Stage: adult
• Reference: [8,40]

• Order Enoplida Schuurmans & Deconing, 1933
• Superfamily Dioctophymatoidea (Railliet, 1915)
• Family Trichuridae (Ransom, 1911) Railliet, 1915
• Capillaria sp. Zeder, 1800
• Hosts: Atelopus aff. bomolochos, Telmatobius jelskii
• Locality: Lima, Piura
• Site of infection: small intestine
• Stage: adult
• References: [2,24,30]

• Order Oxyurida Railliet, 1916
• Superfamily Oxyuroidea Railliet, 1916
• Family Pharyngodonidae Travassos, 1919
• Batracholandros spectatus (Freitas & Ibañez, 1962) Freitas & Ibañez, 1965
• Hosts: Atelopus aff. bomolochos, Osteocephalus taurinus, Rhinella limensis, Rhinella spinulosa, Trachycephalus typhonius
• Locality: Arequipa, Cuzco, Huánuco, Ica, La Libertad, Lima, Piura
• Site of infection: small intestine
• Stage: adult
• References: [2,30,33,38,40,42]

• Parapharyngodon sp. (Chatterji, 1933)
• Hosts: Gastrotheca griswoldi
• Locality: Junín, Huánuco
• Site of infection: intestine
• Stage: adult
• References: [8]

• Order Rhabditida Chitwood, 1933
• Superfamily Rhabditoidea Travassos, 1920
• Family Rhabdiasidae Railliet, 1915
• Rhabdias aff. elegans Gutierrez, 1945
• Host: Gastrotheca griswoldi, Gastrotheca peruana, Rhinella poeppigii
• Locality: Ancash, Amazonas, Junín, Huánuco, La Libertad
• Site of infection: lungs
• Stage: adult
• Reference: [4,8]

• Rhabdias fuelleborni Travassos, 1926
• Host: Rhinella limensis
• Locality: Lima
• Site of infection: lungs
• Stage: adult
• Reference: [5]

• Rhabdias pseudosphaerocephala Kuzmin, Tkach & Brooks, 2007
• Host: Rhinella marina, Rhinella spinulosa
• Locality: Lima, San Martin
• Site of infection: lungs
• Stage: adult
• Reference: [6,9]

• Rhabdias sphaerocephala Goodey, 1924
• Hosts: Atelopus aff. bomolochos, Rhinella arequipensis, Rhinella limensis, Rhinella margaritifera, Rhinella marina, Rhinella spinulosa
• Locality: Arequipa, Cuzco, Huánuco, Ica, Junín, Pasco, Piura
• Site of infection: lungs
• Stage: adult
• References: [2,30,38,40,46,47]

• Order Spirurida Chitwood, 1933
• Superfamily Acuarioidae Molin, 1860
• Family Acuariidae Seurat, 1913
• Acuarioidae gen. sp. Ditwood & Wehr 1934
• Hosts: Dendropsophus rhodopeplus, Edalorhina perezi
• Locality: Cuzco
• Site of infection: stomach
• Stage: larvae
• Reference: [33]

• Superfamily Camallanoidea Chabaud, 1965
• Family Camallanidae Railliet and Henry, 1915
• Camallanus sp. Railliet & Henry, 1915
• Host: Telmatobius jelskii
• Locality: Lima
• Site of infection: intestine
• Stage: larvae
• Reference: [40]

• Superfamily Filarioidea (Weinland, 1858) Stiles, 1907
• Family Filariidae (Weinland, 1858) Cobbold, 1879
• Ochoterenella digiticauda Caballero, 1944
• Hosts: Leptodactylus mystaceus, Rhinella poeppigii
• Locality: Amazonas, Cuzco
• Site of infection: body cavity, subcutaneous
• Stage: adult
• References: [4], Present study: MUSM 2296

• Ochoterenella figueroai Esslinger, 1988
• Hosts: Rhinella poeppigii
• Locality: Amazonas
• Site of infection: body cavity
• Stage: adult
• Reference: [4]

• Ochoterenella vellardi (Travassos, 1929) Esslinger, 1986
• Hosts: Boana fasciata, Osteocephalus taurinus, Rhinella margaritifera, Rhinella marina
• Locality: Cuzco, San Martin
• Site of infection: body cavity
• Stage: adult
• References: [2,9,33]

• Superfamily Habronematoidea Chabaud, 1959
• Family Hedruridae Petter, 1971
• Hedruris juninensis Sarmiento, 1976
• Hosts: Telmatobius brachydactylus, Telmatobius macrostomus
• Locality: Junín
• Site of infection: stomach
• Stage: adult
• References: [2,31,40,48,49,50]

• Hedruris moniezi Ibañez & Cordova, 1976
• Hosts: Rhinella spinulosa, Telmatobius jelskii, Telmatobius marmoratus, Telmatobius peruvianus, Telmatobius culeus, Telmatobius sp.
• Locality: Arequipa, Cuzco, Junín, Lima, Puno
• Site of infection: stomach
• Stage: adult
• References: [2,3,6,38,39,40,45,49,50,51]
• Note: Referred as Hedruris heyeri Bursey & Goldberg, 2007 by Serrano-Martinez et al. [24].

• Hedruris sp. Nitzsch, 1821
• Hosts: Rhinella spinulosa, Telmatobius jelskii
• Locality: Junín, Lima
• Site of infection: stomach
• Stage: adult
• References: [3,40]

• Superfamily Physalopteroidea Sobolev, 1949
• Family Physalopteridae Railliet, 1893 Leiper, 1908
• Physaloptera huascari Ibañez, 1973
• Hosts: Rhinella limensis, Rhinella spinulosa
• Locality: La Libertad
• Site of infection: stomach
• Stage: larvae
• References: [40,52]

• Physaloptera sp. Rud, 1819
• Hosts: Ctenophryne geayi Mocquard, 1904, Dendropsophus leali, Dendropsophus leucophyllatus, Dendropsophus marmoratus, Edalorhina perezi, Hamptophryne boliviana, Boana boans, Boana fasciata, Boana cinerascens, Leptodactylus bolivianus, Leptodactylus leptodactyloides, Leptodactylus mystaceus, Leptodactylus pentadactylus, Leptodactylus rhodonotus, Lithodytes lineatus, Oreobates cruralis, Osteocephalus taurinus, Callimedusa tomopterna, Pristimantis fenestratus, Pseudis paradoxa Linnaeus, 1758, Rhinella limensis, Rhinella marina, Rhinella spinulosa, Scinax ictericus, Scinax ruber (Laurenti, 1768), Trachycephalus coriaceus, Trachycephalus typhonius (Linnaeus, 1758)
• Locality: Cuzco, Junín, San Martin
• Site of infection: stomach
• Stage: larvae
• References: [2,9,33,40,52]

• Physalopteroides venancioi (Lent, Freitas & Proença, 1946) Sobolev, 1949
• Hosts: Adenomera hylaedactyla, Edalorhina perezi, Leptodactylus bolivianus, Leptodactylus mystaceus, Osteocephalus taurinus, Phyllomedusa vaillanti, Scinax ruber, Pristimantis fenestratus, Trachycephalus coriaceus
• Locality: Cuzco
• Site of infection: stomach
• Stage: adult
• References: [2,33]

• Orden Strongylida Molin, 1861
• Superfamily Trichostrongyloidea Cram, 1927
• Family Molineidae Durette-Desset & Chabaud, 1977
• Oswaldocruzia proencai Ben Slimane & Durette-Desset, 1995
• Hosts: Gastrotheca peruana, Rhinella margaritifera
• Locality: Ancash, La Libertad, Ucayali
• Site of infection: intestine
• Stage: adult
• References: [2,8,34]

• Oswaldocruzia subauricularis (Rudolphi, 1819) Travassos, 1917
• Hosts: Leptodactylus rhodonotus, Rhinella margaritifera, Rhinella marina
• Locality: Cuzco, Ucayali
• Site of infection: small intestine
• Stage: adult
• References: [33,40,47]. Present study: MUSM 1316

• Oswaldocruzia lopesi Freitas & Lent, 1938
• Hosts: Ameerega picta, Hamptophryne boliviana, Boana fasciata, Leptodactylus bolivianus, Pristimantis fenestratus, Rhaebo glaberrimus, Rhinella margaritifera, Trachycephalus coriaceus
• Locality: Cuzco
• Site of infection: intestine
• Stage: adult
• References: [2,33]

• Oswaldocrucia manuensis Guerrero, 2013
• Hosts: Rhinella marina
• Locality: Cuzco
• Site of infection: intestine
• Stage: adult
• Reference: [53]

• Oswaldocrucia urubambaensisGuerrero, 2013
• Host: Rhinella marina
• Locality: Cuzco, San Martin
• Site of infection: intestine
• Stage: adult
• Reference: [9,53]

• Schulzia chiribita Durette-Desset, Florindez & Morales, 2000
• Host: Leptodactylus rhodonotus
• Locality: Pasco
• Site of infection: intestine
• Stage: adult
• References: [31,54]

• Schulzia sp. Travasos, 1937
• Host: Leptodactylus rhodonotus
• Locality: Pasco
• Site of infection: large intestine
• Stage: adult
• Reference: [40]

• Family Nicollinidae Durette-Desset & Chabaud, 1981
• Batrachonema bonai Durette-Desset, Baker & Vaucher, 1984
• Host: Rhinella margaritifera
• Locality: Loreto
• Site of infection: large intestine
• Stage: adult
• References: [2,40,55]

• Family Thrichostrongylidae Cram, 1927
• Borrellostrongylus lumbrerasi Tantaleán & Naupay, 1974
• Hosts: Rhinella arequipensis, Rhinella spinulosa
• Locality: Arequipa, Junín
• Site of infection: intestine, cyst in lung tissue
• Stage: adult
• References: [40,56]. Present study (MUSM 1418–1419)

• Phylum Rotifera Cuvier, 1817
• Class Hemirotatoria Markevich, 1989
• Subclass Acanthocephala Koelreuther, 1771
• Order Echinorhynchida Southwell Southwell & MacFie, 1925
• Family Echinorhynchidae Cobbold, 1876
• Pseudoacanthocephalus lutzi (Hamann, 1891)
• Hosts: Rhinella arequipensis, Rhinella limensis, Rhinella marina, Rhinella spinulosa
• Locality: Arequipa, Huánuco, Ica, Lima, Ucayali, San Martin
• Site of infection: intestine
• Stage: adult
• References: [6,57,58,59,60]

• Order Polymorphida Petrochenko, 1956
• Family Centrorhynchidae Van Cleave, 1916
• Centrorhynchus sp. Kuntzi Schmidt and Neiland, 1966
• Hosts: Atelopus aff. bomolochos, Telmatobius jelskii
• Locality: Piura, Lima
• Site of infection: mesentery
• Stage: cystacanth
• References: [2,3,30]

• Oncicola sp. Travassos, 1916
• Hosts: Telmatobius jelskii
• Locality: Lima
• Site of infection: mesentery
• Stage: cystacanth
• Reference: [3]

• Acanthocephala gen. sp. Koelreuther, 1771
• Hosts: Pithecopus palliatus, Rhinella margaritifera, Scinax pedromedinae
• Locality: Cuzco
• Site of infection: mesentery
• Stage: cystacanth
• Reference: [33]

• HOST–PARASITE LIST
• Clase Amphibia
• Order Anura
• Family Bufonidae

• Atelopus aff. bomolochos
• IUCN Red List Status: Critically Endangered (CR)
• Digenea:
• Gorgoderina parvicava
• Cestoda:
• Cylindrotaenia americana
• Nematoda:
• Aplectana hylambatis
• Batracholandros spectatus
• Capillaria sp.
• Rhabdias sphaerocephala
• Acantocephala:
• Centrorhynchus sp.

• Atelopus ignescens
• IUCN Red List Status: Critically Endangered (CR)
• Digenea:
• Rudolphitrema rudolphi

• Atelopus peruensis
• IUCN Red List Status: Critically Endangered (CR)
• Nematoda:
• Cosmocerca cajamarquensis

• Rhaebo glaberrimus
• IUCN Red List Status: Least Concern (LC)
• Nematoda:
• Cosmocerca parva
• Oswaldocruzia lopesi

• Rhinella arequipensis
• IUCN Red List Status: Endangered (EN)
• Digenea:
• Gorgoderina parvicava
• Nematoda:
• Aplectana hylambatis
• Aplectana vellardi
• Rhabdias sphaerocephala
• Cosmocercoides dukae
• Borrellostrongylus lumbrerasi
• Acantocephala:
• Pseudoacanthocephalus lutzi

• Rhinella limensis
• IUCN Red List Status: Least Concern (LC)
• Digenea:
• Gorgoderina parvicava
• Haematoloechus medioplexus
• Mosesia sp.
• Nematoda:
• Cosmocercoides dukae
• Raillietnema gubernaculatum
• Falcaustra máscula
• Falcaustra sp.
• Batracholandros spectatus
• Rhabdias sphaerocephala
• Physaloptera huascari
• Physaloptera sp.
• Acantocephala:
• Pseudoacanthocephalus lutzi

• Rhinella margaritifera
• IUCN Red List Status: Least Concern (LC)
• Monogenea:
• Wetapolystoma almae
• Digenea:
• Gorgoderina parvicava
• Mosesia sp.
• Cestoda:
• Cylindrotaenia americana
• Nematoda:
• Oswaldocruzia proencai
• Aplectana hylambatis
• Cosmocerca parva
• Cosmocerca podicipinus
• Raillietnema ibañezi
• Falcaustra mascula
• Rhabdias sphaerocephala
• Ochoterenella vellardi
• Oswaldocruzia subauricularis
• Oswaldocruzia lopesi
• Batrachonema bonai
• Acantocephala:
• Acantocephala gen. sp.

• Rhinella marina
• IUCN Red List Status: Least Concern (LC)
• Digenea:
• Mesocoelium monas
• Mesocoelium waltoni
• Cestoda:
• Cylindrotaenia americana
• Nematoda:
• Aplectana hylambatis
• Aplectana vellardi
• Cosmocercoides brasiliense
• Cosmocerca parva
• Falcaustra condorcanquii
• Falcaustra sp.
• Rhabdias sphaerocephala
• Rhabdias pseudosphaerocephala
• Physaloptera sp.
• Ochoterenella vellardi
• Oswaldocruzia subauricularis
• Oswaldocruzia manuensis
• Oswaldocruzia urubambaensis
• Acantocephala:
• Pseudoacanthocephalus lutzi
• Rhinella poeppigii
• IUCN Red List Status: Least Concern (LC)
• Nematoda:
• Aplectana hylambatis
• Ochoterenella digiticauda
• Ochoterenella figueroai
• Rhabdias aff. elegans
• Cestoda:
• Cylindrotaenia aff. americana

• Rhinella spinulosa
• IUCN Red List Status: Least Concern (LC)
• Digenea:
• Gorgoderina parvicava
• Phyllostomum (Microlecithus) sp.
• Haematoloechus sp.
• Cestoda:
• Cylindrotaenia americana
• Nematoda:
• Aplectana hylambatis
• Aplectana vellardi
• Raillietnema ibañezi
• Falcaustra máscula
• Batracholandros spectatus
• Rhabdias sphaerocephala
• Hedruris moniezi
• Physaloptera huascari
• Physaloptera sp.
• Acantocephala:
• Pseudoacanthocephalus lutzi

• Family Dendrobatidae

• Allobates femoralis
• IUCN Red List Status: Least Concern (LC)
• Nematoda:
• Cosmocerca podicipinus

• Allobates marchesianus
• IUCN Red List Status: Least Concern (LC)
• Cestoda:
• Cylindrotaenia americana
• Nematoda:
• Cosmocerca podicipinus

• Ameerega trivittata
• IUCN Red List Status: Least Concern (LC)
• Nematoda:
• Cosmocerca parva

• Ameerega picta
• IUCN Red List Status: Least Concern (LC)
• Nematoda:
• Cosmocerca parva

• Family Hemiphractidae

• Gastrotheca marsupiata
• IUCN Red List Status: Least Concern (LC)
• Nematoda:
• Aplectana hylambatis
• Aplectana membranosa

• Gastrotheca excubitor
• IUCN Red List Status: Vulnerable (VU)
• Nematoda:
• Cosmocerca cruzi
• Cosmocercoides lilloi

• Gastrotheca griswoldi
• IUCN Red List Status: Least Concern (LC)
• Nematoda:
• Cosmocercoides lilloi
• Aplectana hylambatis
• Parapharyngodon sp.
• Rhabdias aff. Elegans

• Gastrotheca monticola
• IUCN Red List Status: Least Concern (LC)
• Nematoda:
• Cosmocercoides lilloi

• Gastrotheca peruana
• IUCN Red List Status: Least Concern (LC)
• Nematoda:
• Cosmocercoides lilloi
• Aplectana hylambatis
• Aplectana vellardi
• Oxyascaris sp.
• Oswaldocruzia proencai
• Rhabdias aff. Elegans

• Gastrotheca stictopleura
• IUCN Red List Status: Endangered (EN)
• Nematoda:
• Cosmocerca brasiliensis

• Family Hylidae

• Dendropsophus brevifrons
• IUCN Red List Status: Least Concern (LC)
• Nematoda:
• Cosmocerca brasiliense

• Dendropsophus koechlini
• IUCN Red List Status: Least Concern (LC)
• Nematoda:
• Cosmocerca brasiliense

• Dendropsophus leali
• IUCN Red List Status: Least Concern (LC)
• Nematoda:
• Cosmocerca brasiliensis brasiliense
• Physaloptera sp.

• Dendropsophus leucophyllatus
• IUCN Red List Status: Least Concern (LC)
• Nematoda:
• Cosmocerca brasiliense
• Physaloptera sp.

• Dendropsophus marmoratus
• IUCN Red List Status: Least Concern (LC)
• Nematoda:
• Cosmocerca brasiliense
• Physaloptera sp.

• Dendropsophus parviceps
• IUCN Red List Status: Least Concern (LC)
• Nematoda:
• Cosmocerca brasiliense

• Dendropsophus rhodopeplus
• IUCN Red List Status: Least Concern (LC)
• Nematoda:
• Cosmocerca brasiliense
• Acuarioidea gen. sp.

• Dendropsophus schubarti
• IUCN Red List Status: Least Concern (LC)
• Nematoda:
• Cosmocerca brasiliense

• Dendropsophus sarayacuensis
• IUCN Red List Status: Least Concern (LC)
• Digenea:
• Glypthelmis parva

• Boana boans
• IUCN Red List Status: Least Concern (LC)
• Nematoda:
• Cosmocerca brasiliense
• Physaloptera sp.

• Boana calcarata
• IUCN Red List Status: Least Concern (LC)
• Nematoda:
• Cosmocerca brasiliense

• Boana cinerascens
• IUCN Red List Status: Least Concern (LC)
• Nematoda:
• Brevimulticaecum sp.
• Cosmocerca brasiliense
• Physaloptera sp.

• Boana fasciata
• IUCN Red List Status: Least Concern (LC)
• Nematoda:
• Brevimulticaecum sp.
• Cosmocerca brasiliense
• Ochoterenella vellardi
• Physaloptera sp.
• Oswaldocruzia lopesi

• Osteocephalus taurinus
• IUCN Red List Status: Least Concern (LC)
• Monogenea:
• Wetapolystoma almae
• Nematoda:
• Cosmocerca brasiliense
• Batracholandros spectatus
• Ochoterenella vellardi
• Physaloptera sp.
• Physalopteroides venancioi

• Scarthyla goinorum
• IUCN Red List Status: Least Concern (LC)
• Nematoda:
• Brevimulticaecum sp.
• Cosmocerca brasiliense
• Cosmocerca parva

• Scinax garbei
• IUCN Red List Status: Least Concern (LC)
• Nematoda:
• Cosmocerca brasiliense
• Cosmocerca parva

• Scinax ictericus
• IUCN Red List Status: Least Concern (LC)
• Nematoda:
• Cosmocerca brasiliense
• Cosmocerca parva
• Physaloptera sp.

• Scinax pedromedinae
• IUCN Red List Status: Least Concern (LC)
• Digenea:
• Glypthelmis parva
• Cestoda:
• Cylindrotaenia americana
• Nematoda:
• Cosmocerca brasiliense
• Acantocephala:
• Acantocephala gen. sp.

• Scinax ruber
• IUCN Red List Status: Least Concern (LC)
• Nematoda:
• Physaloptera sp.
• Physalopteroides venancioi

• Sphaenorhynchus lacteus
• IUCN Red List Status: Least Concern (LC)
• Nematoda:
• Brevimulticaecum sp.
• Cosmocerca brasiliense

• Trachycephalus coriaceus
• IUCN Red List Status: Least Concern (LC)
• Digenea:
• Glypthelmis parva
• Nematoda:
• Brevimulticaecum sp.
• Cosmocerca brasiliense
• Physaloptera sp.
• Physalopteroides venancioi
• Oswaldocruzia lopesi

• Trachycephalus typhonius
• IUCN Red List Status: Least Concern (LC)
• Nematoda:
• Batracholandros spectatus
• Physaloptera sp.

• Callimedusa atelopoides
• IUCN Red List Status: Least Concern (LC)
• Nematoda:
• Cosmocerca brasiliense
• Cosmocerca parva

• Pithecopus palliatus
• IUCN Red List Status: Least Concern (LC)
• Nematoda:
• Cosmocercella phyllomedusae
• Acantocephala:
• Acantocephala gen. sp.

• Callimedusa tomopterna
• IUCN Red List Status: Least Concern (LC)
• Nematoda:
• Ophidascaris sp.
• Cosmocerca brasiliense
• Cosmocercella phyllomedusae
• Physaloptera sp.

• Phyllomedusa vaillanti
• IUCN Red List Status: Least Concern (LC)
• Nematoda:
• Cosmocerca brasiliense
• Cosmocercella phyllomedusae
• Physalopteroides venancioi

• Pseudis paradoxa
• IUCN Red List Status: Least Concern (LC)
• Nematoda:
• Physaloptera sp.

• Family Leptodactylidae

• Adenomera andreae
• IUCN Red List Status: Least Concern (LC)
• Nematoda:
• Cosmocerca brasiliense

• Adenomera hylaedactyla
• IUCN Red List Status: Least Concern (LC)
• Nematoda:
• Cosmocerca brasiliense
• Physalopteroides venancioi

• Edalorhina perezi
• IUCN Red List Status: Least Concern (LC)
• Nematoda:
• Cosmocerca brasiliense
• Cosmocerca parva
• Acuarioidea gen. sp.
• Physaloptera sp.
• Physalopteroides venancioi

• Leptodactylus bolivianus
• IUCN Red List Status: Least Concern (LC)
• Nematoda:
• Brevimulticaecum sp.
• Aplectana hylambatis
• Cosmocerca brasiliense
• Physaloptera sp.
• Physalopteroides venancioi
• Oswaldocruzia lopesi

• Leptodactylus leptodactyloides
• IUCN Red List Status: Least Concern (LC)
• Nematoda:
• Aplectana hylambatis
• Cosmocerca parva
• Cosmocerca podicipinus
• Physaloptera sp.

• Leptodactylus mystaceus
• IUCN Red List Status: Least Concern (LC)
• Nematoda:
• Schrankiana larvata
• Schrankiana schrankai
• Cosmocerca parva
• Ocheterenella digicauda
• Physaloptera sp.
• Physalopteroides venancioi
• Leptodactylus pentadactylus
• IUCN Red List Status: Least Concern (LC)
• Nematoda:
• Brevimulticaecum sp.
• Schrankiana larvata
• Schrankianella brasili
• Aplectana hylambatis
• Cosmocerca brasiliense
• Physaloptera sp.

• Leptodactylus petersii
• IUCN Red List Status: Least Concern (LC)
• Nematoda:
• Cosmocerca brasiliense

• Leptodactylus rhodonotus
• IUCN Red List Status: Least Concern (LC)
• Digenea:
• Gorgoderina parvicava
• Nematoda:
• Oswaldocruzia subauricularis
• Schrankiana inconspicata
• Aplectana hylambatis
• Physaloptera sp.
• Schulzia chiribita

• Lithodytes lineatus
• IUCN Red List Status: Least Concern (LC)
• Cestoda:
• Cylindrotaenia americana

• Pleurodema marmoratum
• IUCN Red List Status: Vulnerable (VU)
• Nematoda:
• Aplectana hylambatis
• Oxyascaris sp.

• Family Microhylidae

• Ctenophryne geayi
• IUCN Red List Status: Least Concern (LC)
• Nematoda:
• Physaloptera sp.

• Elachistocleis ovalis
• IUCN Red List Status: Least Concern (LC)
• Nematoda:
• Aplectana hylambatis
• Cosmocerca parva
• Raillietnema gubernaculatum

• Hamptophryne boliviana
• IUCN Red List Status: Least Concern (LC)
• Nematoda:
• Cosmocerca parva
• Physaloptera sp.
• Oswaldocruzia lopesi

• Family Pipidae

• Pipa pipa
• IUCN Red List Status: Least Concern (LC)
• Nematoda:
• Porrocaecum sp.

• Family Ranidae

• Lithobates palmipes
• IUCN Red List Status: Least Concern (LC)
• Digenea:
• Iquitos ceii

• Family Strabomantidae

• Oreobates cruralis
• IUCN Red List Status: Least Concern (LC)
• Nematoda:
• Cosmocerca brasiliense
• Physaloptera sp.

• Oreobates quixensis
• IUCN Red List Status: Least Concern (LC)
• Nematoda:
• Cosmocerca parva

• Pristimantis fenestratus
• IUCN Red List Status: Least Concern (LC)
• Nematoda:
• Cosmocerca brasiliense
• Cosmocerca parva
• Physaloptera sp.
• Physalopteroides venancioi
• Oswaldocruzia lopesi

• Pristimantis imitatrix
• IUCN Red List Status: Least Concern (LC)
• Nematoda:
• Cosmocerca podicipinus

• Pristimantis nephophilus
• IUCN Red List Status: Near Threatened (NT)
• Cestoda:
• Spirometra sp.

• Pristimantis peruvianus
• IUCN Red List Status: Least Concern (LC)
• Nematoda:
• Cosmocerca brasiliense
• Cosmocerca parva

• Pristimantis rhodostichus
• IUCN Red List Status: Least Concern (LC)
• Cestoda:
• Spirometra sp.

• Pristimantis toftae
• IUCN Red List Status: Least Concern (LC)
• Nematoda:
• Cosmocerca brasiliense
• Cosmocerca parva

• Family Telmatobiidae

• Telmatobius brachydactylus
• IUCN Red List Status: Endangered (EN)
• Nematoda:
• Hedruris juninensis
• Digenea:
• Gorgoderina carli
• Gorgoderina chilensis
• Gorgoderina megacysta

• Telmatobius macrostomus
• IUCN Red List Status: Endangered (EN)
• Digenea:
• Gorgoderina chilensis
• Gorgoderina parvicava
• Gorgoderina parvicava minuta
• Gorgoderina sp.
• Cestoda:
• Batrachotaenia sp.
• Nematoda:
• Falcaustra condorcanquii
• Hedruris juninensis

• Telmatobius culeus
• IUCN Red List Status: Endangered (EN)
• Digenea:
• Gorgoderina parvicava

• Telmatobius jelskii
• IUCN Red List Status: Near Threatened (NT)
• Digenea:
• Gorgoderina attenuata
• Gorgoderina parvicava
• Gorgoderina sp.
• Cestoda:
• Cylindrotaenia americana
• Ophiotaenia sp.
• Nematoda:
• Aplectana hylambatis
• Capillaria sp.
• Cosmocercidae gen. sp.
• Falcaustra mascula
• Falcaustra condorcanquii
• Camallanus sp.
• Hedruris moniezi
• Hedruris sp.
• Acantocephala:
• Centrorhynchus sp.
• Oncicola sp.

• Telmatobius marmoratus
• IUCN Red List Status: Endangered (EN)
• Cestoda:
• Cylindrotaenia americana
• Nematoda:
• Aplectana hylambatis
• Falcaustra mascula
• Hedruris moniezi

• Telmatobius peruvianus
• IUCN Red List Status: Vulnerable (VU)
• Digenea:
• Gorgoderina parvicava
• Haematoloechus arequipensis
• Haematoloechus pukinensis
• Nematoda:
• Falcaustra condorcanquii
• Hedruris moniezi
• Telmatobius sp.
• IUCN Red List Status: not applied
• Digenea:
• Gorgoderina alobata

4. Discussion

To date, a total of 83 helminth parasite taxa have been documented in 78 amphibian species in Peru, marking a significant increase of 176.7% in comparison to the earlier findings reported by Campião et al. [2], who registered only 30 taxa in Peru. Nematodes exhibit the highest species richness with 52 taxa, followed by digeneans with 21 taxa. Gonzalez and Hamann [60] reported 47 nematode taxa in 34 species of Argentinean amphibians. In contrast, Fernandez and Kohn [7] cited only 11 species of trematode parasites in Peruvian amphibians. This diversity pattern is consistent with that observed in helminth parasites of other anurans across South American countries [2]. The majority of collected parasites in Peruvian amphibians were identified as mature helminths. Campião et al. [2] noted that adult nematodes are typically found in the gastrointestinal tract and are the most prevalent helminths in South American amphibians. Nematode species have been reported as adults and have dominated the helminth fauna of Argentinean amphibians [60].

Among nematodes, C. brasiliense, Physaloptera sp., A. hylambatis, and C. parva displayed the highest number of hosts, with 34, 27, 18, and 17 hosts, respectively. In South American countries, C. parva, Cosmocerca podicipinus, A. hylambatis, Oswaldocruzia proencai, and G. palmipedis have been documented to exhibit a wide distribution across five countries and infect a greater number of hosts [2,61,62].

In Peru, R. marina, followed by Rhinella margaritifera, hosted the most diverse range of helminth parasites (17 spp. and 16 spp., respectively). In South America, Leptodactylus latrans and Rhinella marina were the hosts with the highest number of reported helminth taxa, with 81 and 47 helminth species, respectively [2]. In Argentina, the bufonid Rhinella schneideri (Werner, 1894) and the leptodactylid Leptodactylus bufonius Boulenger, 1894 displayed the highest species richness of parasitic nematodes (nine spp.) among Argentinean amphibians [60]. In Mexico, anuran host species with the largest number of parasite species are R. marina with 49 species, L. vaillanti with 34, and Lithobates montezumae Baird, 1854, with 30 [13]. Campião et al. [2] indicated that the genus Rhinella (Fitzinger, 1826) had among the highest numbers of helminths in South America [63]. Rhinella marina and R. margaritifera are species complexes of semiaquatic toads found in tropical and subtropical South America and Central America. This species group is poorly delineated, with numerous undescribed cryptic species, which could explain the high diversity of helminth parasites infecting this group [63,64,65,66]. Rhinella marina is known for having a highly diverse diet, primarily consisting of ants and Coleoptera larvae, which may contribute to the high diversity of helminth parasites in this anuran species [67]. In South America, the most prominent families of anurans are Hylidae and Leptodactylidae, comprising nearly 60% of the species studied for helminth parasites [2]. In Peru, Hylidae and Leptodactylidae also have the most hosts evaluated, but Bufonidae and Telmatobiidae are the families with the highest richness of helminth parasites. In Argentina, Bufonidae and Leptodactylidae exhibit the highest species richness of nematodes [60].

Regarding the conservation status of amphibian hosts, it was observed that three families, namely Telmatobiidae, Bufonidae, and Hemiphractidae, contained the highest percentages of host species considered of conservation importance. Understanding the diversity of parasites in amphibians, particularly those species listed as endangered, is crucial. Helminth parasites are integral components of biodiversity, and their extinction can occur in tandem with that of their hosts.

Among the 78 amphibians surveyed for helminths in Peru, there were no caecilian or caudata species. Remarkably, nearly 88% of Peruvian amphibians have not been subjected to studies on helminth parasites. The helminth parasite fauna is a crucial component of the natural history of amphibian hosts and can serve as a valuable and predictive tool for understanding parasite–host interactions and supporting biodiversity conservation research [13,68,69,70,71,72,73,74,75]. However, further research in this area is warranted.

5. Conclusions

This comprehensive update on helminth parasite biodiversity and richness in Peruvian amphibians provides valuable insights into the diversity and distribution of helminth parasites in this region. This research has important implications for conservation, as understanding the diversity and distribution of helminth parasites can aid in the development of conservation strategies and the use of parasites as bioindicators of contamination in specific areas. Additionally, the study highlights the potential threat of extinction faced by helminth parasites if their hosts become threatened. Overall, this study represents a significant contribution to the knowledge of helminth parasite biodiversity in Peruvian amphibians and underscores the need for further research in this field to protect both the amphibian hosts and their associated parasites.