Metazoan Endoparasites of 12 Species of Lizards from Argentina Author(s): Stephen R. Goldberg, Charles R. Bursey, Mariana Morando Source: Comparative Parasitology, 71(2):208-214. 2004. Published By: The Helminthological Society of Washington DOI: http://dx.doi.org/10.1654/4089 URL: http://www.bioone.org/doi/full/10.1654/4089
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Comp. Parasitol. 71(2), 2004, pp. 208–214
Metazoan Endoparasites of 12 Species of Lizards from Argentina STEPHEN R. GOLDBERG,1,4 CHARLES R. BURSEY,2
AND
MARIANA MORANDO,3
1
Department of Biology, Whittier College, Whittier, California 90608, U.S.A. (e-mail:
[email protected]), Department of Biology, Pennsylvania State University, Shenango Campus, Sharon, Pennsylvania 16146, U.S.A. (e-mail:
[email protected]), and 3 CRILAR-CONICET, Mendoza y Entre Rı´os s/n 5301, Anillaco, La Rioja, Argentina. 2
Twenty-five lizards representing 2 species of Leiosauridae, Leiosaurus belli and Leiosaurus catamarcensis, and 10 species of Liolaemidae, Liolaemus andinus, Liolaemus buergeri, Liolaemus chiliensis, Liolaemus elongatus, Liolaemus lemniscatus, Liolaemus neuquensis, Liolaemus pictus argentius, Liolaemus tenuis, Liolaemus vallecurensis, and Phymaturus palluma, from Argentina were examined for endoparasites. One species of Cestoda, Oochoristica travassosi, 3 species of Nematoda, Physaloptera retusa, Parapharyngodon riojensis, and Spauligodon maytacapaci, and 1 species of Pentastomida, Kiricephalus sp. (nymphs), were found. Twelve new host records are reported. KEY WORDS: endoparasites, Cestoda, Nematoda, Pentastomida, Argentina, lizards. ABSTRACT:
and small and large intestines were examined for metazoan endoparasites under a dissecting scope. Nematodes and pentastomes were cleared in a drop of concentrated glycerol on a glass slide and identified from these temporary preparations. Cestodes were regressively stained in Mayer hematoxylin and mounted in Canada balsam for identification. Lizards were deposited in the L. J. Avila and M. Morando herpetology collection housed in Centro Regional de Investigacio´nes Cientı´ficas y Tecnolo´gicas (CRILAR), La Rioja, Argentina. Endoparasites were deposited in the United States National Parasite Collection, Beltsville, Maryland, U.S.A.
Little information is available on the metazoan endoparasites of lizards of Argentina. Khalil and Vogelsang (1930) described the nematode Cruzia fulleborni Khalil and Vogelsang, 1930, from the intestine of Tupinambis teguixin. Cruz et al. (1998) studied the ecology of Tropidurus etheridgei and reported the presence of nematodes they assigned to Physaloptera, Thelandros, and Parapharyngodon. Ramallo et al. (2002a) described the nematode Spauligodon loboi Ramallo, Bursey and Goldberg, 2002, from Liolaemus capillitas and reported it to also occur in Liolaemus ornatus, Liolaemus quilmes, and Liolaemus ramirezae. Ramallo et al. (2002b) described Parapharyngodon riojensis Ramallo, Bursey and Goldberg, 2002, from Phymaturus punae. The purpose of this article is to report 12 new host and locality records for metazoan endoparasites from lizards collected in Argentina.
Leiosaurus belli One specimen (gender not recorded) was collected sometime between 1997 and 1998 near Chichinales, Rı´o Negro Province, General Roca Department, Argentina (388499S; 668479W). Physaloptera retusa Rudolphi, 1819 (¼Physaloptera mucronata Leidy, 1856; Physaloptera largada Sprehn, 1932)
MATERIALS AND METHODS During 1997–1999, 1 Leiosaurus belli, 2 Leiosaurus catamarcensis, 2 Liolaemus andinus, 2 Liolaemus buergeri, 1 Liolaemus chiliensis, 1 Liolaemus elongatus, 1 Liolaemus lemniscatus, 2 Liolaemus neuquensis, 1 Liolaemus pictus argentinus, 1 Liolaemus tenuis, 4 Liolaemus vallecurensis, and 7 Phymaturus palluma were collected by hand in Argentina for studies of the phylogeny and phylogeography of Argentinian lizards and were later made available for parasitological study. Lizards were killed by an overdose of sodium pentobarbital, preserved in 10% neutral buffered formalin, and stored in 70% ethanol until examination. The body cavity was opened by a longitudinal incision from vent to throat, and the digestive tract was removed. The body cavity and liver were examined visually for endoparasites. The digestive tract was opened, and the esophagus, stomach,
4
Prevalence and intensity: One of 1 host infected (100%, 211). Temporal distribution: Collection date unknown, 1997–1998. Site of infection: Stomach. Type host and type locality: Tupinambis teguixin, Brazil (Rudolphi, 1819). Other reported hosts: Ameiva ameiva (Cristofaro et al., 1976; Ribas et al., 1997); Ameiva undulata (Caballero, 1951); Amphisbaena alba (Molin, 1860); Barisia imbricata (Goldberg et al., 1999); Callisaurus draconoides (Telford, 1970); Cnemidophorus
Corresponding author. 208
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burti (Goldberg and Bursey, 1989a); Cnemidophorus lemniscatus (Caballero and Vogelsang, 1947; DiazUngria, 1964; Diaz-Ungria and Gallardo, 1968); Cnemidophorus littoralis (Vrcibradic et al., 2000); Cnemidophorus murinus (Specian and Whittaker, 1980); Cnemidophorus ocellifer (Ribas et al., 1995); Gambelia wislizenii (Telford, 1970); Elgaria multicarinata (Telford, 1970); Eumeces skiltonianus (Telford, 1970); Gerrhonotus ophiurus (Goldberg et al., 1999); Iguana iguana (Diaz-Ungria and Gallardo, 1968); Liolaemus lutzae (Rocha, 1995); Mabuya agilis (Ribas et al., 1998); Mabuya bistriata (recorded as Scincus spixii by Molin [1860]); Sceloporus acanthinus (Caballero, 1951); Sceloporus clarkii (Goldberg et al., 1994); Sceloporus serrifer (Goldberg et al., 1995); Sceloporus formosus (Goldberg et al., 2003); Sceloporus graciosus (Woodbury, 1934; Goldberg and Bursey, 1989b; Goldberg et al., 1997); Sceloporus jarrovii (Goldberg and Bursey, 1990; Bursey and Goldberg, 1991, 1994; Goldberg et al., 1996); Sceloporus magister (Pearce and Tanner, 1973; Walker and Matthias, 1973; Goldberg et al., 1994, 1995); Sceloporus merriami (Goldberg et al., 1995); Sceloporus mucronatus (Goldberg et al., 2003); Sceloporus occidentalis (Grundmann, 1959; Telford, 1970; Pearce and Tanner, 1973; Walker and Matthias, 1973; Goldberg et al., 1998); Sceloporus olivaceus (Goldberg et al., 1995); Sceloporus orcutti (Telford, 1970); Sceloporus parvus (Goldberg et al., 2003); Sceloporus poinsettii (Goldberg et al., 1993, 1995); Sceloporus torquatus (Goldberg et al., 2003); Sceloporus undulatus (Morgan, 1943; Pearce and Tanner, 1973; Goldberg et al., 1994, 1995); Sceloporus variabilis (Goldberg et al., 1995, 2003); Sceloporus virgatus (Goldberg et al., 1994); Tropidurus torquatus (Vicente and Santos, 1967; Cristofaro et al., 1976; Vicente, 1981; Ribas et al., 1998; Vrcibradic et al., 2000); Tropidurus spinulosus (Vicente, 1981); Tupinambis nigropunctatus (Diesing in Morgan, 1943); Tupinambis rufescens (Sprehn, 1932); Tupinambis teguixin (Rudolphi, 1819; Molin, 1860; Ortlepp, 1922; Lent and Freitas, 1948); Uta stansburiana (Telford, 1970). Geographic range: Argentina, Brazil, Mexico, United States, Venezuela, West Indies. Specimens deposited: United States National Parasite Collection 92471 (1 vial). Remarks: The life cycle of P. retusa has not been studied; however, the life cycle studies of Physaloptera hispida, Physaloptera maxillaris, Physaloptera praeputialis, and Physaloptera rara have shown that
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third-stage infective larvae occur in earwigs, crickets, grasshoppers, cockroaches, and beetles (Schell, 1952; Lincoln and Anderson, 1975). Insects scavenging fecal material ingest eggs, which hatch and migrate into body tissue for subsequent development to thirdstage larvae. Leiosaurus belli represents a new host record for P. retusa. Leiosaurus catamarcensis Two specimens (1 male and 1 gender not recorded) were collected 6 km east of Anillaco, La Rioja Province, Castro Barros Department, Argentina (288479S; 668529W), on 13 November 1998 and 2 February 1999, respectively. Physaloptera retusa Rudolphi, 1819 Prevalence and intensity: Hosts infected, 2 of 2 (100%, 2.5 6 1.5, 1–4). Temporal distribution: On 13 November 1998, 1 host with 1; on 2 February 1999, 1 host with 4. Site of infection: Stomach. Specimens deposited: United States National Parasite Collection 92472 (1 vial). Remarks: General information and remarks are reported under L. belli. Leiosaurus catamarcensis represents a new host record for P. retusa. Liolaemus andinus Two specimens (1 male, 1 female) were collected 11 km east of Chaschuil, Catamarca Province, Tinogasta Department, Argentina (278489S; 688079W), on 13 November 1998. Spauligodon maytacapaci (Vicente and Iba´n˜ez, 1968) Barus and Coy Otero, 1974 (¼Parathelandros maytacapaci Vicente and Iba´n˜ez, 1968) Prevalence and intensity: Two of 2 hosts infected (100%, 9.5 6 6.4, 5–14). Temporal distribution: On 13 November 1998, male host with 5; female host with 14. Type host and type locality: Leiocephalus sp.; Peru (Vicente and Iba´n˜ez, 1968). Geographic range: Argentina, Chile, Peru. Other reported hosts: Liolaemus lemniscatus (Goldberg et al., 2001); L. tenuis (Goldberg et al., 2001).
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Specimens deposited: United States National Parasite Collection 92473 (1 vial). Remarks: The life cycle of S. maytacapaci has not been studied, but members of the Oxyurida are strictly monoxenous (Anderson, 2000). Liolaemus andinus represents a new host record for S. maytacapaci. Liolaemus buergeri
Remarks: General information and remarks are reported under L. andinus. Liolaemus chiliensis represents a new host record for S. maytacapaci. Liolaemus elongatus One specimen (gender not recorded) was collected 17.5 km west of Primeros Pinos, Neuque´n Province, Alumine´ Department, Argentina (388539S; 708589W), on 11 January 1999.
Two specimens (gender not recorded) were collected 22 km west of Las Len˜as, Mendoza Province, Malargu¨e Department, Argentina (358089S; 708119W), on 15 January 1999.
Prevalence and intensity: One of 1 host infected (100%, 1).
Parapharyngodon riojensis Ramallo, Bursey and Goldberg, 2002
Temporal distribution: On 11 January 1999, 1 host with 1.
Prevalence and intensity: Two of 2 hosts infected (100%, 21.5 6 16.3 SD, 10–33).
Site of infection: Large intestine.
Temporal distribution: On 15 January 1999, 1 host with 10, 1 host with 33. Infection site: Large intestine. Type host and type locality: Phymaturus punae, Argentina (Ramallo et al., 2002b). Geographic range: Argentina.
Spauligodon maytacapaci (Vicente and Iba´n˜ez, 1968)
Specimens deposited: United States National Parasite Collection 92476 (1 vial). Remarks: General information and remarks are reported under L. andinus. Liolaemus elongatus represents a new host record for S. maytacapaci. Liolaemus lemniscatus
Specimens deposited: United States National Parasite Collection 92474 (1 vial).
One specimen (gender not recorded) was collected in Neuque´n Province, Catan Lil Department, Argentina (398329S; 708579W), 24 km north of Pilolil on 10 January 1999.
Remarks: The life cycle of P. riojensis has not been studied, but members of the Oxyurida are strictly monoxenous (Anderson, 2000). Liolaemus buergeri represents a new host record for P. riojensis.
Kiricephalus sp. (nymphs)
Liolaemus chiliensis One specimen (gender not recorded) was collected 1 km east of Curri Huinca, Neuque´n Province, Lacar Department, Argentina (408109S; 718209W), on 6 January, 1999. Spauligodon maytacapaci (Vicente and Iba´n˜ez, 1968) Prevalence and intensity: One of 1 host infected (100%, 1). Temporal distribution: On 6 January 1999, 1 host with 1. Site of infection: Large intestine. Specimens deposited: United States National Parasite Collection 92475 (1 vial).
Prevalence and intensity: One of 1 host infected (100%, 11). Temporal distribution: On 10 January 1999, 1 host with 11. Site of infection: Cysts in body cavity. Type host: Species of Kiricephalus are parasites of snakes. Reported hosts of nymphs: Kiricephalus coarctatus (Diesing, 1850), a parasite of snakes, is the common pentastomid species found in the Western Hemisphere. Nymphs of K. coarctatus have been reported (Riley and Self, 1980) from Agkistrodon piscivorus, Coluber constrictor, Crotalus adamanteus, Erythrolamprus aesculapii, Erythrolamprus bizona, Lampropeltis getula, Nerodia cyclopion, Nerodia erythrogaster, Nerodia fasciata, Nerodia rhombifer, Nerodia sipedon, Nerodia taxispilota, Pituophis melanoleucus, and Thamnophis sirtalis.
GOLDBERG ET AL.—ENDOPARASITES OF ARGENTINE LIZARDS
Kiricephalus pattoni (Stephens, 1908), also a parasite of snakes, is the common pentastomid species of the Eastern Hemisphere. Nymphs have been reported (Riley and Self, 1980) from Amphiesma sauteri, Amphiesma stolata, Boiga kraepelini, Bungarus multicinctus, Daboia russellii, Deinagkistrodon acutus, Dinodon rufozonatum, Elaphe porphyracea, Enhydris chinensis, Enhydris plumbea, Morelia spilota, Naja naja, Opheodrys major, Psammodynastes pulverulentus, Protobothrops mucrosquamatus, Rhabdophis swinhonis, Sinonatrix annularis, Sinonatrix percarinata, Trimeresurus gramineus, Trimeresurus stejnegeri, Vipera ammodytes, Xenochrophis piscator, Hemidactylus frenatus, Japalura swinhonis, Bufo bankorensis, Bufo bufo, Bufo melanostictus, and Rana limnocharis. Specimens deposited: United States National Parasite Collection 92477 (1 vial). Remarks: Keegan et al. (1969) fed eggs of K. pattoni to an agamid lizard, J. swinhonis, and a toad, B. bankorensis, and recovered encysted nymphs at necropsy 90 d later. Bufo woodhousii and Rana pipiens have also been experimentally infected (Riley and Self, 1980). Riley and Self (1980) suggested a 3-host life cycle, in which amphibians, lizards, and mammals can be directly infected by eggs of Kiricephalus spp., thus becoming first intermediate hosts. Many species of snakes may serve as second intermediate hosts (having nymphs but known to be refractory to infection by egg ingestion); definitive hosts have broad diets, commonly including mammals, reptiles, and amphibians (Riley and Self, 1980). Liolaemus lemniscatus represents a new host record for nymphs of Kiricephalus sp. Liolaemus neuquensis Two specimens (1 male, 1 gender not recorded) were collected 3 km southeast of Termas de Copahue, Neuque´n Province, Norquin Department, Argentina (408299S; 718259W), on 13 January 1999. Physaloptera retusa Rudolphi, 1819 Prevalence and intensity: One of 2 hosts infected (50%, 1). Temporal distribution: On 13 January 1999, male host with 1. Site of infection: Stomach. Specimens deposited: United States National Parasite Collection 92478 (1 vial).
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Remarks: General information and remarks are reported under L. belli. Liolaemus neuquensis represents a new host record for P. retusa. Liolaemus pictus argentinus One male specimen was collected at Pichi Traful, Neuque´n Province, Los Lagos Department, Argentina (408299S; 718259W), on 6 January 1999. Spauligodon maytacapaci (Vicente and Iba´n˜ez, 1968) Prevalence and intensity: One of 1 host infected (100%, 11). Temporal distribution: On 6 January 1999, 1 host with 11. Site of infection: Large intestine. Specimens deposited: United States National Parasite Collection 92479 (1 vial). Remarks: General information and remarks are reported under L. andinus. Liolaemus p. argentinus represents a new host record for S. maytacapaci. Liolaemus tenuis One specimen was collected at Arroyo Remeco´, Neuque´n Province, Alumine´ Department (398029S; 718219W), on 11 January 1999. Spauligodon maytacapaci (Vicente and Iba´n˜ez, 1968) Prevalence and intensity: One of 1 host infected (100%, 1). Temporal distribution: On 11 January 1999, 1 host with 1. Site of infection: Small intestine. Specimens deposited: United States National Parasite Collection 92480 (1 vial). Remarks: General information and remarks are reported under L. andinus. Liolaemus tenuis represents a new host record for S. maytacapaci. Liolaemus vallecurensis Four specimens were collected: 1 female, 1 gender not recorded at the confluence of the Las Taguas River and the Arroyo Amarillo, San Juan Province, Iglesia Department, Argentina (298199S; 708009W), on 21 March 1998; 2 gender not recorded at Campamento Minero Compan˜ia Aguilar, San Juan
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Prevalence and intensity: Three of 4 hosts infected (75%, 3.7 6 4.6, 1–9)
1 cestode, O. travassosi; 5 nematodes, C. fulleborni, P. retusa, S. loboi, S. maytacapaci, and P. riojensis; and 1 pentastomid, Kiricephalus sp. Because Argentina has a diverse and taxonomically complex lizard fauna (Cei, 1986, 1993; Etheridge, 2000; Morando et al., 2003), subsequent helminthological studies should add other metazoan endoparasites to this list.
Temporal distribution: On 21 March 1998, 1 host with 1; on 21 January 1999, 1 host with 1, 1 host with 9.
LITERATURE CITED
Province, Iglesia Department, Argentina (298209S; 698589W), on 21 January 1999. Oochoristica travassosi Rego and Iba´n˜ez, 1965
Site of infection: Small intestine. Type host and type location: Leiocephalus sp., Peru (Rego and Iba´n˜ez, 1965). Other reported hosts: None. Specimens deposited: United States National Parasite Collection 92481 (2 slides). Remarks: The life cycle of O. travassosi has not been studied, but in the related Oochoristica anolis, Conn (1985) found that beetles served as intermediate hosts. Liolaemus vallecurensis represents a new host record for O. travassosi. Phymaturus palluma Seven specimens were collected (gender not recorded): 6 in Neuque´n Province 2 at 16.8 km south of Coli Pilli, Norquin Department, Argentina (378439S; 708299W), on 13 January 1999; 4 at Copahue, Norquin Department, Argentina (378499S; 718059W), on 13 February 1999; 1 at Puente del Inca, Mendoza Province, Las Heras Department, Argentina (328519S; 698469W), on 9 February 1999. Parapharyngodon riojensis Ramallo, Bursey and Goldberg, 2002 Prevalence and intensity: Seven of 7 hosts infected (100%, 16.1 6 19.3 SD, 2–49). Temporal distribution: On 13 January 1999, 4 hosts with 2, 3, 39, 39; on 9 February 1999, 1 host with 7; on 13 February 1999, 2 hosts with 4, 9. Infection site: Large intestine. Specimens deposited: United States National Parasite Collection 92482 (1 vial). Remarks: General information and remarks are reported under L. buergeri. Phymaturus palluma represents a new host record for P. riojensis. DISCUSSION This study increases the number of known metazoan endoparasites of Argentine lizards to 7 species:
Anderson, R. C. 2000. Nematode Parasites of Vertebrates: Their Development and Transmission, 2nd ed. CABI Publishing, Wallingford, Oxon, U.K. 650 pp. Bursey, C. R., and S. R. Goldberg. 1991. Monthly prevalences of Physaloptera retusa in naturally infected Yarrow’s spiny lizard. Journal of Wildlife Diseases 27: 710–715. Bursey, C. R., and S. R. Goldberg. 1994. Persistence of the component parasite community of Yarrow’s spiny lizard, Sceloporus jarrovii, 1967–1991. Journal of the Helminthological Society of Washington 61:141–145. Caballero, E. 1951. Estudios helmintolo´gicos de la regio´n oncocercosa de Me´xico y de la Repu´blica de Guatemala. Nematoda, 6a parte y algunas consideraciones en torno a los ge´neros Onchocerca Diesing, 1841 y Acanthospiculum Skrjabin y Schikhobalowa, 1948. Anales del Instituto de Biologı´a de la Universidad Nacional Auto´noma de Me´xico 20:141–158. Caballero, E., and E. G. Vogelsang. 1947. Fauna helmintolo´gica venezolana. I. Ochetosoma miladelarocai n. sp. de Bothrops atrox L. y hallazgo de Physaloptera retusa (Rud., 1819) en Cnemidophorus lemniscatus lemniscatus. Revista de Medicina Veterinaria y Parasitologia 6:53–62. Cei, J. M. 1986. Reptiles del centro, centro-oeste y sur de la Argentina: Herpetofauna de las zonas a´ridas y semia´ridas. Museo regionale di Scienze Naturali Torino, Monografie IV, Torino, Italy. 527 pp. Cei, J. M. 1993. Reptiles del noroeste, nordeste y este de la Argentina: Herpetofauna de las selvas subtropicales, puna y pampas. Museo Regionale di Scienze Naturali Torino, Monografie XIV, Torino, Italy. 949 pp. Conn, D. B. 1985. Life cycle and postembryonic development of Oochoristica anolis (Cyclophyllidea: Linstowiidae). Journal of Parasitology 71:10–16. Cristofaro, R., J. F. Guimar¼es, and H. de Oliveira Rodrigues. 1976. Alguns Nemato´deos de Tropidurus torquatus (Wied) e Ameiva ameiva (L.)—Fauna Helmintolo´gica de Salvador, Bahia. Atas da Sociedade de Biologia do Rio de Janeiro 18:65–70. Cruz, F. B., S. Silva, and G. J. Scrocchi. 1998. Ecology of the lizard Tropidurus etheridgei (Squamata: Tropiduridae) from the dry Chaco of Salta, Argentina. Herpetological Natural History 6:23–31. Diaz-Ungria, C. 1964. Notas sobre nematodes de reptiles en Venezuela. Revista Veterenaria Venezolana 17:402. Diaz-Ungria, C., and M. F. Gallardo. 1968. Nematodes de reptiles Venezolanos, con descripcio´n de varias especies nuevas. Boletaˆin Sociedad Venezolana de Ciencias Naturales 27:550–570. Etheridge, R. 2000. A review of the lizards of the Liolaemus wiegmannii group (Squamata, Iguania, Tropiduridae), and a history of morphological changes in the
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