Gastrointestinal Helminths from 13 Species of Lizards from Reserva Cuzco Amazónico, Peru

Gastrointestinal Helminths from 13 Species of Lizards from Reserva Cuzco Amazónico, Peru Author(s): Charles R. Bursey, Stephen R. Goldberg, Jeffrey R. Parmelee Source: Comparative Parasitology, 72(1):50-68. 2005. Published By: The Helminthological Society of Washington DOI: http://dx.doi.org/10.1654/4132 URL: http://www.bioone.org/doi/full/10.1654/4132

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Comp. Parasitol. 72(1), 2005, pp. 50–68

Gastrointestinal Helminths from 13 Species of Lizards from Reserva Cuzco Amazo´nico, Peru CHARLES R. BURSEY,1,4 STEPHEN R. GOLDBERG,2

AND

JEFFREY R. PARMELEE3

1

Department of Biology, The Pennsylvania State University, Shenango Campus, Sharon, Pennsylvania 16146, U.S.A. (e-mail: [email protected]), 2 Department of Biology, Whittier college, Whittier, California 90608, U.S.A. (e-mail: [email protected]), and 3 Department of Biology, Simpson College, Indianola, Iowa 50125, U.S.A. (e-mail: [email protected]) ABSTRACT: One hundred sixty-four lizards representing 13 species collected from the Reserva Cuzco Amazo ´ nico, Peru, were examined for helminths. Two species of Cestoda (Oochoristica ameivae and Ophiotaenia flava) and 18 species of Nematoda (adults of Amphibiocapillaria freitaslenti, Cyrtosomum longicaudatum, Oswaldocruzia peruvensis, Oswaldocruzia vitti, Oswaldofilaria azeuedoi, Parapharyngodon scleratus, Physaloptera retusa, Physalopteroides venancioi, Piratuba digiticauda, Piratuba lainsoni, Piratuboides zeae, Rhabdias anolis, Skrjabinelzaia intermedia, Spauligodon oxkutzcabiensis, Spinicauda spinicauda, Strongyluris oscari; larvae of Dujardinascaris sp. and Hastospiculum sp.) were found. In all, 1,617 helminths were collected from 83 (51%) of the 164 lizards examined. Of these, 15 (0.9%) were larval forms, which are thought not capable of reaching maturity in lizards. No infected lizard harbored more than 3 helminth species: 1.70 6 0.08 (
x 6 1 SE) helminth species/infected lizard; 19.45 6 2.82 helminth individuals/infected lizard. No host species harbored more than 7 helminth species: 3.23 6 0.51 helminth species/host species. Twenty-eight new host records and 18 new locality records are reported. KEY WORDS: Cestoda, Nematoda, lizards, Reserva Cuzco Amazo ´ nico, Peru.

1; Tropiduridae: tree runner, Plica plica, 9; Harlequin racerunner, Plica umbra, 14; and the rose whorltail iguana, Stenocercus roseiventris, 12. Reserva Cuzco Amazo´nico is a privately maintained reserve of 10,000 ha on an alluvial plain, 200 m elevation, on the north bank of the Rio Madre de Dios about 15 km ENE of Puerto Maldonado, Departamento Madre de Dios, Amazonian Peru (128359S; 698059W). Duellman and Koechlin (1991) described the reserve in detail, and Duellman and Salas (1991) provided an annotated checklist of lizards in the area. Lizards were collected by hand over a decade long biotic survey by field parties coordinated by William Duellman, University of Kansas. The lizards were fixed in 10% buffered formalin within 6 hr of capture, preserved in 70% ethanol, and shipped to the Museum of Natural History at the University of Kansas, Lawrence, Kansas, U.S.A. Preserved specimens from the University of Kansas herpetological collections were dissected by J.R.P. as part of a study of the community ecology of frogs (Parmelee, 1999) and lizards of Reserva Cuzco Amazo´nico. The body cavity of each lizard was opened by a longitudinal incision from throat to vent, the gastrointestinal tract was slit longitudinally, and stomach and intestinal contents were removed and examined with a dissection microscope. Helminths found in the gastrointestinal tract, lungs, or body cavity were placed in vials of 70% ethanol for later identification, at which time each helminth was placed on a glass slide in a drop of undiluted glycerol for study under a compound microscope. Nematodes were identified from these preparations; cestodes were stained with hematoxylin and mounted in balsam for identification. Voucher specimens of all helminth species were deposited in the United States National Parasite Collection (USNPC), Beltsville, Maryland, U.S.A. Lizards were deposited in the Herpetology Collection (KU) of the Natural History Museum of the University of Kansas, Lawrence, Kansas, U.S.A.

Little information is available on the helminths of Peruvian lizards. Reˆgo and Iba´n˜ez (1965) described the cestode Oochoristica travassosi Reˆgo and Iba´n˜ez, 1965 from specimens taken from an undetermined species of Leiocephalus. Also from an undetermined species of Leiocephalus, Vicente and Iba´n˜ez (1968) described the nematode Parathelandros maytacapaci Vicente and Iba´n˜ez, 1968, which was reassigned to Spauligodon by Barus and Coy Otero (1974). We report helminths collected during the course of an ecological study of the lizard community of Reserva Cuzco Amazo´nico, Peru.

MATERIALS AND METHODS The 164 lizards representing 13 species used in this study were collected between November 1984 and December 1995 during a biotic survey of the Reserva Cuzco Amazo´nico, Peru. Taxa by family and sample sizes were as follows—Gekkonidae: turniptail gecko, Thecadactylus rapicauda, 10; Gymnophthalmidae: ocellated tegu, Cercosaura ocellata, 5; Eigenmann’s prionodactylus, Prionodactylus eigenmanni, 12; Polychrotidae: brown-eared anole, Anolis fuscoauratus, 25; spotted anole, Anolis punctatus, 14; Scincidae: two-striped mabuya, Mabuya bistriata, 11; Teiidae: giant ameiva, Ameiva ameiva, 25; Amazon kentropyx, Kentropyx altamazonicus, 11; forest whiptail, Kentropyx pelviceps, 15; black tegu, Tupinambis teguixin,

4

Corresponding author. 50

BURSEY ET AL.—HELMINTHS OF PERUVIAN LIZARDS

Gekkonidae Thecadactylus rapicauda (Houttuyn, 1782) Ten specimens were collected between December 1984 and December 1995 from tree trunks in Reserva Cuzco Amazo´nico, Peru (128359S; 698059W). Mean snout–vent length (SVL) ¼ 94 mm 6 21 mm SD, range 46–118 mm; 1 juvenile, 6 females, 3 males.

Physaloptera retusa Rudolphi, 1819 (Syn. Spiroptera retusa [Rudolphi, 1819] Dujardin, 1845; Physaloptera mucronata Leidy, 1856; Physaloptera largada Sprehn, 1932.) Prevalence, mean intensity, and range: Four of 10 hosts infected (40%, 8.8 6 8.2, 1–20). Temporal distribution: October 1987, 1 host with 5; February 1990, 1 host with 1; November 1990, 1 host with 20; November 1995, 1 host with 9. Site of infection: Stomach. Additional hosts from Peru: Ameiva ameiva, A. punctatus, K. altamazonicus, K. pelviceps, M. bistriata, P. plica, P. umbra, S. roseiventris, T. teguixin. Type host and type locality: Tupinambis teguixin, Brazil (Rudolphi, 1819). Other reported hosts: Common lesser toad, Bufo granulosus (Gonc¸alves et al., 2002); South American common toad, Bufo typhonius (Gonc¸alves et al., 2002); A. ameiva (Poinar and Vaucher, 1972; Cristofaro et al., 1976; Ribas, Rocha, et al., 1998); rainbow ameiva, Ameiva undulata (Caballero, 1951); red worm lizard, Amphisbaena alba (Molin, 1860); canyon spotted whiptail, Aspidoscelis burti (Goldberg and Bursey, 1989a, reported as Cnemidophorus burti); imbricata alligator lizard, Barisia imbricata (Goldberg et al., 1999); zebratail lizard, Callisaurus draconoides (Telford, 1970); rainbow whiptail, Cnemidophorus lemniscatus (Caballero and Vogelsang, 1947; Diaz-Ungria, 1964; Diaz-Ungria and Gallardo, 1968); Cnemidophorus littoralis (Vrcibradic et al., 2000); Laurent’s whiptail, Cnemidophorus murinus (Specian and Whittaker, 1980); Spix’s whiptail, Cnemidophorus ocellifer (Ribas et al., 1995); southern alligator lizard, Elgaria multicarinata (Telford, 1970); western skink, Eumeces skiltonianus (Telford, 1970); longnose leopard lizard, Gambelia wislizenii (Telford, 1970); Gerrhonotus ophiurus (Goldberg et al., 1999); green iguana,

51

Iguana iguana (Diaz-Ungria and Gallardo, 1968); Bell’s anole, Leiosaurus belli (Goldberg et al., 2004); Leiosaurus catamarcensis (Goldberg et al., 2004); Lutz’s tree iguana, Liolaemus lutzae (Rocha, 1995); Liolaemus neuquensis (Goldberg et al., 2004); Mabuya agilis (Ribas, Teixeira-Filho, et al., 1998); twostriped mabuya, M. bistriata (recorded as Scincus spixii by Molin, 1860); Bocourt’s spiny lizard, Sceloporus acanthinus (Caballero, 1951); Clark’s spiny lizard, Sceloporus clarkii (Goldberg et al., 1994); blue spiny lizard, Sceloporus serrifer (Goldberg, Bursey, and McAllister, 1995); emerald spiny lizard, Sceloporus formosus (Goldberg et al., 2003); sagebrush lizard, Sceloporus graciosus (Woodbury, 1934; Goldberg and Bursey, 1989b; Goldberg et al., 1997); Yarrow’s spiny lizard, Sceloporus jarrovii (Goldberg and Bursey, 1990; Bursey and Goldberg, 1991, 1994; Goldberg, Bursey, and Bezy, 1995; Goldberg, Bursey, and Bezy, 1996); desert spiny lizard, Sceloporus magister (Pearce and Tanner, 1973; Walker and Matthias, 1973; Goldberg et al., 1994; Goldberg, Bursey, and McAllister, 1995); canyon lizard, Sceloporus merriami (Goldberg, Bursey, and McAllister, 1995); southern crevice lizard, Sceloporus mucronatus (Goldberg et al., 2003); western fence lizard, Sceloporus occidentalis (Grundmann, 1959; Telford, 1970; Pearce and Tanner, 1973; Walker and Matthias, 1973; Goldberg et al., 1998); Texas spiny lizard, Sceloporus olivaceus (Goldberg, Bursey, and McAllister, 1995); granite spiny lizard, Sceloporus orcutti (Telford, 1970); bluebelly lizard, Sceloporus parvus (Goldberg et al., 2003); crevice spiny lizard, Sceloporus poinsettii (Goldberg et al., 1993; Goldberg, Bursey, and McAllister, 1995); crevice swift lizard, Sceloporus torquatus (Goldberg et al., 2003); eastern fence lizard, Sceloporus undulatus (Morgan, 1943; Pearce and Tanner, 1973; Goldberg et al., 1994; Goldberg, Bursey, and McAllister, 1995); rosebelly lizard, Sceloporus variabilis (Goldberg, Bursey, and McAllister, 1995; Goldberg et al., 2003); striped plateau lizard, Sceloporus virgatus (Goldberg et al., 1994); black lava lizard, Tropidurus melanopleurus (Roca, 1997); Amazon lava lizard, Tropidurus torquatus (Vicente and Santos, 1967; Cristofaro et al., 1976; Vicente, 1981; Ribas, Rocha, et al., 1998; Vrcibradic et al., 2000); spiny lava lizard, Tropidurus spinulosus (Vicente, 1981); black spotted tegu, Tupinambis nigropunctatus (Diesing in Morgan, 1943); red tegu, Tupinambis rufescens (Sprehn, 1932); T. teguixin (Rudolphi, 1819; Molin, 1860; Ortlepp, 1922; Lent and Freitas, 1948; Schuurmans-Stekhoven, 1950); side-blotched lizard, Uta stansburiana (Telford, 1970).

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COMPARATIVE PARASITOLOGY, 72(1), JANUARY 2005

Geographic range: Argentina, Brazil, Mexico, United States, Venezuela, West Indies (see Baker [1987]); Peru. Specimen deposited: USNPC 93264 (1 vial). Remarks: Four species of Physaloptera occur in South America, Physaloptera liophis Vicente and Santos, 1974, Physaloptera lutzi Cristofaro, Guimaraes and Rodrigues, 1976, Physaloptera obtusissima Molin, 1860, and P. retusa. In P. liophis the vulva is located at midbody; in P. lutzi, slightly anterior to the anus; and, in the remaining 2 species the vulva is located in the anterior third of the body. In P. obtusissima, the 3 pairs of caudal papillae are equidistant from one another; in P. retusa, the middle pair of the caudal papillae is closer to the terminal pair. The life cycle of P. retusa has not been studied, but life cycle studies of Physaloptera hispida Schell, 1950, Physaloptera maxillaris Molin, 1860, Physaloptera praeputialis Linstow, 1889, and Physaloptera rara Hall and Wigdor, 1918 demonstrate that 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, which are infective to both definitive and paratenic hosts. Thecadactylus rapicauda represents a new host record, and Peru is a new location record for P. retusa. Physalopteroides venancioi (Lent, Freitas and Proenc¸a, 1946) Sobolev, 1949 (Syn. Physaloptera venancioi Lent, Freitas and Proenc¸a, 1946; Thubunaea dactyluris Karve, 1938 sensu Fabio and Rolas, 1974.) Prevalence, mean intensity, and range: Three of 10 hosts infected (30%, 5.7 6 0.6, 5–6). Temporal distribution: October 1987, 1 host with 6; November 1990, 1 host with 5; November 1995, 1 host with 6. Site of infection: Stomach. Additional hosts from Peru: Napo’s tropical bullfrog Adenomera hylaedactyla, Perez’s snouted frog Edalorhina perezi, Rı´o Mamore robber frog Eleutherodactylus fenestratus, Bolivian white-lipped frog Leptodactylus bolivianus, basin white-lipped frog Leptodactylus mystaceus, Manaus slender-legged treefrog Osteocephalus taurinus, Surinam golden-

eyed treefrog Phrynohyas coriacea, white-lined leaf frog Phyllomedusa vaillanti, red snouted treefrog Scinax ruba (Bursey et al., 2001); C. ocellata, K. altamazonicus, K. pelviceps, M. bistriata, T. teguixin. Type host and type locality: Cururu toad, Bufo paracnemis, Assunc¸a˜o, Paraguay (Lent et al., 1946). Other reported hosts: Ameiva ameiva (Fabio and Rolas, 1974; Vrcibradic, Vicente, and Bursey, 2000); M. agilis (Vrcibradic, Cunha-Barros, et al., 2000; Vrcibradic et al., 2002); Hoge’s mabuya, Mabuya macrorhyncha (Vrcibradic, Cunha-Barros, et al., 2000; Vrcibradic et al., 2001, 2002); T. torquatus (Vrcibradic, Cunha-Barros, et al., 2000). Geographic range: Brazil (Fabio and Rolas, 1974); Paraguay (Lent et al., 1946); Peru (Bursey et al., 2001). Specimen deposited: USNPC 93265 (1 vial). Remarks: Physalopteroides venancioi is the only species of the genus Physalopteroides known from the South American continent (Vrcibradic et al., 2000). The life cycle of P. venancioi has not been studied, but the physalopterines generally require an insect intermediate host (Anderson, 2000). Thecadactylus rapicauda represents a new host record for P. venancioi.

Spauligodon oxkutzcabiensis (Chitwood, 1938) Skrjabin, Schikhobalova and Lagodovskaja, 1960 (Syn. Pharyngodon oxkutzcabiensis Chitwood, 1938.) Prevalence, mean intensity, and range: Two of 10 hosts infected (20%, 13.5 6 16.2, 2–25). Temporal distribution: November 1994, 1 host with 25; November 1995, 1 host with 2. Site of infection: Large intestine. Additional hosts from Peru: None. Type host and type locality: Thecadactylus rapicauda, Yucatan, Mexico (Chitwood, 1938). Other reported hosts: Green spiny lizard, Sceloporus malachiticus (Goldberg and Bursey, 1992). Geographic range: Costa Rica (Goldberg and Bursey, 1992); Mexico (Chitwood, 1938); Peru. Specimen deposited: USNPC 93266 (1 vial). Remarks: Five species of Spauligodon occur in South America, Spauligodon garciaprietoi Jime´nez-Ruiz, Leo´n-Re`gagnon and Campbell, 2003, Spauligodon

BURSEY ET AL.—HELMINTHS OF PERUVIAN LIZARDS

maytacapaci (Vicente and Iba´n˜ez, 1968), Spauligodon loboi Ramallo, Bursey and Goldberg, 2002, S. oxkutzcabiensis, and Spauligodon viracochai (Freitas, Vicente and Iba´n˜ez, 1968). In S. garciaprietoi the tail of the female tapers to a point; in S. maytacapaci the tail of the female is spike-like. The other South American species of Spauligodon have filiform tails. In S. loboi the tail of the female is smooth; in S. viracochai the tail of the female has 2 spines; and, in S. oxkutzcabiensis the tail of the female has 13–15 spines. A key to the species of Spauligodon in the Western Hemisphere was provided by Jime´nez-Ruiz et al. (2003). The life cycle of S. oxkutzcabiensis has not been studied, but oxyurids are strictly monoxenous (Anderson, 2000). Peru is a new locality record for S. oxkutzcabiensis.

Gymnophthalmidae Cercosaura ocellata Wagler, 1830 Five specimens were collected between January 1986 and June 1989 from leaf litter in Reserva Cuzco Amazo´nico; SVL ¼ 46 mm 6 14 mm SD, range 22– 54 mm; 1 juvenile, 4 females. Physalopteroides venancioi (Lent, Freitas and Proenc¸a, 1946) Sobolev, 1949 Prevalence and intensity: One of 5 hosts infected (20%, 15). Temporal distribution: July 1993, 1 host with 15. Site of infection: Stomach. Specimen deposited: USNPC 93241 (1 vial). Remarks: Cercosaura ocellata represents a new host record for P. venancioi. General information and remarks are reported under T. rapicauda. Prionodactylus eigenmanni Griffin, 1917 Twelve specimens were collected between January 1986 and December 1991 from leaf litter in Reserva Cuzco Amazo´nico; SVL ¼ 30 mm 6 10 mm SD, range ¼ 18–49 mm; 8 juveniles, 2 females, 2 males. Oswaldocruzia vitti Bursey and Goldberg, 2004 Prevalence and intensity: One of 12 hosts infected (8%, 1).

53

Temporal distribution: February 1990, 1 host with 1. Site of infection: Small intestine. Additional hosts from Peru: None. Type host and type locality: Prionodactylus eigenmanni, Amazonas Province, Brazil (Bursey and Goldberg, 2004). Other reported hosts: O’Shaughnessy’s prionodactylus, Prionodactylus oshaughnessyi (Bursey and Goldberg, 2004). Geographic range: Brazil, Ecuador (Bursey and Goldberg, 2004); Peru. Specimen deposited: USNPC 93259 (1 vial). Remarks: Twenty-one species of Oswaldocruzia occur in South America (Ben Slimane et al., 1996). Of these, 4 are parasites of lizards: Oswaldocruzia bainae Ben Slimane and Durette-Desset, 1996, Oswaldocruzia brasiliensis Lent and Freitas, 1935, Oswaldocruzia peruensis Ben Slimane, Verhaagh and Durette-Desset, 1995, and O. vitti. These species are initially distinguished by caudal bursa type: type I in O. brasiliensis and O. vitti, type II in O. peruensis, type III in O. bainae (see Ben Slimane et al. [1996]); and then by cervical alae: absent in O. brasiliensis but present in O. vitti. The life cycles of species of Oswaldocruzia parasitic in lizards have not been studied, but the life cycle of Oswaldocruzia pipiens Walton, 1929, a parasite of toads, was studied by Baker (1978). He reported a 2-phase life cycle comprising a free-living preinfection phase and a parasitic phase. Infection of adult toads is accomplished by penetration of skin by third-stage larvae (Anderson, 2000). Peru is a new locality record for O. vitti.

Polychrotidae Anolis fuscoauratus D’Orbigny, 1837 Twenty-five specimens were collected between November 1983 and August 1989 from ground or bushes (,1.5 m in height) in Reserva Cuzco Amazo´nico, Peru: SVL ¼ 41 mm 6 3 mm SD, range 32–44 mm; 1 juvenile, 16 females, 8 males. Strongyluris oscari Travassos, 1923 (Syn. Strongyluris sai Travassos, 1926; Strongyluris freitasi Alho, 1969; Strongyluris travassosi Alho, 1969.)

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Prevalence and intensity: One of 25 hosts infected (4%, 1). Temporal distribution: October 1987, 1 host with 1. Site of infection: Large intestine. Additional hosts from Peru: Anolis punctatus, P. plica, P. umbra, S. roseiventris. Type host and type locality: Tropidurus sp., Mato Grosso, Brazil (Travassos, 1923). Other reported hosts: Ameiva ameiva (Travassos, 1926; Alho, 1969; Ribas, Rocha, et al., 1998); M. agilis (Ribas, Teixeira-Filho, et al., 1998); T. melanopleurus (Roca, 1997); T. torquatus (Pereira, 1935; Alho, 1969, 1970; Kohn et al., 1973; Cristofaro et al., 1976; Vicente, 1981; Ribas, Teixeira-Filho, et al., 1998; Vrcibradic, Cuhna-Barros, et al., 2000); T. spinulosus (Vicente, 1981). Geographic range: Brazil (Travassos, 1926; Pereira, 1935; Alho, 1969, 1970; Kohn et al., 1973; Cristofaro et al., 1976; Vicente, 1981; Ribas, Teixeira-Filho, 1998; Vrcibradic et al., 2000); Bolivia (Roca, 1997); Peru. Specimen deposited: USNPC 93233 (1 vial). Remarks: Strongyluris oscari is the only species of the genus Strongyluris known to occur in South America. The life cycle of S. oscari has not been studied, but heterakoids are monoxenous (Anderson, 2000). Anolis fuscoauratus represents a new host record and Peru a new locality for S. oscari. Anolis punctatus Daudin, 1802 Fourteen specimens were collected between July 1979 and January 1994 from bushes, tree limbs, or tree trunks in Reserva Cuzco Amazo´nico, Peru: SVL ¼ 71 mm 6 12 mm SD, range 50–88 mm; 3 juveniles, 4 females, 7 males. Cyrtosomum longicaudatum Brenes and Bravo Hollis, 1960 (Syn. Atractis longicaudatum [Brenes and Bravo Hollis, 1960] Baker, 1987. Prevalence and intensity: One of 14 hosts infected (7%, 94). Temporal distribution: January 1986, 1 host with 94. Site of infection: Large intestine. Additional hosts from Peru: None.

Type host and type locality: Black iguana, Ctenosaura similis, Costa Rica (Brenes and Bravo Hollis, 1960). Other reported hosts: Baracoa anole, Anolis baracoae (Coy Otero and Barus, 1979); knight anole, Anolis equestris (Barus and Coy Otero, 1968, 1969; Coy Otero, 1970; Coy Otero and Barus, 1973, 1979); Jamaica giant anole, Anolis garmani (Bundy et al., 1987); white-throated anole, Anolis luteogularis (Coy Otero and Barus, 1979); false chameleon, Chamaeleolis chamaeleonides (Coy Otero and Barus, 1979); Cope’s false chameleon, Chamaeleolis porcus (Barus and Coy Otero, 1969; Coy Otero, 1970; Coy Otero and Barus, 1979); Brooks gecko, Hemidactylus brooki (Coy Otero and Barus, 1973, 1979); northern curlytail lizard, Leiocephalus carinatus (Barus and Coy Otero, 1969; Coy Otero, 1970; Coy Otero and Barus, 1979); Cuban curlytail lizard, Leiocephalus cubensis (Coy Otero, 1970); Cabo Corrientes curlytail lizard, Leiocephalus stictigaster (Coy Otero and Barus, 1979). Geographic range: Cuba (Barus and Coy Otero, 1968, 1969; Coy Otero, 1970; Coy Otero and Barus, 1973, 1979); Costa Rica (Brenes and Bravo Hollis, 1960); Honduras (Caballero DeLoya, 1971); Jamaica (Bundy et al., 1987); Mexico (Caballero DeLoya, 1971; Peru). Specimen deposited: USNPC 93234 (1 vial). Remarks: In their revision of the genus Atractis, Bursey and Flanagan (2002) listed no species of Cyrtosomum from South America. Four Western Hemisphere species are known: Cyrtosomum longicaudatum from Costa Rica, Cyrtosomum mega Bowie and Franz, 1974 from the West Indies, Cyrtosomum penneri Gambino, 1957 from the United States, and Cyrtosomum scelopori Gedoelst, 1919 from Mexico. Of these, C. scelopori has equal spicules, the others have unequal spicules. The long spicule of C. mega is over 375 lm in length; the long spicule of C. penneri and C. longicaudatum is less than 150 lm in length. The tail of C. longicaudatum is long, that of C. penneri is short. Species of Cyrtosomum are autoinfective (Anderson, 2000). Anolis punctatus represents a new host record, and Peru is a new locality record for C. longicaudatum. Oswaldocruzia peruvensis Ben Slimane, Verhaagh and Durette-Desset, 1995 Prevalence and intensity: One of 14 hosts infected (7%, 1).

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Temporal distribution: July 1993, 1 host with 1.

55

Specimen deposited: USNPC 93235 (1 vial).

caudal papillae and spicule length can be used to separate these species. Piratuba digiticauda has 4 large terminal caudal papillae, the other species have smaller and more numerous termal caudal papillae. Piratuba lainsoni has subequal spicules; P. scaffi and P. shawi have spicules of distinctly differing lengths. Piratuba scaffi has 10 or fewer terminal caudal papillae; P. shawi has 20 or more terminal caudal papillae. The filaroids use arthropods as intermediate hosts, most of which release third-stage juveniles on the skin of the definitive host when they feed (Anderson, 2000). Peru is a new locality record for P. lainsoni.

Remarks: Anolis punctatus represents a new host record for O. peruvensis. See remarks on O. vitti under P. eigenmanni.

Rhabdias anolis Bursey, Goldberg and Telford, 2003

Site of infection: Large intestine. Additional hosts from Peru: Stenocercus roseiventris (Ben Slimane et al., 1995). Type host: Stenocercus roseiventris, Huanuco, Peru (Ben Slimane et al., 1995). Other reported hosts: None. Geographic range: Peru (Ben Slimane et al., 1995).

Physaloptera retusa Rudolphi, 1819 Prevalence and intensity: One of 14 hosts infected (7%, 1).

Prevalence, mean intensity, and range: Two of 14 hosts infected (14%, 3.5 6 2.1, 2–5). Temporal distribution: November 1990, 1 host with 2; February 1994, 1 host with 5. Site of infection: Lungs.

Temporal distribution: December 1984, 1 host with 1.

Additional hosts from Peru: None.

Site of infection: Stomach.

Type host and type locality: Anolis frenatus, Panama Province, Panama (Bursey et al., 2003).

Specimen deposited: USNPC 93236 (1 vial). Remarks: Anolis punctatus represents a new host record for P. retusa. General information and remarks are reported under T. rapicauda. Piratuba lainsoni Bain, 1974 Prevalence and intensity: One of 14 hosts infected (7%, 1). Temporal distribution: July 1993, 1 host with 1. Site of infection: Body cavity. Additional hosts from Peru: None. Type host and type locality: Many-colored bush anole, Polychrus marmoratus, Belem, Brazil (Bain, 1974). Other reported hosts: Anolis punctatus (Bain, 1974). Geographic range: Brazil (Bain, 1974); Peru. Specimen deposited: USNPC 93237 (1 vial). Remarks: Four species of Piratuba occur in South America, Piratuba digiticauda Lent and Freitas, 1941, P. lainsoni, Piratuba scaffi Bain, 1974, and Piratuba shawi Bain, 1974. The species are best separated on the basis of microfilaria (Bain, 1974), but the pattern of

Other reported hosts: None. Geographic range: Panama (Bursey et al., 2003); Peru. Specimen deposited: USNPC 93238 (1 vial). Remarks: Of the 46 nominal species comprising Rhabdias, only 3 are known from lizards, Rhabdias chamaeleonis (Skrjabin, 1916) and Rhabias gemellipara Chabaud, Brygoo and Petter, 1961 from the Ethiopian biogeographical realm and R. anolis from the Neotropical realm (see Bursey et al. [2003]). Rhabdias of undetermined species have been reported from Caribbean lizards including the crested anole Anolis cristatellus, Evermann’s anole Anolis evermanni, Gundlach’s anole Anolis gundlachi, stripefoot anole Anolis lineatopus, and Watt’s anole Anolis wattsi (Torres Ortiz, 1980; Bundy et al., 1987; Dobson et al., 1992). Species of Rhabdias infecting reptiles presumably infect the host orally (Anderson, 2000). Anolis punctatus represents a new host record, and Peru is a new locality record for R. anolis. Skrjabinelazia intermedia (Freitas, 1940) Chabaud, 1973 (Syn. Salobrella intermedia Freitas, 1940.)

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COMPARATIVE PARASITOLOGY, 72(1), JANUARY 2005

Prevalence and intensity: One of 14 hosts infected (7%, 1). Temporal distribution: November 1990, 1 host with 1. Site of infection: Stomach. Additional hosts from Peru: None. Type host and type locality: Tropidurus spinulosus, Mato Grosso, Brazil (Freitas, 1940). Other reported hosts: Tropidurus torquatus (Vicente, 1981). Geographic range: Brazil (Freitas, 1940; Vicente, 1981).

Reserva Cuzco Amazo´nico, Peru: SVL ¼ 78 mm 6 17 mm SD, range 53–97; 1 juvenile, 5 females, 5 males.

Parapharyngodon scleratus (Travassos, 1923) Freitas, 1957 (Syn. Thelandros scleratus Travassos, 1923; Pseudothelandros sceleratus [Travassos, 1923] Read, Amrein and Walton, 1952.) Prevalence and intensity: One of 11 hosts infected (9%, 1). Temporal distribution: November 1992, 1 host with 1.

Specimen deposited: USNPC 93239 (1 vial).

Site of infection: Large intestine.

Remarks: Two species of Skrjabinelazia occur in South America, Skrjabinelazia galliardi Chabaud, 1973 and S. intermedia. These species can be diagnosed using egg morphology: the surface of the eggs of S. galliardi are covered with numerous bosses; the surface of the eggs of S. intermedia are smooth. Chabaud et al. (1988) reported that females produced 2 types of eggs: a thin-shelled egg containing a thirdstage larva, which was probably autoinfective, and thick-shelled eggs with third-stage larva that pass from the host and were infective to crickets. Anolis punctatus represents a new host record, and Peru is a new location record for S. intermedia.

Additional hosts from Peru: Ameiva ameiva, K. pelviceps.

Strongyluris oscari Travassos, 1923 Prevalence, mean intensity, and range: Five of 14 hosts infected (36%, 3.8 6 3.1, 1–9). Temporal distribution: November 1990, 1 host with 1; November 1992, 2 hosts with 2, 9, respectively; July 1993, 1 host with 4; January 1994, 1 host with 3. Site of infection: Large intestine. Specimen deposited: USNPC 93240 (1 vial). Remarks: Anolis punctatus represents a new host record for S. oscari. General information and remarks are reported under A. fuscoauratus.

Scincidae Mabuya bistriata (Spix, 1825) Eleven specimens were collected between January 1986 and August 1993 from the ground or logs in

Type host and type locality: Tropidurus torquatus, Manguinhos, Brazil (Travassos, 1923). Other reported hosts: Ameiva ameiva (Rodrigues and Pinto, 1967; Ribas, Rocha, et al., 1998); C. ocellifer (Ribas et al., 1995); Hemidactylus mabouia (Alho and Rodrigues, 1963; Rodrigues, 1970); M. agilis (Van Sluys et al., 1997; Ribas, Teixeira-Filho, et al., 1998; Vrcibradic et al., 2001, 2002); Mabuya frenata (Vrcibradic et al., 1999); M. macrorhyncha (Ribas, Teixeira-Filho, et al., 1998; Vrcibradic et al., 2001, 2002; Tropidurus albemarlensis (Barus, 1973); Tropidurus itambere (Van Sluys et al., 1994); T. melanopleurus (Roca, 1997); T. torquatus (Travassos, 1923; Pereira, 1935, 1937; Vicente, 1981; Ribas, Teixeira-Filho, et al., 1998); T. spinulosus (Vicente, 1981); Tropidurus sp. (reported as Tapinurus scutipunctatus, Pereira, 1935, 1937). Geographic range: Brazil (Travassos, 1923; Pereira, 1935, 1937; Alho and Rodrigues, 1963; Rodrigues and Pinto, 1967; Vicente, 1981; Van Sluys et al., 1997; Ribas, Roca, et al., 1998; Vrcibradic et al., 2001, 2002); Bolivia (Roca, 1997); Galapagos Islands (Barus, 1973); Peru. Specimen deposited: USNPC 93249 (1 vial). Remarks: Six species of Parapharyngodon occur in South America, Parapharyngodon alvarengai Freitas, 1957, Parapharyngodon largitor Alho and Rodrigues, 1963, Parapharyngodon riojensis Ramallo, Bursey, and Goldberg, 2002, P. scleratus, Parapharyngodon senisfaciecaudus Freitas, 1957, Parapharyngodon verrucosus Freitas and Dobbin, 1959. Ramallo et al. (2002) provide a key to the

BURSEY ET AL.—HELMINTHS OF PERUVIAN LIZARDS

neotropical species of Parapharyngodon. Mabuya bistriata represents a new host record, and Peru is a new locality record for P. scleratus. Physaloptera retusa Rudolphi, 1819 Prevalence, mean intensity, and range: Four of 11 hosts infected (36%, 5.5 6 4.2, 1–10). Temporal distribution: December 1984, 2 hosts with 1, 8, respectively; November 1992, 1 host with 3; June 1993, 1 host with 10.

57

Remarks: Piratuboides zeae is the only species of Piratuboides occurring in South America (Baker, 1987). Peru is a new locality record for P. zeae. Teiidae Ameiva ameiva (Linnaeus, 1758) Twenty-five specimens were collected between December 1984 and March 1994 from the ground or logs in Reserva Cuzco Amazo´nico, Peru: SVL ¼ 100 mm 6 41 mm SD, range ¼ 48–158 mm; 9 juveniles, 6 females, 10 males.

Site of infection: Stomach. Specimen deposited: USNPC 93250 (1 vial).

Oochoristica ameivae (Beddard, 1914) Baer, 1924

Remarks: General information and remarks are reported under T. rapicauda.

(Syn. Linstowia ameivae Beddard, 1914; Oochoristica brasiliensis Fuhrmann, 1927; Oochoristica fuhrmanni Hughes, 1949.)

Physalopteroides venancioi (Lent, Freitas and Proenc¸a, 1946) Sobolev, 1949

Prevalence, mean intensity, and range: Three of 25 hosts infected (12%, 1.7 6 1.2, 1–3).

Prevalence, mean intensity, and range: Two of 11 hosts infected (18%, 5.0 6 5.7, 1–9). Temporal distribution: February 1984, 1 host with 9; July 1993, 1 host with 1. Site of infection: Stomach. Specimen deposited: USNPC 93251 (1 vial). Remarks: Mabuya bistriata represents a new host record for P. venancioi. General information and remarks are reported under T. rapicauda. Piratuboides zeae (Bain, 1974) Bain and Sulahian, 1974 (Syn. Piratuba zeae Bain, 1974.) Prevalence and intensity: One of 11 hosts infected (9%, 2). Temporal distribution: June 1993, 1 host with 2. Site of infection: Body cavity. Additional hosts from Peru: None. Type host and type locality: Mabuya bistriata (recorded as M. mabouya), Belem, Brazil (Bain, 1974). Other reported hosts: None. Geographic range: Brazil (Bain, 1974); Peru. Specimen deposited: USNPC 93252 (1 vial).

Temporal distribution: December 1984, 1 host with 1; January 1990, 2 hosts with 1, 3, respectively. Site of infection: Small intestine. Additional hosts from Peru: None. Type host and type locality: Ameiva ameiva, Brazil (Beddard, 1914). Other reported hosts: Mabuya agilis (Vrcibradic et al., 2002); M. macrorhyncha (Vrcibradic et al., 2001). Geographic range: Brazil (Beddard, 1914; Vrcibradic et al., 2001, 2002); Peru. Specimen deposited: USNPC 93228 (1 slide, 1 vial). Remarks: Six species of Oochoristica occur in South America, O. ameivae (Beddard, 1914), Oochoristica bresslaui Fuhrmann, 1927, Oochoristica iguanae Bursey and Goldberg, 1996, Oochoristica insulaemargaritae Lopez-Neyra and Diaz-Ungria, 1957, Oochoristia travassoi Rego and Iba´n˜ez, 1965, Oochoristica vanzolinii Rego and Oliveira-Rodrigues (see Bursey and Goldberg [1996]). These species can be grouped on the basis of sucker shape: O. ameivae, O. bresslaui, and O. vanzolinii have circular suckers but O. iguanae, O. insulaemargaritae, and O. travassosi have oval suckers. Among those with circular suckers, O. vanzolinii has less than 30 testes, O. ameivae and O. bresslaui more than 30; O. ameivae has a wide scolex (450–650 lm), O. bresslaui

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COMPARATIVE PARASITOLOGY, 72(1), JANUARY 2005

a narrow scolex (140 lm). Among those with oval suckers, O. insulaemargaritae has more than 50 testes, O. iguanae and O. travassosi have less than 50 testes; O. travassosi has a wide scolex (600 lm), O. iguanae a narrow scolex (175–240 lm). Peru is a new locality record for O. ameivae.

Temporal distribution: December 1984, 5 hosts with 1, 1, 2, 2, 4, respectively; January 1990, 1 host with 3; January 1994, 1 host with 2; February 1994, 1 host with 3. Site of infection: Stomach. Specimen deposited: USNPC 93231 (1 vial).

Amphibiocapillaria freitasilenti (Araujo and Gandra, 1941) Moravec, 1986 (Syn. Capillaria freitasilenti Araujo and Gandra, 1941.) Prevalence and intensity: One of 25 hosts infected (4%, 1). Temporal distribution: February 1994, 1 host with 1. Site of infection: Small intestine. Additional hosts from Peru: None. Type host and type locality: Tropidurus torquatus, Goias, Brazil (Araujo and Gandra, 1941). Other reported hosts: None. Geographic range: Brazil (Araujo and Gandra, 1941); Peru. Specimen deposited: USNPC 93229 (1 vial). Remarks: Amphibiocapillaria freitasilenti is the only capillariid currently known to parasitize lizards (Moravec, 1986). The life cycle of A. freitasilenti has not been described; but species of the Capillariinae are known to produce autoinfective eggs, directly infect the definitive host and indirectly infect the definitive host (Anderson, 2000). Ameiva ameiva represents a new host record for A. freitasilenti. Peru is a new location record. Parapharyngodon scleratus (Travassos, 1923) Freitas, 1957 Prevalence and intensity: One of 25 hosts infected (4%, 1). Temporal distribution: February 1990, 1 host with 1. Site of infection: Large intestine. Specimen deposited: USNPC 93230 (1 vial). Remarks: General information and remarks are reported under M. bistriata. Physaloptera retusa Rudolphi, 1819 Prevalence, mean intensity, and range: Eight of 25 hosts infected (32%, 2.3 6 1.0, 1–4).

Remarks: General information and remarks are reported under T. rapicauda.

Spinicauda spinicauda (Olfers in Rudolphi, 1819) Travassos, 1923 (Syn. Heterakis turgida Schneider, 1866, Spinicauda amarali Pereira, 1935, Heterakis campanula Linstow, 1899.) Prevalence, mean intensity, and range: Fifteen of 25 hosts infected (60%, 13.7 6 17.2, 1–66). Temporal distribution: December 1984, 7 hosts with 1, 3, 3, 6, 11, 12, 27, respectively; January 1990, 4 hosts with 5, 5, 20, 66; February 1990, 2 hosts with 1, 15; January 1994, 1 host with 1; February 1994, 1 host with 29. Site of infection: Large intestine. Additional hosts from Peru: None. Type host and type locality: Tupinambis teguixin, Brazil (Rudolphi, 1819). Other reported hosts: Ameiva ameiva (Travassos, 1923; Pereira, 1935; Diaz-Ungria, 1964; Everard, 1975; Rodrigues and Feijo´, 1976); Anolis armouri (Goldberg, Bursey, and Cheam, 1996); Anolis leachi (Dobson et al., 1992); Anolis oculatus (Goldberg, Bursey, and Cheam, 1996). Geographic range: Antigua (Dobson et al., 1992); Brazil (Travassos, 1920, 1923; Pereira, 1935; Rodrigues and Feijo´, 1976); Dominica (Goldberg and Bursey, 1996); Haiti (Goldberg and Bursey, 1996); Trinidad (Everard, 1975); Venezuela (Diaz-Ungria, 1964); Peru. Specimen deposited: USNPC 93232 (1 vial). Remarks: Spinicauda spinicauda is the only species of Spinicauda to occur in South America (Baker, 1987); it is monoxenous, eggs containing the infective stage are ingested by the host (Anderson, 2000). Peru is a new locality record for S. spinicauda.

BURSEY ET AL.—HELMINTHS OF PERUVIAN LIZARDS

Kentropyx altamazonicus Cope, 1876 Eleven specimens were collected between December 1984 and February 1986 from the ground in Reserva Cuzco Amazo´nico, Peru: SVL ¼ 66 mm 6 14 mm SD, range ¼ 36–87 mm; 1 juvenile, 6 females, 4 males. Physaloptera retusa Rudolphi, 1819 Prevalence, mean intensity, and range: Three of 11 hosts infected (27%, 2.0 6 1.7, 1–4). Temporal distribution: December 1984, 2 hosts with 1, 4, respectively; February 1986, 1 host with 1. Site of infection: Stomach. Specimen deposited: USNPC 93242 (1 vial). Remarks: Kentropyx altamazonicus represents a new host record for P. retusa. General information and remarks are reported under T. rapicauda. Physalopteroides venancioi (Lent, Freitas and Proenc¸a, 1946) Sobolev, 1949 Prevalence and intensity: One of 11 hosts infected (9%, 2). Temporal distribution: December 1984, 1 host with 2. Site of infection: Stomach. Specimen deposited: USNPC 93243 (1 vial). Remarks: Kentropyx altamazonicus represents a new host record for P. venancioi. General information and remarks are reported under T. rapicauda. Kentropyx pelviceps Cope, 1868 Fifteen specimens collected January 1985 and July 1993 from the ground or logs in Reserva Cuzco Amazo´nico, Peru: SVL ¼ 93 mm 6 15 mm SD, range 60–109 mm; 1 juvenile, 5 females, 9 males. Ophiotaenia flava Rudin, 1917 Prevalence, mean intensity, and range: Four of 15 hosts infected (27%, 3.0 6 2.3, 1–5). Temporal distribution: January 1990, 1 host with 5; February 1990, 2 hosts with 1, 5, respectively; June 1993, 1 host with 1.

59

Site of infection: Small intestine. Additional hosts from Peru: None. Type host and type locality: Coluber sp., Brazil (Rudin, 1917). Other reported hosts: None. Geographic range: Brazil (Rudin, 1917); Peru. Specimen deposited: USNPC 93244 (2 slides, 1 vial). Remarks: Seven species of Ophiotaenia have been reported from South American reptiles, namely, Ophiotaenia calmetti (Barrois, 1898), Ophiotaenia elongata Fuhrmann, 1927, O. flava, Ophiotaenia hyalina Rudin, 1917, Ophiotaenia jarara Fuhrmann, 1927, Ophiotaenia macrobothra Rudin, 1817, Ophiotaenia racemosa (Rudolphi, 1819), (see Freze [1965]). We have assigned our specimens to Ophiotaenia flava based on the definitive characters used in the key developed by Freze (1965): the uterine stem gives off only 1 layer of diverticula, strobulation begins immediately behind the scolex, and the cirrus pouch reaches the middle of the proglottid. No additional host has been reported for O. flava since the original description. Kentropyx pelviceps represents a new host record and Peru a new locality record for O. flava. Dujardinascaris sp. (third-stage larvae) Prevalence and intensity: One of 15 hosts infected (7%, 7). Temporal distribution: January 1990, 1 host with 7. Site of infection: Stomach. Additional hosts from Peru: Tupinambis teguixin (this report). Specimen deposited: USNPC 93248 (1 vial). Remarks: Adults of species of Dujardinascaris are parasites of fishes and aquatic reptiles (Sprent, 1977). Two species, Dujardinascaris longispicula Travassos, 1933 and Dujardinascaris paulista Travasssos, 1933, have been reported from Brazil and Venezuela (Travassos, 1933; Diaz-Ungria and Gallardo, 1968; Sprent, 1977), both from the narrow-snouted spectacled caiman, Caiman crocodilus. Baylis (1928) found larvae of Dujardinascaris in the mesenteries of an African fish. Brandt (1936) observed third-stage larvae in the body cavity of the bullfrog, Rana catesbeiana, and in the Florida leopard frog, Rana

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COMPARATIVE PARASITOLOGY, 72(1), JANUARY 2005

sphenocephala, from North Carolina, U.S.A. Arya (1980) found fourth-stage larvae of Dujardinascaris in the mesenteries of an Indian fish and reported infections in a 5-mo-old salt water crocodile, Crocodylus porosus. Arya (1980) also collected second-stage larvae from the body cavity of small fishes and crustaceans, concluding that they serve as intermediate hosts. This is the first report of larvae of Dujardinascaris sp. in lizards and the first report of the genus Dujardinascaris from Peru. Parapharyngodon scleratus (Travassos, 1923) Freitas, 1957 Prevalence and intensity: One of 15 hosts infected (7%, 1).

Site of infection: Stomach. Specimen deposited: USNPC 93247 (1 vial). Remarks: Kentropyx pelviceps represents a new host record for P. venancioi. General information and remarks are reported under T. rapicauda. Tupinambis teguixin (Linnaeus, 1758) One female specimen was collected February 1990 from the ground in Reserva Cuzco Amazo´nico, Peru: SVL ¼ 202 mm. Dujardinascaris sp. third-stage larvae)

Temporal distribution: February 1990, 1 host with 1.

Prevalence and intensity: One of 1 hosts infected (100%, 8).

Site of infection: Large intestine.

Temporal distribution: February 1990, 1 host with 8.

Specimen deposited: USNPC 93245 (1 vial).

Site of infection: Stomach.

Remarks: Kentropyx pelviceps represents a new host record for P. scleratus. General information and remarks are reported under M. bistriata.

Specimen deposited: USNPC 93269 (1 vial).

Physaloptera retusa Rudolphi, 1819

Physaloptera retusa Rudolphi, 1819

Prevalence, mean intensity, and range: Nine of 15 hosts infected (60%, 8.2 6 9.9, 1–32).

Prevalence and intensity: One of 1 hosts infected (100%, 72).

Temporal distribution: January 1985, 1 host with 1; June 1989, 1 host with 14; January 1990, 3 hosts with 4, 9, 32, respectively; February 1990, 2 hosts with 1, 2; June 1993, 2 hosts with 2, 9.

Temporal distribution: February 1990, 1 host with 72.

Site of infection: Stomach.

Specimen deposited: USNPC 93267 (1 vial).

Specimen deposited: USNPC 93246 (1 vial).

Remarks: General information and remarks are reported under T. rapicauda.

Remarks: Kentropyx pelviceps represents a new host record for P. retusa. General information and remarks are reported under T. rapicauda. Physalopteroides venancioi (Lent, Freitas and Proenc¸a, 1946) Sobolev, 1949 Prevalence, mean intensity, and range: Seven of 15 hosts infected (47%, 19.0 6 37.7, 1–104). Temporal distribution: January 1985, 1 host with 1; June 1989, 1 host with 2; January 1990, 1 host with 1; February 1990, 3 hosts with 4, 8, 104, respectively; June 1993, 1 host with 13.

Remarks: General information and remarks are reported under K. pelviceps.

Site of infection: Stomach.

Physalopteroides venancioi (Lent, Freitas and Proenc¸a, 1946) Sobolev, 1949 Prevalence and intensity: One of 1 hosts infected (100%, 2). Temporal distribution: February 1990, 1 host with 2. Site of infection: Stomach. Specimen deposited: USNPC 93268 (1 vial). Remarks: Tupinambis teguixin represents a new host record for P. venancioi. General information and remarks are reported under T. rapicauda.

BURSEY ET AL.—HELMINTHS OF PERUVIAN LIZARDS

Tropiduridae Plica plica (Linnaeus, 1758) Nine specimens were collected between December 1984 and March 1994 from tree branches and tree trunks in Reserva Cuzco Amazo´nico, Peru: SVL ¼ 102 mm 6 31 mm SD, range 47–141 mm; 1 juvenile, 8 females.

61

Hastospiculum sp. (third-stage larvae) Prevalence and intensity: One of 14 hosts infected (7%, 2). Temporal distribution: February 1994, 1 host with 2. Site of infection: Coelom, under peritoneum. Additional hosts from Peru: None. Specimen deposited: USNPC 93258 (1 vial).

Physaloptera retusa Rudolphi, 1819 Prevalence, mean intensity, and range: Six of 9 hosts infected (67%, 4.8 6 4.4, 1–12). Temporal distribution: December 1984, 2 hosts with 6, 12, respectively; November 1990, 2 hosts with 1, 7; February 1994, 1 host with 2; January 1994, 1 host with 1. Site of infection: Stomach. Specimen deposited: USNPC 93253 (1 vial). Remarks: Plica plica represents a new host record for P. retusa. General information and remarks are reported under T. rapicauda. Strongyluris oscari Travassos, 1923 Prevalence, mean intensity, and range: Five of 9 hosts infected (56%, 15.6 6 14.3, 2–37). Temporal distribution: December 1984, 1 host with 2; November 1990, 2 hosts with 7, 9 respectively; January 1994, 1 host with 37; February 1994, 1 host with 23. Site of infection: Large intestine. Specimen deposited: USNPC 93254 (1 vial). Remarks: Plica plica represents a new host record for S. oscari. General information and remarks are reported under A. fuscoauratus.

Remarks: Adults of species of Hastospiculum are parasites of lizards and snakes (Baker, 1987). Three species of Hastospiculum have been reported from South American snakes: Hastospiculum digiticaudum Freitas, 1955 from the Brazilian green racer, Phylodryas aestivus, collected in Brazil (Freitas, 1955); Hastospiculum onchocercum Chitwood, 1932, originally described from specimens taken from the intestinal peritoneum of a boa constrictor, Boa constrictor, that died in the National Zoological Park, Washington, D.C. (Chitwood, 1932), and from the same host in Mexico (Caballero, 1947) and Trinidad (Everard, 1975), neotropical rattlesnake, Crotalus durissus, from Brazil (Desportes, 1941, Araujo, 1970), rainbow boa, Epicrates cenchria, from Trinidad (Everard, 1975), P. aestivus from Brazil (Vicente et al., 1993), and tropical rat snake, Spilotes pullatus, also from Brazil (Vicente and Jardim, 1980); and Hastospiculum venezuelense Vado, 1972 from an undetermined species of Dryadophis and bushmaster, Lachesis muta, from Venezuela (Vado, 1972). The life cycle has not been described, but filarids generally require arthropod vectors. It is unknown whether South American lizards can serve as definitive hosts for species of Hastospiculum, but varanid lizards are known to harbor species of Hastospiculum (see Baker [1987]). Peru is a new location record for the genus Hastospiculum. Physaloptera retusa Rudolphi, 1819 Prevalence, mean intensity, and range: Nine of 14 hosts infected (64%, 24.1 6 30.0, 1–94).

Plica umbra (Linnaeus, 1758) Fourteen specimens were collected between December 1985 and March 1994 from tree trunks in Reserva Cuzco Amazo´nico, Peru: SVL ¼ 102 mm 6 31 mm SD, range 47–141 mm; 3 juveniles, 6 females, 5 males.

Temporal distribution: December 1985, 1 host with 6; January 1990, 1 host with 8; February 1990, 1 host with 94; November 1990, 1 host with 6; October 1992, 1 host with 30; December 1993, 1 host with 1; February 1994, 3 hosts with 7, 18, 47, respectively. Site of infection: Stomach.

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COMPARATIVE PARASITOLOGY, 72(1), JANUARY 2005

Specimen deposited: USNPC 93255 (1 vial). Remarks: Plica umbra represents a new host record for P. retusa. General information and remarks are reported under T. rapicauda.

Stenocercus roseiventris Dume´ril and Bibron, 1837 Twelve specimens collected October 1987 and January 1996 from the ground and logs in Reserva Cuzco Amazo´nico, Peru: SVL ¼ 81 mm 6 20 mm SD, range 38–101 mm; 1 juvenile, 5 females, 9 males.

Piratuba digiticauda Lent and Freitas, 1941 Prevalence, mean intensity, and range: Two of 14 hosts infected (14%, 4.5 6 4.9, 1–8).

Oswaldocruzia peruvensis Ben Slimane, Verhaagh and Durette-Desset, 1995

Temporal distribution: December 1993, 1 host with 8; February 1994, 1 host with 1.

Prevalence and intensity: One of 12 hosts infected (8%, 12).

Site of infection: Body cavity.

Temporal distribution: October 1987, 1 host with 12.

Additional hosts from Peru: None.

Site of infection: Small intestine.

Type host and type locality: Unidentified lizard, Brazil (Lent and Freitas, 1941).

Specimen deposited: USNPC 93260 (1 vial).

Other reported hosts: Anolis baracoae (Barus and Coy Otero, 1969), two-marked anole, Anolis biporcatus (Bursey et al., 2003); A. equestris (Barus and Coy Otero, 1969); C. porcus (Barus and Coy Otero, 1969); T. torquatus (Vicente and Jardim, 1980; Vicente, 1981); and T. spinulosus (Vicente and Jardim, 1980; Vicente, 1981). Geographic range: Brazil (Lent and Freitas, 1941; Vicente and Jardim, 1980; Vicente, 1981); Cuba (Barus and Coy Otero, 1969); Panama (Bursey et al., 2003); Peru. Specimen deposited: USNPC 93256 (1 vial). Remarks: Plica plica represents a new host record and Peru is a new locality record for P. digiticauda. See remarks under A. punctatus.

Remarks: General information and remarks are reported under A. punctatus. Oswaldofilaria azevedoi Bain, 1974 Prevalence, mean intensity, and range: Four of 12 hosts infected (33%, 2.0 6 0.8, 1–3) Temporal distribution: October 1987, 1 host with 1; November 1990, 1 host with 2; November 1992, 1 host with 2; July 1993, 1 host with 3. Site of infection: Body cavity. Additional hosts from Peru: None. Type host and type locality: Polychrus marmoratus, Belem, Brazil (Bain, 1974). Other reported hosts: None.

Strongyluris oscari Travassos, 1923

Geographic range: Brazil (Bain, 1974); Peru.

Prevalence, mean intensity, and range: Eight of 14 hosts infected (57%, 5.0 6 4.7, 1–16).

Specimen deposited: USNPC 93261 (1 vial).

Temporal distribution: December 1985, 1 host with 4; February 1990, 1 host with 1; December 1993, 1 host with 5; January 1994, 1 host with 3; February 1994, 4 hosts with 2, 3, 6, 16, respectively. Site of infection: Large intestine. Specimen deposited: USNPC 93257 (1 vial). Remarks: Plica umbra represents a new hosts record for S. oscari. General information and remarks are reported under A. fuscoauratus.

Remarks: Eight species of Oswaldofilaria occur in South America, Oswaldofilaria bacillaris (Molin, 1858) and Oswaldofilaria medemi Marinkelle, 1981, in crocodilians; Oswaldofilaria carinii (Vaz and Pereira, 1935) in snakes; Oswaldofilaria azevedoi, Oswaldofilaria belemensis Bain and Dulahian, 1974, Oswaldofilaria brevicaudata (Rodhain and Vuylsteke, 1937), Oswaldofilaria petersi Bain and Sulahian, 1974, and Oswaldofilaria spinosa Bain and Sulahian, 1974, in lizards (see Baker [1987]). The species in lizards can be separated into 2 groups using spicule length ratios: O. azevedoi and O. brevicaudata

BURSEY ET AL.—HELMINTHS OF PERUVIAN LIZARDS

have a 1:2 or less short to long spicule ratio, the other species have 1:3 or greater ratios. Oswaldofilaria brevicaudata has 4 large terminal caudal papillae; O. azevedoi has 4 tiny terminal caudal papillae. Oswaldofilaria spinosa has an unpaired median ventral tail papilla; O. belemensis has 4 large terminal caudal paipllae; and O. petersi has 4 very small terminal caudal papillae. The life cycles of several species of Oswaldofilaria have been examined (Bain and Chabaud, 1975). Mosquito vectors transfer larvae to definitive hosts. Stenocercus roseiventris represents a new host record, and Peru is a new locality record for O. azevedoi. Physaloptera retusa Rudolphi, 1819 Prevalence, mean intensity, and range: Six of 12 hosts infected (50%, 4.2 6 3.1, 1–8). Temporal distribution: November 1990, 2 hosts with 3, 8, respectively; June 93, 1 host with 2; July 1993, 1 host with 8; February 1994, 1 host with 3; December 1995, 1 host with 1. Site of infection: Stomach. Specimen deposited: USNPC 93262 (1 vial). Remarks: Stenocercus roseiventris represents a new host record for P. retusa. General information and remarks are reported under T. rapicauda. Strongyluris oscari Travassos, 1923 Prevalence and intensity: Eleven of 12 hosts infected (92%, 35.9 6 31.3, 1–98). Temporal distribution: October 1987, 1 host with 98; January 1990, 1 host with 1; November 1990, 2 hosts with 20, 38, respectively; November 1992, 1 host with 38; June 1993, 1 host with 14; July 1993, 1 host with 35; January 1994, 2 hosts with 5, 19; February 1994, 1 host with 39; December 1995, 1 host with 88. Site of infection: Large intestine. Specimen deposited: USNPC 93262 (1 vial). Remarks: Stenocercus roseiventris represents a new host record for S. oscari. General information and remarks are reported under A. fuscoauratus. DISCUSSION In all, 1, 617 helminths were collected from 83 (51%) of the 164 lizards examined. Of these, 15

63

(0.9%) were larval forms not capable of reaching maturity in lizards. There were 20 helminth species represented in the sample, but no individual host harbored more than 3 helminth species. Of the infected lizards, 36 (43%) harbored 1 species of helminth; 35 (42%) harbored 2 species; and 12 (15%) harbored 3 species. There were 1.70 6 0.08 (
x61 SE) helminth species/infected lizard and 19.45 6 2.82 helminth individuals/infected lizard. No host species harbored more than 7 helminth species: 3 lizard species (23%) harbored 1 species of helminth; 2 species (15%) harbored 2; 2 species (15%) harbored 3, 3 species (23%) harbored 4, 2 species (15%) harbored 5; and 1 species (7%) harbored 7. There were 3.23 6 0.51 helminth species/host species. Aho (1990) compiled distributional patterns for lizard helminths in general and reported the mean (6SE) total number of helminth species per host species as 2.06 6 0.13 (range 0–5) with a mean species richness per host individual of 0.63 6 0.06 (range 0–2.5). The values reported here are larger than those of Aho (1990), perhaps reflecting differences in biome or regional (tropical vs. temperate or tropical vs. worldwide) helminth distribution patterns. Of the 20 helminth species found in this study, only 4 had prevalences greater than 2%; P. retusa was most prevalent, occurring in 51 of 164 lizards (31%), followed by S. oscari (30 of 164 [18%]), P. venancioi (15 of 164 [9%]), and S. spinicauda (15 of 164 [9%]). Abundance values for these 4 species were 3.04, 3.25, 1.09, and 1.25, respectively. Bursey et al. (2001) introduced the concept of importance (I), an estimate of the influence of a species within a community calculated as I ¼ relative prevalence þ relative abundance 3 100. Using this measure, the most important helminth in the lizard community under study is P. retusa (I ¼ 68), followed by S. oscari (I ¼ 55), S. spinicauda (I ¼ 23), and P. venancioi (I ¼ 22). Physaloptera retusa and P. venancioi are heteroxenous. Stongyluris oscari and S. spinicauda are monoxenous. Helminth species requiring mosquito vectors, P. digiticauda, P. lainsoni, P. zeae, and O. azevedoi, have importance values of 1.9, 0.8, 0.8, and 3.3, respectively. The data presented here suggest that Peruvian lizards are infected by helminth generalists (helminths not restricted to a single host species). All helminths examined in this study are known to infect at least 2 host species. This observation parallels that of an earlier study on anurans from Reserva Cuzco Amazo´nico (Bursey et al., 2001) that also reported generalist helminth communities. Interestingly, only 1 species, P. venancioi, was found to infect both frogs

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(9 species) and lizards (6 species) in Reserva Cuzco Amazo´nico. However, the importance of P. venancioi to the anuran community was I ¼ 7.5 as compared with I ¼ 22 for the lizard community. The life cycle of P. venancioi has not been studied, but physalopterines require an insect intermediate host (Anderson, 2000); thus, some common element in the diet of these species may be inferred. ACKNOWLEDGMENTS Collections by W. E. Duellman and other herpetologists who worked at Reserva Cuzco Amazo´nico, Peru, made this study possible. J.R.P. thanks William Duellman and the Museum of Natural History at the University of Kansas for financial support when examining the diet of these tropical lizards. Fieldwork was supported by grants from the National Geographic Society (3196-85, 3405-86, and 4016-89; W. E. Duellman, principal investigator). Permits were issued by Ing. G. B. Mejı´a Mun˜oz and Blgo. Jose´ Purisaca of the Direccio´n General Forestal y de Fauna, Lima, Peru.

LITERATURE CITED Aho, J. M. 1990. Helminth communities of amphibians and reptiles: comparative approaches to understanding patterns and processes. Pages 157–195 in G. W. Esch, A. O. Bush, and J. M. Aho, eds. Parasite Communities: Patterns and Processes. Chapman and Hall, London. Alho, C. J. R. 1969. Oxyurata de largartos do Planalto Central. Soˆbre o geˆnero Strongyluris Mueller, 1894 com descric¸a˜o de duas espe´cies novas. Revista Brasileira de Biologia 29:65–74. Alho, C. J. R. 1970. Frequency of infestation by Strongyluris freitasi Alho, 1969 in Tropidurus troquatus (Wied). Revista Brasileira de Biologia 30:539–542. Alho, C. J. R., and H. O. Rodrigues. 1963. Soˆbre noˆvo hospedeiro de Parapharyngodon scleratus (Travassos, 1923) Freitas, 1957, com redescric¸a˜o da espe´cie (Nematoda, Oxyuroidea). Atas Sociedade de Biologia do Rio de Janeiro 7:4–6. Anderson, R. M. 2000. Nematode Parasites of Vertebrates: Their Development and Transmission, 2nd ed. CABI Publishing, Wallingford, Oxon, U.K. 650 pp. Araujo, P. 1970. Considerac¸~oes soˆbre Hastospiculum onchocercum major Desportes, 1941, parasita de serpente Crotalus durissus terrificus. Revista de farmacia e gioquimica da Universidade de Sa˜o Paulo 8:121–124. Araujo, T. L., and V. Gandra. 1941. Soˆbre uma nova espe´cie do geˆnero Capillaria e observac¸~oes helmintolo´gicas. Revista da Faculdade de Medicina Veterinaria 2:29–32. Arya, S. N. 1980. On the larval forms of a nematode, Dujardinascaris cybii Arya and Johnson, 1978. Revista Brasileira de Biologia 40:751–754. Bain, O. 1974. Description de nouvelles filaires Oswaldofilariinae de le´zards sud-ame´ricains; hypothe`se sur

l’e´volution des filaires de Reptiles. Me´moires du Muse´um National d’Histoire Naturelle Series 3 138: 169–200. Bain, O., and A.-G. Chabaud. 1975. De´veloppement chez des moustiques de trois filaires de le´zards sudame´ricains du genre Oswaldofilaria. Annales de Parasitologie (Paris) 50:209–221. Baker, M. R. 1978. Development and transmission of Oswaldocruzia pipiens Walton, 1929 (Nematoda: Trichostrongylidae) in amphibians. Canadian Journal of Zoology 56:1026–1031. Baker, M. R. 1987. Synopsis of the Nematoda parasitic in amphibians and reptiles. Memorial University of Newfoundland, Occasional Papers in Biology 11: 1–325. Barus, V. 1973. Some remarks on the neotropical species of the genera Parapharyngodon and Batracholandros (Oxyuridae). Folia Parasitologica (Praha) 20:131–139. Barus, V., and A. Coy Otero. 1968. Freitasia teixeirai gen. n. et sp. n. and other nematodes parasitizing Anolis equestris (Squamata: Iguanidae). Folia Parasitologica (Praha) 15:41–45. Barus, V., and A. Coy Otero. 1969. Systematic survey of nematodes parasitizing lizards (Sauria) in Cuba. Helminthologia 10:329–346. Barus, V., and A. Coy Otero. 1974. Nematodes of the genera Spauligodon, Skrjabinodon, and Pharyngodon (Oxyuridae) parasitizing Cuban lizards. Vestnik Ceskoslovenske Spolecunosti Zoologicke 38:1–12. Baylis, H. A. 1928. Some parasitic worms mainly from fishes from Lake Tanganyika. Annals and Magazine of Natural History Series 10 4:552–562. Beddard, F. E. 1914. Contributions to the anatomy and systematic arrangement of the Cestoidea. XIII. On two new species belonging to the genera Oochoristica and Linstowia, with remarks upon those genera. Proceedings of the Zoological Society of London 1914: 263–282. Ben Slimane, B., A.-G. Chabaud, and M.-C. DuretteDesset. 1996. Les ne´matodes Trichostrongylina parasites d’amphibiens et de reptiles: proble`mes taxonomiques, phyle´tiques et bioge´ographiques. Systematic Parasitology 35:179–206. Ben Slimane, B., M. Verhaagh, and M.-C. DuretteDesset. 1995. Oswaldocruzia peruensis n. sp. (Nematoda, Trichostrongylina, Molineoidea), parasite d’un iguanidae du Pe´rou. Me´moires du Muse´um National d’Histoire Naturelle de Paris 17:77–82. Brandt, B. B. 1936. Parasites of certain North Carolina Salientia. Ecological Monographs 6:491–532. Brenes, R.-R., and M. Bravo Hollis. 1960. Helmintos de la Republica de Costa Rica IX. Nematoda 3. Algunos nema´todos de reptiles, con descripcio´n de dos nuevas especies: Atractis caballeroi n. sp. y Cyrtosomum longicaudatum n. sp. Pages 451–464 in E. Caballero, ed. Libro Homenaje al Docter Eduardo Caballero y Caballero; Jubileo 1930–1950. Escuela Nacional de Ciencias Biolo´gicas, Mexico. Bundy, D. A. P., P. Vogel, and E. A. Harris. 1987. Helminth parasites of Jamaican anoles (Reptilia: Iguanidae): a comparison of the helminth fauna of 6 Anolis species. Journal of Helminthology 61:77–83. Bursey, C. R., and J. P. Flanagan. 2002. Atractis marquezi n. sp. (Nematoda: Atractidae) and a revision of Atractis

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Coy Otero, A. 1970. Contribucion al conocimiento de la helmintofauna de los saurios cubanos. Ciencias 4:1–58. Coy Otero, A., and V. Barus. 1973. Notes on nematodes of the genus Cyrtosomum (Atractidae) parasitic in Cuban lizards (Sauria). Folia Parasitologica (Praha) 20: 297–305. Coy Otero, A., and V. Barus. 1979. Nematodes parasitizing Cuban reptiles. Acta Scientiarum Naturalium Academiae Scientiarum Bohemoslovacae Brno 13:1–43. Cristofaro, R., J. F. Guimara˜es, and H. de Oliveira Rodrigues. 1976. Alguns nemato´deos de Tropidurus torquatus (Wied) e Ameiva ameiva (L.)—Fauna Helmintolo´gica de Salvador, Bahia. Atas Sociedade de Biologia de Rio de Janeiro 18:65–70. Desportes, C. 1941. Sur un Hastospiculum parasite du crotale. Annales de Parasitologie Humaine et Compare´e 18:198–208. Diaz-Ungria, C. 1964. Notas sobre nematodes de reptiles en Venezuela. Revista Veterinaria 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. Dobson, A. P., S. V. Pacala, J. D. Roughgarden, E. R. Carper, and E. A. Harris. 1992. The parasites of Anolis lizards in the northern Lesser Antilles. I. Patterns of distribution and abundance. Oecologia 91:110–117. Duellman, W. E., and J. E. Koechlin. 1991. The Reserva Cuzco Amazonico, Peru: biological investigations, conservation, and ecotourism. Occasional Papers of the Museum of Natural History, University of Kansas 142:1–38. Duellman, W. E., and A. W. Salas. 1991. Annotated checklist of the amphibians and reptiles of Cuzco Amazonico, Peru. Occasional Papers of the Museum of Natural History, University of Kansas 143:1–13. Everard, C. O. R. 1975. Endoparasites of some Amphibia, reptiles and small mammals from Trinidad. Journal of the Trinidad and Tobago Field Naturalists’ Club 1975:72–79. de Fabio, S. P., and F. J. T. Rolas. 1974. Contribuic¸a˜o ao conhecimento de Thubuneae dactyluris Karve, 1938 (Nematoda, Spiruroidea). Memo´rias do Instituto Oswaldo Cruz 72:283–290. Freitas, J. F. T. 1940. Sobre um interessante nematodeo parasito de reptil (Spiruroidea). Memo´rias do Instituto Oswaldo Cruz 35:603–605. Freitas, J. F. T. 1955. Nota soˆbre a fauna helmintolo´gica de re´pteis Brasileiros. Revista Brasileira de Biologia 15: 279–284. Freze, V. I. 1965. Proteocephalata in Fish, Amphibians, and Reptiles. Academy of Sciences of the USSR. Helminthological Laboratory. IPST Press, Jerusalem. 597 pp. (Translated 1969 by Israel Program for Scientific Translations.) Goldberg, S. R., and C. R. Bursey. 1989a. Helminths of the giant spotted whiptail, Cnemidophorus burti stictogrammus (Sauria: Teiidae). Proceedings of the Helminthological Society of Washington 56:86–87. Goldberg, S. R., and C. R. Bursey. 1989b. Physaloptera retusa (Nematoda, Physalopteridae) in naturally infected sagebrush lizards, Sceloporus graciosus (Iguanidae). Journal of Wildlife Diseases 25:425–429. Goldberg, S. R., and C. R. Bursey. 1990. Gastrointestinal

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FIRST ANNOUNCEMENT OF JOINT MEETINGS The Helminthological Society of Washington Southeastern Society of Parasitologists Animal Disease Research Workers in the Southern States Southern Conference on Animal Parasites Southern Conference of Researchers in Aquatic Diseases

6–9 April 2005 DONALDSON-BROWN HOTEL AND CONFERENCE CENTER VIRGINIA TECH BLACKSBURG, VIRGINIA Abstracts due: 12 February 2005 Contact: David S. Lindsay ([email protected]) Additional information available at http://www.gettysburg.edu/;shendrix/helmsoc.html