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BioOne sees sustainable scholarly publishing as an inherently collaborative enterprise connecting authors, nonprofit publishers, academic institutions, research libraries, and research funders in the common goal of maximizing access to critical research. 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 BioOne (www.bioone.org ) is a nonprofit, online aggregation of core research in the biological, ecological, and environmental sciences. BioOne provides a sustainable online platform for over 170 journals and books published by nonprofit societies, associations, museums, institutions, and presses. Your use of this PDF, the BioOne Web site, and all posted and associated content indicates your acceptance of BioOne’s Terms of Use, available at www.bioone.org/page/ terms_of_use . Usage of BioOne content is strictly limited to personal, educational, and non-commercial use. Commercial inquiries or rights and permissions requests should be directed to the individual publisher as copyright holder.
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Gastrointestinal Helminths from 13 Species of Lizards from Reserva Cuzco Amazónico, Peru

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Page 1: Gastrointestinal Helminths from 13 Species of Lizards from Reserva Cuzco Amazónico, Peru

BioOne sees sustainable scholarly publishing as an inherently collaborative enterprise connecting authors, nonprofitpublishers, academic institutions, research libraries, and research funders in the common goal of maximizing access tocritical research.

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

BioOne (www.bioone.org) is a nonprofit, online aggregation of core research in thebiological, ecological, and environmental sciences. BioOne provides a sustainable onlineplatform for over 170 journals and books published by nonprofit societies, associations,museums, institutions, and presses.

Your use of this PDF, the BioOne Web site, and all posted and associated contentindicates your acceptance of BioOne’s Terms of Use, available at www.bioone.org/page/terms_of_use.

Usage of BioOne content is strictly limited to personal, educational, and non-commercialuse. Commercial inquiries or rights and permissions requests should be directed to theindividual publisher as copyright holder.

Page 2: Gastrointestinal Helminths from 13 Species of Lizards from Reserva Cuzco Amazónico, Peru

Gastrointestinal Helminths from 13 Species of Lizards fromReserva Cuzco Amazonico, 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]), and3 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 Amazonico, 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 Amazonico, Peru.

Little information is available on the helminths of

Peruvian lizards. Rego and Ibanez (1965) described

the cestode Oochoristica travassosi Rego and Ibanez,

1965 from specimens taken from an undetermined

species of Leiocephalus. Also from an undetermined

species of Leiocephalus, Vicente and Ibanez (1968)

described the nematode Parathelandros maytacapaciVicente and Ibanez, 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 Amazonico, Peru.

MATERIALS AND METHODS

The 164 lizards representing 13 species used in this studywere collected between November 1984 and December1995 during a biotic survey of the Reserva CuzcoAmazonico, Peru. Taxa by family and sample sizes wereas follows—Gekkonidae: turniptail gecko, Thecadactylusrapicauda, 10; Gymnophthalmidae: ocellated tegu, Cerco-saura ocellata, 5; Eigenmann’s prionodactylus, Prionodac-tylus 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; Amazonkentropyx, Kentropyx altamazonicus, 11; forest whiptail,Kentropyx pelviceps, 15; black tegu, Tupinambis teguixin,

1; Tropiduridae: tree runner, Plica plica, 9; Harlequinracerunner, Plica umbra, 14; and the rose whorltail iguana,Stenocercus roseiventris, 12. Reserva Cuzco Amazonico isa privately maintained reserve of 10,000 ha on an alluvialplain, 200 m elevation, on the north bank of the Rio Madrede Dios about 15 km ENE of Puerto Maldonado,Departamento Madre de Dios, Amazonian Peru (128359S;698059W). Duellman and Koechlin (1991) described thereserve in detail, and Duellman and Salas (1991) providedan annotated checklist of lizards in the area.

Lizards were collected by hand over a decade long bioticsurvey 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 atthe University of Kansas, Lawrence, Kansas, U.S.A.Preserved specimens from the University of Kansasherpetological collections were dissected by J.R.P. as partof a study of the community ecology of frogs (Parmelee,1999) and lizards of Reserva Cuzco Amazonico. The bodycavity of each lizard was opened by a longitudinal incisionfrom throat to vent, the gastrointestinal tract was slitlongitudinally, and stomach and intestinal contents wereremoved and examined with a dissection microscope.Helminths found in the gastrointestinal tract, lungs, or bodycavity were placed in vials of 70% ethanol for lateridentification, at which time each helminth was placed ona glass slide in a drop of undiluted glycerol for study undera compound microscope. Nematodes were identified fromthese preparations; cestodes were stained with hematoxylinand mounted in balsam for identification. Voucher speci-mens of all helminth species were deposited in the UnitedStates National Parasite Collection (USNPC), Beltsville,Maryland, U.S.A. Lizards were deposited in the Herpetol-ogy Collection (KU) of the Natural History Museum of theUniversity of Kansas, Lawrence, Kansas, U.S.A.4 Corresponding author.

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

50

Page 3: Gastrointestinal Helminths from 13 Species of Lizards from Reserva Cuzco Amazónico, Peru

Gekkonidae

Thecadactylus rapicauda(Houttuyn, 1782)

Ten specimens were collected between December

1984 and December 1995 from tree trunks in Reserva

Cuzco Amazonico, 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 retusaRudolphi, 1819

(Syn. Spiroptera retusa [Rudolphi, 1819] Dujardin,

1845; Physaloptera mucronata Leidy, 1856; Phys-aloptera 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. bis-triata, P. plica, P. umbra, S. roseiventris, T. teguixin.

Type host and type locality: Tupinambis teguixin,

Brazil (Rudolphi, 1819).

Other reported hosts: Common lesser toad, Bufogranulosus (Goncalves et al., 2002); South American

common toad, Bufo typhonius (Goncalves 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 Cnemidophorusburti); imbricata alligator lizard, Barisia imbricata(Goldberg et al., 1999); zebratail lizard, Callisaurusdraconoides (Telford, 1970); rainbow whiptail,

Cnemidophorus lemniscatus (Caballero and Vogel-

sang, 1947; Diaz-Ungria, 1964; Diaz-Ungria and

Gallardo, 1968); Cnemidophorus littoralis (Vrcibra-

dic et al., 2000); Laurent’s whiptail, Cnemidophorusmurinus (Specian and Whittaker, 1980); Spix’s

whiptail, Cnemidophorus ocellifer (Ribas et al.,

1995); southern alligator lizard, Elgaria multicari-nata (Telford, 1970); western skink, Eumecesskiltonianus (Telford, 1970); longnose leopard

lizard, Gambelia wislizenii (Telford, 1970); Gerrho-notus ophiurus (Goldberg et al., 1999); green iguana,

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); Ma-buya agilis (Ribas, Teixeira-Filho, et al., 1998); two-

striped mabuya, M. bistriata (recorded as Scincusspixii 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 (Gold-

berg, 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, Sceloporusmucronatus (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, Sceloporusorcutti (Telford, 1970); bluebelly lizard, Sceloporusparvus (Goldberg et al., 2003); crevice spiny lizard,

Sceloporus poinsettii (Goldberg et al., 1993; Gold-

berg, 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, Tropidurustorquatus (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 (Tel-

ford, 1970).

BURSEY ET AL.—HELMINTHS OF PERUVIAN LIZARDS 51

Page 4: Gastrointestinal Helminths from 13 Species of Lizards from Reserva Cuzco Amazónico, Peru

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, Guimar-

aes and Rodrigues, 1976, Physaloptera obtusissimaMolin, 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, Phys-aloptera praeputialis Linstow, 1889, and Physalop-tera 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 third-

stage larvae, which are infective to both definitive

and paratenic hosts. Thecadactylus rapicauda re-

presents a new host record, and Peru is a new location

record for P. retusa.

Physalopteroides venancioi(Lent, Freitas and Proenca, 1946)Sobolev, 1949

(Syn. Physaloptera venancioi Lent, Freitas and

Proenca, 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 Eda-lorhina perezi, Rıo Mamore robber frog Eleuther-odactylus 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, Bufoparacnemis, Assuncao, 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, Mabuyamacrorhyncha (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 rapicaudarepresents a new host record for P. venancioi.

Spauligodon oxkutzcabiensis(Chitwood, 1938) Skrjabin, Schikhobalovaand 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 rapi-cauda, Yucatan, Mexico (Chitwood, 1938).

Other reported hosts: Green spiny lizard, Sceloporusmalachiticus (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 Jimenez-Ruiz,

Leon-Regagnon and Campbell, 2003, Spauligodon

52 COMPARATIVE PARASITOLOGY, 72(1), JANUARY 2005

Page 5: Gastrointestinal Helminths from 13 Species of Lizards from Reserva Cuzco Amazónico, Peru

maytacapaci (Vicente and Ibanez, 1968), Spauligo-don loboi Ramallo, Bursey and Goldberg, 2002, S.oxkutzcabiensis, and Spauligodon viracochai (Frei-

tas, Vicente and Ibanez, 1968). In S. garciaprietoi the

tail of the female tapers to a point; in S. maytacapacithe 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 Jimenez-Ruiz

et al. (2003). The life cycle of S. oxkutzcabiensis has

not been studied, but oxyurids are strictly monoxe-

nous (Anderson, 2000). Peru is a new locality record

for S. oxkutzcabiensis.

Gymnophthalmidae

Cercosaura ocellataWagler, 1830

Five specimens were collected between January

1986 and June 1989 from leaf litter in Reserva Cuzco

Amazonico; SVL ¼ 46 mm 6 14 mm SD, range 22–

54 mm; 1 juvenile, 4 females.

Physalopteroides venancioi(Lent, Freitas and Proenca, 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 eigenmanniGriffin, 1917

Twelve specimens were collected between January

1986 and December 1991 from leaf litter in Reserva

Cuzco Amazonico; SVL ¼ 30 mm 6 10 mm SD,

range ¼ 18–49 mm; 8 juveniles, 2 females, 2 males.

Oswaldocruzia vittiBursey and Goldberg, 2004

Prevalence and intensity: One of 12 hosts infected

(8%, 1).

Temporal distribution: February 1990, 1 host with 1.

Site of infection: Small intestine.

Additional hosts from Peru: None.

Type host and type locality: Prionodactylus eigen-manni, Amazonas Province, Brazil (Bursey and

Goldberg, 2004).

Other reported hosts: O’Shaughnessy’s prionodacty-

lus, 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 Oswaldocruziaoccur in South America (Ben Slimane et al., 1996).

Of these, 4 are parasites of lizards: Oswaldocruziabainae 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 inO.brasiliensis andO. vitti, type II inO. peruensis, type III

in O. bainae (see Ben Slimane et al. [1996]); and then

by cervical alae: absent inO. brasiliensis but present in

O. vitti. The life cycles of species of Oswaldocruziaparasitic in lizards have not been studied, but the life

cycle of Oswaldocruzia pipiens Walton, 1929, a par-

asite 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 fuscoauratusD’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

Amazonico, Peru: SVL¼ 41 mm 6 3 mm SD, range

32–44 mm; 1 juvenile, 16 females, 8 males.

Strongyluris oscariTravassos, 1923

(Syn. Strongyluris sai Travassos, 1926; Strongylurisfreitasi Alho, 1969; Strongyluris travassosi Alho,

1969.)

BURSEY ET AL.—HELMINTHS OF PERUVIAN LIZARDS 53

Page 6: Gastrointestinal Helminths from 13 Species of Lizards from Reserva Cuzco Amazónico, Peru

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 punctatusDaudin, 1802

Fourteen specimens were collected between July

1979 and January 1994 from bushes, tree limbs, or

tree trunks in Reserva Cuzco Amazonico, Peru:

SVL ¼ 71 mm 6 12 mm SD, range 50–88 mm; 3

juveniles, 4 females, 7 males.

Cyrtosomum longicaudatumBrenes 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, Cteno-saura similis, Costa Rica (Brenes and Bravo Hollis,

1960).

Other reported hosts: Baracoa anole,Anolis baracoae(Coy Otero and Barus, 1979); knight anole, Anolisequestris (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, Chamaele-olis 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, Leio-cephalus 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 long-icaudatum from Costa Rica, Cyrtosomum megaBowie 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. longicaudatumis 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 peruvensisBen Slimane, Verhaagh andDurette-Desset, 1995

Prevalence and intensity: One of 14 hosts infected

(7%, 1).

54 COMPARATIVE PARASITOLOGY, 72(1), JANUARY 2005

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Temporal distribution: July 1993, 1 host with 1.

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).

Specimen deposited: USNPC 93235 (1 vial).

Remarks: Anolis punctatus represents a new host

record for O. peruvensis. See remarks on O. vittiunder P. eigenmanni.

Physaloptera retusaRudolphi, 1819

Prevalence and intensity: One of 14 hosts infected

(7%, 1).

Temporal distribution: December 1984, 1 host with 1.

Site of infection: Stomach.

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 lainsoniBain, 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 PiratubashawiBain, 1974. The species are best separated on the

basis of microfilaria (Bain, 1974), but the pattern of

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. Piratubalainsoni has subequal spicules; P. scaffi and P. shawihave spicules of distinctly differing lengths. Piratubascaffi has 10 or fewer terminal caudal papillae;P. shawihas 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.

Rhabdias anolisBursey, Goldberg and Telford, 2003

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.

Additional hosts from Peru: None.

Type host and type locality: Anolis frenatus, Panama

Province, Panama (Bursey et al., 2003).

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, Rhabdiaschamaeleonis (Skrjabin, 1916) and Rhabias gemelli-para 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 ever-manni, Gundlach’s anole Anolis gundlachi, stripefoot

anole Anolis lineatopus, and Watt’s anole Anoliswattsi (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.)

BURSEY ET AL.—HELMINTHS OF PERUVIAN LIZARDS 55

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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).

Specimen deposited: USNPC 93239 (1 vial).

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 third-

stage larva, which was probably autoinfective, and

thick-shelled eggs with third-stage larva that pass from

the host and were infective to crickets. Anolispunctatus represents a new host record, and Peru is

a new location record for S. intermedia.

Strongyluris oscariTravassos, 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

Reserva Cuzco Amazonico, 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; Pseu-dothelandros 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.

Site of infection: Large intestine.

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

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); Tro-pidurus 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 scuti-punctatus, 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 Frei-

tas, 1957, Parapharyngodon largitor Alho and

Rodrigues, 1963, Parapharyngodon riojensis Ram-

allo, 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

56 COMPARATIVE PARASITOLOGY, 72(1), JANUARY 2005

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neotropical species of Parapharyngodon. Mabuyabistriata represents a new host record, and Peru is

a new locality record for P. scleratus.

Physaloptera retusaRudolphi, 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.

Site of infection: Stomach.

Specimen deposited: USNPC 93250 (1 vial).

Remarks: General information and remarks are

reported under T. rapicauda.

Physalopteroides venancioi(Lent, Freitas and Proenca, 1946)Sobolev, 1949

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 asM.mabouya), Belem, Brazil (Bain, 1974).

Other reported hosts: None.

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

Specimen deposited: USNPC 93252 (1 vial).

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 Amazonico, Peru: SVL ¼ 100

mm 6 41 mm SD, range ¼ 48–158 mm; 9 juveniles,

6 females, 10 males.

Oochoristica ameivae(Beddard, 1914) Baer, 1924

(Syn. Linstowia ameivae Beddard, 1914; Oochoris-tica brasiliensis Fuhrmann, 1927; Oochoristicafuhrmanni Hughes, 1949.)

Prevalence, mean intensity, and range: Three of 25

hosts infected (12%, 1.7 6 1.2, 1–3).

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; Vrcibra-

dic 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), Oochoristicabresslaui Fuhrmann, 1927, Oochoristica iguanaeBursey and Goldberg, 1996, Oochoristica insulae-margaritae Lopez-Neyra and Diaz-Ungria, 1957,

Oochoristia travassoi Rego and Ibanez, 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. ameivaehas a wide scolex (450–650 lm), O. bresslaui

BURSEY ET AL.—HELMINTHS OF PERUVIAN LIZARDS 57

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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.

Amphibiocapillaria freitasilenti(Araujo and Gandra, 1941) Moravec, 1986

(Syn.Capillaria freitasilentiAraujo 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. freitasilentihas not been described; but species of the Capillar-

iinae are known to produce autoinfective eggs,

directly infect the definitive host and indirectly infect

the definitive host (Anderson, 2000). Ameiva ameivarepresents 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 retusaRudolphi, 1819

Prevalence, mean intensity, and range: Eight of 25

hosts infected (32%, 2.3 6 1.0, 1–4).

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).

Remarks: General information and remarks are

reported under T. rapicauda.

Spinicauda spinicauda(Olfers in Rudolphi, 1819)Travassos, 1923

(Syn. Heterakis turgida Schneider, 1866, Spinicaudaamarali Pereira, 1935, Heterakis campanula Lin-

stow, 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; Ro-

drigues 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 (Ander-

son, 2000). Peru is a new locality record for

S. spinicauda.

58 COMPARATIVE PARASITOLOGY, 72(1), JANUARY 2005

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Kentropyx altamazonicusCope, 1876

Eleven specimens were collected between December

1984 and February 1986 from the ground in Reserva

Cuzco Amazonico, Peru: SVL¼66 mm 6 14 mm SD,

range ¼ 36–87 mm; 1 juvenile, 6 females, 4 males.

Physaloptera retusaRudolphi, 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 Proenca, 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 pelvicepsCope, 1868

Fifteen specimens collected January 1985 and July

1993 from the ground or logs in Reserva Cuzco

Amazonico, Peru: SVL ¼ 93 mm 6 15 mm SD,

range 60–109 mm; 1 juvenile, 5 females, 9 males.

Ophiotaenia flavaRudin, 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.

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), Ophiotaeniaelongata Fuhrmann, 1927, O. flava, Ophiotaeniahyalina Rudin, 1917, Ophiotaenia jarara Fuhrmann,

1927, Ophiotaenia macrobothra Rudin, 1817, Ophio-taenia 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. Kentropyxpelviceps 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 Travas-

sos, 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 specta-

cled 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, Ranacatesbeiana, and in the Florida leopard frog, Rana

BURSEY ET AL.—HELMINTHS OF PERUVIAN LIZARDS 59

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sphenocephala, from North Carolina, U.S.A. Arya

(1980) found fourth-stage larvae of Dujardinascarisin 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).

Temporal distribution: February 1990, 1 host with 1.

Site of infection: Large intestine.

Specimen deposited: USNPC 93245 (1 vial).

Remarks: Kentropyx pelviceps represents a new host

record for P. scleratus. General information and

remarks are reported under M. bistriata.

Physaloptera retusaRudolphi, 1819

Prevalence, mean intensity, and range: Nine of 15

hosts infected (60%, 8.2 6 9.9, 1–32).

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.

Site of infection: Stomach.

Specimen deposited: USNPC 93246 (1 vial).

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 Proenca, 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.

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 Amazonico, Peru:

SVL ¼ 202 mm.

Dujardinascaris sp.third-stage larvae)

Prevalence and intensity: One of 1 hosts infected

(100%, 8).

Temporal distribution: February 1990, 1 host with 8.

Site of infection: Stomach.

Specimen deposited: USNPC 93269 (1 vial).

Remarks: General information and remarks are

reported under K. pelviceps.

Physaloptera retusaRudolphi, 1819

Prevalence and intensity: One of 1 hosts infected

(100%, 72).

Temporal distribution: February 1990, 1 host with

72.

Site of infection: Stomach.

Specimen deposited: USNPC 93267 (1 vial).

Remarks: General information and remarks are

reported under T. rapicauda.

Physalopteroides venancioi(Lent, Freitas and Proenca, 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.

60 COMPARATIVE PARASITOLOGY, 72(1), JANUARY 2005

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Tropiduridae

Plica plica(Linnaeus, 1758)

Nine specimens were collected between December

1984 and March 1994 from tree branches and tree

trunks in Reserva Cuzco Amazonico, Peru: SVL ¼102 mm 6 31 mm SD, range 47–141 mm; 1 juvenile,

8 females.

Physaloptera retusaRudolphi, 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 oscariTravassos, 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 re-

ported under A. fuscoauratus.

Plica umbra(Linnaeus, 1758)

Fourteen specimens were collected between De-

cember 1985 and March 1994 from tree trunks in

Reserva Cuzco Amazonico, Peru: SVL ¼ 102 mm 6

31 mm SD, range 47–141 mm; 3 juveniles, 6 females,

5 males.

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).

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 digiticaudumFreitas, 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, Spilotespullatus, 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 retusaRudolphi, 1819

Prevalence, mean intensity, and range: Nine of 14

hosts infected (64%, 24.1 6 30.0, 1–94).

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.

BURSEY ET AL.—HELMINTHS OF PERUVIAN LIZARDS 61

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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.

Piratuba digiticaudaLent and Freitas, 1941

Prevalence, mean intensity, and range: Two of 14

hosts infected (14%, 4.5 6 4.9, 1–8).

Temporal distribution: December 1993, 1 host with

8; February 1994, 1 host with 1.

Site of infection: Body cavity.

Additional hosts from Peru: None.

Type host and type locality: Unidentified lizard,

Brazil (Lent and Freitas, 1941).

Other reported hosts: Anolis baracoae (Barus and

Coy Otero, 1969), two-marked anole, Anolis bipor-catus (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.

Strongyluris oscariTravassos, 1923

Prevalence, mean intensity, and range: Eight of 14

hosts infected (57%, 5.0 6 4.7, 1–16).

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.

Stenocercus roseiventrisDumeril and Bibron, 1837

Twelve specimens collected October 1987 and

January 1996 from the ground and logs in Reserva

Cuzco Amazonico, Peru: SVL¼81 mm 6 20 mm SD,

range 38–101 mm; 1 juvenile, 5 females, 9 males.

Oswaldocruzia peruvensisBen Slimane, Verhaagh andDurette-Desset, 1995

Prevalence and intensity: One of 12 hosts infected

(8%, 12).

Temporal distribution: October 1987, 1 host with 12.

Site of infection: Small intestine.

Specimen deposited: USNPC 93260 (1 vial).

Remarks: General information and remarks are

reported under A. punctatus.

Oswaldofilaria azevedoiBain, 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.

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

Specimen deposited: USNPC 93261 (1 vial).

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 Vuyl-

steke, 1937), Oswaldofilaria petersi Bain and Sula-

hian, 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

62 COMPARATIVE PARASITOLOGY, 72(1), JANUARY 2005

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have a 1:2 or less short to long spicule ratio, the other

species have 1:3 or greater ratios. Oswaldofilariabrevicaudata has 4 large terminal caudal papillae; O.azevedoi has 4 tiny terminal caudal papillae. Oswal-dofilaria 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 retusaRudolphi, 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 oscariTravassos, 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

(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 (�x 6 1

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 (I¼ 68), followed by S. oscari (I¼55), S. spinicauda (I¼ 23), and P. venancioi (I¼ 22).

Physaloptera retusa and P. venancioi are heteroxe-

nous. 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 (hel-

minths 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

Amazonico (Bursey et al., 2001) that also reported

generalist helminth communities. Interestingly, only

1 species, P. venancioi, was found to infect both frogs

BURSEY ET AL.—HELMINTHS OF PERUVIAN LIZARDS 63

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(9 species) and lizards (6 species) in Reserva Cuzco

Amazonico. However, the importance of P. venancioito the anuran community was I ¼ 7.5 as compared

with I¼22 for the lizard community. The life cycle of

P. venancioi has not been studied, but physalopter-

ines 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 herpe-

tologists who worked at Reserva Cuzco Amazonico,

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 Geo-

graphic Society (3196-85, 3405-86, and 4016-89; W.

E. Duellman, principal investigator). Permits were

issued by Ing. G. B. Mejıa Munoz and Blgo. Jose

Purisaca of the Direccion General Forestal y de

Fauna, Lima, Peru.

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FIRST ANNOUNCEMENT OF JOINT MEETINGS

The Helminthological Society of WashingtonSoutheastern Society of Parasitologists

Animal Disease Research Workers in the Southern StatesSouthern 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

68 COMPARATIVE PARASITOLOGY, 72(1), JANUARY 2005