Upload
jeffrey-r
View
223
Download
7
Embed Size (px)
Citation preview
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.
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
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
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
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
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
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
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
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
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
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
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
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
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
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
(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.
LITERATURE CITED
Aho, J. M. 1990. Helminth communities of amphibians andreptiles: comparative approaches to understandingpatterns 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 PlanaltoCentral. Sobre o genero Strongyluris Mueller, 1894com descricao de duas especies novas. RevistaBrasileira de Biologia 29:65–74.
Alho, C. J. R. 1970. Frequency of infestation by Strong-yluris freitasi Alho, 1969 in Tropidurus troquatus(Wied). Revista Brasileira de Biologia 30:539–542.
Alho, C. J. R., and H. O. Rodrigues. 1963. Sobre novohospedeiro de Parapharyngodon scleratus (Travassos,1923) Freitas, 1957, com redescricao da especie(Nematoda, Oxyuroidea). Atas Sociedade de Biologiado Rio de Janeiro 7:4–6.
Anderson, R. M. 2000. Nematode Parasites of Vertebrates:Their Development and Transmission, 2nd ed. CABIPublishing, Wallingford, Oxon, U.K. 650 pp.
Araujo, P. 1970. Considerac~oes sobre Hastospiculumonchocercum major Desportes, 1941, parasita de ser-pente Crotalus durissus terrificus. Revista de farmacia egioquimica da Universidade de Sao Paulo 8:121–124.
Araujo, T. L., and V. Gandra. 1941. Sobre uma novaespecie do genero Capillaria e observac~oes helminto-logicas. Revista da Faculdade de Medicina Veterinaria2:29–32.
Arya, S. N. 1980. On the larval forms of a nematode,Dujardinascaris cybii Arya and Johnson, 1978. RevistaBrasileira de Biologia 40:751–754.
Bain, O. 1974. Description de nouvelles filaires Oswaldo-filariinae de lezards sud-americains; hypothese sur
l’evolution des filaires de Reptiles. Memoires duMuseum National d’Histoire Naturelle Series 3 138:169–200.
Bain, O., and A.-G. Chabaud. 1975. Developpement chezdes moustiques de trois filaires de lezards sud-americains du genre Oswaldofilaria. Annales de Para-sitologie (Paris) 50:209–221.
Baker, M. R. 1978. Development and transmission ofOswaldocruzia pipiens Walton, 1929 (Nematoda:Trichostrongylidae) in amphibians. Canadian Journalof Zoology 56:1026–1031.
Baker, M. R. 1987. Synopsis of the Nematoda parasitic inamphibians and reptiles. Memorial University ofNewfoundland, Occasional Papers in Biology 11:1–325.
Barus, V. 1973. Some remarks on the neotropical species ofthe 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 Anolisequestris (Squamata: Iguanidae). Folia Parasitologica(Praha) 15:41–45.
Barus, V., and A. Coy Otero. 1969. Systematic survey ofnematodes parasitizing lizards (Sauria) in Cuba.Helminthologia 10:329–346.
Barus, V., and A. Coy Otero. 1974. Nematodes of thegenera Spauligodon, Skrjabinodon, and Pharyngodon(Oxyuridae) parasitizing Cuban lizards. Vestnik Ce-skoslovenske Spolecunosti Zoologicke 38:1–12.
Baylis, H. A. 1928. Some parasitic worms mainly fromfishes from Lake Tanganyika. Annals and Magazine ofNatural History Series 10 4:552–562.
Beddard, F. E. 1914. Contributions to the anatomy andsystematic arrangement of the Cestoidea. XIII. On twonew species belonging to the genera Oochoristica andLinstowia, with remarks upon those genera. Proceed-ings of the Zoological Society of London 1914:263–282.
Ben Slimane, B., A.-G. Chabaud, and M.-C. Durette-Desset. 1996. Les nematodes Trichostrongylinaparasites d’amphibiens et de reptiles: problemestaxonomiques, phyletiques et biogeographiques. Sys-tematic Parasitology 35:179–206.
Ben Slimane, B., M. Verhaagh, and M.-C. Durette-Desset. 1995. Oswaldocruzia peruensis n. sp. (Nem-atoda, Trichostrongylina, Molineoidea), parasite d’uniguanidae du Perou. Memoires du Museum Nationald’Histoire Naturelle de Paris 17:77–82.
Brandt, B. B. 1936. Parasites of certain North CarolinaSalientia. Ecological Monographs 6:491–532.
Brenes, R.-R., and M. Bravo Hollis. 1960. Helmintos de laRepublica de Costa Rica IX. Nematoda 3. Algunosnematodos de reptiles, con descripcion de dos nuevasespecies: Atractis caballeroi n. sp. y Cyrtosomumlongicaudatum n. sp. Pages 451–464 in E. Caballero,ed. Libro Homenaje al Docter Eduardo Caballero yCaballero; Jubileo 1930–1950. Escuela Nacional deCiencias Biologicas, 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 6Anolis species. Journal of Helminthology 61:77–83.
Bursey, C. R., and J. P. Flanagan. 2002. Atractis marquezin. sp. (Nematoda: Atractidae) and a revision of Atractis
64 COMPARATIVE PARASITOLOGY, 72(1), JANUARY 2005
Dujardin, 1945 sensu Baker, 1987. Journal of Parasi-tology 88:320–324.
Bursey, C. R., and S. R. Goldberg. 1991. Monthly pre-valences of Physaloptera retusa in naturally infectedYarrow’s spiny lizard. Journal of Wildlife Diseases 27:710–715.
Bursey, C. R., and S. R. Goldberg. 1994. Persistenceof the component parasite community of Yarrow’sSpiny Lizard, Sceloporus jarrovii, 1967–1991. Journalof the Helminthological Society of Washington 61:141–145.
Bursey, C. R., and S. R. Goldberg. 1996. Oochoristicamaccoyi n. sp. (Cestoda: Linstowiidae) from Anolisgingivinus (Sauria: Polychrotidae) collected in An-guilla, Lesser Antilles. Caribbean Journal of Science32:390–394.
Bursey, C. R., and S. R. Goldberg. 2004. Cosmocercavrcibradici n. sp. (Ascaridida, Cosmocercidae), Oswal-docruzia vitti n. sp. (Strongylida, Molineoidae), andother helminths from Prionodactylus eigenmanni andPrionodactylus oshaughnessyi (Sauria, Gymnophthal-midae) from Brazil and Ecuador. Journal of Parasitol-ogy 90:140–145.
Bursey, C. R., S. R. Goldberg, and J. R. Parmelee. 2001.Gastrointestinal helminths of 51 species of anuransfrom Reserva Cuzco Amazonico, Peru. ComparativeParastiology 68:21–35.
Bursey, C. R., S. R. Goldberg, and S. R. Telford, Jr.2003. Rhabdias anolis n. sp. (Nematoda: Rhabdiasidae)from the lizard, Anolis frenatus (Sauria: Polychrotidae),from Panama. Journal of Parasitology 89:113–117.
Caballero, E. 1947. Algunas filarias de mamıferos y dereptiles de la Republica de Colombia y Panama. Analesdel Instituto de Biologıa, Universidad Nacional Auton-oma de Mexico 18:169–188.
Caballero, E. 1951. Estudios helmintologicos de la regiononcocercosa de Mexico y de la Republica deGuatemala. Nematoda. 6a parte y algunas consider-aciones en torno a los generos Onchocerca Diesing,1841 y Acanthospiculum Skrjabin y Schikhobalowa,1948. Anales del Instituto de Biologıa, UniversidadNacional Autonoma de Mexico 20:141–158.
Caballero, E., and E. G. Vogelsang. 1947. Faunahelmintologica venezolana. I. Ochetosoma miladelar-ocai n. sp., de Bothrops atrox L. y hallazgo dePhysaloptera retusa (Rud., 1819) en Cnemidophoruslemniscatus lemniscatus. Revista de Medicina Veter-inaria y Parasitologia 6:53–62.
Caballero DeLoya, J. 1971. Redescripcion de Cyrtosomumlongicaudatum Brenes y Bravo, 1960 (Nematoda:Oxyuroidea). Anales del Instituto de Biologıa, Uni-versidad Nacional Autonoma de Mexico 42:29–34.
Chabaud, A. G., O. Bain, and G. O. Poinar. 1988.Skrjabinelazia galliardi (Nematoda, Seuratoidea):complements morphologiques and cycle biologique.Annales de Parasitologie Humaine et Comparee 63:278–284.
Chitwood, B. G. 1932. A review of the nematodes of thegenus Hastospiculum, with descriptions of two newspecies. Proceedings of the U.S. National Museum 80:1–9.
Chitwood, B. G. 1938. Some nematodes from the caves ofYucatan. Publication of the Carnegie Institute ofWashington 491:51–66.
Coy Otero, A. 1970. Contribucion al conocimiento de lahelmintofauna de los saurios cubanos. Ciencias 4:1–58.
Coy Otero, A., and V. Barus. 1973. Notes on nematodes ofthe genus Cyrtosomum (Atractidae) parasitic in Cubanlizards (Sauria). Folia Parasitologica (Praha) 20:297–305.
Coy Otero, A., and V. Barus. 1979. Nematodes parasitiz-ing Cuban reptiles. Acta Scientiarum Naturalium Aca-demiae Scientiarum Bohemoslovacae Brno 13:1–43.
Cristofaro, R., J. F. Guimaraes, and H. de OliveiraRodrigues. 1976. Alguns nematodeos de Tropidurustorquatus (Wied) e Ameiva ameiva (L.)—FaunaHelmintologica de Salvador, Bahia. Atas Sociedadede Biologia de Rio de Janeiro 18:65–70.
Desportes, C. 1941. Sur un Hastospiculum parasite ducrotale. Annales de Parasitologie Humaine et Com-paree 18:198–208.
Diaz-Ungria, C. 1964. Notas sobre nematodes de reptilesen Venezuela. Revista Veterinaria Venezolana 17:402.
Diaz-Ungria, C., and M. F. Gallardo. 1968. Nematodes dereptiles Venezolanos, con descripcion de variasespecies nuevas. Boletain Sociedad Venezolana deCiencias Naturales 27:550–570.
Dobson, A. P., S. V. Pacala, J. D. Roughgarden, E. R.Carper, and E. A. Harris. 1992. The parasites ofAnolis lizards in the northern Lesser Antilles. I. Patternsof distribution and abundance. Oecologia 91:110–117.
Duellman, W. E., and J. E. Koechlin. 1991. The ReservaCuzco Amazonico, Peru: biological investigations,conservation, and ecotourism. Occasional Papers ofthe Museum of Natural History, University of Kansas142:1–38.
Duellman, W. E., and A. W. Salas. 1991. Annotatedchecklist of the amphibians and reptiles of CuzcoAmazonico, Peru. Occasional Papers of the Museum ofNatural History, University of Kansas 143:1–13.
Everard, C. O. R. 1975. Endoparasites of some Amphibia,reptiles and small mammals from Trinidad. Journal ofthe Trinidad and Tobago Field Naturalists’ Club1975:72–79.
de Fabio, S. P., and F. J. T. Rolas. 1974. Contribuicao aoconhecimento de Thubuneae dactyluris Karve, 1938(Nematoda, Spiruroidea). Memorias do Instituto Os-waldo Cruz 72:283–290.
Freitas, J. F. T. 1940. Sobre um interessante nematodeoparasito de reptil (Spiruroidea). Memorias do InstitutoOswaldo Cruz 35:603–605.
Freitas, J. F. T. 1955. Nota sobre a fauna helmintologica derepteis Brasileiros. Revista Brasileira de Biologia 15:279–284.
Freze, V. I. 1965. Proteocephalata in Fish, Amphibians, andReptiles. Academy of Sciences of the USSR. Helmin-thological Laboratory. IPST Press, Jerusalem. 597 pp.(Translated 1969 by Israel Program for ScientificTranslations.)
Goldberg, S. R., and C. R. Bursey. 1989a. Helminths ofthe giant spotted whiptail, Cnemidophorus burtistictogrammus (Sauria: Teiidae). Proceedings of theHelminthological Society of Washington 56:86–87.
Goldberg, S. R., and C. R. Bursey. 1989b. Physalopteraretusa (Nematoda, Physalopteridae) in naturally in-fected sagebrush lizards, Sceloporus graciosus (Igua-nidae). Journal of Wildlife Diseases 25:425–429.
Goldberg, S. R., and C. R. Bursey. 1990. Gastrointestinal
BURSEY ET AL.—HELMINTHS OF PERUVIAN LIZARDS 65
helminths of the yarrow spiny lizard, Sceloporusjarrovii jarrovii Cope. American Midland Naturalist124:360–365.
Goldberg, S. R., and C. R. Bursey. 1992. Gastrointestinalhelminths of the lizard, Sceloporus malachiticus(Sauria: Iguanidae) from Coast Rica. Journal of theHelminthological Society of Washington 59:125–126.
Goldberg, S. R., and C. R. Bursey. 1996. Gastrointestinalhelminths of the anole Anolis oculatus (Polychridae)from Dominica, Lesser Antilles. Journal of the Hel-minthological Society of Washington 63:125–128.
Goldberg, S. R., C. R. Bursey, and R. L. Bezy. 1995.Helminths of isolated montane populations of Yarrow’sspiny lizard, Sceloporus jarrovii (Phrynosomatidae).Southwestern Naturalist 40:330–333.
Goldberg, S. R., C. R. Bursey, and R. L. Bezy. 1996.Gastrointestinal helminths of Yarrow’s spiny lizardSceloporus jarrovii (Phrynosomatidae) in Mexico.American Midland Naturalist 135:299–309.
Goldberg, S. R., C. R. Bursey, and J. L. Camarillo-Rangel. 1999. Helminths of two lizards, Barisiaimbricata and Gerrhonotus ophiurus (Sauria: Angui-dae), from Mexico. Journal of the HelminthologicalSociety of Washington 66:205–208.
Goldberg, S. R., C. R. Bursey, and J. L. Camarillo-Rangel. 2003. Gastrointestinal helminths of sevenspecies of sceloporine lizards from Mexico. South-western Naturalist 48:208–217.
Goldberg, S. R., C. R. Bursey, and H. Cheam. 1996.Gastrointestinal helminths of six anole species, Anolisarmouri, A. barahonae, A. bahorucoensis, A. brevir-ostris, A. chlorocyanus and A. coelestinus (Polychro-tidae) from Hispaniola. Caribbean Journal of Science32:112–115.
Goldberg, S. R., C. R. Bursey, and H. Cheam. 1997.Persistence and stability of the component helminthcommunity of the sagebrush lizard, Sceloporus gra-ciosus (Phrynosomatidae) from Los Angeles County,California, 1972–1973, 1986–1996. American MidlandNaturalist 138:418–421.
Goldberg, S. R., C. R. Bursey, and H. Cheam. 1998.Composition of helminth communities in montane andlowland populations of the western fence lizard,Sceloporus occidentalis from Los Angeles County,California. American Midland Naturalist 140:186–191.
Goldberg, S. R., C. R. Bursey, and C. T. McAllister.1995. Gastrointestinal helminths of nine species ofSceloporus lizards (Phrynosomatidae) from Texas.Journal of the Helminthological Society of Washington62:188–196.
Goldberg, S. R., C. R. Bursey, and M. Morando.2004 Metazoan endoparasites of twelve species of li-zards from Argentina. Comparative Parasitology 71:208–214.
Goldberg, S. R., C. R. Bursey, and R. Tawil. 1993.Gastrointestinal helminths of the crevice spiny lizard,Sceloporus poinsettii (Phrynosomatidae). Journal of theHelminthological Society of Washington 60:263–265.
Goldberg, S. R., C. R. Bursey, and R. Tawil. 1994.Gastrointestinal helminths of Sceloporus lizards (Phry-nosomatidae) from Arizona. Journal of the Helmintho-logical Society of Washington 61:73–83.
Goncalves, A. Q., J. J. Vicente, and R. M. Pinto. 2002.Nematodes of Amazonian vertebrates deposited in the
Helminthological Collection of the Oswaldo CruzInstitute with new records. Revista Brazileira deZoologia 19:453–465.
Grundmann, A. W. 1959. Parasites recovered from sixspecies of Utah lizards. Journal of Parasitology 45:394.
Jimenez-Ruiz, F. A., V. Leon-Regagnon, and J. A.Campbell. 2003. A new species of Spauligodon(Nematoda: Pharyngodonidae) parasite of Cnemido-phorus spp. (Lacertilia: Teiidae) from southern Mexico.Journal of Parasitology 89:351–355.
Kohn, A., R. M. Pinto, and B. M. M. Fernandes. 1973.Contribuicao ao conhecimento de Strongyluris oscariTravassos, 1923 (Nematoda, Subuluroidea). Memoriasdo Instituto Oswaldo Cruz 71:219–225.
Lent, H., and J. F. T. Freitas. 1941. Sobre os filarideosparasitos de lacertıdeos neotropicos. Revista Brasileirade Biologia 1:383–386.
Lent, H., and J. F. T. Freitas. 1948. Uma colecao denematodeos de vertebrados, do Museu de Historia Na-tural de Montevideo. Memorias do Instituto OswaldoCruz 46:1–71.
Lent, H., J. F. T. Freitas, and M. C. Proenca. 1946.Alguns helmintos de batraquios colecionados no Para-guai. Memorias do Instituto Oswaldo Cruz 44:195–214.
Lincoln, R. C., and R. C. Anderson. 1975. Developmentof Physaloptera maxillaris (Nematoda) in the commonfield cricket (Gryllus pennsylvanicus). Canadian Jour-nal of Zoology 53:385–390.
Molin, R. 1860. Una monografia del genre Physaloptera.Sitzungsberichte Akademii Wissenschafter, Wien 39:637–672.
Moravec, F. 1986. Review of capillariid nematodes(Capillariinae) parasitic in amphibians and reptiles.Part 2. Genus Amphibiocapillaria. Vestnik Ceskoslo-venske Spolecunosti Zoologicke 50:217–230.
Morgan, B. B. 1943. The Physaloptera (Nematoda) ofreptiles. Le Naturaliste Canadien 70:179–185.
Ortlepp, R. J. 1922. The nematode genus PhysalopteraRud. Proceedings of the Zoological Society of London72:999–1107.
Parmelee, J. R. 1999. Trophic ecology of a tropical anuranassemblage. Scientific Papers of the Natural HistoryMuseum of the University of Kansas 11:1–59.
Pearce, R. C., and W. W. Tanner. 1973. Helminths ofSceloporus lizards in the Great Basin and UpperColorado plateau of Utah. Great Basin Naturalist 33:1–18.
Pereira, C. 1935. Os oxyurata parasitos de lacertilia donordeste Brasileiro. Archivos do Instituto Biologico 6:5–27.
Pereira, C. 1937. On Thelandros scleratus Travassos, 1923.Pages 463–466 in R. S. Shul’ts, m. P. Gnedina, andK. I. Skriabin, eds. Raboty po gel’mintologii. Izd-voVsesoiuznoi Akademii, Moscow, Russia.
Poinar, G. O., and C. Vaucher. 1972. Cycle larvaire dePhysaloptera retusa Rudolphi, 1819 (Nematoda, Phys-alopteridae), parasite d’un lezard sud-americain. Bul-letin du Museum National D’Histoire Naturelle,Zoologie 74:1321–1327.
Ramallo, G., C. R. Bursey, and S. R. Goldberg. 2002.Parapharyngodon riojensis n. sp. (Nematoda: Phar-yngodonidae) from the lizard Phymaturus punae
66 COMPARATIVE PARASITOLOGY, 72(1), JANUARY 2005
(Squamata: Iguania: Liolaemidae) from northwesternArgentina. Journal of Parasitology 88:979–982.
Rego, A. A., and N. Ibanez. 1965. Duas novas especies deOochoristica, parasitas de lagartixas do Peru (Cestoda,Anoplocephalidae). Memorias do Instituto OswaldoCruz 63:67–73.
Ribas, S. C., C. F. D. Rocha, P. F. Teixeira-Filho, andJ. J. Vicente. 1995. Helminths (Nematoda) of thelizard Cnemidophorus ocellifer (Sauria: Teiidae):Assessing the effect of rainfall, body size and sexin the nematode infection rates. Ciencia e Cultura 47:88–91.
Ribas, S. C., C. F. D. Rocha, P. F. Teixeira-Filho, andJ. J. Vicente. 1998. Nematode infection in twosympatric lizards (Tropidurus torquatus and Ameivaameiva) with different foraging tactics. Amphibia-Reptilia 19:323–330.
Ribas, S. C., P. F. Teixeira-Filho, C. R. D. Rocha, and J.J. Vicente. 1998. Parasitismo por nematoideos em duasespecies simpatricas de Mabuya (Lacertilia: Scincidae)na restinga de Barra de Marica, RJ. Anais do VIIISeminario Regional de Ecologia 8:883–894.
Roca, V. 1997. Tropidurus melanopleurus (NCN) parasites.Herpetological Review 28:204.
Rocha, C. F. D. 1995. Nematode parasites of the Braziliansand lizard, Liolaemus lutzae. Amphibia-Reptilia 16:412–415.
Rodrigues, H. O. 1970. Estudo da fauna helmintologicade Hemidactylus mabouia (M. de J.) nno estado daGuanabara. Atas Sociedade de Biologica de Rio deJaneiro 12:15–23.
Rodrigues, H. O., and L. M. F. Feijo. 1976. Redescricaode Spinicauda spinicauda (Olfers, 1919) Travassos,1920 (Nematoda, Oxyuroidea). Atas Sociedade deBiology de Rio de Janeiro 18:59–63.
Rodrigues, H. O., and R. M. Pinto. 1967. Ocorrencia dedois faringodonıdeos em Ameiva ameiva (L.) no estadoda Guanabara (Nematoda, Oxyuroidea). Atas Socie-dade de Biologia Rio de Janeiro 11:123–125.
Rudin, E. 1917. Die Ichthyotaenien der Reptilien. Revuesuisse de Zoologie 25:179–381.
Rudolphi, C. 1819. Entozoorum synopsis, cui acceduntmantissa duplex et indices locupletissimi. AugustRucker, Berlin, Germany. 811 pp.
Schell, S. C. 1952. Studies on the life cycle of Physalopterahispida Schell (Nematoda: Spiruroidea) a parasite ofthe cotton rat (Sigmodon hispidus littoralis Chapman).Journal of Parasitology 38:462–472.
Schuurmans-Stekhoven, J. H. 1950. Nematodos para-sitarios del chaco Paraguayo y de Argentina delMuseo de Estocolmo. Acta Zoologica Lilloana 9:325–345.
Specian, R. D., and F. H. Whittaker. 1980. A survey of thehelminth fauna of Cnemidophorus murinus from theIsland of Curacao. Proceedings of the HelminthologicalSociety of Washington 47:275–276.
Sprehn, C. 1932. Uber einige von Dr. Eisentraut in Boliviengesammelte nematoden. Zoologischer Anzeiger 100:273–284.
Sprent, J. F. A. 1977. Ascaridoid nematodes of amphibiansand reptiles: Dujardinascaris. Journal of Helminthol-ogy 51:253–287.
Telford, S. R., Jr. 1970. A comparative study of endo-parasitism among some southern California lizard
populations. American Midland Naturalist 83:516–554.
Torres Ortiz, R. A. 1980. Otro hospedador final paraRhabdias sp. (Nematoda) e implicaciones evolutivas delas especies del hospedador. Caribbean Journal ofScience 16:109–114.
Travassos, L. 1920. Contribuic~oes para o conhecimento dafauna helmintolojica brazileira. IX. Sobre as especiesdo genero Spinicauda n. g. Memorias do InstitutoOswaldo Cruz 12:41–50.
Travassos, L. 1923. Informac~oes sobre a fauna helmintho-logica de Mato Grosso. Folha Medica (Rio de Janeiro)4:58–60.
Travassos, L. 1926. Sobre mais uma especie dogenero Strongyluris Mueller, 1894, encontrada noBrasil. Anais de Faculdade de Medicena Sao Paulo 1:83–84.
Travassos, L. 1933. Ascarides des crocodiles sud-amer-icains. Comptes Rendus des Seances de la Societe deBiologie, Paris 114:833–834.
Vado, E. Y. 1972. Etude de huit nematodes parasites devertebres du Venezuela et de la Colombie. Bulletin duMuseum national d’Histoire naturelle, Zoologie 35:477–498.
Van Sluys, M., C. F. D. Rocha, H. G. Bergallo, D.Vrcibradic, and S. C. Ribas. 1997. Nematodeinfection in three sympatric lizards in an isolatedfragment of restinga habitat in southeastern Brazil.Amphibia-Reptilia 18:442–446.
Van Sluys, M., C. F. D. Rocha, and S. C. Ribas.1994. Nematodes infecting the lizard Tropidurusitambere in southeastern Brazil. Amphibia-Reptilia15:405–408.
Vicente, J. J. 1981. Helmintos de Tropidurus (Lacertilia,Iguanidae) da colecao helmintologica do InstitutoOswaldo Cruz. II. Nematoda. Atas Sociedade deBiologia do Rio de Janeiro 22:7–18.
Vicente, J. J., and H. N. Ibanez. 1968. Nova especie dogenero Parathelandros Baylis, 1930 (Nematoda, Oxy-uroidea). Atas Sociedade de Biologia do Rio de Janeiro11:185–187.
Vicente, J. J., and C. R. Jardim. 1980. Filarıdeos daColecao helmintologica do Instituto Oswaldo Cruz. I.Peixes, anfıbios e repteis. Atas Sociedade de Biologiado Rio de Janeiro 21:47–57.
Vicente, J. J., H. O. Rodrigues, D. C. Gomes, and R. M.Pinto. 1993. Nematoides do Brasil. Parte III: Nem-atoides de repteis. Revista brasileria de Zoologica 10:19–168.
Vicente, J. J., and E. Santos. 1967. Ocorrencia dePhysaloptera retusa Rudolphi, 1819 em novo hospe-deiro (Nematoda, Physalopteridae). Atas Sociedade deBiologia do Rio de Janeiro 11:75–77.
Vrcibradic, D., M. Cunha-Barros, J. J. Vicente, C. A. C.Galdino, F. H. Hatano, M. Van Sluys, and C. F. D.Rocha. 2000. Nematode infection patterns in foursympatric lizards from a restinga habitat (Jurubatiba) inRio de Janeiro state, southeastern Brazil. Amphibia-Reptilia 21:307–316.
Vrcibradic, D., C. F. D. Rocha, C. R. Bursey, and J. J.Vicente. 2002. Helminth communities of two sympat-ric skinks (Mabuya agilis and Mabuya macrorhyncha)from two ‘restinga’ habitats in southeastern Brazil.Journal of Helminthology 76:355–361.
BURSEY ET AL.—HELMINTHS OF PERUVIAN LIZARDS 67
Vrcibradic, D., C. F. D. Rocha, S. C. Ribas, and J. J.Vicente. 1999. Nematodes infecting the skink Mabuyafrenata in Valinhos, Sao Paulo State, southeasternBrazil. Amphibia-Reptilia 20:333–339.
Vrcibradic, D., C. F. D. Rocha, and M. Van Sluys. 2001.Mabuya macrorhyncha (NCN) Endoparasites. Herpe-tological Review 32:256.
Vrcibradic, D., J. J. Vicente, and C. R. Bursey. 2000.
Thubunaea dactyluris sensu Fabio and Rolas, a syno-nym of Physalopteroides venancioi (Spirurida, Phys-alopteridae). Journal of Parasitology 86:1163–1165.
Walker, K. A., and D. V. Matthias. 1973. Helminths ofsome northern Arizona lizards. Proceedings of theHelminthological Society of Washington 40:168–169.
Woodbury, L. A. 1934. Notes on some parasites of threeUtah reptiles. Copeia 1934:51–52.
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