Gastrointestinal Helminths from Six Species of Frogs and Three

Species of Lizards, Sympatric in Pará State, Brazil

Author(s): Stephen R. Goldberg, Charles R. Bursey, Janalee P. Caldwell, Laurie
J. Vitt, and Gabriel C. Costa
Source: Comparative Parasitology, 74(2):327-342. 2007.
Published By: The Helminthological Society of Washington
DOI: 10.1654/4268.1
URL: http://www.bioone.org/doi/full/10.1654/4268.1

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Comp. Parasitol.
74(2), 2007, pp. 327–342

Gastrointestinal Helminths from Six Species of Frogs and Three Species of

Lizards, Sympatric in Pará State, Brazil

STEPHEN R. GOLDBERG,1,4 CHARLES R. BURSEY,2 JANALEE P. CALDWELL,3 LAURIE J. VITT,3 AND

GABRIEL C. COSTA3
1
Department of Biology, Whittier College, Whittier, California 90608, U.S.A. (e-mail: sgoldberg@whittier.edu),
2
Department of Biology, Pennsylvania State University, Shenango Campus, Sharon, Pennsylvania 16146, U.S.A.
(e-mail: cxb@psu.edu), and
3
Sam Noble Oklahoma Museum of Natural History and Zoology Department, University of Oklahoma, Norman,
Oklahoma 73072, U.S.A. (e-mails: caldwell@ou.edu, vitt@ou.edu, costage@ou.edu).

ABSTRACT: Forty-three frogs representing 6 species (Dendropsophus cachimbo, Scinax fuscomarginatus, Leptodactylus
fuscus, Leptodactylus martinezi, Leptodactylus mystaceus, and Leptodactylus rhodomystax) and 35 lizards representing 3
species (Kentropyx calcarata, Leposoma osvaldoi, and Potamites ecpleopus) collected in the Brazilian state of Pará were
examined for helminths. One species of Trematoda, Brachycoelium salamandrae, and 12 species of Nematoda, adults of
Capillaria recondita, Cosmocerca brasiliense, Cosmocerca podicipinus, Falcaustra belemensis, Falcaustra maculata,
Kentropyxia sauria, Oswaldocruzia vaucheri, Physaloptera retusa, Schrankiana formulosa, Schrankiana fuscus,
Schrankiana schranki, and juveniles of Acuariidae gen. sp., were found. Only B. salamandrae occurred in both frogs and
lizards. There were 1.53 6 0.13 (x 6 1 SE) helminth species/infected frogs and 28.52 6 11.7 helminth individuals/infected
frogs and 1.12 6 0.08 helminth species/infected lizards and 6.47 6 1.58 helminth individuals/infected lizards. Thirteen new
host records and 2 new locality records are reported.
KEY WORDS: Trematoda, Nematoda, Anura, Sauria, Pará, Brazil.

The Amazonian-Guiana region of South America
has a diverse anuran fauna of at least 305 amphibian
species (Duellman, 1999) and at least 89 lizard species
(Avila-Pires, 1995). The purpose of this study is to
report gastrointestinal helminths from 6 species of
frogs and 3 species of lizards from the Brazilian
state of Pará, the northeastern portion of the Amazo-
nian Guiana biogeographical region: Cachimbo tree-
frog, Dendropsophus cachimbo; brown-bordered
snouted treefrog, Scinax fuscomarginatus; whistling
frog, Leptodactylus fuscus; Martinez’s thin-toed frog,
Leptodactylus martinezi; rose-lipped thin-toed frog,
Leptodactylus rhodomystax; and Spix’s kentropyx,
Kentropyx calcarata; Oswald’s leposoma, Leposoma
osvaldoi; and the common stream lizard, Potamites
ecpleopus.
were later transferred to the Sam Noble Oklahoma Museum
of Natural History (OMNH), University of Oklahoma,
Norman, Oklahoma. Specimens were fixed in 10% neutral
buffered formalin and preserved in 70% ethanol. The
abdominal cavity of each specimen was opened and the
digestive tract was removed. The digestive tract was opened
and examined for helminths under a dissecting microscope.
Trematodes were regressively stained in hematoxylin,
cleared in xylene, and mounted in Canada balsam for
identification. Nematodes were placed on a glass slide,
cleared in a drop of lactophenol solution, covered with
a cover slip, and identified. Helminths were deposited in
the United States National Parasite Collection (USNPC),
Beltsville, Maryland. Amphibian taxonomy is in accordance
with Frost et al. (2006); lizard taxonomy is in accordance
with Avila-Pires (1995) and Doan and Castoe (2005),
however, we have indicated the taxonomic assignment of
a host at the time of its collection.

MATERIALS AND METHODS
Forty-three frogs (Dendropsopus cachimbo, n ¼ 10),
(S. fuscomarginatus, n ¼ 12), (L. fuscus, n ¼ 12),
(L. martinezi, n ¼ 3), (Leptodactylus mystaceus, n ¼ 1),
(L. rhodomystax, n ¼ 5) and 35 lizards (L. osvaldoi, n ¼ 5),
(P. ecpleopus, n ¼ 13), (K. calcarata, n ¼ 17) were col-
lected in Pará, Novo Progresso Municipality, Brazil in
October 2004 and initially deposited in the herpetology
collection of the University of Brası́lia. These specimens
4
Corresponding author.
Hylidae
Dendropsophus cachimbo
(Napoli and Caramaschi, 1999)
Ten specimens were collected 21 October 2004 at
Campo de Provas Brigadeiro Veloso, Novo Progresso
Municipality, Pará, Brazil (9821928.80S, 54854920.20W)
and deposited in OMNH as 40195–40204.
Cosmocerca brasiliense Travassos, 1925
Prevalence and intensity: One of 10 hosts infected
(10%, 1).

327
328 COMPARATIVE PARASITOLOGY, 74(2), JULY 2007
Temporal distribution: 21 October 2004, 1 host with
1 male nematode.
Site of infection: Large intestine.
Type host and type locality: Chaunaus crucifer
(¼Bufo crucifer), Angra do Reis, Brazil (Travassos,
1925).
Additional hosts from Brazil: Guenther’s robber
frog, Eleutherodactylus guentheri, (Travassos, 1925);
smooth horned frog, Proceratophrys appendiculata
(Boquimpani-Freitas et al., 2001); rock river frog,
Thoropa miliaris (Travassos, 1925).
Other reported hosts: Lowland thin-toed frog, Lep-
todactylus andreae (¼Adenomera andreae; Bursey
et al., 2001); Napo’s thin-toed frog, Leptodactylus
hylaedactylus (¼Adenomera hylaedactyla; Bursey
et al., 2001); marine toad, Chaunus marinus (¼Bufo
marinus; Bursey et al., 2001); ruby poison frog,
Epipedobates parvulus (¼Dendrobates parvulus;
Dyer and Altig, 1976, 1977); Perez’s snouted frog,
Edalorhina perezi (Bursey et al., 2001); Amazon
robber frog, Eleutherodactylus altamazonicus (Dyer
and Altig, 1976, 1977); La Paz robber frog,
Eleutherodactylus cruralis (Bursey et al., 2001);
Rı́o Mamore robber frog, Eleutherodactylus fenes-
tratus (Bursey et al., 2001); metallic robber frog,
Eleutherodactylus lanthanites (Dyer and Altig, 1976,
1977); Peru robber frog, Eleutherodactylus peruvia-
nus (Bursey et al., 2001); Pachitea robber frog,
Eleutherodactylus toftae (Bursey et al., 2001); bleat-
ing frog, Hamptophryne boliviana (Dyer and Altig,
1976, 1977); giant gladiator treefrog, Hypsiboas
boans (¼Hyla boans; Dyer and Altig, 1976, 1977;
Bursey et al., 2001); short-headed treefrog, Dendrop-
sophus brevifrons (¼Hyla brevifrons; Bursey et al.,
2001); Troeschel’s treefrog, Hypsiboas calcaratus
(¼Hyla calcarata; Bursey et al., 2001); banded
treefrog, Hypsiboas fasciatus (¼Hyla fasciata; Bursey
et al., 2001); map treefrog, Hypsiboas geographicus
(¼Hyla geographica; Dyer and Altig, 1976, 1977);
Demerara Falls treefrog, Hypsiboas granosus (¼Hyla
granosa; Bursey et al., 2001); Koechlin’s treefrog,
Dendroposphus koechlini (¼Hyla koechlini; Bursey
et al., 2001); basin treefrog, Hypsiboas lanciformis
(¼Hyla lanciformis; Dyer and Altig, 1976, 1977);
Leal’s treefrog, Dendropsophus leali (¼Hyla leali;
Bursey et al., 2001); Beireis’ treefrog, Dendropso-
phus leucophyllatus (¼Hyla leucophyllata; Dyer and
Altig, 1976, 1977; Bursey et al., 2001); marbled
treefrog, Dendropsophus marmoratus (¼Hyla mar-
morata; Dyer and Altig, 1976, 1977; Bursey et al.,
2001); small-headed treefrog, Dendropsophus parvi-
ceps (¼Hyla parviceps; Bursey et al., 2001); red-
robed treefrog, Dendropsophus rhodopeplus (¼Hyla
rhodopepla; Bursey et al., 2001); Sarayacu treefrog,
Dendropsophus sarayacuensis (¼Hyla sarayacuen-
sis; Dyer and Altig, 1976, 1977); Schubart’s treefrog,
Dendropsophus schubarti (¼Hyla schubarti; Bursey
et al., 2001); common big-headed frog, Ischnocnema
quixensis (Dyer and Altig, 1976, 1977); Bolivian
white-lipped frog, Leptodactylus bolivianus (Bursey
et al., 2001); South American bullfrog, Leptodactylus
pentadactylus (Bursey et al., 2001); Peter’s frog,
Leptodactylus petersii (Bursey et al. 2001); Manaus
slender-legged treefrog, Osteocephalus taurinus
(Bursey et al., 2001); Surinam golden-eyed treefrog,
Trachycephalus coriaceus (¼Phrynohyas coriacea;
Bursey et al., 2001); toady leaf frog, Phyllomedusa
atelopoides (Bursey et al., 2001); jaguar leaf frog,
Phyllomedusa palliata (Bursey et al., 2001); brown-
belly leaf frog, Phyllomedusa tarsius (Dyer and
Altig, 1976, 1977); tiger-striped leaf frog, Phyllome-
dusa tomopterna (Bursey et al., 2001); white-lined
leaf frog, Phyllomedusa vaillanti (Bursey et al.,
2001); Tarauaca snouted treefrog, Scarthyla goino-
rum (¼Scarthyla ostinodactyla; Bursey et al., 2001);
Eirunepe snouted treefrog, Scinax garbei (Bursey
et al., 2001); yellow-snouted treefrog, Scinax icter-
icus (Bursey et al., 2001); Henle’s snouted treefrog,
Scinax pedromedinai (Bursey et al., 2001); Orinoco
lime treefrog, Sphaenorhynchus lacteus (Bursey
et al., 2001).
Geographic range: Brazil (Travassos, 1925), Ecua-
dor (Dyer and Altig, 1976, 1977), Peru (Bursey et al.,
2001).
Specimen deposited: USNPC 98183; 1 vial.
Remarks: Dendropsophus cachimbo represents a new
host record for Cosmocerca brasiliense.
Scinax fuscomarginatus
(Lutz, 1925)
Twelve specimens were collected from 21–31
October 2004. Seven collected on 31 October were
from 46.9 km W Campo de Provas Brigadeiro
Veloso, Novo Progresso Municipality, Pará, Brazil
(9822950.10S, 55820925.70W) and deposited in OMNH
as 40216–40222. Four collected on 21 October were
from Campo de Provas Brigadeiro Veloso, Novo
Progresso Municipality, Pará, Brazil (9821928.80S,
54854921.80W) OMNH 40211–40214. One collected
on 24 October was from 3 km W, 6.5 km N Campo
de Provas Brigadeiro Veloso, Novo Progresso
 GOLDBERG ET AL.—HELMINTHS OF FROGS AND LIZARDS IN BRAZIL
Municipality, Pará, Brazil (9821928.80S, 54856936.00W)
OMNH 40215.
Cosmocerca podicipinus Baker and
Vaucher, 1984
Prevalence and intensity: One of 12 hosts infected
(8%, 2).
Temporal distribution: 24 October 2004, 1 host with
1 male and 1 female nematodes.
Site of infection: Large intestine.
Type host and type locality: Dark-bellied frog,
Leptodactylus podicipinus, Capitan Bado, Amambay
Province, Paraguay (Baker and Vaucher, 1984).
Additional hosts from Brazil: None.
Other reported hosts: Pacific fat sleeper, Dormitator
latifrons (Garrido-Olvera et al., 2004); Condoto
stubfoot toad, Atelopus spurrelli (Goldberg and
Bursey, 2003); South American leaf toad, Rhinella
margaritifer (¼Bufo typhonius; Bursey et al., 2001);
dull rocket frog, Colostethus marchesianus (Bursey
et al., 2001); harlequin poison frog, Dendrobates
histrionicus (Goldberg and Bursey, 2003); imitator
robber frog, Eleutherodactylus imitatrix (Bursey
et al., 2001); polymorphic robber frog, Eleutherodacty-
lus rhodopis (Goldberg et al., 2002); brillant-thighed
poison frog, Allobates femoralis (¼Epipedobates femo-
ralis; Bursey et al., 2001); marbled white-lipped frog,
Leptodactylus elenae (Baker and Vaucher, 1984);
whistling frog, Leptodactylus fuscus (Baker and
Vaucher, 1984); stripe-thighed frog, Leptodactylus
leptodactyloides (Bursey et al., 2001); Miranda’s
white-lipped frog, Leptodactylus macrosternum (Baker
and Vaucher, 1984); Sabinal frog, Leptodactylus
melanonotus (Goldberg and Bursey, 2002; Goldberg
et al., 2002); Forrer’s grass frog, Lithobates forreri
(¼Rana forreri; Goldberg and Bursey, 2002); Lithobates
cf. forreri (¼Rana cf. forreri; Bursey and Goldberg,
2005); blue-spotted Mexican treefrog, Smilisca cyanos-
tica (Goldberg et al., 2002).
Geographic range: Brazil (this study), Colombia
(Goldberg and Bursey, 2003), Costa Rica (Bursey
and Goldberg, 2005), Mexico (Goldberg and Bursey,
2002), Paraguay (Baker and Vaucher, 1984), Peru
(Bursey et al., 2001).
Specimens deposited: USNPC 98194; 1 vial.
Remarks: Scinax fuscomarginatus represents a new
host record for Cosmocerca podicipinus. Brazil is
a new locality record.
329
Acuariidae gen. sp. ( juveniles in cysts)
Prevalence and intensity: One of 12 hosts infected
(8%, 2).
Temporal distribution: 21 October 2004, 1 host with
2 cysts.
Site of infection: Stomach wall.
Type host and type locality: Adult acuariid nematodes
are parasites of aquatic birds (Anderson, 2000).
Additional hosts of juveniles from Brazil: agile
mabuya, Mabuya agilis (Vrcibradic et al., 2000;
Vrcibradic et al., 2002); Caissara mabuya, Mabuya
caissara (Rocha and Vrcibradic, 2003); Hoge’s
mabuya, Mabuya macrorhyncha (Vrcibradic et al.,
2000; Vrcibradic et al., 2002); Amazon lava lizard,
Tropidurus torquatus (Vrcibradic et al., 2000).
Other reported hosts of juveniles: Edalorhina perezi
(Bursey et al., 2001); Hyla rhodopepla (Bursey et al.,
2001). Acuariid juveniles have also been reported
from the border anole, Norops limifrons from Costa
Rica (Bursey and Goldberg, 2003) and from the
western fence lizard, Sceloporus occidentalis from
California (Goldberg et al., 1998).
Geographic range: The family Acuariidae has
cosmopolitan distribution (Yamaguti, 1961).
Specimens deposited: USNPC 98195; 1 vial.
Remarks: Acuariid nematodes typically mature in
aquatic birds; they require an arthropod intermediate
host to complete the life cycle; frogs may serve as para-
tenic hosts (Anderson, 2000). Scinax fuscomarginatus
represents a new host record for acuariid juveniles.
Leptodactylidae
Leptodactylus fuscus
(Schneider, 1799)
Twelve specimens were collected from 21 October–
11 November 2004. Eleven were collected on 29, 31
October and 1–3, 9, 11 November from 46.9 km W
Campo de Provas Brigadeiro Veloso, Novo Progresso
Municipality, Pará, Brazil (9822950.10S, 55820925.70W)
and deposited in OMNH as 40229–40239. One
was collected on 21 October at Campo de Provas
Brigadeiro Veloso, Novo Progresso Munici-
pality, Pará, Brazil (9821928.80S, 54854921.80W)
and deposited as OMNH 40228.
Oswaldocruzia vaucheri Ben Slimane and
Durette-Desset, 1993
Prevalence and intensity: One of 12 hosts infected
(8%, 2).
330 COMPARATIVE PARASITOLOGY, 74(2), JULY 2007
Temporal distribution: 31 October, 2004, 1 host
with 2.
Site of infection: Small intestine.
Type host and type locality: common big-headed
frog, Ischnocnema quixensis, San Pablo, Napo Prov-
ince, Ecuador (Ben Slimane and Durette-Desset,
1993).
Additional hosts from Brazil: None.
Other reported hosts: None.
Geographic range: Ecuador (Ben Slimane and
Durette-Desset, 1993), Brazil (this study).
Specimens deposited: USNPC 98187; 1 vial.
Remarks: Leptodactylus fuscus represents a new host
record for Oswaldocruzia vaucheri. Brazil is a new
locality record for O. vaucheri.
Schrankiana formulosa Freitas, 1959
Prevalence, mean intensity, and range: Eight of 12
hosts infected (67%, 12.8 6 13.9 SD, 2–38).
Temporal distribution: 29, 31 October. 2, 3, 9, 11
November 2004, 8 hosts with 2, 2, 3, 8, 9, 9, 31, 38,
respectively.
Site of infection: Large intestine.
Type host and type locality: Leptodactylus fuscus,
Itaguai, Rio de Janeiro, Brazil (Freitas, 1959).
Additional hosts from Brazil: None.
Other reported hosts: Leptodactylus elenae (Baker
and Vaucher, 1988).
Geographic range: Brazil (Freitas, 1959), Paraguay
(Baker and Vaucher, 1988).
Specimens deposited: USNPC 98188; 1 vial.
Remarks: This is the third study to report Schranki-
ana formulosa in Leptodactylus fuscus (Freitas, 1959;
Baker and Vaucher, 1988; this study).
Schrankiana fuscus Baker and
Vaucher, 1988
Prevalence, mean intensity, and range: Nine of 12
hosts infected (75%, 11.33 6 6.5 SD, 1–19).
Temporal distribution: 31 October, 2, 3, 9, 11
November 2004, 9 hosts with 2, 4, 5, 9, 13, 16, 17,
17, 19, respectively.
Site of infection: Large intestine.
Type host and type locality: Leptodactylus fuscus,
Alto Parana Province, Paraguay (Baker and Vaucher,
1988).
Additional hosts from Brazil: None.
Other reported hosts: None.
Geographic range: Brazil (this study); Paraguay
(Baker and Vaucher, 1988).
Specimens deposited: USNPC 98189; 1 vial.
Remarks: Brazil is a new locality record for S. fuscus.
Leptodactylus martinezi
(Bokermann, 1956)
Three specimens were collected on 9 November
2004 from 46.9 km W Campo de Provas Brigadeiro
Veloso, Novo Progresso Municipality, Pará, Brazil
(9822950.10S, 55820925.70W) and deposited in OMNH
as 40225–40227.
Brachycoelium salamandrae (Frolich, 1789)
Dujardin, 1845
Synonymy: Fasciola salamandrae Froelich, 1789;
Distoma crassicolle Rudolphi, 1809; Distomum
flavocinctum Linstow, 1879; Brachycoelium hospi-
tale Stafford, 1903; Brachycoelium obesum Nicoll,
1914; Brachycoelium trituri Holl, 1928; Brachycoe-
lium daviesi Harwood, 1932; Brachycoelium mer-
idionalis Harwood, 1932; Brachycoelium storeriae
Harwood, 1932; Brachycoelium dorsale Byrd, 1937;
Brachycoelium georgianum Byrd, 1937; Brachycoe-
lium louisianae Byrd, 1937; Brachycoelium mesor-
chium Byrd, 1937; Brachycoelium ovale Byrd, 1937;
Brachycoelium elongatum Cheng, 1958; Brachycoe-
lium mesocoeliiformis Freitas, 1961; Brachycoelium
ambystomae Couch, 1966.
Prevalence, mean intensity, and range: Three of 3
hosts infected (100%, 3.7 6 2.3 SD, 1–5).
Temporal distribution: 9 November 2004, 3 hosts
with 1, 5, 5, respectively.
Site of infection: Small intestine.
Type host and type locality: alpine salamander,
Salamandra atra, Germany (Frölich, 1789).
Additional hosts from Brazil: goldenscale anole,
Anolis nitens (¼A. chrysolepis but reported as Anolis
scypheus; Freitas, 1961); Leptodactylus rhodomystax
 GOLDBERG ET AL.—HELMINTHS OF FROGS AND LIZARDS IN BRAZIL
(this study); Leptodactylus rhodomystax (this study);
Leposoma osvaldoi (this study).
Other reported hosts: Nearctic Realm: northern cricket
frog, Acris crepitans (¼Acris gryllus; Najarian, 1955);
American toad, Anaxyrus americanus (¼Bufo amer-
icanus; Rosen and Manis, 1976); western toad,
Anaxyrus boreas (¼Bufo boreas; Lehmann, 1965);
Fowler’s toad, Anaxyrus fowleri (¼Bufo fowleri;
Brandt, 1936; Rankin, 1938; Hering and Murad,
1969); southern toad, Anaxyrus terrestris (¼Bufo
terrestris; Manter, 1938); green treefrog, Hyla
cinerea (Harwood, 1932); crawfish frog, Lithobates
areolatus (¼Rana areolata; Manter, 1938); pickerel
frog, Lithobates palustris (¼Rana palustris; Rankin,
1938, 1945); southern leopard frog, Lithobates
sphenocephalus (¼Rana sphenocephala; Harwood,
1932; Brandt, 1936; Byrd, 1937; Manter, 1938;
Rankin, 1938; Parker, 1941); wood frog, Lithobates
sylvaticus (¼Rana sylvatica; Odlaug, 1954; Najarian,
1955; Prudhoe and Bray, 1982; McAllister, Trauth
and Bursey, 1995a; McAllister, Upton et al., 1995);
Brimley’s chorus frog, Pseudacris brimleyi (Brandt,
1936; Rankin, 1938); spring peeper, Pseudacris
crucifer (Brandt, 1936; Rankin, 1938); ornate chorus
frog, Pseudacris ornata (¼Pseudacris occidentalis;
Byrd, 1937); Pacific chorus frog, Pseudacris regilla
(¼Hyla crucifer; Lehmann, 1965); western chorus
frog, Pseudacris triseriata (Harwood, 1932; Odlaug,
1954); American bullfrog, Lithobates catesbeianus
(¼Rana catesebiana; Parker, 1941; Prudhoe and
Bray, 1982); green frog, Lithobates clamitans
(¼Rana clamitans; Fantham and Porter, 1948);
northern leopard frog, Lithobates pipiens (¼Rana
pipiens; Lehmann, 1965; Rosen and Manis, 1976);
red-legged frog, Rana aurora (Lehmann, 1965);
spotted frog, Rana pretiosa (Lehmann, 1965);
Jefferson salamander, Ambystoma jeffersonianum
(Rankin, 1938, 1945); blue-spotted salamander,
Ambystoma laterale (Muzzall and Schinderle, 1992);
long-toed salamander, Ambystoma macrodactylum
(Waitz, 1961); spotted salamander, Ambystoma
maculatum (Rankin, 1937, 1938, 1945; Rabalais,
1970; Dyer and Brandon, 1973; Rosen and Manis,
1976; Coggins and Sajdak, 1982; Bolek and Coggins,
1998); marbled salamander, Ambystoma opacum
(Byrd, 1937; Rankin, 1937, 1938; Parker, 1941;
Couch, 1966; Rabalais, 1970; Dyer and Brandon,
1973; Joy and Mills, 1975); mole salamander,
Ambystoma talpoideum (Dyer and Brandon, 1973);
small-mouthed salamander, Ambystoma texanum
(Harwood, 1932; Rosen and Manis, 1976); three-
toed amphiuma, Amphiuma tridactylum (Bennett and
331
Humes, 1938); southern dusky salamander, Des-
mognathus auriculatus (Bennett, 1938); Ouachita
dusky salamander, Desmognathus brimleyorum
(McAllister, Bursey et al., 1995); northern dusky
salamander, Desmognathus fuscus (Byrd, 1937;
Rankin, 1937, 1938, 1945; Parker, 1941; Odlaug,
1954; Fischthal, 1955a, b; Cheng, 1958; Cheng and
Chase, 1961; Rabalais, 1970; Rosen and Manis,
1976; Bogitsh and Ryckman, 1982); seal salamander,
Desmognathus monticola (Rankin, 1937; Cheng,
1958); Allegheny Mountain dusky salamander,
Desmognathus ochrophaeus (Rankin, 1937); black-
bellied salamander, Desmognathus quadramaculatus
(Rankin, 1937; Cheng, 1958); California giant sala-
mander, Dicamptodon ensatus (Pratt and McCauley,
1961); ensatina, Ensatina eschscholtzii (Lehmann,
1954; Pratt and McCauley, 1961); northern two-lined
salamander, Eurycea bislineata (Rankin, 1937, 1945;
Parker, 1941; Fischthal, 1955a, b; Rabalais, 1970);
long-tailed salamander, Eurycea longicauda (Rankin,
1937; Dyer and Brandon, 1973); cave salamander,
Eurycea lucifuga (Dyer and Brandon, 1973; Dyer and
Peck, 1975; Castle et al., 1987); many-ribbed
salamander, Eurycea multiplicata (McAllister, Trauth
and Bursey, 1995b); grotto salamander, Eurycea
spelaea (¼Typhlotriton spelaeus; Dyer, 1975); spring
salamander, Gyrinophilus porphyriticus (Fischthal,
1955b; Catalano et al., 1982); four-toed salamander,
Hemidactylum scutatum (Rankin, 1938); black-spotted
newt, Notophthalmus meridionalis (Harwood,
1932; Rankin, 1937; Manter, 1938); eastern newt,
Notophthalmus viridescens (Stafford, 1900, 1903;
Holl, 1928; Rankin, 1937, 1938; Russell, 1951;
Fischthal, 1955b; Cheng, 1958; Jackson and Beau-
doin, 1967; Rabalais, 1970); Red Hills salamander,
Phaeognathus hubrichti (Brandon, 1965); eastern
red-backed Salamander, Plethodon cinereus (Stafford,
1903, 1905; Rankin, 1937, 1938, 1945; Fischthal,
1955a, b; Cheng, 1958; Cheng and Chase, 1961;
Coggins and Sajdak, 1982; Muzzall, 1990; Bursey
and Schibli, 1995; Bolek and Coggins, 1998);
northern zigzag salamander, Plethodon dorsalis
(Dyer and Brandon, 1973); northern slimy salaman-
der, Plethodon glutinosus (Byrd, 1937; Rankin, 1937,
1938; Parker, 1941; Fischthal, 1955b; Cheng, 1958;
Cheng and Chase, 1961; Brandon, 1965; Rabalais,
1970; Dyer and Brandon, 1973; Brooks, 1979);
Jordan’s salamander, Plethodon jordani (Rankin,
1937, 1938; Dyer, 1983); western red-backed sala-
mander, Plethodon vehiculum (Panitz, 1969); Yo-
nahlossee salamander, Plethodon yonahlossee
(Rankin, 1937); red salamander, Pseudotriton ruber
(Rankin, 1937, 1938; Parker, 1941; Catalano et al.,
332 COMPARATIVE PARASITOLOGY, 74(2), JULY 2007
1982); rough-skinned newt, Taricha granulosa
(Lehmann, 1954; Pratt and McCauley, 1961; Weath-
erly and Canaris, 1961; Moravec, 1984); Texas alli-
gator lizard, Gerrhonotus liocephalus (¼Gerrhonotus
ophiurus; Goldberg et al., 1999); eastern glass
lizard, Ophisaurus ventralis (Byrd, 1937); little brown
skink, Scincella lateralis, (Harwood, 1932; Byrd, 1937);
rough green snake, Opheodrys aestivus (Nicoll, 1911);
DeKay’s brown snake, Storeria dekayi (Harwood,
1932); eastern box turtle, Terrapene carolina (Rausch,
1947; Stuart and Miller, 1993).
Palaearctic Realm: common European toad, Bufo
bufo (Lühe, 1909; André, 1912, 1917; Freund, 1934);
European common frog, Rana temporaria (Dujardin,
1845; Freund, 1934); Japanese firebelly newt,
Cynops pyrrhoghaster (Rankin, 1938); Corsican
Mountain salamander, Euproctus montanus (Combes
and Knoepffler, 1968; Timon-David and Giudicelli,
1969); Caucasian salamander, Mertensiella caucasica
(Yildirimhan et al., 2005); alpine salamander, Sala-
mandra atra (Frölich, 1789; Stossich, 1889);
European fire salamander, Salamandra salamandra
(Rudolphi, 1819; Dujardin, 1845; Creplin, 1846;
Stossich, 1889; Lühe, 1909; André, 1912, 1917;
Lopez-Neyra, 1916; Freund, 1934; Grabda and
Grabda, 1953; Szabó, 1961; Barus et al., 1963;
Vojtková and Vojtek, 1972; Prokopic and Krivanec,
1975; Prudhoe and Bray, 1982; Bertman, 1986);
spectacled salamander, Salamandrina terdigitata
(Parona, 1896; Sonsiono, 1896); Laurenti’s alpine
newt, Triturus alpestris (Stossich, 1889; Lühe, 1909;
Freund, 1934; Callot, 1946; Buttner, 1951; Barus and
Groschaft, 1962; Vojtková and Vojtek, 1972); north-
ern crested newt, Triturus cristatus (Lühe, 1909;
Parona, 1896; Freund, 1934; Vojtková and Vojtek,
1972); palmate newt, Triturus helveticus (André,
1917; Buttner, 1951); marbled newt, Triturus mar-
moratus (Pontallié, 1852; Stossich, 1889); smooth
newt, Triturus vulgaris (Diesing, 1850; Lühe, 1909;
Freund, 1934; Barus et al., 1963; Frandsen, 1974);
slowworm, Anguis fragilis (Baylis, 1928; Callot,
1946).
Geographic range: Europe, North America, South
America (Prudhoe and Bray, 1982).
Specimens deposited: USPNC 98186; 2 slides.
Remarks: There are considerable differences of opin-
ion concerning the number of species assigned to the
genus Brachycoelium. Rankin (1938) concluded that
heavy infections produce many small flukes, and light
infections produce large individuals and reduce all
known species to B. salamandrae, the type species.
However, Parker (1941) and Cheng (1958) did not
accept the synonym and recognized 7 and 10 species,
respectively, and later, Cheng and Chase (1960) and
Couch (1966) described additional species to bring
the number of species assigned to the genus to 13.
Rabalais (1970) preferred the single species ap-
proach; McAllister et al. (1995) proposed the use
of a conservative approach suggesting that until an
exhaustive revision of the genus is completed,
B. salamandrae should represent all individuals as-
signed to the genus. Bolek and Coggins (1998) and
Goldberg et al. (1999) followed the conservative ap-
proach and assigned their specimens to B. salaman-
drae. Rankin (1938) reported the egg size range of
B. salamandrae as 39–55 lm 3 27–37 lm.
Moravec and Huffman (2000) described Brachy-
coelium longleyi from specimens fixed in situ in
endemic plethodontid salamanders collected in cen-
tral Texas and differentiated it from B. salamandrae
by egg size, 21–27 lm 3 12–15 lm in B. longleyi,
39–55 lm 3 27–37 lm in B. salamandrae. Species
assigned to the genus Langeronia, a species also
infecting amphibians, have an egg size similar to
that reported for B. longleyi: Langeronia burseyi (23–
28 lm 3 12–15 lm); Langeronia macrocirra (18–
26 lm 3 7–11 lm); Langeronia parva (14–16 lm 3
9–10 lm); Langeronia provitellaria (19–22 lm 3
10–11 lm) (data from Caballero and Bravo Hollis,
1949; Sacks, 1952; Christian, 1970; Dailey and
Goldberg, 2000). Brachycoelium and Langeronia
have similar morphological characters, but a major
difference is the position of the genital pore: median
in Brachycoelium and to the left of midline in
Langeronia. Fig. 1C of Moravec and Huffman (2000)
suggests the possibility of a nonmidline position for
the genital pore of B. longleyi. Given the small egg
size and the position of the genital pore we are hes-
itant to assign Moravec and Huffman’s specimen to
Brachycoelium, although we would agree that it is not
synonymous with B. salamandrae. We will continue
the conservative approach and assign our specimens
to B. salamandrae.
Leptodactylus martinezi represents a new host rec-
ord for B. salamandrae. Four South American hosts
are now known for B. salamandrae: L. martinezi,
L. rhodomystax, Anolis nitens, L. osvaldoi. (Freitas,
1961; this study).
Leptodactylus mystaceus
(Spix, 1824)
One specimen was collected on 24 October 2004
at 3 km W, 6.5 km N Campo de Provas Brigadeiro
 GOLDBERG ET AL.—HELMINTHS OF FROGS AND LIZARDS IN BRAZIL
Veloso, Novo Progresso Municipality, Pará, Brazil
(9817959.30S, 54856936.00W) and deposited as
OMNH 40240.
Schrankiana freitasi Baker, 1982
Prevalence and intensity: One of 1 host infected
(100%, 234).
Temporal distribution: 24 October 2004, 1 host with
234 nematodes.
Site of infection: Large intestine.
Type host and type locality: Leptodactylus pentadac-
tylus, Exu, Pernambuco, Brazil (Baker, 1982).
Additional hosts from Brazil: None.
Other reported hosts: None.
Geographical range: Brazil (Baker, 1982).
Specimen deposited: USNPC 98190; 1 vial.
Remarks: Leptodactylus mystaceus represents a new
host record for Schrankiana freitasi.
Leptodactylus rhodomystax
Boulenger, 1884
Five specimens were collected from 28 October to
7 November 2004 at 46.9 km W Campo de Provas
Brigadeiro Veloso, Novo Progresso Municipality,
Pará, Brazil (9822950.10S, 55820925.70W) and de-
posited as OMNH 40241–40245.
Brachycoelium salamandrae (Frölich, 1789)
Prevalence and intensity: One of 5 hosts infected
(20%, 1).
Temporal distribution: 29 October, 2004, 1 host
with 1.
Site of infection: Small intestine.
Specimens deposited: USNPC 98191, 1 host with 1.
Remarks: General information and additional remarks
are reported under L. martinezi. Leptodactylus
rhodomystax represents a new host record for B.
salamandrae.
Capillaria recondita Freitas and Lent, 1942
Prevalence and intensity: One of 5 hosts infected
(20%, 4).
Temporal distribution: 29 October, 2004, 1 host
with 4.
333
Site of infection: Small intestine.
Type host and type locality: Brazilian spiny-thumbed
frog, Crossodactylus gaudichaudii, Brazil (Freitas
and Lent, 1942).
Additional hosts from Brazil: None.
Other reported hosts: None.
Geographic range: Brazil (Freitas and Lent, 1942).
Specimens deposited: USNPC 98192; 1 vial.
Remarks: Leptodactylus rhodomystax represents a
new host record for Capillaria recondita.
Falcaustra maculata (Rudolphi, 1819)
Freitas and Lent, 1941
(Syn. Ascaris mascula Rudolphi, 1819; Floren-
cioia nitida Travassos, 1920).
Prevalence, mean intensity, and range: Three of 5
hosts infected (60%, 3.7 6 3.8 SD, 1–8).
Temporal distribution: 28, 29 October, 2004, 3 hosts
with 1, 2, 8, respectively.
Site of infection: Small intestine.
Type host and type locality: Rio tropical racer,
Mastigodryas bifossatus (¼Coluber lichtensteini),
Brazil (Rudolphi, 1819).
Additional hosts from Brazil: yellow Cucuru toad,
Chaunus ictericus (¼Bufo ictericus; Rodrigues et al.,
1982); Crossodactylus gaudichaudii (Gomes and
Vicente, 1966); blacksmith gladiator frog, Hypiboas
faber (¼Hyla faber; Freitas and Lent, 1941); Criolla
frog, Leptodactylus ocellatus (¼Leptodactylus caligi-
nosus; Freitas and Lent, 1941; Vicente and Santos,
1976; Fabio, 1982; Rodrigues et al., 1982); Lep-
todactylus pentadactylus (Freitas, 1955; Guimaraes
et al., 1976; Rodrigues et al., 1982); Santa Catarina
frog, Hylodes nasus (¼Elosia nasus; Freitas and Lent,
1941); Wied’s groundsnake, Liophis poecilogyrus
(¼Leimadophis poecilogyrus; Freitas, 1955).
Other reported hosts: Schneider’s toad, Chaunus
schneideri (¼Bufo paracnemis; Lent et al., 1946).
Geographic range: Brazil (Rudolphi, 1819), Para-
guay (Lent et al., 1946).
Specimens deposited: USNPC 98193; 1 vial.
Remarks: Leptodactylus rhodomystax represents
a new host record for Falcaustra maculata.
334 COMPARATIVE PARASITOLOGY, 74(2), JULY 2007
Schrankiana schranki (Travassos, 1925)
Strand, 1942
(Syn. Schrankia schranki Travassos, 1925).
Prevalence, mean intensity, and range: Two of 5
hosts infected (40%, 65.5 6 67.2 SD, 16–111).
Temporal distribution: 7 November 2004, 2 hosts
with 16, 111, respectively.
Site of infection: Large intestine.
Type host and type locality: Leptodactylus pentadac-
tylus, Brazil (Travassos, 1925)
Additional hosts from Brazil: Leptodactylus labyrin-
thicus (Fahel, 1952).
Other reported hosts: Leptodactylus mystaceus
(Dyer, 1990).
Geographic range: Brazil (Travassos, 1925), Ecua-
dor (Dyer and Altig, 1977).
Specimens deposited: USNPC 98194; 1 vial.
Remarks: Leptodactylus rhodomystax represents
a new host record for Schrankiana schranki.
Teiidae
Kentropyx calcarata Spix, 1825
Seventeen specimens were collected 3–12 Novem-
ber 2004 at 46.9 km W Campo de Provas Brigadeiro
Veloso, Novo Progresso Municipality, Pará, Brazil
(9822950.10S, 55820925.70W) and deposited as
OMNH 41812–41828.
Kentropyxia sauria Baker, 1982
Prevalence, mean intensity, and range: Seven of 17
hosts infected (41%, 4.9 6 3.3 SD, 1–11).
Temporal distribution: 3, 4, 6, 8, 11, 12 November,
2004, 7 hosts with 1, 3, 3, 4, 5, 7, 11, respectively.
Site of infection: Small intestine.
Type host and type locality: Kentropyx calcarata,
Belem, Brazil (Baker, 1982).
Additional hosts from Brazil: None.
Other reported hosts: None.
Geographic range: Brazil (Baker, 1982).
Specimens deposited: USNPC 98198; 1 vial.
Remarks: This is only the second report of Kentro-
pyxia sauria.
Physaloptera retusa Rudolphi, 1819
(Syn. Spiroptera retusa [Rudolphi, 1819] Dujar-
din, 1845; Physaloptera mucronata Leidy, 1856;
Physaloptera largada Sprehn, 1932).
Prevalence, mean intensity, and range: Two of 17
hosts infected (12%, 1.5 6 0.71 SD, 1–2).
Temporal distribution: 7 November 2004, 2 hosts
with 1, 2, respectively.
Site of infection: Stomach.
Type host and type locality: golden tegu, Tupinambis
teguixin, Brazil (Rudolphi, 1819).
Additional hosts from Brazil: common lesser toad,
Chaunus granulosus (¼Bufo granulosus; Gonçalves
et al., 2002); South American leaf toad, Rhinella
margaritifer (¼Bufo typhonius; Gonçalves et al.,
2002); giant ameiva, Ameiva ameiva (Poinar and
Vaucher, 1972; Cristofaro et al., 1976; Ribas, Rocha
et al., 1998); red worm lizard, Amphisbaena alba,
(Molin, 1860); sand dune lizard, Cnemidophorus
abaetensis (Dias et al., 2005); green tailed lizard,
Cnemidophorus littoralis (Vrcibradic et al., 2000); no
common name, Cnemidophorus nativo (Menezes
et al., 2004); Spix’s whiptail, Cnemidophorus
ocellifer (Ribas et al., 1995); Lutz’s tree iguana,
Liolaemus lutzae (Rocha, 1995); agile mabuya,
Mabuya agilis (Ribas, Teixeira-Filho et al., 1998);
two-striped mabuya, Mabuya bistriata (recorded as
Scincus spixii by Molin, 1860); Paraguay mabuya,
Mabuya dorsivittata (Rocha et al., 2003); spiny lava
lizard, Tropidurus spinulosus (Vicente, 1981); Am-
azon lava lizard, Tropidurus torquatus (Vicente and
Santos, 1967; Cristofaro et al., 1976; Vicente, 1981;
Ribas, Rocha et al., 1998; Vrcibradic et al., 2000);
golden tegu, Tupinambis teguixin (¼Tupinambis
nigropunctatus; Diesing, 1851; Rudolphi, 1819;
Molin, 1860; Ortlepp, 1922).
Other reported hosts: rainbow ameiva, Ameiva
undulata (Caballero, 1951); spotted anole, Anolis
punctatus (Bursey et al., 2005); canyon spotted
whiptail, Aspidoscelis burti (¼Cnemidophorus burti;
Goldberg and Bursey, 1989a); imbricate alligator
lizard, Barisia imbricata (Goldberg et al., 1999);
zebra-tailed lizard, Callisaurus draconoides (Telford,
1970); rainbow whiptail, Cnemidophorus lemniscatus
(Caballero and Vogelsang, 1947; Diaz-Ungria, 1964;
Diaz-Ungria and Gallardo, 1968); Laurent’s whiptail,
Cnemidophorus murinus (Specian and Whittaker,
1980); southern alligator lizard, Elgaria multicari-
nata (Telford, 1970); San Lucan alligator lizard,
 GOLDBERG ET AL.—HELMINTHS OF FROGS AND LIZARDS IN BRAZIL
Elgaria paucicarinata (Goldberg, Bursey and
Beaman, 2004); western skink, Pleistodon skiltonia-
nus (¼Eumeces skiltonianus), (Telford, 1970); long-
nosed leopard lizard, Gambelia wislizenii (Telford,
1970); Gerrhonotus liocephalus (¼Gerrhonotus
ophiurus; Goldberg et al., 1999); common green
iguana, Iguana iguana (Diaz-Ungria and Gallardo,
1968); Amazon kentropyx, Kentropyx altamazonica
(Bursey et al., 2005); forest kentropyx, Kentropyx
pelviceps (Bursey et al., 2005); Bell’s anole,
Leiosaurus bellii (Goldberg et al., 2004); no common
name, Leiosaurus catamarcensis (Goldberg et al.,
2004); no common name, Liolaemus neuquensis
(Goldberg et al., 2004); copper anole, Norops
cupreus (Goldberg and Bursey, 2004); tree runner,
Plica plica (Bursey et al., 2005); harlequin race-
runner, Plica umbra (Bursey et al., 2005); Bocourt’s
spiny lizard, Sceloporus acanthinus (Caballero,
1951); Clark’s spiny lizard, Sceloporus clarkii
(Goldberg et al., 1994); blue spiny lizard, Sceloporus
cyanogenys (Goldberg et al., 1995); emerald spiny
lizard, Sceloporus formosus (Goldberg et al., 2003);
common sagebrush lizard, Sceloporus graciosus
(Woodbury, 1934), mesquite lizard, Sceloporus
grammicus (Goldberg et al., 2003); Yarrow’s spiny
lizard, Sceloporus jarrovii (Goldberg and Bursey,
1990; Bursey and Goldberg, 1991, 1994; Goldberg
et al., 1995; Goldberg et al., 1996); desert spiny
lizard, Sceloporus magister (Pearce and Tanner,
1973; Walker and Matthias, 1973; Goldberg et al.,
1994; Goldberg et al., 1995); canyon lizard, Scelo-
porus merriami (Goldberg et al., 1995); southern
crevice spiny lizard, Sceloporus mucronatus (Gold-
berg et al., 2003); Sceloporus occidentalis (Grund-
mann, 1959; Telford, 1970; Pearce and Tanner, 1973;
Walker and Matthias, 1973; Goldberg et al., 1998);
Texas spiny lizard, Sceloporus olivaceus (Goldberg
et al., 1995); granite spiny lizard, Sceloporus orcutti
(Telford, 1970); blue-bellied lizard, Sceloporus
parvus (Goldberg et al., 2003); crevice spiny lizard,
Sceloporus poinsettii (Goldberg et al., 1993; Gold-
berg et al., 1995); blue spiny lizard, Sceloporus
serrifer (Goldberg et al., 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 et al., 1995); southern sagebrush lizard,
Sceloporus vandenburgianus (¼Sceloporus gracio-
sus; Goldberg and Bursey, 1989b; Goldberg et al.,
1997); rose-bellied lizard, Sceloporus variabilis
(Goldberg et al., 1995; Goldberg et al., 2003); striped
plateau lizard, Sceloporus virgatus (Goldberg et al.,
1994); rose whorl-tailed iguana, Stenocercus rose-
335
iventris (Bursey et al., 2005); turnip-tailed gecko,
Thecadactylus rapicauda (Bursey et al., 2005); black
lava lizard, Tropidurus melanopleurus (Roca, 1997);
red tegu, Tupinambis rufescens (Sprehn, 1932); com-
mon side-blotched lizard, Uta stansburiana (Telford,
1970); eastern hog-nosed snake, Heterodon platirhi-
nos (Ortlepp, 1922); American alligator, Alligator
mississippiensis (Walton, 1927).
Geographic range: Argentina, Brazil, Mexico, United
States, Venezuela, West Indies (Baker 1987); Peru
(Bursey et al., 2005).
Specimens deposited: USNPC 98199; 1 vial.
Remarks: Kentropyx calcarata represents a new host
record for P. retusa.
Gymnopthalmidae
Leposoma osvaldoi Avila-Pires, 1995
Five specimens were collected from 2–6 Novem-
ber 2004 at 46.9 km W Campo de Provas Brigadeiro
Veloso, Novo Progresso Municipality, Pará, Brazil,
(9822950.10S, 55820925.70W) and deposited as
OMNH 41787–41791.
Brachycoelium salamandrae (Frölich, 1789)
Prevalence and intensity: One of 5 hosts infected
(20%, 1).
Temporal distribution: 5 November 2004, 1 host
with 1.
Site of infection: Small intestine.
Specimens deposited: USNPC 98195; 1 slide.
Remarks: General information and additional remarks
are reported under L. martinezi. Leposoma osvaldoi
represents a new host record for B. salamandrae.
Potamites ecpleopus
(Cope 1876)
Thirteen specimens were collected on 23 October
2004 at Cachoeira do Curúa, Altamira Municipality,
Pará, Brazil (884498.20S, 54857949.00W). Lizards
were deposited in OMNH as 41792–41804.
Falcaustra belemensis Baker and Bain, 1981
Prevalence, mean intensity, and range: Eight of 13
hosts infected (62%, 8.968.2 SD, 2–26).
Temporal distribution: 23 October, 2004, 8 hosts
with 2, 3, 4, 5, 6, 9, 16, 26. respectively.
336 COMPARATIVE PARASITOLOGY, 74(2), JULY 2007
Site of infection: Small intestine.
Type host and type locality: Two-faced neusticurus,
Neusticurus bicarinatus (Baker and Bain, 1981).
Additional hosts from Brazil: None.
Other reported hosts: None.
Geographic range: Brazil (Baker and Bain, 1981).
Specimens deposited: USNPC 98196; 1 vial.
Remarks: Potamites ecpleopus represents a new host
record for Falcaustra belemensis.
Physaloptera retusa Rudolphi, 1819
Prevalence and intensity: One of 13 hosts infected
(7%, 1).
Temporal distribution: 23 October 2004, 1 host with 1.
Site of infection: Stomach.
Specimens deposited: USNPC 98197; 1 vial.
Remarks: General information and additional remarks
are reported under K. calcarata. Potamites ecpleopus
represents a new host record for P. retusa.
DISCUSSION
A total of 709 helminths was collected: 599 from
21 (49%) of 43 frogs and 110 from 17 (49%) of 35
lizards. Of these, 2 nematodes (0.2%) from frogs
were juveniles not capable of reaching maturity in
herptiles. There were 14 helminth species represented
in the sample, but no individual host harbored more
than 3 species. Only one helminth species, B.
salamandrae, occurred in both frogs and lizards. Of
the infected frogs, 11 (50%) harbored 1 species of
helminth; 9 (45%) harbored 2 species; and 1 (5%)
harbored 3 species. There were 1.53 6 0.13 (x 6 1
SE) helminth species/infected frogs and 28.52 6 11.7
helminth individuals/infected frogs. No frog species
harbored more than 4 helminth species: 3 frog species
harbored 1 helminth species; 1 frog species harbored
2 helminth species; 1 frog species harbored 3
helminth species; and one frog species harbored 4
helminth species. There were 2.0 6 0.52 helminth
species/frog host species. Of the infected lizards, 15
(88%) harbored 1 species of helminth; 2 (12%)
harbored 2 species. There were 1.12 6 0.08 helminth
species/infected lizard and 6.47 6 1.58 helminth
individuals/infected lizard. Aho (1990) compiled
distributional patterns for frog and lizard helminths
in general and reported the mean 6 1 SE total
number of helminth species per host frog species as
3.54 6 0.24 (range 0–9) and per lizard species as
2.06 6 0.13 (range 0–5). The values reported here
are within the ranges reported by Aho (1990),
although the mean infective species for frogs is lower
than that reported. This may be a reflection of
regional differences in frog infection rates. The values
for lizards are similar to the values of Aho (1990).
Of the 14 species of helminths only B. salaman-
drae infected both frog and lizard species, L. fuscus,
L. rhodomystax, and L. osvaldoi. The remaining 13
helminth species, with the exception of P. retusa,
were found to infect a single host species.
ACKNOWLEDGMENTS
We thank Jessica Carlson, Cassie Ho, and Sean
Kark for assistance with dissections. Reptile and
amphibian specimens were collected as part of
a collaborative project between the Sam Noble
Oklahoma Museum of Natural History and the
University of Brası́lia (through Dr. Guarino R. Colli)
approved by the Ministério de Ciência e Tecnologia
(Portaria No. 649) and the Instituto Brasileiro do
Meio Ambiente e dos Recorsos Naturaı̂s Renováveis
(IBAMA, Permit No. 0217/2004-CGFAU/LIC).
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