BioOne sees sustainable scholarly publishing as an inherently collaborative enterprise connecting authors, nonprofit publishers, academic institutions, research libraries, and research funders in the common goal of maximizing access to critical research. Gastrointestinal Helminths from 13 Species of Lizards from Reserva Cuzco Amazónico, Peru Author(s): Charles R. Bursey, Stephen R. Goldberg, Jeffrey R. Parmelee Source: Comparative Parasitology, 72(1):50-68. 2005. Published By: The Helminthological Society of Washington DOI: http://dx.doi.org/10.1654/4132 URL: http://www.bioone.org/doi/full/10.1654/4132 BioOne (www.bioone.org ) is a nonprofit, online aggregation of core research in the biological, ecological, and environmental sciences. BioOne provides a sustainable online platform for over 170 journals and books published by nonprofit societies, associations, museums, institutions, and presses. Your use of this PDF, the BioOne Web site, and all posted and associated content indicates your acceptance of BioOne’s Terms of Use, available at www.bioone.org/page/ terms_of_use . Usage of BioOne content is strictly limited to personal, educational, and non-commercial use. Commercial inquiries or rights and permissions requests should be directed to the individual publisher as copyright holder.
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Gastrointestinal Helminths from 13 Species of Lizards from Reserva Cuzco Amazónico, Peru
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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.
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
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);
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
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
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.
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FIRST ANNOUNCEMENT OF JOINT MEETINGS
The Helminthological Society of WashingtonSoutheastern Society of Parasitologists
Animal Disease Research Workers in the Southern StatesSouthern Conference on Animal Parasites
Southern Conference of Researchers in Aquatic Diseases