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Turtles of the Tiputini Biodiversity Station with remarks on the diversity and distribution of the Testudines from Ecuador

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Ten species of Testudines, from four different families: Chelidae, Kinosternidae, Podocnemididae, and Testudinidae,are reported from the Tiputini Biodiversity Station (Universidad San Francisco de Quito), northeastern Amazonian Ecuador;including the second report from Ecuador of Batrachemys heliostemma. These ten species are sympatric but not completelysyntopic, thus I comment on their diversity and habitat preferences patterns. The general distribution of some turtles fromAmazonian Ecuador is discussed, providing additional records and distribution maps for Chelus fimbriatus, Platemysplatycephala, Batrachemys heliostemma, Batrachemys raniceps, Mesoclemmys gibba, Phrynops geoffroanus, Kinosternonscorpioides, and Geochelone denticulata. The presence of Peltocephalus dumerilianus in Amazonian Ecuador is con-firmed; based on re-examination of specimens previously reported by Dr. Gustavo Orcés in 1949. The total Ecuadorianrichness of Testudines species includes 31 living taxa, and I provide an annotated list of the turtles and tortoises of mainlandand insular Ecuador. Much information is still needed in order to understand the ecological relationships of the Testudinesfrom Ecuador and future research should focus on long-term studies designed to answer several questions about theirbiology and ecology. Several threats are currently jeopardizing the long-term conservation of turtle populations in Ecuador,and we must increase our knowledge on these amazing animals in order to protect and preserve them.
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Turtles of the Tiputini Biodiversity Station with remarks on the diversity and
distribution of the Testudines from Ecuador
1
College of Biological and Environmental Sciences, Universidad San Francisco de Quito, Casilla Postal 17-12-841 Quito,
Ecuador. E-mail: diegofrancisco_cisneros@yahoo.com
Abstract
Cisneros-Heredia, D.F. Turtles of the Tiputini Biodiversity Station with remarks on the diversity and distribution of the
Testudines from Ecuador. Biota Neotrop. Jan/Abr 2006, vol. 6, no. 1 http://www.biotaneotropica.org.br/v6n1/pt/
abstract?inventory+bn00906012006. ISSN 1676-0611
Ten species of Testudines, from four different families: Chelidae, Kinosternidae, Podocnemididae, and Testudinidae,
are reported from the Tiputini Biodiversity Station (Universidad San Francisco de Quito), northeastern Amazonian Ecuador;
including the second report from Ecuador of Batrachemys heliostemma. These ten species are sympatric but not completely
syntopic, thus I comment on their diversity and habitat preferences patterns. The general distribution of some turtles from
Amazonian Ecuador is discussed, providing additional records and distribution maps for Chelus fimbriatus, Platemys
platycephala, Batrachemys heliostemma, Batrachemys raniceps, Mesoclemmys gibba, Phrynops geoffroanus, Kinosternon
scorpioides, and Geochelone denticulata. The presence of Peltocephalus dumerilianus in Amazonian Ecuador is con-
firmed; based on re-examination of specimens previously reported by Dr. Gustavo Orcés in 1949. The total Ecuadorian
richness of Testudines species includes 31 living taxa, and I provide an annotated list of the turtles and tortoises of mainland
and insular Ecuador. Much information is still needed in order to understand the ecological relationships of the Testudines
from Ecuador and future research should focus on long-term studies designed to answer several questions about their
biology and ecology. Several threats are currently jeopardizing the long-term conservation of turtle populations in Ecuador,
and we must increase our knowledge on these amazing animals in order to protect and preserve them.
Key words: Reptilia, Testudines, Chelidae, Kinosternidae, Podocnemididae, Testudinidae, Chelus fimbriatus, Platemys
platycephala, Batrachemys heliostemma, Batrachemys raniceps, Mesoclemmys gibba, Phrynops geoffroanus, Kinosternon
scorpioides, Geochelone denticulata, Podocnemis expansa, Podocnemis unifilis, Peltocephalus dumerilianus, diversity,
habitat preferences, distribution, checklist, Amazonia, Ecuador
Resumen
Cisneros-Heredia, D.F. Las Tortugas de la Estación de Biodiversidad Tiputini con notas sobre la distribución de alginos
Testudines de Ecuador. Biota Neotrop. Jan/Abr 2006, vol. 6, no. 1http://www.biotaneotropica.org.br/v6n1/pt/
abstract?inventory+bn00906012006. ISSN 1676-0611
Diez especies de Testudines, de cuatro familias diferentes: Chelidae, Kinosternidae, Podocnemididae, y Testudinidae,
son reportadas de la Estación de Biodiversidad Tiputini (Universidad San Francisco de Quito), en la Amazonía Nororiental
de Ecuador; incluyendo el segundo reporte para Ecuador de Batrachemys heliostemma. Estas diez especies son simpátricas
pero no completamente sintópicas, por lo que comento sobre sus patrones de diversidad y preferencias de hábitat. La
distribución general de algunas especies de tortugas de la Amazonía de Ecuador es discutida, proveyendo registros
adicionales y mapas de distribución para Chelus fimbriatus, Platemys platycephala, Batrachemys heliostemma, Batrachemys
raniceps, Mesoclemmys gibba, Phrynops geoffroanus, Kinosternon scorpioides, y Geochelone denticulata. La presencia
de Peltocephalus dumerilianus en la Amazonía de Ecuador es confirmada; basada en la reexaminación de especímenes
previamente reportados por el Dr. Gustavo Orcés en 1949. La riqueza total de especies de tortugas en Ecuador incluye 31 taxa
vivientes; proveo una lista anotada de las especies de tortugas de Ecuador continental e insular. Mucha información es aún
requerida para poder comprender las relaciones ecológicas de los Testudines de Ecuador e investigaciones futuras deberían
enfocarse en estudios a largo plazo diseñados para responder a varias preguntas sobre su biología y ecología. Algunos
peligros amenazan la conservación a largo plazo de las poblaciones de tortugas de Ecuador y debemos incrementar nuestro
conocimiento sobre estos sorprendentes animales para poder protegerlos y conservarlos.
Palabras-clave:Reptilia, Testudines, Chelidae, Kinosternidae, Podocnemididae, Testudinidae, Chelus fimbriatus,
Platemys platycephala, Batrachemys heliostemma, Batrachemys raniceps, Mesoclemmys gibba, Phrynops geoffroanus,
Kinosternon scorpioides, Geochelone denticulata, Podocnemis expansa, Podocnemis unifilis, Peltocephalus
dumerilianus, diversidad, preferencias de hábitat, distribución, lista, Amazonia, Ecuador
Biota Neotropica v6 (n1) –http://www.biotaneotropica.org.br/v6n1/pt/abstract?inventory+bn00906012006
Diego F. Cisneros-Heredia
1
Date Received 05/23/2005 - Revised 10/18/2005 - Accepted 01/01/2006
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Cisneros-Heredia, D.F. - Biota Neotropica, v6 (n1) - BN00906012006
Introduction
Thirty taxa of living Testudines have been recog-
nized for Ecuador (Miyata 1982, Pritchard & Trebbau
1984, Almendáriz 1991, Coloma et al. 2000-2004); four
species are marine turtles, 11 taxa correspond to the en-
demic giant tortoises of the Galapagos Archipelago
(McFarland et al. 1974), and 15 species occur in main-
land Ecuador. Five mainland taxa are restricted to the
Pacific lowlands of Ecuador, including one member of
the family Kinosternidae (Kinosternon leucostomum
Duméril & Bibron) and all Ecuadorian members of the
families Chelydridae (1 sp.: Chelydra serpentina
[Linnaeus]) and Geoemydidae (3 spp.: Rhinoclemmys
annulata [Gray], R. melanosterna [Gray], and R. nasuta
[Boulenger]). The greatest diversity of Testudines in
mainland Ecuador occurs in the Amazonian lowlands;
unfortunately little information is available on most
turtles from mainland Ecuador, and although most Ama-
zonian taxa are conceived as widespread, little data ex-
ists on its distribution and ecology. The objective of
this paper is to make an initial approach oT the testudines
diversity in Ecuador, with some remarks on the distribu-
tion of the Testudines from Amazonian Ecuador, compil-
ing every record from Amazonian Ecuador, presenting
some specific data on the diversity and habitat prefer-
ences of the turtles and tortoise from the Tiputini
Biodiversity Station; and an annotated list of the turtles
and tortoises of mainland and insular Ecuador.
Materials and Methods
Field work was done at the Tiputini Biodiversity
Station, (TBS - 00°37’05" S, 76°10’19" W, 190 – 270 m
a.s.l.), a research station located in the province of
Orellana, Republic of Ecuador, ca. 280 km ESE from Quito;
in the northern bank of the Tiputini River, part of the
Napo and Amazonas Rivers Basins. TBS was established
in 1995 by the Universidad San Francisco de Quito
(Ecuador), in cooperation with Boston University (USA),
as a center of education, research and conservation. TBS
is adjacent to the Yasuní National Park and it is part of
the Yasuní Biosphere Reserve.
TBS preserves a tract of 650 hectares which mostly
includes primary Terra Firme forest (Lowland Evergreen
Forest), but there is a rather narrow belt of flooded vegeta-
tion of Varzea (Lowland Evergreen Forest flooded by white-
waters) and Igapó (Lowland Evergreen Forest flooded by
black-waters) towards the river, streams, and around an small
oxbow lake. Along the Tiputini River, several beaches are
uncovered during the dry season, but all are short, never
greater than 100 m.
Herpetological work at TBS was conducted be-
tween 1997 and 2001, with 11 human/months. Turtles
were surveyed by day and night with through
encounters technique (Heyer et al. 1994), especially on the
wetlands (swamps, rivulets, streams, oxbow lake, and river).
Some specimens were examined and later released to confirm
their identities. Individuals not captured (especially those
observed while basking over logs in the lake and river) were
examined using 10x42 Swarovski binoculars in order to deter-
mine its identification. Additional specimens of Ecuadorian
Testudines were examined from the collection of the Fundación
Herpetológica G. Orcés, Quito (FHGO).
Results and Discussion
The family Chelidae is represented at TBS by six spe-
cies, including Chelus fimbriatus (Schneider), Platemys
platycephala (Schneider), Batrachemys heliostemma
McCord, Ouni & Lamar, Batrachemys raniceps (Gray),
Mesoclemmys gibba (Schweigger), and Phrynops
geoffroanus (Schweigger). The last four species correspond
to the commonly named “toadhead” turtles, all previously
classified under the genus Phrynops until the recent re-
evaluation by McCord et al. (2001).
Batrachemys heliostemma was recorded twice at TBS;
one juvenile was observed in August 1999 and a second
juvenile was found on February 2001, both at a shallow
swamp located behind the oxbow lake of TBS by day. Both
individuals showed the yellow facial marks characteristic of
the species. The second individual was caught, examined
and released; it had a carapace length of 121 mm, maximum
plastral length of 100 mm, and carapace width of 92 mm. The
habitat where both individuals were observed coincides with
descriptions provided by McCord et al. (2001) for Peruvian
populations, and supports their hypothesis about the ap-
parent preference of B. heliostemma for high non-flooded
areas. Batrachemys heliostemma is a recently described
species known from the upper Amazon Basin with records
in Venezuela, Colombia, Ecuador, Brazil, and Peru (McCord
et al. 2001). This species was known in Amazonian Ecuador
from only one locality, “Mariam” in the province of
Sucumbíos, Ecuador, collected in October 1983 (McCord et
al. 2001). TBS is the second locality for the species in Ecua-
dor, and the westernmost record of the species (Figure 1).
Batrachemys raniceps was recorded twice at TBS,
both on the border of the small drainage connecting the
oxbow lake and the Tiputini River by day (both exam-
ined and later released). One individual of Mesoclemmys
gibba was found at TBS while walking ca. 10 m away
from the Tiputini River, on an ephemeral marsh created
by a recent flooding; it released a strong-smelling musk
when captured (later released). Phrynops geoffroanus
was the most common toadhead turtle recorded at TBS;
several individuals were observed basking over logs
partially submerged at the mouth of a small affluent of
the Tiputini River, and along the river itself. In the ox-
bow lake, at least two individuals were seen partially
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Figure 1. Known records of Batrachemys heliostemma (squares) and Batrachemys raniceps (circles) in the Republic of Ecuador. TBS =
Tiputini Biodiversity Station, closed symbols = material studied, open symbols = literature records (see text), numbers correspond to the
following provinces in Amazonian Ecuador: 1 = Sucumbíos, 2 = Napo, 3 = Orellana, 4 = Pastaza, 5 = Morona-Santiago, 6 = Zamora-
Chinchipe.
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Figure 2. Known records of Chelus fimbriatus (squares) and Kinosternon scorpioides (circles) in the Republic of Ecuador. For symbols
equivalence see Figure 1.
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Figure 3. Known records of Mesoclemmys gibba (circles) in the Republic of Ecuador. For symbols equivalence see Figure 1.
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Figure 4. Known records of Phrynops geoffroanus (circles) in the Republic of Ecuador. For symbols equivalence see Figure 1.
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Figure 5. Known records of Platemys platycephala (circles) in the Republic of Ecuador. For symbols equivalence see Figure 1.
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Figure 6. Known records of Geochelone denticulata (circles) in the Republic of Ecuador. For symbols equivalence see Figure 1.
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immersed in a marsh area on the border of the lake, and
at least three were usually observed while basking over
logs partially submerged. One individual of P.
geoffroanus released a strong-smelling musk when
captured (later released).
Platemys platycephala was recorded in five occa-
sions, always walking in non-flooded and seasonally flooded
forest by day. At non-flooded forest, it was usually near
shallow streams. One individual was found in 09 August
1999 during day among the leaf-litter near to a small rivulet
in primary non-flooded forest; it was examined and released,
having a carapace length of 117 mm, maximum plastral length
of 109 mm, carapace width of 80 mm, head length of 41 mm,
and weight of 139.5 g. Individuals observed coincides well
with the description of the subspecies Platemys
platycephala melanonota by Ernst (1983). Chelus
fimbriatus was observed once at TBS during late afternoon
on August 2000, moving in a shallow swamp located behind
the oxbow lake of TBS.
Few localities for members of the family Chelidae
in Ecuador have been mentioned in the literature, so in
addition to the TBS records, I report herein some new
localities. One specimen of Batrachemys raniceps was
collected at Shushufindi (00º12’43" S, 76º39’29" W, 220
m), province of Sucumbíos, by J.-M. Touzet on 02 Octo-
ber 1996 (FHGO 1481). Batrachemys raniceps was first
reported in Ecuador by Miyata (1982) without mention-
ing a locality, and I could not locate additional published
records for the species in Ecuador; thus the species is
apparently reported in Ecuador just from the provinces
of Sucumbíos and Orellana (Figure 1). Two specimens of
Chelus fimbratus were collected at Laguna Grande,
Cuyabeno Reserve (00º05’ S,76º10’ W, 220 m), province
of Sucumbíos, by F. Campos on 25 December 1989 (FHGO
206) and 25 February 1990 (FHGO 207). Chelus fimbriatus
was reported by Orcés (1949) from localities in the prov-
ince of Zamora-Chinchipe and Sucumbíos; by Duellman
(1978) from the province of Sucumbíos, and Iverson (1992)
Figure 7. Schematic map of a section of the Tiputini River (T.R.) at the Tiputini Biodiversity Station (station laboratory = star), province of
Orellana, Ecuador; indicating habitat preferences by ten chelonian species. O.L. = oxbow lake; dotted area next to oxbow lake = non-
seasonally flooded shallow swamp; gray shadow areas = seasonally flooded forest. B.h. = Batrachemys heliostemma: B.r. = Batrachemys
raniceps, C.f. = Chelus fimbratus, G.d. = Geochelone denticulata, K.s. = Kinosternon scorpioides, M.g. = Mesoclemmys gibba, P.g. =
Phrynops gibbus, P.p, = Platemys platycephala, P.e. = Podocnemis expansa, P.u. = Podocnemis unifilis.
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plotted a locality in the province of Pastaza; therefore
the distribution of C. fimbratus apparently comprises
the entire Amazonian Ecuador, with confirmed records
from the provinces of Sucumbíos, Orellana, Pastaza, and
Zamora-Chinchipe (Figure 2). One specimen of
Mesoclemmys gibba was collected at Shushufindi
(00º12’43" S, 76º39’29" W, 220 m), province of Sucumbíos,
by J.-M. Touzet on 20 September 1987 (FHGO-live 037),
and one individual was observed at the Laguna Grande,
Cuyabeno Reserve (00º05’ S,76º10’ W, 220 m), province
of Sucumbíos, by the author on August 2000.
Messoclemys gibba has been reported from the prov-
ince of Sucumbíos (Duellman 1978, Pritchard & Trebbau
1984), from the province of Orellana and Pastaza
(Pritchard & Trebbau 1984, Iverson 1992), and more re-
cently from the province of Napo (Deforce et al. 2004).
Therefore, Messoclemys gibba has confirmed records in
northern Amazonian Ecuador, in the provinces of Napo,
Sucumbíos, Orellana and Pastaza (Figure 3). One speci-
men of Phrynops geoffroanus was collected at the Hollín
River (Hollín-Loreto road), near Tena (ca. 00º41’ S, 77º40’
W), province of Napo, by M. Mamallacta on 15 July 1998
(FHGO-live 2188). Orcés (1949) cited this species as “P.
hilari” from the province of Napo (localities also men-
tioned by Pritchard & Trebbau 1984), and was reported
from the province of Sucumbíos by Duellman (1978) and
Pritchard & Trebbau (1984). Thus, Phrynopus
geoffroanus has been recorded in Ecuador from the prov-
inces of Napo, Sucumbíos, and Orellana (Figure 4). The
absence of records of M. gibba and P. geoffroanus in
southern Amazonian Ecuador could reflect the paucity
of surveys in these area rather than real absence or rar-
ity. One specimen of Platemys platycephala was col-
lected at San Pablo de Kantesiaya (00º15’00" S, 76º25’30"
W, 240 m), province of Sucumbíos, by J.-M. Touzet on
01 February 1987 (FHGO-live 122), and another was col-
lected at the Shiripuno River (sector Ñoneno) (01º05’ S,
76º50’ W), province of Pastaza, by J.-M. Touzet on 30
August 1997 (FHGO 2261). Platemys platycephala was
reported by Orcés (1949) from the province of Orellana
and Pastaza, Duellman (1978) reported the species from
the province of Sucumbíos, and Pritchard & Trebbau
(1984) mentioned records from the provinces of Orellana,
Napo, Sucumbíos, Pastaza, and Morona-Santiago
(Pritchard & Trebbau [1984] cited the locality of Macuma
(=Makuma) in the province of Zamora-Chinchipe, how-
ever it is located in the province of Morona-Santiago).
Almendáriz (1987) reported the species from the prov-
ince of Pastaza and Iverson (1992) and Ernst (1983) plot-
ted several localities, especially in southeastern Ecua-
dor. Platemys platycephala has the widest reported dis-
tribution in eastern Ecuador, with confirmed records at
every Amazonian province: Napo, Sucumbíos, Orellana,
Pastaza, Morona-Santiago and Zamora-Chinchipe (Figure 5).
The family Kinosternidae is represented at TBS by
one species, Kinosternon scorpioides (Linnaeus). One K.
scorpioides was observed among the border vegetation at
the drainage of a rivulet, tributary of the Tiputini River, and
a second turtle was captured (late released) while walking
amidst leaf-litter on the margin of the same rivulet. Orcés
(1949) reported the species from the province of Orellana,
and Duellman (1978) and Pritchard & Trebbau (1984) from
the province of Sucumbíos. Kinosternon scorpioides has
confirmed records along northern Amazonian Ecuador, in
the provinces of Sucumbíos and Orellana (Figure 2).
Two species of the family Podocnemididae occur at
TBS, Podocnemis expansa (Schweigger) and P. unifilis
Troschel. Podocnemis expansa was the rarest among them,
with just two direct observations of two big-size adults bask-
ing on a partially submerged tree at the Tiputini River, near
the beach, plus the observation of at least two nests along
the beaches of the Tiputini River on August 1999. The cara-
pace of an adult P. expansa (kept at the laboratory of the
station) had a length of 780 mm and was obtained ca. 1995
near TBS. Podocnemis unifilis was the most observed turtle
along the Tiputini River and at the oxbow lake. Groups up to
12 adult or juvenile individuals were frequently observed
basking over partially submerged logs or along vegetation
near the border of the river or lake. Several nests were ob-
served at the beaches of the Tiputini River. On one occa-
sion, up to four nests were attacked at the same beach by a
couple of Black Caracaras (Daptrius ater) and by a Tegu
Lizard (Tupinambis teguixin). Eggs of P. unifilis (and prob-
ably P. expansa) were extracted occasionally from nests by
native inhabitants, even though this practice is illicit in that
section of the Tiputini River.
The only tortoise at TBS is Geochelone denticulata
(Linnaeus). This tortoise was rather common, especially at
non-flooded primary forest, but also at flooded primary for-
est (during low flood periods). Five to eight individual
records were obtained each year between 1997 and 2001.
Individuals were found walking usually during early and
late afternoon. Most tortoises had ticks on the carapace.
Five individuals found in year 2000 (January, April and Au-
gust) were measured, showing a mean carapace length of
410.8 ± 75.0 mm (range 475.0 – 310.0 mm), mean carapace
width of 327.5 ± 85.4 mm (range 380.0 – 200.0 mm), and maxi-
mum plastral length of 287.7 ± 55.5 mm (range 336.0 – 227.0
mm). Orcés (1949) reported this species from the provinces
of Sucumbíos, Orellana, and Pastaza; Duellman (1978) re-
ported it from the province of Sucumbíos; Pritchard &
Trebbau (1984) reported G. denticulata from the provinces
of Sucumbíos and Morona-Santiago; and Almendáriz (1987)
reported it from the province of Pastaza. I have observed
this species at several localities near Tarapoa (ca. 00º08’ S,
76º24’ W, ca. 300 m), province of Sucumbíos, near Puyo (ca.
01º28’ S, 77º59’ W, ca. 950 m), province of Pastaza, and on
the Macas-Puyo road (ca. 02º19’ S, 78º07’ W), province of
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Morona-Santiago. Thus, Geochelone denticulata has been
reported from almost every province in Amazonian Ecuador
(Napo, Sucumbíos, Orellana, Pastaza and Morona-Santiago)
except for the province of Zamora-Chinchipe (Figure 6).
Although two species of the genus Podocnemis (P.
unifilis and P. expansa) are usually regarded as the only
members of the family Podocnemididae in Ecuador, Orcés
(1949) reported Peltocephalus dumerilianus (Schweigger)
from Ecuador based on five specimens from the province of
Pastaza, and one juvenile with uncertain locality. The iden-
tity of those specimens has never been confirmed, nor the
specimens examined again; and just Iverson (1992) and
Iverson & Vogt (2002) accepted those records. Miyata (1982),
Almendáriz (1991) and Coloma et al. (2000-2004) did not in-
cluded P. dumerilianus in their lists of Ecuadorian reptiles.
Two out of six specimens reported by Orcés (1949) were
located at the collection of the Colegio Mejia, Quito; they
correspond well with P. dumerilianus and confirmed the
presence of this species in the Republic of Ecuador, with
records at least in the province of Pastaza. Medem (1960)
reported P. dumerilianus from the Apaya lagoon, near
Puerto Leguizamo, on the Putumayo River, not far from the
border of Colombia with Ecuador (ca. 20 km); therefore, P.
dumerilianus could also exist in the wetlands of eastern
Sucumbíos, Ecuador. The Ecuadorian richness of Chelo-
nian species increases to 31 living taxa with the recognition
of Peltocephalus dumerilianus, with 11 species distributed
in the Amazonian lowlands. As several nomenclatural and
systematic changes have occurred affecting the classifica-
tion of Ecuadorian turtles and tortoises, I present an anno-
tated list of the turtles and tortoises from mainland and in-
sular Ecuador in Appendix 1.
The total chelonian diversity at the Tiputini
Biodiversity Station comprises ten species, from four differ-
ent families: Chelidae, Kinosternidae, Podocnemididae, and
Testudinidae. Data presented herein suggest that although
most chelonians at TBS are sympatric, they are not com-
pletely syntopic; this case is especially evident in the four
toadhead turtles, supporting hypotheses presented by
McCord et al. (2001) on habitat preferences. The shallow
swamps in non-flooded areas (behind oxbow lake) were
occupied in syntopy by Batrachemys heliostemma and
Chelus fimbriatus. Small drainages and ephemeral marshes
in flooded areas were the habitat of Batrachemys raniceps
and Mesoclemmys gibba. Non-flooded and seasonally
flooded forests, including hill areas far from the Tiputini
River, were inhabited by Platemys platycephala and
Geochelone denticulata. Open waters of the oxbow lake
were inhabited by Phrynopus geoffroanus and Podocnemis
unifilis, while at small drainages P. geoffroanus was in
syntopy with Kinosternon scorpioides; and at the river with
both members of the family Podocnemididae, Podocnemis
expansa and P. unifilis. Podocnemis expansa is an exclu-
sive inhabitant of the river (Figure 7).
However, much information is still needed in order to
understand the ecological relationships of these sympatric
Testudines. Little information is available on the life cycles
of the Amazonian species, including egg deposition sites,
nesting periods, environmental requirements for success-
ful egg development, predation effects over nests, emer-
gence period of neonates, behavior of neonates, ontoge-
nic changes in habitat, microhabitat and time use patterns,
movement patterns, maturity time for males and females,
sex ratio, survivorship and longevity (Gibbons 1990, Burke
et al. 2000). Future research at TBS and other research sta-
tions along Amazonian Ecuador should focused on long-
term studies designed to answer these biological and eco-
logical questions.
Several threats are currently jeopardizing the long-
term conservation of turtle populations along the planet.
Habitat alteration, pollution, human exploitation, diseases
and introduced species are causal factors for the declines
of several turtles (Burke et al. 2000). Podocnemis expansa
is an explicit case of the fragility of the turtle populations in
Amazonian Ecuador. The extreme and uncontrolled
overexploitation of this giant river turtle has reduced its
numbers to the point that its long-term survivorship is un-
certain. Several conservation efforts, including sustainable
harvesting, protection and management of nests, have been
established in Ecuador for the economically-relevant spe-
cies of the genus Podocnemis, but research and conserva-
tion efforts focused on every turtle taxon must be estab-
lished in Ecuador. We must increase our knowledge on
these amazing animals in order to protect and preserve them.
Acknowledgments
I am grateful to the staff of the Tiputini
Biodiversity Station, Universidad San Francisco de Quito,
especially to David Romo, Consuelo Barriga de Romo,
Kelly Swing, Mayer Rodríguez and Franklin Narváez, for
their continuous support during my field work; and to
Jean-Marc Touzet and Ana María Velasco for allowing
access to material and information at the FHGO. I thank
Ana Almendáriz, John Iverson, Stephen Karl, the Sa-
vanna River Ecology Laboratory, the Charles Darwin
Foundation, and the Department of Vertebrate Zoology,
Smithsonian Institution, for providing useful literature
or access to their libraries. I am extremely grateful to
María Elena Heredia and Laura Heredia for their con-
tinuous moral and financial support. Field work was par-
tially financed by the Tiputini Biodiversity Station,
Universidad San Francisco de Quito, lab work by the
Universidad San Francisco de Quito, and work at the
Smithsonian Institution by the 2002 Research Training
Program, National Museum of Natural History and
Smithsonian’s Women Committee.
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Cisneros-Heredia, D.F. - Biota Neotropica, v6 (n1) - BN00906012006
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Title: Turtles of the Tiputini Biodiversity Station with
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Author: Diego F. Cisneros-Heredia
Biota Neotropica, Vol. 6 ( number 1): 2006
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Date Received 05/23/2005 - Revised 10/18/2005
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ISSN 1676-0611
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APPENDIX 1.
Annotated Checklist of the living Testudines (turtles
and tortoises) from mainland and insular Ecuador.
FAMILY CHELIDAE
Notes: All species of toad-head turtles were included
in the genus Phrynops until the revision of McCord et al.
(2001) whom resurrected the genera Batrachemys,
Mesoclemmys and Rhinemys from its synonymy.
1. Batrachemys heliostemma McCord, Ouni &
Lamar, 2001
Distribution in Ecuador: Eastern Tropical (Amazonia).
Provinces of Sucumbíos and Orellana.
2. Batrachemys raniceps (Gray, 1855)
Distribution in Ecuador: Eastern Tropical (Amazonia).
Provinces of Sucumbíos and Orellana.
Notes: Previously under the synonymy of
Batrachemys (Phrynops) nasuta.
3. Chelus fimbriatus (Schneider, 1783)
Distribution in Ecuador: Eastern Tropical
(Amazonia). Provinces of Sucumbíos, Orellana, Pastaza,
and Zamora-Chinchipe.
4. Mesoclemmys gibba (Schweigger, 1812)
Distribution in Ecuador: Eastern Tropical (Amazonia).
Provinces of Napo, Sucumbíos, Orellana and Pastaza.
5. Platemys platycephala (Schneider, 1792)
Distribution in Ecuador: Eastern Tropical (Amazonia).
Provinces of Napo, Sucumbíos, Orellana, Pastaza, Morona-
Santiago, and Zamora-Chinchipe.
Notes: Ecuadorian populations are assigned to the
subspecies Platemys platycephala melanonota Ernst.
6. Phrynops geoffroanus (Schweigger, 1812)
Distribution in Ecuador: Eastern Tropical (Amazonia).
Provinces of Napo, Sucumbíos, and Orellana.
FAMILY CHELONIIDAE
7. Chelonia mydas Bocourt, 1868
Distribution in Ecuador: Pacific coast of Ecuador (in-
cluding surrounding islands) and Galápagos Archipelago.
Notes: The Ecuadorian populations have been as-
signed to the subspecies Chelonia mydas agassizii Bocourt
1868, which is considered by some authors (eg. Pritchard
1999) as a species, different from mydas: Chelonia agassizii.
However, there is still much controversy regarding the real
differentiation of populations assigned to the taxon agassizii
from those assigned to mydas, and its real taxonomic status
(see Bowen & Karl 1997, 1999, Karl & Bowen 1999, Pritchard
1999, Bowen & Karl 2000). As the black turtle agassizii is
considered “unique in some respect and is likely to be an
emerging evolutionary lineage” (Bowen & Karl 2000), I herein
maintained the subspecific status (as Chelonia mydas
agassizii) for the East Pacific populations (including Ecua-
dor), pending further studies.
8. Eretmochelys imbricata (Linnaeus, 1766)
Distribution in Ecuador: Pacific coast of Ecuador (in-
cluding surrounding islands) and Galápagos Archipelago.
Notes: The name Eretmochelys imbricata bissa
(Rüppell) (with squamata Agassiz as a synonym) has been
assigned by several authors to the Indo-Pacific populations,
including the Ecuadorian (see Pritchard and Trebbau 1984).
The intra-specific variation of Eretmochelys imbricata is
still insufficiently known, and the subspecific status has
been questioned by some authors (see Diamond 1976).
Okayama et al. (1999) found that the distribution of the
mtDNA haplotype diversity is consistent with the funda-
mental division between the Atlantic and Pacific popula-
tions, but also found significant lineage segregation within
the Indo-Pacific stock (Okayama et al., 1999). The subspe-
cies division is herein maintained until more studies pro-
vide a more complete understanding of the species varia-
tion and systematics.
9. Lepidochelys olivacea (Eschscholtz, 1829)
Distribution in Ecuador: Pacific coast of Ecuador (in-
cluding surrounding islands) and Galápagos Archipelago.
FAMILY CHELYDRIDAE
10. Chelydra serpentina (Linnaeus, 1758)
Distribution in Ecuador: Western Tropical. Provinces
of Esmeraldas, Manabí, Guayas, Los Ríos, Pichincha.
Notes: Ecuadorian populations are assigned to the
subspecies C. s. acutirostris Peters. However, the taxonomic
status of the different names under Chelydra serpentina is
still controversial. Currently four subspecies are recognized
for C. serpentina: C. s. serpentina, C. s. acutirostris, C. s.
osceola, and C. s. rossignonii. The North American taxa
(nominal subspecies and osceola) have been considered
either as synonyms or as subspecies but further studies are
needed (Phillips et al. 1996, Walker et al. 1998, Sites &
Crandall 1997). Much controversy still exists on the status
of the Central and South American populations; Phillips et
al. (1996) suggested specific status for both taxa (acutirostris
and rossignonii) based on patterns of geographic variation
in isozyme and mitochondrial DNA restriction fragment pat-
tern data; but Sites & Crandall (1997) presented an alterna-
tive interpretation of their data and concluded that species
status may not be guaranteed for Central and South Ameri-
can taxa, but that the Ecuadorian population may deserve
species status if the fixation of a unique allele at two nuclear
isozyme loci (M-Icdh and S-Icdh) is showed to be present
with more extensive sampling; however additional studies
have not been performed yet and the subspecies status is
maintained for the Ecuadorian population.
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Several specimens of Chelydra serpentina have been
found in places outside its previously reported distribution
range (ej. Los Frailes, Machalilla National Park, province of
Manabí – FHGO 713; La Maná, province of Cotopaxi – not
preserved). However, the validity of these records is uncer-
tain because they could correspond to animals transported
by illegal traffic, rather than a real extension in the distribu-
tion range of the species (ej., Geochelone denticulata re-
ported by Orcés [1949] from western Ecuador). Illegal traffic
of turtles in Ecuador has involved several thousands of
freshwater turtles and tortoises over the years. The exten-
sive commerce towards the biggest cities (ej., Quito and
Guayaquil) was stopped (or at least minimized) in the late-
90’s through a public campaign to stop wildlife traffic; how-
ever, several dozens are still captured for the pet markets in
small-medium towns. Illegal traffic involves almost any main-
land species, but the most common-captured species are:
Geochelone denticulata, Chelydra serpentina,
Kinosternon leucostomum, and Rhinoclemmys spp. Fur-
ther, thousands of specimens of Trachemys spp. are ex-
ported into Ecuador for the legal pet market, and several of
those turtles are released into Ecuadorian wetlands when
people are unable to care them. It is unknown if there is any
population of Trachemys spp. established in Ecuador and
the consequences on the native wildlife.
FAMILY DERMOCHELYIDAE
11. Dermochelys coriacea (Vandelli, 1761)
Distribution in Ecuador: Pacific coast of Ecuador (in-
cluding surrounding islands) and Galápagos Archipelago.
Notes: Some authors recognized two subspecies in-
side Dermochelys coriacea, the nominal and schlegelii
(Garman) (with augusta [Philippi] as a synonym); with the
nominal subspecies for the Atlantic populations and
schlegelii assigned to the Indo-Pacific populations. Al-
though differences between Eastern Pacific and Atlantic
populations have been mentioned (Pritchard and Trebbau
1984), low genetic variation was showed in analyses of
mtDNA sequence divergence between Pacific and Atlantic
populations (Dutton et al. 1996); and currently Dermochelys
coriacea is recognized as monotypic, without subspecific
divisions (Pritchard 1980, Pritchard and Trebbau 1984).
FAMILY GEOEMYDIDAE
Notes: The family Geoemydidae (with Bataguridae
as a junior synonym) was previously considered a subfam-
ily inside Emydidae, however currently both are considered
as separate families; with Emydidae as the sister group of a
Geoemydidae/Testudinidae clade (Spinks et al. 2004).
12. Rhinoclemmys annulata (Gray, 1860)
Distribution in Ecuador: Western Tropical. Provinces
of Esmeraldas, Manabí, Guayas, Los Ríos, Imbabura, Pichincha.
Notes: This species was separated into the resur-
rected genus Chelopus by Yasukawa et al. (2001), how-
ever recent analysis strongly suggest that the genus
Rhinoclemmys (including annulata) is monophyletic and
the recognition of the genus Chelopus is not justified
(Spinks et al. 2004).
13. Rhinoclemmys melanosterna (Gray, 1861)
Distribution in Ecuador: Northwestern Tropical. Prov-
ince of Esmeraldas.
Notes: Records of Rhinoclemmys melanosterna
(ej. Puerto Cayo-Puerto Nuevo road, Manabí – FHGO
1335, Manta-Eloy Alfaro road, Manabí – FHGO 1478)
and R. nasuta (record from the Machalilla National Park
reported by Almendáriz & Carr 1992) from outside their
currently known distribution ranges must be evaluated
carefully, as they can be product of artificial introduc-
tions rather than reflect the real distribution of the spe-
cies (see notes under Chelydra serpentina).
14. Rhinoclemmys nasuta (Boulenger, 1902)
Distribution in Ecuador: Northwestern Tropical. Prov-
inces of Esmeraldas and Pichincha.
Notes: See notes under Rhinoclemmys melanosterna.
FAMILY KINOSTERNIDAE
15. Kinosternon leucostomum Duméril & Bibron, 1851
Distribution in Ecuador: Western Ecuador. Provinces
of Esmeraldas, Guayas, Los Ríos, Pichincha, and Cañar.
Notes: Ecuadorian population is assigned to the sub-
species Kinosternon leucostomum postinguinale (Cope).
Records of Kinosternon leucostomum from outside its cur-
rently known distribution range must be evaluated care-
fully, as they can be product or artificial introductions rather
than reflect the real distribution of the species (see notes
under Chelydra serpentina).
16. Kinosternon scorpioides (Linnaeus, 1766)
Distribution in Ecuador: Eastern Tropical (Amazonia).
Provinces of Sucumbíos and Orellana.
Notes: Ecuadorian populations are assigned to the sub-
species Kinosternon scorpioides scorpioides (Linnaeus).
FAMILY PODOCNEMIDIDAE
Notes: The family Podocnemididae was previously
considered a subfamily inside Pelomedusidae, but currently
the Malagasy and American species are in the family
Podocnemididae and just the African species in the family
Pelomedusidae (see Meylan 1996)
17. Podocnemis expansa (Schweigger, 1812)
Distribution in Ecuador: Eastern Tropical (Amazonia).
Provinces of Sucumbíos, Napo, Orellana, and Pastaza.
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Notes: Ecuadorian populations are critically endan-
gered and on the brink of extinction.
18. Podocnemis unifilis Troschel, 1848
Distribution in Ecuador: Eastern Tropical (Amazonia).
Provinces of Sucumbíos, Napo, Orellana, Pastaza and
Morona-Santiago.
19. Peltocephalus dumerilianus (Schweigger, 1812)
Distribution in Ecuador: Eastern Tropical (Amazonia).
Province of Pastaza.
Notes: First reported from Ecuador by Orcés (1949).
FAMILIA TESTUDINIDAE
Notes: All Ecuadorian species of the family
Testudinidae are members of the subgenus Chelonoidis,
which has been accorded generic status by some authors
(ej. Bour 1980). The taxonomic arrangement of the Galapagos
tortoises is not completely settled:
(1) Some authors relegate all Galapagos tortoise taxa
as subspecies of a single species, usually named G. nigra
or G. elephantopus.
(2) The populations from southern Isla Isabela (Dar-
win and Alcedo Volcanoes, Cerro Azul, Sierra Negra) are
problematic. Pritchard (1996) suggested that the four south-
ern Isla Isabela taxa do not warrant separate status. Caccone
et al. (1999) found that those populations lack genetic dif-
ferentiation; but latter, Caccone et al. (2002) found that
microphyes (Volcan Darwin) and vandenburghi (Volcan
Alcedo) are genetically distinct, and deserve separation;
but they can not refute the suggestion that the two south-
ern most subspecies are not genetically distinct.
(3) It was recently recognized that the Isla San Cruz
populations, formerly recognized as the single taxon nigrita
(with porteri as a synonym), include three distinct lineages,
with at least one deserving its recognition as a new and
endangered taxon (Russello et al. 2005).
20. Geochelone abingdonii (Günther, 1877)
Distribution in Ecuador: Galápagos Archipelago,
Isla Pinta.
Notes: Only survivor (Lonesome George) lives at the
Charles Darwin Research Station.
21. Geochelone becki (Rothschild, 1901)
Distribution in Ecuador: Galápagos Archipelago, Isla
Isabela (Wolf volcano).
22. Geochelone chathamensis (Van Denburgh, 1907)
Distribution in Ecuador: Galápagos Archipelago,
Isla San Cristobal.
23. Geochelone darwini (Van Denburgh, 1907)
Distribution in Ecuador: Galápagos Archipelago,
Isla San Salvador.
24. Geochelone denticulata (Linnaeus, 1766)
Distribution in Ecuador: Eastern Tropical
(Amazonia). Provinces of Sucumbíos, Napo, Orellana,
Pastaza and Morona-Santiago.
Notes: Records from Western Ecuador are certainly
specimens transported by illegal traffic (Carr & Almendáriz
1989, also see notes under Chelydra serpentina).
It has been suggested that Geochelone
carbonaria (Spix) could occur in Ecuador (Vanzolini in
Miyata 1982, P. Salvador pers. comm.), however no
voucher specimens are known.
25. Geochelone ephippium (Günther, 1875)
Distribution in Ecuador: Galápagos Archipelago,
Isla Pinzón.
26. Geochelone guntheri (Baur, 1890)
Distribution in Ecuador: Galápagos Archipelago, Isla
Isabela (Sierra Negra).
27. Geochelone hoodensis (Van Denburgh, 1907)
Distribution in Ecuador: Galápagos Archipelago,
Isla Española.
28. Geochelone microphyes (Günther, 1875)
Distribution in Ecuador: Galápagos Archipelago,
Isla Isabela (Darwin volcano).
29. Geochelone nigrita (Duméril & Bibron, 1835)
Distribution in Ecuador: Galápagos Archipelago,
Isla Santa Cruz.
30. Geochelone vandenburghi (DeSola, 1930)
Distribution in Ecuador: Galápagos Archipelago,
Isla Isabela (Alcedo volcano).
31. Geochelone vicina (Günther, 1875)
Distribution in Ecuador: Galápagos Archipelago,
Isla Isabela (Cerro Azul).
... Aún cuando Mesoclemmys heliostemma ocurre en simpatría con M. raniceps, tal vez no se encuentran microsimpatricamente, dado que mientras la no inundables, en donde el agua es clara, la segunda habita la parte baja de los caños, en aguas más profundas y turbias y penetra en la varzéa y áreas de inundación de los mismos (CISNEROS-HEREDIA, 2006). Especie de actividad nocturna que se alimenta de peces pequeños y animales de cuerpo blando. ...
... El complejo de las tortugas gigantes de las Islas Galápagos ha tenido un amplio debate taxonómico, dado que muchos autores las han tratado como 1992; PRITCHARD, 1996;CACCONE et al., 1999CACCONE et al., , 2002BEHEREGARAY et al., 2004;VETTER, 2005;FRITZ & HAVAŠ ERNST et al., 2000;CISNEROS-HEREDIA, 2006), sobre la base del aislamiento reproductivo que existe entre las diferentes subespecies y a la inviabilidad de la descendencia de las entidades que se entrecruzan. También hay datos genéticos indicando la posibilidad de cambios en la taxonomía et al., RUSSELLO et al., 2005;CIOFI et al., 2006). ...
... In Venezuela, it occurs in the Yagua, Atacavi (Pérez-Emán and Paolillo 1997), Autana and Sipapo rivers (Barrio-Amorós and Narbaiza 2008). In Ecuador it occurs in the Province of Pastaza, and possibly in the Province of Sucumbíos (Cisneros-Heredia 2006). In French Guyana it is uncommon and restricted to the far east of the country (Fretey 1987;Bonin et al. 2006;Böhm 2010). ...
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