Status and Distribution of Nonmarine Turtles of the Philippines

Transcription

Chelonian Conservation and Biology, 2008, 7(2): 157–177
Ó 2008 Chelonian Research Foundation
Status and Distribution of Nonmarine Turtles of the Philippines
ARVIN C. DIESMOS1,3,4, RAFE M. BROWN2,3,4, ANGEL C. ALCALA3,4,
AND
ROGELIO V. SISON1
1
Herpetology Section, Zoology Division, National Museum of the Philippines, Padre Burgos Avenue, Ermita 1000, Manila,
Philippines [[email protected]];
2
Natural History Museum and Biodiversity Research Center and Department of Ecology and Evolutionary Biology, University of
Kansas, Dyche Hall 1345, Jayhawk Boulevard, Lawrence, Kansas 66045-7561 USA [[email protected]];
3
Silliman University-Angelo King Center for Research and Environmental Management, Marine Laboratory Building, Bantayan,
Dumaguete City 6200, Philippines [[email protected]];
4
Wildlife Conservation Society of the Philippines, Institute of Biology, University of the Philippines Diliman 1101, Quezon City,
Philippines
ABSTRACT. – We present updated accounts on the status and distribution of all known species of
freshwater turtles in the Philippines, including information on taxonomy, natural history,
exploitation, and conservation status. Six species of native nonmarine turtles (Dogania subplana,
Pelochelys cantorii, Cuora amboinensis, Cyclemys dentata, Heosemys spinosa, and Siebenrockiella
leytensis) are recognized from the Philippines, 2 of which are trionychids and 4 are geoemydids.
One of these (S. leytensis) is an endemic species. There is some evidence to suggest that Philippine
populations of P. cantorii, C. amboinensis, C. dentata, and H. spinosa may warrant further
taxonomic study. The fauna also includes 3 introduced species (Pelodiscus sinensis, Trachemys
scripta elegans, and Chrysemys picta). Pelodiscus sinensis, D. subplana, C. amboinensis, and S.
leytensis feature prominently in the domestic food and pet trade. Preliminary information
indicates that these species are also illegally exported to supply international animal markets. The
lack of basic information on the natural history of freshwater turtles in the Philippines stands as
an obstacle to the implementation of effective conservation and management measures. We
recommend that a national program for Philippine freshwater turtles be established to address
priorities for research, conservation, and management of this threatened and poorly studied
faunal assemblage.
KEY WORDS. – Reptilia; Testudines; Geoemydidae; Trionychidae; turtles; conservation; status;
distribution; illegal wildlife trade; Southeast Asia; Philippines
Gray (1844) provided the earliest description of the
testudine fauna of the Philippine Islands (Map 1). He
reported on the presence of 3 freshwater and 1 marine
species from the archipelago. Gray (1864) also published
the description of Pelochelys cumingii, a Philippine
endemic taxon that was later synonymized with P. cantorii
(Boulenger 1889; see Species Accounts). Subsequent
publications on Philippine turtles dealt largely with species
lists based on museum collections and new distribution
records (e.g., Günther 1879; Boulenger 1884; de Elera
1895; Siebenrock 1909). The work of de Elera (1895)
contributed important records to the turtle fauna despite
the questionable nature of some of the taxonomic
decisions and species identifications (e.g., Taylor 1921;
Das 1996b; Gaulke and Fritz 1998).
In the first modern review of Philippine turtles, Taylor
(1920) recognized a total of 8 species comprised of 4
marine turtles and 4 freshwater turtles, including the
endemic Heosemys leytensis (¼ Siebenrockiella leytensis;
Diesmos et al. 2005), which he formally described in that
monograph. Taylor (1920, 1921) also commented on the
depauperate freshwater turtle fauna of the Philippines
when compared with other regions of Asia but highlighted
the possibility of discovering new species from unexplored
freshwater ecosystems in the archipelago.
Current knowledge of the diversity and distribution of
Philippine marine turtles is far more complete than that of
the freshwater turtle fauna. For instance, field studies
undertaken across many regions of the Philippines since
the 1950s have increased significantly our understanding
of the ecology, conservation status, and management of
marine turtles (e.g., Domantay 1953; Alcala 1980; De
Veyra 1997; Torres et al. 2004). Five species of marine
turtles are known in Philippine waters, namely, Caretta
caretta, Lepidochelys olivacea, Chelonia mydas, Eretmochelys imbricata (Cheloniidae), and Dermochelys
coriacea (Dermochelyidae).
In contrast, information on Philippine nonmarine
turtles is now outdated and has not been systematically
reviewed in many decades. Consequently, the Philippines
has one of the least known nonmarine turtle faunas in
Asia. Even basic information on species richness and
distribution remains uncertain (Das 1996b; van Dijk et al.
2000; Brown et al. 2002). The possibility of additional
undocumented and/or undescribed Philippine taxa has
been suggested by recent workers (Das 1996a; Gaulke and
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CHELONIAN CONSERVATION
AND
BIOLOGY, Volume 7, Number 2 – 2008
data to this effect (see Acknowledgments). This work is
naturally far from complete, and we hope that our effort to
synthesize available information will help facilitate muchneeded conservation work and stimulate field studies on
this poorly known group of turtles.
METHODS
Map 1. The Philippine Islands (darkly shaded islands) with
tracings of 120-m submarine contours that define Pleistocene
Aggregate Island Complexes (PAIC) indicated by light shading.
Islands and channels mentioned in text are included and the
capital city (Manila) is indicated with a star.
Fritz 1998), but no systematic studies have been
forthcoming.
As with most other regions in Southeast Asia,
information on the fossil records of Philippine turtles is very
limited, and this potentially rich source of data on diversity
and biogeography of this group remains poorly studied. For
instance, archaeological explorations of Cagayan Valley in
northeastern Luzon Island have yielded apparent trionychine
and testudinid fossils (de Vos and Bautista 2003). This
discovery has significant implications as no extant species of
Testudinidae is known from the Philippines (Alcala 1986;
Ernst and Barbour 1989; Iverson 1992).
Here we review the status of all species of freshwater
turtles that are currently known from the Philippines and
provide updated information on their distribution, natural
history, and conservation. For most species, we present
recommendations on what we consider to be the most
essential areas for scientific research and conservation
efforts. The information we present is a result of our
fieldwork throughout the Philippines and includes direct
observations of turtles in the wild and examinations of
voucher materials from numerous natural history collections. In addition, field biologists working on other aspects
of Philippine biodiversity generously contributed valuable
General distributional data (i.e., names of islands and
general geographic features such as mountains, lakes, or
rivers) are provided in the species accounts, with more
detailed locality data (based on vouchers, field observations, collectors’ notes, or photographic evidence) summarized in the Appendix. Philippine Pleistocene
Aggregate Island Complexes (biogeographical subprovince, or PAICs) harbor unique and often nonoverlapping
fauna and correspond to paleoislands that existed during
periods of reduced sea level during the Pleistocene (Brown
and Diesmos 2002). Thus, we often refer to the PAIC,
where we suspect a species might be found. Information
on ecology and natural history of species is limited to
Philippine populations; for more information on these
subjects, refer to Ernst and Barbour (1989), Lim and Lim
(1992), Cox et al. (1998), Lim and Das (1999), and Ernst
et al. (2000). The global conservation status of species was
based on the International Union for Conservation of
Nature Red List of Threatened Animals (IUCN 2007;
www.iucnredlist.org). We also included the legal status of
internationally traded species based on the Convention on
International Trade of Endangered Species of Wild Fauna
and Flora (CITES 2005; www.cites.org). Vernacular
names and respective dialects (in parentheses) are
preceded by common English names.
NATIVE SPECIES
Trionychidae
Dogania subplana (Geoffroy Saint-Hilaire, 1809)
(Plate 1, Map 2)
Common Names. — Malayan softshell turtle, stream
softshell turtle, hillstream soft-shelled turtle, pawikan
(Tagalog), labi, labe (Pala’wan).
Distribution. — In the Philippines, D. subplana is
known from the islands of Luzon, Palawan, Mindoro, and
Mindanao. This highly disjunct distribution suggests that
the species may yet be recorded from other islands in the
country. Outside the Philippines, this species is known
from southern Myanmar, Thailand, West Malaysia,
Singapore, Sumatra, Borneo, Java, and associated islands
in the Sunda Shelf (Iverson 1992; Cox et al. 1998; Lim
and Das 1999; Fritz and Havas 2007).
Remarks. — De Elera (1895) first reported the
occurrence of D. subplana in the Philippines and listed it
from the islands of Mindanao and ‘‘Paragua’’ (¼ Palawan).
Later authors (e.g., Siebenrock 1906, 1909; de Rooij 1915)
subsequently summarized and included these records in
DIESMOS
ET AL.
— Philippine Nonmarine Turtles
159
Plate 1. (A) Pelodiscus sinensis from Lake Taal (photo J. Holden/Fauna and Flora International). (B) A captive bred P. sinensis from
Cebu City (photo A.C. Diesmos). (C and D) Dogania subplana from Rizal, Palawan Island (photo S. Schoppe). (E and F) Pelochelys
cantorii (PNM 8487) from Quirino Province, Luzon Island (photo M. de Guia).
several monographs. At that time, the materials that form
the basis of this account were deposited at the Museo de
Santo Tomas of the University of Santo Tomas, Manila
(de Elera 1895). Taylor (1920, 1921) doubted the
authenticity of much of de Elera’s (1895) records and
did not include D. subplana in his monograph and listings
of Philippine turtle fauna, apparently because of his
inability to locate voucher materials. Similarly, the species
was excluded from more recent checklists and monographs of Philippine amphibians and reptiles (e.g., Brown
and Alcala 1970; Alcala 1986).
We have encountered D. subplana in the field since
the early 1990s and have examined living and preserved
specimens from public and private zoos and zoological
reference collections of universities. Field biologists
conducting biodiversity surveys have likewise called our
attention to their own observations of the species from
different parts of the country. Gavino and Schoppe (2004)
and Lopez and Schoppe (2004) gave detailed accounts of
their field observations and morphological studies of D.
subplana on Palawan.
Natural History. — Dogania subplana has been found
in streams, rivers, and natural and artificial lakes and
ponds in pristine and degraded forests and in nonforested
habitats at elevations of near sea level to about 500 m. It
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AND
Map 2. Distribution of Dogania subplana in the Philippines. All
records are recent and confirmed.
has occasionally been found in irrigation canals, ditches,
and flooded rice fields.
Exploitation and Conservation. — This species is
legally protected in the Philippines through Republic Act
9147 (Philippines Wildlife Act). The illicit trade in D.
subplana within the Philippines is extensive. The species
is also exported to several countries in Asia (Gavino and
Schoppe 2004). Our data indicate that many wild-caught
D. subplana, especially from Palawan, end up in the
markets and restaurants in the cities of Manila and Cebu.
These turtles are also popular in pet shops and are used for
traditional medicine. Based on interviews gathered from
local respondents, commercial breeding farms of D.
subplana are presently operating in Palawan, Mindoro,
and Manila in order to sustain the trade. Although some
populations of D. subplana on Palawan appear to be
thriving, the species is potentially threatened by a
combination of overexploitation and habitat destruction
(Regodos and Schoppe 2005).
Research Priorities. — Information on population
estimates, life history, and the dynamics of illegal trade of
the species are needed. Field surveys are also needed to
gather additional information on the species’ true distribution in the Philippines.
Pelochelys cantorii Gray, 1864
(Plate 1, Map 3)
Map 3. Distribution of Pelochelys cantorii in the Philippines.
Circles ¼ recent confirmed records; squares ¼ historical records;
triangles ¼ unconfirmed reports.
Common Names. — Asian giant softshell turtle, frogfaced giant softshell turtle, frog-headed giant softshell,
Cantor’s giant softshell, antipa (Tagalog), kagot (Ilokano
and Ibanag), dinata, danata (Manobo).
Distribution. — Philippine populations of P. cantorii
are known thus far from the islands of Luzon (Cagayan
River and Ilaguen River) and Mindanao (Agusan Marsh
Wildlife Sanctuary and the Panabo River, Davao del Norte
Province). Other areas on Mindanao where its occurrence
has been reported to us but for which we lack verification
include Liguasan Marsh (North Cotabato Province) and
Laac (Compostella Valley Province). It may occur on
other major islands of the Philippines and is likely to be
found in the Sulu archipelago (Webb 2002). The species is
historically known from Balabac in the Palawan island
group (de Elera 1895; Siebenrock 1903) and from the
following localities on Luzon: San Miguel River, Bulacan
Province (Taylor 1920, 1921); Laguna de Bay, Laguna
Province (SMF 8069; examined by I. Das, in litt., March
2006; also mentioned in Webb 2002), and ‘‘Manille’’ (¼
Manila) (MNHN 579; examined by I. Das, in litt., March
2006). No information, however, is forthcoming to
indicate that the species still occurs in these localities.
Pelochelys cantorii ranges from southern India through
Bangladesh, southern China, Myanmar, Vietnam, Cambo-
DIESMOS
ET AL.
dia, Laos, Thailand, Malaysia, and Kalimantan, Indonesia
(Lim and Das 1999; Ernst et al. 2000; Webb 1995, 2002;
Stuart and Platt 2004; Fritz and Havas 2007).
Remarks. — The status and distribution of this large
freshwater softshell in the Philippines has been shrouded
in confusion for many years. Gray (1864) reported a large
specimen and several small ones of a Pelochelys collected
in the Philippines by Hugh Cuming (who visited the
islands during the period 1836–1839), an English amateur
conchologist and prolific collector of botanical and
zoological specimens; Gray (1864) named the species as
new, P. cumingii. This form was characterized as having
‘‘the head olive, minutely black-dotted; the throat olive,
minutely white-speckled’’ (Gray 1864). Pelochelys cumingii was subsequently synonymized with P. cantorii by
Boulenger (1889), an arrangement adopted by later authors
(e.g., de Rooij 1915; Taylor 1920, 1921; Iverson 1992). In
his extensive review, Webb (2002) doubted the authenticity of most Philippine records of P. cantorii because of the
spurious collection data of some past workers, the lack of
voucher materials from the Philippines, and the failure of
recent workers to relocate the species in the wild. Webb
(2002), however, accepted a record from the island of
Balabac (Siebenrock 1903) in the Palawan island group,
western Philippines, and suggested that it might occur on
other nearby (Philippine) islands north of Borneo. He also
emphasized that exhaustive field surveys with extensive
trapping were needed to confirm that Pelochelys does
occur in the Philippines.
We confirm the continued existence of P. cantorii in
the Philippines based on voucher materials, our own field
sightings of live animals, and examinations of shells and
photographs of the species. We have observed the species
at Ilaguen River, Isabela Province, in 1998 and at Cagayan
River, Cagayan Province, in 2001 and 2006, both in
northern Luzon. We have examined shells in the
possession of villagers from turtles that were collected at
the Ilaguen River, where the species is collected
occasionally for food. In addition, we have examined
photographs and specimens of P. cantorii from Agusan
Marsh Wildlife Sanctuary, a national protected area in
western Mindanao, and from the upper Cagayan River
basin in Quirino Province, northern Luzon. In all
instances, we recognized the turtles as P. cantorii; our
identification was later verified by R.G. Webb (in litt., 10
September 2003).
Natural History. — Pelochelys cantorii inhabits
freshwater marshes, lakes, and rivers. Adult and juvenile
turtles have reportedly been incidentally caught in fishing
nets placed across such habitats. An adult turtle was taken
in 2002 from a 3-m-wide tributary of the Panabo River,
Davao del Norte Province, Mindanao, during a dredging
operation. Local residents near the rivers of Ilaguen and
Cagayan on Luzon reportedly collect eggs of the species
from sandy shores and sandbars, especially during the start
of the rainy season (i.e., June or July). Turtles that were
intended for consumption by collectors fed on fish,
161
shrimps, and aquatic algae while in captivity. The general
scarcity of confirmed records from field surveys and the
lack of specimens indicate that this species may be
uncommon to rare in the Philippines.
Exploitation and Conservation. — The most important threat to P. cantorii appears to be habitat disturbance
and conversion, specifically the draining of wetlands for
agricultural plantations and the widespread quarrying of
gravel and sand from riverine habitats. The species is also
collected for local consumption. Indigenous peoples
groups on Luzon (e.g., Ibanag) and on Mindanao (e.g.,
Mandaya) make use of the carapace in traditional
medicine. Local beliefs also dictate that keeping of body
parts of the species brings good luck. Quantitative data on
exploitation for the pet and food trade are not available,
and this subject needs to be investigated. Pelochelys
cantorii is currently listed as endangered (IUCN 2007) and
is an Appendix II species (CITES 2005). It is legally
protected under the Philippines Wildlife Act.
Research Priorities. — The taxonomic status of P.
cumingii needs to be reevaluated. There is a possibility
that this taxon may be a distinct, endemic form (Das
1996b; Ernst et al. 2000). Significant morphological
differences pertaining to skull dimensions and neural bone
counts had previously been reported between Philippine
and mainland Asian populations of P. cantorii (Baur 1891;
Taylor 1920, 1921). There is unquestionably a need for an
in-depth taxonomic study, preferably with the use of
molecular and genetic methods. The taxonomic status of
Philippine Pelochelys cannot be reliably assessed without
the study of additional material (live or preserved
specimens from known localities; see Lehn et al. 2007).
At present, we are aware of the existence of only 2
preserved specimens of P. cantorii from the Philippines
(PNM 8487 and an uncataloged specimen). A quantitative
and comparative study of the various populations of P.
cantorii across the Asian region is also warranted. Further,
information on its population biology and life history need
to be collected. An assessment of the impact of habitat
degradation and exploitation of the species should be
considered a priority research action.
Geoemydidae
Cuora amboinensis (Daudin, 1801)
(Plate 2, Map 5)
Common Names. — Malayan box turtle, Southeast
Asian box turtle, Amboina box turtle, pagong (Tagalog),
siroman, bayu-o (Pala’wan), bakuku (bakoko), bari,
krusan (Cuyonin), ba-o (Visayan), kumaw (Waray), pannu (Bagobo), kula-kula (Tausug).
Distribution. — Cuora amboinensis is widely distributed in the Philippines and is expected to be found on
other islands that are not listed here. Details on localities
are enumerated in the Appendix. In the Philippines, 2
subspecies are distributed as follows: Cuora a. amboinensis is found on Luzon (recorded from numerous
162
AND
Plate 2. (A and B) Cuora amboinensis kamaroma from Quezon, Palawan Island (photo: A.C. Diesmos). (C) Juvenile C. a. kamaroma
from Rizal, Palawan Island (photo: A.C. Diesmos). (D and E) Cuora a. amboinensis from Polillo Island, Luzon PAIC (photo: T. Devitt).
(F) Cuora a. amboinensis from Devao City, Mindanao (photo: A.C. Diesmos).
localities across the island, originally reported as C.
amboinensis, e.g., Taylor 1920, 1921; Alcala 1986; Brown
et al. 1996, 2000; Diesmos et al. 2004a), Polillo, Verde
(Sison and Diesmos 2008), Mindoro, Sibuyan, Palawan,
Dumaran, Panay (Ferner et al. 2000), Guimaras, Pan de
Azucar, Negros, Cebu, Masbate, Leyte, Samar, Bohol,
Dinagat, and Mindanao. Cuora a. kamaroma has been
recorded from Sanga-sanga, Tawi-Tawi, Jolo (Gaulke
1995a, 1995b), Dumaran, and Palawan (PNM specimens;
see Appendix). Nominate C. amboinensis ranges from
eastern India, Bangladesh, Burma, Thailand, Vietnam,
Malaysia, Timor, Borneo, Sulawesi, Ceram, and Amboina,
Indonesia (Iverson 1992; Lim and Das 1999; Ernst et al.
2000; Fritz and Havas 2007).
Remarks. — Cuora amboinensis is the most wideranging freshwater turtle in the Philippines, being found on
nearly all major islands of the archipelago, including small
satellite islands (Taylor 1920, 1921; Alcala 1986; Ernst
and Barbour 1989; Iverson 1992). Four subspecies of C.
amboinensis are presently recognized (Rummler and Fritz
1991; McCord and Philippen 1998; Fritz and Havas 2007),
and 2 of these occur in the Philippines, C. a. amboinensis
and C. a. kamaroma (Gaulke 1995a; Gaulke and Fritz
1998). The 2 forms can be differentiated morphologically
by the following characters: C. a. amboinensis has a
relatively flat carapace with vertebral keels, and the
plastron has prominent black blotches; C. a. kamaroma
has a high-domed, relatively smooth carapace, and the
plastron lacks or has few black markings. Gaulke (1995a)
DIESMOS
ET AL.
Map 5. Distribution of Cuora amboinensis in the Philippines.
Circles ¼ C. a. amboinensis; squares ¼ C. a. kamaroma. All
and Gaulke and Fritz (1998) asserted that these 2
subspecies occupy disparate biogeographical zones in the
Philippines and do not overlap in geographic distribution.
Although C. a. kamaroma is apparently restricted to the
Sulu and Palawan island groups (regions that are biogeographically traditionally associated with Sundaland: Heaney 1986; Brown et al. 2002; Diesmos et al. 2005; but see
Brown and Diesmos, in press), recent herpetological
surveys of Palawan have recorded both C. a. kamaroma
and C. a. amboinensis in the same habitats (Diesmos et al.
2004b; Diesmos and Brown, unpubl. data). We have
observed both forms occurring in sympatry in forest
habitats on the island of Dumaran (in the Palawan island
group) and in swamp forests on the island of Palawan. The
hypothesis that Palawan populations represent a morphologically intermediate and currently undescribed taxon
warrants additional study (S. Schoppe, pers. comm., 2007).
Natural History. — his species inhabits a wide variety
of aquatic habitats, including marshes, swamps, rivers,
streams, lakes, and ponds, and is occasionally found in
man-made habitats, such as reservoirs, irrigation ditches,
flooded rice fields, and even water buffalo wallows. Its
recorded elevational distribution is from sea level to about
1000 m (Alcala 1986; Diesmos et al. 2004a). Both
subspecies of C. amboinensis appear to be fairly common
in areas where they are known to occur (pers. obs.). Alviola
163
et al. (2003) observed that subadult C. amboinensis on
Polillo Island dispersed more widely compared with
juveniles and adults. In general, very few ecological studies
of the species have been undertaken despite its apparent
abundance and wide distribution in the Philippines.
Exploitation and Conservation. — This box turtle is
extremely popular in pet shops in the cities of Manila,
Cebu, and Davao. Both juvenile and adult turtles are sold
with prices ranging from 50 to 250 pesos (about $1 to $5)
per individual. We have received reports of wholesale wild
collecting of this species on Sibuyan Island, where it was
previously plentiful. One of us (RMB) interviewed
residents of the island who reported that 10–15 50-kg rice
sacks of live C. amboinensis were harvested weekly from
the 1980s to the early 1990s for sale to Chinese and
Taiwanese traders and that the species was now much less
plentiful on Sibuyan Island compared to 20–30 years ago.
Based on interviews with wildlife collectors, we
believe that most if not all turtles sold in animal markets
are wild caught. Its meat is considered a delicacy in many
regions of the country; hence, the turtle is regularly
harvested for consumption. On the other hand, there is a
prevalent belief that the species brings wealth and good
luck, and animals are commonly kept as pets in many rural
villages across the Philippines. Cuora amboinensis is
listed as vulnerable (IUCN 2007) and is an Appendix II
species (CITES 2005). It is a protected species under the
Philippines Wildlife Act.
Cyclemys dentata (Gray, 1831)
(Plate 3, Map 6)
Common Names. — Asian leaf turtle, brown stream
terrapin, bakuku (bakoko), kuritan (Cuyonin), bayu-o,
kalumbagis (Pala’wan).
Distribution. — In the Philippines, C. dentata is
restricted to the Palawan island group (Calauit, Busuanga,
Culion, Dumaran, Palawan, and Balabac), the Sulus, and
Tawi-Tawi. A specimen is known from the island of Siasi
in the Sulu archipelago (MNHN 584; examined by I. Das,
in litt., March 2006). As currently understood, C. dentata
(sensu lato) ranges from eastern India through Myanmar,
Vietnam, Laos, Cambodia, Thailand, Malaysia, Singapore
(doubtful record: Lim and Lim 1992), Sumatra, Java, and
Borneo (Iverson 1992; Lim and Das 1999; Ernst et al.
2000; Fritz and Havas 2007).
Remarks. — Cyclemys dentata has been reported from
the island of Leyte (Taylor 1920, 1921; Alcala 1986; Ernst
and Barbour 1989; Iverson 1992; Gaulke and Fritz 1998).
However, herpetological surveys of Leyte and adjacent
islands have repeatedly failed to find the species (e.g.,
Denzer et al. 1994; Gaulke 1994; Diesmos et al. 2004c).
Because there is compelling evidence that the original Leyte
account stems from a mislabeling of specimens (Diesmos et
al. 2004b, 2004c) and no substantive subsequent records
have been forthcoming in the intervening years, we suspect
that, like Siebenrockiella leytensis (see the following
discussion), the Leyte record for C. dentata is in error
164
AND
Plate 3. (A and B) Cyclemys dentata from Dumaran Island, Palawan PAIC. (C) Juvenile C. dentata from Quezon, Palawan Island. (D)
Cyclemys dentata from Puerto Princesa, Palawan Island (all preceding photos A.C. Diesmos). (E and F) Cyclemys dentata from TawiTawi Island, Sulu PAIC (photo E.L. Rico/Haribon Foundation).
and should not be included further in faunal distribution
lists and checklists of species on the island.
Natural History. — The species inhabits small streams
in primary and secondary hill forests and freshwater
swamps and is rarely found in artificial habitats and
agricultural areas. During the day, when it is generally
inactive, individuals can be found under rocks, buried
under sand and gravel, or in crevices of boulders on the
banks of streams or rivers. In the island of Dumaran, we
observed adults and juveniles of the species feeding on
ripe figs (Ficus spp.) that fell into streams. Turtles in
captivity took pieces of meat, fruits, and the ubiquitous
aquatic plant Ipomoea aquatica.
Exploitation and Conservation. — Cyclemys dentata
is collected by indigenous peoples and rural villagers as a
protein source and is commonly consumed because of the
supposed medicinal properties of its meat. Wildlife traders
from Palawan revealed that C. dentata is among the turtles
that are regularly (and illegally) exported to supply the pet
markets of Asia and Europe. The trade in C. dentata is
extensive within Palawan but appears to be negligible
elsewhere in the Philippines. This species can be
occasionally seen in the pet shops of Manila (pers. obs.).
Cyclemys dentata is fairly common in the Palawan island
group, and many populations can be found inside protected
areas (e.g., Puerto Princesa Subterranean River Natural
Park, Puerto Princesa City). Gaulke (1995a) reported that it
is ‘‘not rare’’ on Tawi-Tawi Island. This species is legally
protected under the Philippines Wildlife Act.
DIESMOS
ET AL.
Map 6. Distribution of Cyclemys dentata in the Philippines. All
Research Priorities. — Cyclemys dentata is probably
a complex of several species (Fritz 1997; Iverson and
McCord 1997; Guicking et al. 2002). Philippine Cyclemys
may represent more than 1 lineage, particularly given the
fact that they occur in the Palawan, Sulu, and Tawi-Tawi
PAICs (Plate 3E–F) (P.P. van Dijk, in litt., December
2007). Taxonomic studies, preferably with the use of
molecular data, could help resolve uncertainty concerning
the true identity of Philippine Cyclemys.
Heosemys spinosa (Gray, 1831)
(Plate 4, Map 7)
Common Names. — Spiny hill turtle, spiny turtle,
kula-kula (Tausug), pagong (Tagalog).
Distribution. — Tawi-Tawi Island in southern Philippines. It is likely to occur in other forested islands within
the Sulu and Tawi-Tawi PAIC. We have demonstrated
(see the following discussion) that the record of H. spinosa
from Mindanao (Das 1996a) is in error; although, we
strongly underscore the need for thorough faunal inventories of the island to gain a better understanding of its
testudine fauna. Heosemys spinosa ranges from southern
Thailand, Myanmar, Malaysia, Sumatra, Borneo, and
many other Indonesian islands (Iverson 1992; Lim and
Das 1999; Ernst et al. 2000).
165
Remarks. — Das (1996a) first reported Heosemys
spinosa from the Philippines based on 2 preserved
specimens at the National Museum of the Philippines that
allegedly originated from Mindanao. In fact, the specimens
in question originated from Tawi-Tawi and were taken
during a faunal inventory of the island by biologists of the
National Museum of the Philippines and the Cincinnati
Museum of Natural History in 1991 (R.S. Kennedy, in
litt., June 2003; PNM herpetological registry). Fritz (1997)
independently validated the occurrence of H. spinosa on
Tawi-Tawi, while a more recent ‘‘rapid biodiversity
assessment’’ by biologists from Haribon Foundation (a
Philippine-based conservation nongovernmental organization) confirmed the continued existence of the species
there through photographic records (E.L. Rico, in litt.,
December 2004).
Natural History. — This turtle has been observed near
streams and ponds in primary and logged-over lowland
forest below 300 m elevation. Juveniles have also been
found in these same habitats during the wetter months of
the year (October–November). Seasoned wildlife hunters
and collectors from Tawi-Tawi claimed they rarely
encounter this turtle in their collecting trips, indicating
that the species may naturally occur at low densities on the
island or that its population may be declining because of
continued habitat destruction.
Exploitation and Conservation. — Recent field
reports from biologists suggest that the consumption of
this turtle for food is not prevalent and that it is only
occasionally collected for the wildlife trade. Overall, the
level of exploitation of the species appears to be
negligible, but more research is needed on this subject.
Heosemys spinosa is globally threatened (endangered;
IUCN 2007) and is an Appendix II species (CITES 2003).
It is legally protected under the Philippines Wildlife Act.
Research Priorities. — Very little information is
available on the status and distribution of the Philippine
population of H. spinosa. Priority research activities must
include habitat utilization studies, reproductive biology,
population assessments, and distribution surveys, particularly in the Sulu island group. The unstable sociopolitical
situation and perceived security risk in the region have
successfully prevented biologists from conducting fieldwork there for over 30 years (see duPont and Rabor 1973).
Biodiversity studies in this PAIC have been sporadic at
best and limited to the Tawi-Tawi archipelago (e.g., Taylor
1923; Gaulke 1995b). The biologically unknown Sulu
archipelago may possibly harbor a remarkable and unique
herpetofauna (Brown et al. 2002) and possibly may
support other unrecorded turtle species.
Siebenrockiella leytensis (Taylor, 1920)
(Plate 5, Map 8)
Common Names. — Leyte pond turtle, Philippine
pond turtle, Philippine freshwater turtle, Philippine forest
turtle, tumanog (Tagbanua), panyaen, bayu-o (Pala’wan),
bakuku (bakoko) suyan (Cuyonin).
166
AND
Plate 4. (A and B) Heosemys spinosa (PNM 2232, male) from Tawi-Tawi Island, Sulu PAIC (photo A.C. Diesmos). (C) Juvenile H.
spinosa from Tawi-Tawi Island, Sulu PAIC (photo E.L. Rico/Haribon Foundation).
Distribution. — Siebenrockiella leytensis is confined
to the Palawan island group. It has so far been found only
in the northern regions of Palawan (Taytay, San Vicente,
Puerto Princesa) and from the island of Dumaran (Diesmos
et al. 2004b, 2004c). This species may have a much wider
distribution within this PAIC than is presently known and
may eventually be found (if suitable habitats exist) on the
islands of Busuanga, Coron, Culion, Linapacan, and
possibly Balabac (Diesmos et al. 2004b).
Remarks. — The Philippine forest turtle formerly was
one of the most enigmatic species of freshwater turtles in
Southeast Asia. It has been surrounded with more
confusion and misconception than any other geoemydid/
batagurid turtle known from the region (Diesmos et al.
2004b). For over 80 years, its true geographic distribution
in the Philippines remained a mystery; its apparent rarity
and the rudimentary knowledge of its natural history had
bestowed on it a mythical reputation (van Dijk et al. 2000).
The decades-long belief that Siebenrockiella leytensis
occurred on the island of Leyte (Taylor 1920) was finally
shown to be erroneous, resulting from a mislabeling of the
type specimens (Diesmos et al. 2004b). Siebenrockiella
leytensis is confined to the Palawan PAIC (Timmerman
and Auth, 1988; Diesmos et al. 2004b, 2005), which is
biologically distinct from the rest of the Philippines and
considered by some to be biogeographically affiliated with
Sundaland (Heaney 1985, 1986) or composed of a mixed
fauna with both Sundaic and oceanic Philippine origins
(McGuire and Alcala 2000; Brown and Guttman 2002;
Evans et al. 2003; Brown and Diesmos, in press).
Regardless, this PAIC harbors high concentrations of
endemic biodiversity not found in Southeast Asia
(Widmann 1998; Diesmos and Palomar 2004).
Taxonomy. — Taylor (1920) described S. [orig.
Heosemys] leytensis based on 2 specimens retrieved from
a zoological laboratory at the University of the Philippines
in Manila. He placed this taxon in the genus Heosemys on
the basis of an absence of a temporal arch. Wermuth and
Mertens (1961) suppressed Heosemys and allocated most
of its member species to Geoemyda, a placement that was
rejected by McDowell (1964), who cited the distinctive
skull anatomy of turtles within this group. Nonetheless,
DIESMOS
ET AL.
Map 7. Distribution of Heosemys spinosa in the Philippines.
This species is now known from only 2 confirmed localities on a
single island (circles).
subsequent papers listed leytensis as a species of Geoemyda (Ernst and Barbour 1989; but see Buskirk 1989).
McCord et al. (2000) proposed that leytensis remain in the
genus Heosemys until more robust information on the
systematics of these taxa indicated otherwise. A recent
morphological and molecular phylogenetic study (Diesmos et al. 2004b, 2005) showed that leytensis is the sister
lineage to the previously monotypic Sundaic turtle genus
Siebenrockiella. Thus, leytensis is the sister species to S.
crassicollis, which occurs from the middle Mekong to
Borneo. Nonetheless, the high genetic divergence (ca. 13%
cytb) between S. leytensis and S. crassicollis and a suite of
morphological differences prompted Diesmos et al. (2005)
to erect the subgenus Panyaenemys to accommodate the
distinct Philippine lineage.
Natural History. — The Philippine forest turtle is
nocturnal and semiaquatic. Hatchings and small juveniles
appear to rely on aquatic habitats more than adults (S.
Schoppe, pers. comm., 2007). This species inhabits
riparian habitats (streams, creeks, medium-sized rivers)
in pristine lowland forest but has also been found in
disturbed forest fragments and second-growth vegetation
near remnant original forests. It is known from freshwater
swamps (Fidenci 2004) and may occur in inland lakes.
Siebenrockiella leytensis has been recorded at elevations
from near sea level to about 300 m. During the driest
167
months of the year (November–April) when most streams
and medium-sized rivers have dried up, it hides under
limestone boulders or inside limestone caves, presumably
to escape the heat and/or reduce dehydration (Diesmos et
al. 2004c). On Dumaran Island, we found S. leytensis
ensconced under large rocks and deep burrows on
riverbanks during the day. At night, we observed them
foraging on freshwater fish, crabs, shrimps (e.g., Macrobrachium spp.), algae, and ripe figs (Ficus spp.) along
stream banks and deep stationary pools. In captivity, S.
leytensis is crepuscular and takes leafy vegetables, soft
fruits (banana, papaya), pieces of meat (chicken, pork,
fish), and even dog food pellets (Diesmos 2004c; Yuyek
2004). Data on reproduction are still fragmentary. Egg
laying has been recorded during the period June–August.
Two eggs laid by a female turtle measured 45.5 3 23.2 mm
and 57.8 3 24.3 mm and weighed 20 and 30 g,
respectively. The eggs are brittle-shelled, oblong, and pale
pink in color (Diesmos et al. 2004c; Yuyek 2004).
Hatching appears to occur between January and April (S.
Schoppe, unpubl. data). Wildlife trappers on Palawan
claim that it oviposits in deep burrows on near-vertical
banks of streams and rivers. Siebenrockiella leytensis is
the largest and the heaviest geoemydid turtle known from
the Philippines. Old adults have a carapace length of more
than 300 mm and a plastron length of over 220 mm and
can weigh well over 5 kg. We expect that data on this
subject will be produced quickly, given the current
research efforts of biologists including, regrettably, the
(entirely illegal) proliferation of S. leytensis in the hands of
wildlife collectors and herpetofaunal enthusiasts.
Exploitation and Conservation. — Siebenrockiella
leytensis is currently listed as critically endangered by the
IUCN Red List and is included in Appendix II of CITES
(CITES 2005; IUCN 2007). It is a nationally protected
species in the Philippines through the Wildlife Act, the
violation of which carries a penalty of heavy fines,
including imprisonment (PAWB 2004a, 2004b).
The greatest threats facing S. leytensis are overcollection for the illegal wildlife trade and the continuing
destruction of its primary habitat. The former is presently
the most serious threat and is certainly the most damaging
to its populations. As was predicted previously (van Dijk
et al. 2000), the rediscovery of S. leytensis has spurred a
collecting frenzy among wildlife trappers and traders to
supply illegal domestic and international markets. We
have gathered compelling evidence that its populations in
the region of Taytay and San Vicente in northern Palawan
are declining rapidly because of indiscriminate collection
(Diesmos et al. 2004c). Along with other wild vertebrate
species, poachers sell S. leytensis collected from Taytay
and San Vicente to collectors, middlemen, and traders. A
majority of the turtles are purchased and smuggled into
Manila, Cebu, or Davao. These cities serve as transshipment points where wildlife traffickers transport the animals
to several destinations in Asia and Europe (Diesmos et al.
2004c; Gavino and Schoppe 2004; Yuyek 2004; Plate 6).
168
AND
Plate 5. (A and B) Siebenrockiella leytensis from Dumaran Island, Palawan PAIC. (C) Juvenile S. leytensis from San Vicente, Palawan
Island (photo P. Widmann). (D and E) Captive S. leytensis from Taytay, Palawan Island. (F) Siebenrockiella leytensis photographed
from a stream on Dumaran Island, Palawan PAIC (photos A–B and D–F A.C. Diesmos).
As with other turtles in the Philippines, S. leytensis is
consumed for its meat, which is believed to possess
medicinal properties. The carapace and plastron are also
fashioned into indigenous musical instruments and various
curios. Members of ethnic groups from Palawan (particularly the Cuyonin and Tagbanua) and the economically
marginalized populace often carry out this form of
utilization.
Research Priorities. — Priority research topics
include population assessments, demography, and breeding biology and seasonality. In particular, baseline data on
population size need to be established to have an accurate
assessment of the impacts of habitat degradation and of
collection for wildlife trade and local consumption.
Additional field surveys, conducted with adequate documentation (Lehn et al. 2007), are essential to determine the
extent of its distribution within the Palawan PAIC.
INTRODUCED SPECIES
Trionychidae
Pelodiscus sinensis (Wiegmann, 1835)
(Plate 1, Map 4)
Common Names. — Chinese softshell turtle, pawikang luno (Tagalog).
DIESMOS
ET AL.
169
Map 8. Distribution of Siebenrockiella leytensis in the
Philippines. Circles ¼ recent confirmed; squares ¼ unconfirmed
but plausible reports; triangles ¼ unconfirmed less plausible
reports (Diesmos et al. 2004b, 2004c). Star ¼ type locality (in
error; see text).
Distribution. — Luzon, Mindoro, Panay, Cebu,
Bohol, Leyte, and Mindanao. Pelodiscus sinensis is likely
to be found in numerous other localities and islands across
the Philippines, chiefly from the deliberate introduction for
commercial farming. This species is native to mainland
Asia and ranges from Russia, China, Taiwan, Hainan,
Korea, Japan, and northern Vietnam. It has been
introduced into many countries, including Hawaii, Guam,
Indonesia, and Malaysia (Iverson 1992; Cox et al. 1998;
Lim and Das 1999; Ernst et al. 2000; Fritz and Havas
2007).
Remarks. — The history of introduction of P. sinensis
in the Philippines is undocumented; although, de Elera
(1895) did report the species from the Batanes island
group off northern Luzon. Later authors, however, did not
include the species in their listings of Philippine
herpetofauna (e.g., Taylor 1920, 1921; Brown and Alcala
1975; Alcala 1986) because of a lack of voucher materials
and the suspect nature of much of the information
provided by de Elera (1895; see comments by Das
1996b). A series of herpetofaunal inventories of the
Babuyan and Batanes islands conducted in the late 1980s
(e.g., Ota and Crombie 1989; Lazell 1992) and between
2004 and 2006 (Oliveros and Broad, unpubl. data; Oliver
Plate 6. (A) Through the initiative of Palawan-based conservation groups, such as the KATALA Foundation in partnership
with the local government, stakeholders such as this family from
Dumaran Island are actively engaged in conservation work on the
rich biodiversity of their island. (B) Dozens of live Siebenrockiella leytensis, Cuora amboinensis, and Cyclemys dentata turtles
on exhibition in an artificial pond, Puerto Princesa City, Palawan
Island. The keeping of turtles is a common practice in the
Philippines, as it is believed that turtles bring wealth and good
luck (photos A–B A.C. Diesmos). (C) At present, the most
significant threat to S. leytensis is overcollection for the illegal
wildlife trade. This photograph was downloaded from the Web
site of a wildlife trader based in Japan; specimens of S. leytensis
were advertised (July 2003) for sale at a price of US$1350 per
animal.
et al., unpubl. data) did not find the species. This softshell
may certainly have been brought into the Philippines
chiefly as a source of meat. Either escaping from
confinement or intentionally released into the wild, feral
populations of P. sinensis have now become successfully
170
AND
Map 4. Distribution of Pelodiscus sinensis in the Philippines.
Circles ¼ recent confirmed records; square (Batan Isl.) ¼
historical record.
established in many regions of the country. Numerous
distribution records for the species exist, and we have also
examined many specimens from public and private zoos
and pet shops.
Natural History. — This species occurs in both natural
and artificial lakes and ponds, reservoirs, flooded rice
fields, and irrigation ditches and has also been found in
streams inside natural forests. It buries itself in mud during
most of the day and is most active at night. Collectors and
turtle breeders claim that it feeds on frogs, tadpoles, fish,
crustaceans, worms, and the aquatic plant ‘‘kangkong’’
(water spinach, Ipomoea aquatica). Pelodiscus sinensis is
common to abundant in areas where it has been recorded.
Breeders indicate that it is a hardy species and requires a
minimum of effort to farm in commercial quantities. Its
ecological impact on native fauna is unknown and needs to
be investigated.
Exploitation and Conservation. — The trade in P.
sinensis for the food and pet market is widespread in the
Philippines. This turtle is also popular in pet shops of
Manila and Cebu. Reptile dealers sell each turtle from
US$4–11 per kilogram (Sy et al. 2004); although, heavier
and larger adults may command a premium price. Many
local people ascribe various medicinal properties to this
turtle, and, as a consequence, the consumption of its meat
by rural villagers is common. Chinese softshell turtles are
Map 9. Distribution of Trachemys scripta elegans (circles) and
Chrysemys picta (squares) in the Philippines.
also served in the restaurants of Manila and Cebu (and
likely in other cities) that specialize in turtle meat. Smallscale breeding facilities of the species are operating in
Luzon, Mindoro, and Cebu; apparently, this industry is
economically beneficial to local communities of these
regions (Arevalo 2003). Interviews with turtle breeders
and dealers revealed that ‘‘large numbers’’ of softshell
turtles bred or collected from feral populations in the
Philippines are exported to Asian countries, such as
Taiwan, Japan, and Hong Kong. It is a vulnerable species
(IUCN 2007) based on the status of remaining wild
populations. As with all other species of Philippine
freshwater turtles, P. sinensis is afforded legal protected
status under the Philippines Wildlife Act.
Emydidae
Trachemys scripta elegans (Schoepff, 1792)
Chrysemys picta (Schneider, 1783)
(Map 9)
Common Names. — Trachemys scripta elegans: redeared slider, common slider, pond slider, pagong (Tagalog); Chrysemys picta: painted turtle, painted terrapin,
pagong (Tagalog).
Distribution. — We have observed feral populations
of T. scripta elegans and C. picta from Luzon (La Mesa
Dam reservoir, Ipo Dam reservoir, Laguna de Bay, and
DIESMOS
ET AL.
Lake Taal), Cebu (several localities in and around Cebu
City), and Mindanao (Malagos and Davao City). They are
probably more widespread than is presently known.
Trachemys scripta is widely introduced in many countries
in Central and South America, Europe, Asia, and
Southeast Asia (Iverson 1992; Lim and Das 1999; Ernst
et al. 2000).
Remarks. — Both T. scripta elegans and C. picta are
native to North America. These colorful turtles were
introduced to the Philippines for the pet trade and have
become extremely popular among herpetoculturalists.
Feral populations of both species can now be found in
parts of the country because of deliberate releases of
unwanted pets or from escaped individuals. Compared
with C. picta, T. scripta elegans appears to have adapted
well to the local environment and has also successfully
established breeding populations in some localities.
Natural History. — The 2 species are aquatic,
inhabiting natural and man-made lakes, ponds, and
reservoirs, and are occasionally seen in tributaries and
ditches in the cities. Our field observations also show that
feral populations of T. scripta elegans in the Philippines
are not as sizable as those reported from other regions of
Southeast Asia (Lim and Das 1999). Their impact on
native fauna is unknown.
Exploitation and Conservation. — These turtles are
ubiquitous in pet shops across the country. Feral turtles are
collected occasionally for consumption by rural villagers.
Both species are afforded legal protection under the
Philippines Wildlife Act.
DISCUSSION
The nonmarine turtles of the Philippines are one of the
least understood chelonian faunas of Asia. Information on
life history and natural distribution is lacking for most
indigenous species; what little information is available is
based largely on non-Philippine populations. We have
attempted to fill in some gaps in our knowledge of this
fauna, but much remains to be done. A full understanding
of the systematics and ecology of Philippine freshwater
turtles is crucial in view of the prevailing Asian turtle crisis
and the high conservation priority placed on many
chelonian species from the region (van Dijk et al. 2000;
Stuart and Thorbjarnarson 2003; IUCN 2007).
With intensive field surveys particularly of unexplored river systems, wetlands, and lakes, additional
species may be recorded from the Philippines. Among
the species that could possibly exist in these unexplored
regions might be wide-ranging, secretive aquatic turtles
such as Amyda cartilaginea and species of Chitra and/or
Cyclemys. We recommend that thorough field surveys be
conducted in Cagayan River, Chico River, Abra River,
Lake Buhi, and Lake Bato on Luzon; Lake Manguao on
Palawan; Jalaud River on Panay; Lake Danao on Leyte;
and Ligwasan Marsh, Lake Lanao, Lake Mainit, Agusan
River, and Pulangi River on Mindanao. Relatively well-
171
studied wetland and lake ecosystems (e.g., Naujan Lake on
Mindoro; Lake Taal, Lake Buhi, and Laguna de Bay on
Luzon; and Agusan Marsh on Mindanao) may harbor
previously undetected indigenous populations of turtles
and hence should be reinvestigated. Basic studies on
population ecology, reproductive biology, and life history,
including focused research on phylogenetic relationships,
are needed especially for globally threatened taxa. We
highlight the importance of collecting well-documented
voucher specimens of Philippine freshwater turtles from
known localities (whole individuals, body parts, tissues,
and/or photographs) and deposition of specimens in
internationally accredited museums (Leviton et al. 1985;
Brown et al. 2002; Simmons 2002; Lehn et al. 2007).
Because most indigenous species of this fauna are
underrepresented—or even completely lacking—in museum collections, the necessity of obtaining specimens
cannot be overemphasized. For instance, the uncertainty
surrounding the taxonomy and natural distribution of the
Philippine endemic S. leytensis for over 80 years was
unraveled only after fresh museum material and genetic
samples became available (Diesmos et al. 2005). This
example also demonstrates the advantage of combining
molecular tools with active fieldwork and conventional
morphological techniques in resolving some of the most
perplexing issues at the interface of systematics and
conservation.
Nonnative species of freshwater turtles have already
established feral populations in various localities in the
Philippines because of either deliberate or accidental
releases through the pet trade. No comprehensive studies
on the ecological impact of nonnative species to Philippine
biodiversity currently exist, and this deficiency needs to be
addressed with urgency. The introduction of nonindigenous species into natural habitats can be prevented through
the enforcement (by wildlife authorities) of the national
policies on biodiversity and active monitoring of the
burgeoning pet trade industry. Educating consumers on the
probable hazards of introduction of alien invasive species
must complement these activities. Two nonnative species
have established breeding populations in the Philippines
(T. s. elegans and P. sinensis). One possibility, worthy of
consideration by relevant government agencies, is the
potential promotion of the Chinese softshell (P. sinensis)
as an income-generating species suitable for commercial
farming. The value of farming this species for its meat has
already been demonstrated. Pelodiscus sinensis could be
an important source of income for many rural villagers.
We encourage the commercial utilization of this species
because 1) it is nonnative to the Philippines and has the
potential to negatively impact native populations of turtles
if left unchecked in the wild, 2) this species has the
potential to generate much-needed revenue (as already
shown by the black market) that is otherwise lost to illegal
trade, and 3) farming of this locally nonthreatened and
commercially viable nonnative species may help ease the
pressure of exploitative harvesting of native species (see
172
AND
van Dijk et al. 2000). However, because farming of exotic
turtles often results in unintentional releases of animals
into habitats of native species, the impact of commercial
operations must be thoroughly evaluated before undertaking such high-risk endeavor.
Of the 6 species of indigenous freshwater turtles now
known from the Philippines, at least 4 are found in the
Palawan PAIC. Because this region is undergoing
persistent destruction of its natural habitats and illegal
trade of freshwater turtles is rampant (Widmann 1998;
Anda and Tabangay-Balderra 2004; Diesmos and Palomar
2004; Diesmos et al. 2004b, 2004c; Gavino and Schoppe
2004; Regodos and Schoppe 2005), ongoing conservation
efforts must center on curbing these factors. In particular,
widespread mining activities (now in every municipality of
the island) may severely threaten freshwater ecosystems of
Palawan. We urge relevant environmental authorities of
Palawan to develop a systematic program to actively
monitor known populations of threatened turtles, determine the effects of mining on Palawan’s wild turtle
populations, and halt the burgeoning black market trade
that stems from this island. Further, in order to be more
successful, this monitoring program should eventually be
integrated with the numerous environmental programs of
various government agencies and nongovernmental conservation groups operating in the region.
The unchecked exploitation of freshwater turtles and
other wildlife species in the illegal trade stems from a near
complete lack of enforcement and implementation of the
relevant national and international laws on wildlife
conservation and trade by Philippine wildlife authorities.
There are indications that government authorities are
generally unaware of the extent of illegal wildlife trade in
the country and are ill equipped and undertrained to
monitor and control this activity (Diesmos et al. 2004c;
Lepiten-Tabao and Tabaranza 2004; Regodos and
Schoppe 2005). Although data currently are not available
on the volume of freshwater turtles that are caught from
the wild and smuggled out of the Philippines, we
anticipate the figures to be astounding. For instance, only
a few months following the rediscovery of S. leytensis, we
noted the presence of live animals for sale at exorbitant
prices (US$1000–2000 per animal) throughout Asia and
Europe. A careful investigation of the dynamics of the
illegal trade and exploitation of freshwater turtles (and
other wildlife species) is critical for conservation and
management purposes. One recommendation might be to
elevate the protected status of S. leytensis such that it
would be included under Appendix I of CITES. We might
expect that this action would offer another level of
protection to wild populations and reduce illegal international trade of the species. However, given that trade and
export of CITES Appendix II species is already prohibited
by Philippine law (and yet trade continues), elevation to
higher protection levels may not produce the desired
effect. At present, we do not recommend captive breeding
as a conservation action for threatened freshwater turtles in
the Philippines. We believe that limited conservation
resources should instead be directed to strategies of higher
priority, such as in situ protection of species and their
natural habitat. However, we support the establishment of
‘‘assurance colonies’’ comprised of threatened species that
are presently being held in captivity by private wildlife
collectors and government-confiscated animals from the
illegal trade. Unfortunately, most of the animals that we
know of that have been confiscated in recent years have
died in captivity because wildlife law enforcers are
untrained and poorly equipped to care for animals.
Finally, we recommend the creation of a national
research and conservation program for Philippine freshwater turtles. Such a program could be responsible for
designing a management and conservation action plan for
species and the protection of critical habitats where
important species are known to occur. Further, a strategic
conservation plan of this sort could coordinate and
implement research, conservation programs, and public
education activities. We also believe that the conservation
of threatened turtles in the Philippines can be achieved if
wildlife collectors, traders, poachers, and other stakeholders are all considered as likely partners in such
conservation efforts.
ACKNOWLEDGMENTS
We thank the Protected Areas and Wildlife Bureau of
the Philippine Department of Environment and Natural
Resources for facilitating research protocols and granting
permits necessary for this and related studies. Support for
our herpetological work in the Philippines beginning in the
1990s was generously provided by the Society of
Systematic Biologists, US National Science Foundation
(DEB 0743491), the Explorer’s Club, Society for the
Study of Amphibians and Reptiles, American Society of
Ichthyologists and Herpetologists, Texas Memorial Museum, Charles Stearns Fellowships, the Conservation &
Environmental Grants of Ford Motor Company Philippines (‘‘Herp Watch Luzon 2000’’), BP Conservation
Programme (‘‘Herp Watch Palawan 2001’’ Project No.
1554), Declining Amphibians Population Task Force Seed
Grant (2002), Turtle Conservation Fund, Royal Melbourne
Zoological Gardens, North of England Zoological Society,
Conservation International (Philippines), Fauna and Flora
International, Rufford Small Grant for Nature Conservation (No. 171/07/04), the Natural History Museum and
Biodiversity Center of the University of Kansas, Silliman
University, and the National Museum of the Philippines.
We are indebted to Mae Diesmos, Genevieve Gee-Das,
Carlos Infante, Aldrin Mallari, Allan Altamirano, Nonito
Antoque, Peter and Indira Widmann, Sabine Schoppe,
Cameron Siler, Jason Diesmos, Jeffrey Diesmos, Siegfred
Diaz, Debbie Villafuerte, Rolito Dumalag, Snapper Poche,
Cling Peneyra, Flor Lopez, Cathy Gavino, Jude Dimalibot,
Sol Pedregosa, Juliet Ceniza, Paciencia Milan, Emerson
Sy, Boying Fernandez, and Enteng Yngente for the
DIESMOS
ET AL.
valuable assistance extended to us in the field. We thank
Indraneil Das, Roberto Puentespina, Maren Gaulke,
Michael de Guia, Edmund Leo Rico, Aloy Duya, Jayson
Ibañez, Sabine Schoppe, Peter Widmann, Carlo Custodio,
Esteven Toledo, Virgil Sagun, Jeremy Holden, and
William Oliver for providing important information and
photographs or sharing data. Bryan Stuart, Robert Webb,
James Parham, and Peter Paul van Dijk assisted us with
species identifications. For the loans of specimens and
provision of working space during museum visits, we
thank Jens Vindum, Michelle Koo (CAS), Alan Resetar,
Harold Voris, Robert Inger (FMNH), Travis La Duc,
David Cannatella (TNHC), and Kelvin Lim (ZRC). We
thank P.P. van Dijk, J. Parham, M. Gaulke, S. Schoppe, I.
Das, M. Diesmos, and Jen Weghorst for providing critical
reviews of the manuscript.
LITERATURE CITED
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hawksbill turtle in the central Visayas, Philippines. Fisheries
Research Journal of the Philippines 5(2):45–52.
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APPENDIX
Data on species distributions are partly based on specimens
housed at the National Museum of the Philippines (PNM), Field
Museum of Natural History (FMNH), California Academy of
Sciences (CAS), Raffles Museum of Biodiversity Research
(ZRC), Texas Natural History Collection of the Texas Memorial
Museum (TNHC), Natural History Museum of the University of
Kansas (KU), Cincinnati Museum of Natural History (CMNH),
Natur-Museum und Forschungs-Institut Senckenberg (SMF),
Muséum National d’Histoire Naturelle (MNHN), and the
zoological reference collections of the following Philippine
universities: Silliman University (Dumaguete City), University of
the Philippines (at Los Baños City and Davao City), University
of Santo Tomas (Manila), and Mindanao State University (at
Iligan City and Marawi City).
Dogania subplana. — Luzon Island: Camarines Sur
Province, Naga City, Barangay Concepcion Pequena (FMNH
251711); Laguna Province, Municipality of San Pablo, Lake
Bunot (ACD field obs., April 1999). Palawan Island: Palawan
Province, Municipality of Rizal, Barangay Campung Ulay, Sitio
Lolbokan (PNM 8230); Municipality of Quezon (unverified
report of Fidenci 2005); Municipality of Bataraza (ACD and
RVS field obs., November 2003). Mindoro Island: Oriental
Mindoro Province, Municipality of Socorro, near Lake Naujan
(PNM photo voucher); Municipality of Bongabong (PNM photo
voucher). Mindanao Island: Compostella Valley Province, Laac
(PNM photo voucher).
Pelochelys cantorii. — Luzon Island: Quirino Province,
Municipality of Nagtipunan, Barangay Sangbay, Sitio Maat-ad,
Upper Cagayan River Basin (PNM 8487); Cagayan Valley
Province, Municipality of Gattaran, Cagayan River (ACD field
obs., June 2006); Isabela Province, Ilaguen River (ACD field
obs., March 1995). Mindanao Island: Agusan del Sur Province,
Agusan Marsh Wildlife Sanctuary (PNM photo voucher); Davao
del Norte Province, Panabo City, Alejal (PNM photo voucher;
uncataloged specimen with Roberto Puentespina, D.V.M., Davao
City).
Pelodiscus sinensis. — Luzon Island: Bataan Province,
Municipality of Abucay (ACD field obs., March 1996); Bulacan
Province, Ipo Dam (PNM photo voucher); Metro Manila, Quezon
City, Parkway, Kamuning creek (PNM 892); Laguna Province,
Lake Bunot (zoological collection, University of the Philippines
176
AND
at Los Baños); Cavite Province (PNM photo voucher); Albay
Province, Municipality of Guinobatan (Arevalo, 2003); Camarines Sur Province, San Francisco River (Arevalo, 2003);
Batangas Province, Municipality of Talisay, Barangay Quiling,
Lake Taal (PNM 8260). Mindoro Island: Occidental Mindoro
Province; Siburan (ACD field obs., February 2002); Oriental
Mindoro Province, Polar River (PNM photo voucher). Panay
Island: Dinginan, Roxas City, Capiz Province (records from
Protected Areas and Wildlife Bureau, Department of Environment and Natural Resources). Cebu Island: Cebu Province,
Municipality of Talisay, Barangay Calumbangan (PNM 1427).
Leyte Island: San Juan and Saint Bernard, Southern Leyte
Province (ACD & RVS field obs., November 2003). Bohol
Island: Tagbilaran City (ACD field obs., April 2004). Mindanao
Island: Zamboanga City (PNM 8263).
Cuora amboinensis. — Luzon Island: Nueva Viscaya
Province, Municipality of Quezon, Barangay Maddiangat, Sitio
Dayog: KU 308932; Ilocos Norte Province, Municipality of
Burgos, Barangay Pao (PNM 917–922); ‘‘Zambales Province’’
(CAS 15376–82); Zambales Province, Municipality of Olongapo,
SBMA Naval Base, ‘‘Nav-mag’’ area, Nabasan River (TNHC
62989); Zambales Province, Municipality of Olongapo, ‘‘Subic
Bay’’ (CAS 15375); Cagayan Valley Province, Municipality of
Solana, Barrio Nabbutuan (PNM 6730); Isabela Province,
Municipality of San Mariano, Barangay Disulap, (PNM 658);
Municipality of Palanan, Barangay Didian, Northern Sierra
Madre Natural Park (PNM 6499); Aurora Province, Municipality
of San Luis and Maria Aurora, Barangay Villa Aurora, Aurora
National Park, Kabatangan River (PNM 5784); Aurora National
Park, Kamantis River (PNM 5661, 5784); Bulacan Province,
Municipality of Norzagaray, Barangay San Mateo, Sitio
Anginan, Ipo Dam (PNM 3578); Municipality of Los Baños
(CAS 61175–78); Laguna Province, Municipality of Los Baños,
Mt. Maquiling (TNHC 54903); Camarines Sur Province,
Municipality of Labo, Mt. Labo, Barangay Tulay na Lupa (KU
305164, 305166, 305561–62 plus 2 uncataloged specimens:
RMB 9876–77); Municipality of Jovellar, Barangay Bautista
(PNM 5005–06); Municipality of Bacacay, Barangay Nahapunan
(PNM 6072–73); Sitio Cauayan (PNM 4156); Sitio Limasan
(PNM 4231); Barangay Manangcas, Sitio Inuringan (PNM
4324); Municipality of Bulusan, Barangay San Roque (PNM
3495, 3557, 3573); Municipality of Sorsogon, Barangay
Guinajon (KU 300459, 300460). Polillo Island: Quezon
Province, Municipality of Burdeos, ‘‘nursery area’’ (PNM
5485–89); Municipality of Polillo, Polillo Town Watershed,
Barangay Pinaglubayan (KU 30056–58, 305144–47, 307449–
50); ‘‘Polillo Island’’ (CAS 62438–44). Cagraray Island: Albay
Province, Municipality of Bacacay, Barangay Manaet (PNM
6110, 6197–99). Verde Island: (ACD & RVS field obs.,
September 2003); Mindoro Island: Oriental Mindoro Province,
Municipality of Baco, Barangay Lantuyan, Dulangan River
(PNM 5640–41). Busuanga Island: Palawan Province (CAS
60549). Palawan Island: Palawan Province, Puerto Princesa
City, ‘‘vicinity of Malabosog Creek’’ (CAS 157295, 157466);
Municipality of Babuyan, ‘‘near sea level’’ (FMNH 52799);
Palawan Province, ‘‘NE of Puerto Princesa’’ (CAS 157467);
Municipality of Quezon, Barangay Carumpuro (ACD & RVS
field obs., November 2003). Panay Island: Aklan Province,
Municipality of Tangalan, Barangay Panayakan, Sitio Kalibungan (PNM 1565–67); Sitio Bongol (PNM 1890–93); Barangay
Lanyega, Sitio Buho’t-buwaya (PNM 1888–89, 1894–95);
Municipality of Ibajay, Barangay Bugtong-bato (PNM 913–
914); Antique Province, Municipality of Culasi, Barangay
Bagacay (PNM 1288), Municipality of Pandan, Barangay
Duyong (KU 300446); Guimaras Island: Iloilo Province, Daan
Banwa (CAS 185507); Municipality of Buenavista, Old
Poblacion (KU 30447–48); Municipality of Jordan, Barangay
Rizal (KU 30449); Pan de Azucar Island: Iloilo Province,
Talutoan (CAS 153872); Semirara Island: Antique Province,
Municipality of Caluya, Barangay Tinoboc (KU 300450–54);
Tablas Island: Romblon Province, Municipality of Odiongan,
Barangay Anahaw (2 uncataloged specimens deposited at KU:
RMB 8753, 8802); Municipality of Calatrava, Barangay Balogo,
Sitio Piqueno: 2 uncataloged specimen at KU (RMB 8807,
8857); Sibuyan Island: Romblon Province, Municipality of
Magdiwang, Barangay Silum: KU 300460, 305173; Barangay
Poblacion (KU 300456); Negros Island: Negros Oriental
Province, Municipality of Zamboanguita (CAS 133090–92);
Negros Oriental Province, Municipality of Dauin (CAS 134330);
Negros Oriental Province, ‘‘Tolong, Naliong’’ (FMNH 62907);
Negros Occidental Province, Municipality of Cuayan, Barangay
Camalandaan: 2 uncataloged specimens at KU (CDS GS 009,
101). Cebu Island: ‘‘Cebu Province’’ (CAS-SU 11356, 11437–
38). Bohol Island: Bohol Province, Municipality of Sierra
Bullones, Barangay Magsaysay (PNM 915–916); Bohol Province, Municipality of Sierra Bullones, Barangay Dusita (CAS
147526). Samar Island: Eastern Samar Province, Municipality
of Mercedes (CAS-SU 11442); Municipality of Bantayan, Sitio
Cadapnan (PNM 107); Municipality of Catbalogan (KU 305563);
Municipality of Taft, Barangay San Rafael (KU 310427–31).
Leyte Island: ‘‘Leyte Island’’ (KU 47173); Leyte Province,
Municipality of Palo (FMNH 44984); Municipality of Baybay,
Barangay Pilim, Sitio San Viscente (one uncatloged specimen at
KU: CDS 3473); Southern Leyte Province, Municipality of
Cabalian (CAS 60926–29). Dinagat Island: Surigao del Sur
Province, Municipality of Loreto, Barangay Panamaon, Sitio San
Ramon (PNM 2838), Barangay Esperanza (KU 305165, 305167,
305557–58, 310179). Camiguin Norte Island: Cagayan Province, Municipality of Calayan, Barangay Balatubat, area ¼
‘‘Limandok’’ (KU 304656–59); area ¼ ‘‘Mambit’’ (KU 304773–
74, 304778–79); Mindanao Island: ‘‘Zamboanga’’ (FMNH
14331; CAS-SU 11439–40); Lanao del Sur Province, Marawi
City, Lake Lanao (uncataloged specimens in Mindanao State
University natural history collection); ‘‘Misamis Occidental
Province’’ (CAS-SU 11324–37); North Cotabato Province,
Kidapawan City, Katituan (FMNH 52789); South Cotabato
Province, Municipality of Lake Sebu (David et al. 2006);
‘‘Cotabato Province, Buayan, Sputon’’ (FMNH 52794–96);
‘‘Cotabato Province, Bugasan, near sea level’’ (FMNH: 52764–
65); ‘‘Cotabato Province, Saub’’ (FMNH 121879–80); Davao del
Norte Province, Tagum City, Madaum, ‘‘near sea level’’ (FMNH
52766–86); Maco, ‘‘near sea level’’ (FMNH 52756–60); Davao
del Sur Province, Municipality of Malita, Lacayon, ‘‘near sea
level’’ (FMNH: 52797–98); Davao Oriental Province, Municipality of Mati, ‘‘sea level’’ (FMNH 52790–93); Mt. Hamiguitan
(PNM 108); Agusan del Sur Province, Agusan Marsh Wildlife
Sanctuary (PNM photo voucher).
Cyclemys dentata. — Busuanga Island: Palawan Province,
Dimaniang (FMNH 51600). Culion Island: Palawan Province,
San Pedro (PNM 891, 893). Dumaran Island: Palawan
Province, Municipality of Dumaran, Barangay Santo Tomas,
Sitio Umoy (Diesmos et al. 2004c). Palawan Island: Municipality of Taytay, Barangay Poblacion, near Lake Manguao (ACD
field obs., September 2002); Palawan Province, Puerto Princesa
City, Mt. Bloomfield (PNM 6276–77, 6281); Barangay Cabayugan, Sitio Martafe (PNM 6302); Barangay Tagabinit, Kayasan,
Lamod River (PNM 6388); Barangay Iwahig, Baguio (FMNH
51597); Barangay Iwahig (CAS 134332); Barangay Iwahig,
Iwahig watershed (KU 309079–80); S. slope of Thumb Peak,
Malatgan River (CAS 134331); Barangay Irawan, Bunton River
DIESMOS
ET AL.
(PNM 8261); Malabosog creek (CAS 157280); ‘‘Puerto Princesa’’
(FMNH 51596, 51598–99); Palawan Province, Municipality of
Narra, Barangay Estrella Falls: 4 uncataloged specimens
deposited in CMNH. Tawi-Tawi Island: ‘‘Languyan’’ (Gaulke
and Fritz, 1998).
Heosemys spinosa. — Tawi-Tawi Island: Tawi-tawi
Province Municipality of Languyan, Barrio Marang-Marang
(PNM 2232–33).
Siebenrockiella leytensis. — Palawan Island: Palawan
177
Province, Municipality of Taytay (PNM 8488); reportedly from
the Municipality of Aborlan (salvaged specimen: KU 309175);
Dumaran Island: Palawan Province, Municipality of Dumaran,
Barangay Santo Tomas, Sitio Umoy (Diesmos et al. 2004c).
‘‘Leyte Island: Municipality of Cabalian’’ (¼ type locality, in
error: Diesmos et al., 2004b) (CAS 60930; Neotype).
Received: 4 September 2006
Revised and Accepted: 2 September 2008

Status and Distribution of Nonmarine Turtles of the Philippines

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