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Feral Populations of Amphibians and Reptiles
in the Ryukyu Archipelago, Japan
Hidetoshi OTA1, Mitsuhiko TODA2, Gen MASUNAGA1, Akira KIKUKAWA3,
and Mamoru TODA1,4
1

Tropical Biosphere Research Center, University of the Ryukyus,
e-mail: ota@sci.u-ryukyu.ac.jp
2
Japan Wildlife Research Center
3
Kanagusuku Junior High School
4
Present address: Department of Zoology, Graduate School of Science, Kyoto University

Abstract
The Ryukyu Archipelago is an assemblage of many continental islands and a few oceanic islands
scattered between Kyushu of Japan, and Taiwan. Recent changes in geographic ranges were reviewed
for feral amphibian and reptile populations in this archipelago, chiefly on the basis of data and information becoming available after 1995. Six species of amphibians (Rana catesbeiana, Fejervarya sp., Polypedates leucomystax, Microhyla ornata, Bufo gargarizans miyakonis, B. marinus) and 14 reptiles(Chinemys reevesii, Cuora flavomarginata, Mauremys mutica, Trachemys scripta elegans, Pelodiscus
sinensis, Hemidactylus frenatus, Hemiphyllodactylus typus typus, Lepidodactylus lugubris, Anolis carolinensis, Ramphotyphlops braminus, Elaphe quadrivirgata, E. taeniura, Protobothrops elegans, P.
mucrosquamatus) are considered to have one or more feral populations in the Ryukyu Archipelago atpresent. Some biological aspects of these populations are also reviewed. Possible effects of such feral
amphibians and reptiles upon the indigenous biota, including a number of endemic and endangered taxa,
are briefly discussed.
Key words: amphibians, artificial transportation, feral population, Japan, reptiles, Ryukyu Archipelago

1. Introduction
The Ryukyu Archipelago is an assemblage ofapproximately 70 inhabited islands and some uninhabited islets located in the southwestern part of Japan.
The main part of this archipelago is composed of anisland chain extending from the southwestern side of
Kyushu to the northeastern side of Taiwan. This
Ryukyu Chain is usually divided into six island groups,
the Osumi Group, the Tokara Group, the Amami
Group, the Okinawa Group, the Miyako Group, andthe Yaeyama Group, in order from northeast to southwest (Kizaki, 1985). Supposedly, the land corresponding to this region first emerged as an eastern extensionof the Eurasian Continent during the Late Mesozoic
and then insularized with the subduction of the
Ryukyu Trough. This process is considered to havecommenced no later than the Late Miocene (Kizaki &Oshiro, 1980).
The remaining part of the archipelago includes the
Senkaku Group, an assemblage of several uninhabitedislands ca. 150 km north of the Yaeyama Group, andthe Daito Group, which is located ca. 320 km southGlobal Environmental Research8(2)/2004: 133-143
printed in Japan

east of the Amami Group (Kizaki, 1985). Of these, theSenkaku Group is situated on the eastern margin of
the current continental shelf and thus is considered tohave been connected to the continent during the Late
Pleistocene Glacial Maxima (ca. 15-20 thousand years
ago: Ota et al., 1993; Ota, 1998; but see Ota (2004a)
for a different view). In contrast, islands composing
the Daito Group are of pure oceanic origin and havenever been involved in a dry-land connection to otherland masses (Kizaki, 1985).
The fauna of the Ryukyu Archipelago is characterized by a high ratio of endemic taxa and geneticallydiverged populations, most of which have supposedly
been isolated from their relatives as a result of island
formation (e.g., Ikehara, 1996; Ota, 1998, 2000a). At
present, many of such Ryukyu endemics are seriously
threatened. For example, eight out of 17 species/
subspecies of amphibians (47.1%) and 13 out of 42
reptiles (31.0%) endemic to the Ryukyus are assigned
to one of the endangered categories (i.e., Endangered
IA, IB, and II) in the latest version of the Red List ofJapan (Environment Agency of Japan, 2000: also see
Ota (2000b)).
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Artificial habitat destruction and commerciallybased collecting have generally been considered themost serious threats to the survival of such endangered taxa of the Ryukyus (see Environment Agencyof Japan (2000) for cases of amphibians and reptiles).
However, several recent studies suggest that ecological and genetic impacts from artificially introduced
alien species also constitute another sort of serious
threats to the indigenous endangered taxa there (e.g.,
Ito et al., 2000; Murakami & Washitani, 2002).
Numerous alien organisms have been recorded in
the Ryukyu Archipelago, many of which have alreadyestablished breeding populations (Takehara et al.,
1997; Murakami & Washitani, 2002). The number ofsuch established alien species in the Ryukyus is everincreasing, and in many cases, their geographic ranges
also seem to be rapidly extending within thearchipelago. Data regarding such on-going dynamics
of the alien species are essential to the prediction of
their impact upon the native biodiversity and elaboration of effective measures for their control.
Some amphibians and reptiles, such as Bufo marinus and Boiga irregularis, are highly invasive in areas
where they have arrived by artificial means, having
serious negative impacts on native ecosystem andindigenous biodiversity (Rodda et al., 1999; Lever,2001). Ota (1999) reviewed the status of nonnative
amphibian and reptile populations of the Ryukyu Archipelago on the basis of data available at the end of
1995. In this article, we provide additional data forferal amphibian and reptile populations in this archipelago that have become available mostly from the
beginning of 1996 to the end of 2004, with remarks
chiefly regarding their impacts on the conservation ofnative biota. We also provide critical comments onand corrections to recent reviews as needed (Ota,
1999; Lever, 2003).
Following Ota (1999), we assumed establishmentof a feral population (i.e., a non-captive breeding colony deriving from artificially introduced individuals)of a given bisexual species on a given island if morethan ten individuals were found thereon during the last
30 years; more than five individuals including gravid
females or hatchlings were found on the island during
the last 30 years; or more than one individual was
found on the island during the last 30 years and morethan one inhabitant of the island, apparently having
reliable knowledge on local animals, verified that the
species had recently been occurring there. With respect to parthenogenetic species, successful colonization of a given island was assumed if more than one
animal was found there during the last 30 years.

2. Species Accounts
2.1 Amphibia
2.1.1 Rana catesbeiana
On the basis of data and information obtained by
the end of 1995, Ota (1999) referred to this large car-nivorous North American frog (e.g., Bury & Whelan,1984) as being possibly established on two islands of
the Amami Group, seven islands of the Okinawa
Group, and two islands of the Yaeyama Group (seeAppendix). However, our recent surveys on
Tokashikijima (in 2001 and 2002: H. Ota, unpublisheddata) and Tonakijima (in 1996, 2000, and 2001: H.
Ota, unpublished data) of the Okinawa Group, andIshigakijima (in 1996-2004: H. Ota, unpublished data)of the Yaeyama Group failed to obtain any evidence ofthe persistence of R. catesbeiana, such as its characteristic calls and tadpoles, nor was there informationfrom inhabitants regarding recent observations of thisprominently gigantic frog. It is thus likely that populations of this frog have already disappeared on theseislands. On the other hand, several individuals wereobserved and calls heard on Sesokojima of the
Okinawa Group (in 1997: Takahashi & Miyahira,1998; H. Ota, unpublished observation), and thisstrongly suggests that this island also has a feral population of this frog.
Referring to Ikehara (1974) as a source of information, Lever (2003) listed Kerama Retto as an island in
addition to Tokashikijima and six other islands of the
Okinawa Group (see above) having a population of R.
catesbeiana. Kerama Retto, in fact, however, refers toa small subgroup of islands within the Okinawa Group,in which R. catesbeiana has been recorded only from
Tokashikijima (Ota & Masunaga, 2004).
Recently Ota (2004b) confirmed an occasional
syntopy of this frog with a critically endangered snake,
Opisthotropis kikuzatoi, on Kumejima of the Okinawa
Group (Ota, 2000b). Outside Japan, there are a number of records of predation by R. catesbeiana uponsuch small-sized snakes as O. kikuzatoi (see Bury &Whelan (1984) for a review). One immature frog, collected from a site where O. kikuzatoi is frequently
observed (Ota, 2004b), had two small individuals of
the endangered freshwater crab, Candidiopotamon
kumejimense, in its stomach (H. Ota, unpublisheddata). Because the freshwater crab is the sole known
food item for O. kikuzatoi (Ota, 2004b), there are concerns that R. catesbeiana has been having a negative
effect on this critically endangered snake not only
through direct predation but also through competition
for food since its initial introduction to Kumejima in
1953 (Ota, 1999). Intensive field studies are strongly
desirable to examine this problem and devise effective
measures for eradication of R. catesbeiana from thenatural habitat of O. kikuzatoi.
2.1.2 Fejervarya sp.
This frog, presently called the ‘Sakishima numa
gaeru’ in Japanese (meaning ‘southern Ryukyu marsh
frog’), was formerly considered identical to thebroadly distributed paddy frog, Fejervarya limnocharis. However, because recent genetic studies have revealed distinct divergence of the southern Ryukyu
populations from the remaining East Asian popula-

Feral Populations of Amphibians and Reptiles in the Ryukyu Archipelagotions (Toda et al., 1997; Toda, 1999), Maeda andMatsui (1999) recognized this form as a distinct species but did not officially describe it.
On the basis of data available at the end of 1995,
Ota (1999) recognized feral populations of Fejervarya
limnocharis (sensu lato, as Rana limnocharis then: see
Dubois & Ohler (2000) for a discussion relevant to its
recent separation from Rana) on two islands of the
Daito Group, one island of the Miyako Group, andone island of the Yaeyama Group (see Appendix).
Toyama (1998) clarified that the frog, currently occurring on Yonagunijima of the Yaeyama Group in highdensities, is also nonnative, originating from an introduction from Ishigakijima in or before the early 1920s.
All these nonnative populations actually belong to thesouthern Ryukyu F. sp., not F. limnocharis sensu
stricto (Maeda & Matsui, 1999).
Ota (1999) surmised that this frog was first introduced to the Daito Group in the early 1970s. However,according to additional information becoming available after 1995 (Osawa & Osawa, 1997, personalcommunication; Okudo, 2000), F. sp. seems to havealready been introduced to the Daito Group before theWorld War II, most likely as early as 1921.
Lever (2003), citing from Ota (1999), stated that
“The nominate form of R. limnocharis (actually F. sp.:see above) was brought to Kohamajima andIriomotejima prior to the 1930s by tourists as souvenirs to attract inhabitants’ attention.” There is no evidence, however, of an exotic origin of these two island
populations nor such a statement in Ota (1999).
2.1.3 Polypedates leucomystax
Ota (1999) recognized establishment of feral populations of this originally Southeast Asian tree frog onfive islands of the Okinawa Group at the end of 1995
(see Appendix). Since then, this frog has been recorded from Iheyajima (first in 1999: Ota, unpublished data), Izenajima (in 1996: H. Ota, unpublisheddata), Iejima (in 2000: Kawauchi, 2002), and Tonakijima (in 2001: H. Ota and Ma. Toda, unpublishedobservation) of the Okinawa Group, Miyakojima (in1997: Iwanaga, 1998; Nohina et al., 1998), Irabujima(in 2002: Yoshigou et al., 2003), Kurimajima (sometime between November 1997 and March 2001:
Nakata, 2001) and Taramajima (in 2002: Shiroma &Ota, 2004) of the Miyako Group, and Ishigakijima(some time between 1995 and 2001: Japan Wildlife
Research Center, 2001) of the Yaeyama Group. Of
these islands, Ishigakijima may not have an established population yet, because the record derives froma single observation (Japan Wildlife Research Center,2001), and no other recent surveys on this island haveyielded any evidence of its occurrence (Mi. Toda, unpublished data). On the other islands, a number of individuals including calling males, or more than one
egg mass characterized by a foamy outer layer, or both,have been observed, and it is almost certain that the
species has already established breeding colonies onthese islands.

135

The impact of this species upon the native ecosystem remains to be investigated. Hasegawa (1993)
found helminth parasites in the gonads of male P.
leucomystax from Okinawajima that were completely
unknown in the native species despite recent intensive
surveys of the helminth fauna parasitizing Ryukyu
amphibians by his colleagues and himself (seeHasegawa & Asakawa (2004) for a review). There isconcern that this parasite most likely entered the
Ryukyus from Southeast Asia through an accidental
introduction of P. leucomystax in the early 1960s (Ota,
1999) and will have negative effects on native Ryukyu
amphibians through infection.
2.1.4 Microhyla ornata
Ota (1999) recognized a feral population of thisspecies on Suwanosejima of the Tokara Gruop,
Taramajima of the Miyako Group, and Kuroshima of
the Yaeyama Group. Referring to Ota (1983, 1999),
Lever (2003) wrote that M. ornata was also successfully introduced to Yonagunijima of the Yaeyama
Group. Actually, however, there are no such records inthese and other publications (Ota, 1999). Moreover,recent surveys of Yonagunijima by H. Ota yielded noevidence of the occurrence of M. ornata (H. Ota, unpublished data collected in 1998 and 2001). Thus,
Lever’s (2003) statement as above is obviously inerror.
2.1.5 Bufo gargarizans miyakonis
Ota (1999) recognized feral populations of B. g.
miyakonis in the northern part of Okinawajima, as
well as on two islands of the Daito Group (see Appendix). Since 1999, however, no individuals have beenfound at the northern Okinawajima site, where thistoad was observed in fairly high densities in the
mid-1980s (M. Toyama & Y. Chigira, private communication; H. Ota, unpublished data). Such a recent decline is probably a consequence of the enthusiastic
eradication effort by M. Toyama, Y. Chigira, and theircolleagues and inhabitants from the late 1980s to the
middle 1990s. Indeed, a large number of toads were
hand-captured, and eggs and larvae removed frombreeding ponds several times (M. Toyama & Y.
Chigira, private communication). On the two islandsof the Daito Group, B. g. miyakonis seems even now
to maintain dense populations (H. Ota, unpublisheddata).
2.1.6 Bufo marinus
Ota (1999) recognized established populations of
this highly invasive toad (Lever, 2001) from two
islands of the Daito Group, and Ishigakijima of the
Yaeyama Group (see Appendix). A survey on Hatomajima of the Yaeyama Group in 2001 by G. Masunaga
yielded four specimens of adult B. marinus (given to
the Zoological Collection of Kyoto University
Museum as KUZ R52784-787). Results of interviewsof seven inhabitants then indicated that Hatomajima
had an established population of this toad and that it

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originated from an introduction of no more than ten
individuals from Ishigakijima in the late 1980s or
early 1990s by a school teacher. Information from inhabitants also suggests that the population density,
though once remarkably raised, has been dropping,most likely as a result of inhabitants’ efforts to control
it by killing individuals when encountered and minimizing surface water available for its reproduction.
Bufo marinus has also been found on Iriomotejimaof the Yaeyama Group no less than 30 times involving
at least 24 individuals, since the capture of one individual at the mouth of the Urauchigawa River in 1986
(Japan Wildlife Research Center, 2003, 2004). These
records are considered to represent accidental entries
of individuals, most likely in association with construction materials transported from Ishigakijima. To
date, there has been no evidence of reproduction ofthis toad on this island. From the cases at twoneighboring islands (Ishigakijima and Hatomajima:
Ota, 1999; see above), however, establishment ofdense population of this toad on Iriomotejima ishighly likely once even a small number of individuals
start to reproduce successfully.
In some areas outside Japan where B. marinus has
successfully colonized, considerable impacts uponnative ecosystems are reported (Lever, 2001). Relevant data have yet to be collected and analyzed withrespect to those islands of the Ryukyus that alreadyhave feral populations of this toad. A mass death ofdomestic ducks, once occurring on Hatomajima, isbelieved by some inhabitants of the island to havebeen caused by poison of toadlets that were eaten by
ducks around a breeding pond (T. Toji, private communication). This case deserves particular attention in
the context of economic damage caused by this toad,although the effect of toadlet poison on ducks stillneeds careful verification.
2.2 Reptilia
2.2.1 Chinemys reevesii
This freshwater species is originally known only
from the main islands of Japan, Taiwan, and the eastern continent (Iverson, 1992). However, Masuno et al.
(1998) collected five individuals, including two hatchlings, of C. reevesii from a pond in the south-central
part of Okinawajima, and also referred to another individual observed in a nearby river. Also, two individuals were collected from the same pond in 2003 (H. Ota,
unpublished data). It is thus highly likely that this species has already established a breeding population, atleast in the south-central part of Okinawajima. A few
individuals of C. reevesii have also been found on
Kumejima of the Okinawa Group (Ota & Hamaguchi,
2003), but its establishment of a breeding colony on
this island remains to be studied.
Masuno et al. (1998) also demonstrated that on
Okinawajima hatchlings of this species are commonly
sold as pets. They thus surmised that the putative feral
population there had been derived from such pet indi-viduals, as in the case of Trachemys scripta elegans(see below). The impact of this omnivorous freshwater
turtle upon the native ecosystem of Okinawajima remains to be studied.
2.2.2 Cuora flavomarginata
Two subspecies, C. f. flavomarginata from Taiwan
and the eastern continent, and C. f evelynae from the
Yaeyama Group, are recognized for this species(Yasukawa & Ota, 1999). However, because the rangeof morphological variation more or less overlaps
between these subspecies (Yasukawa & Ota, 1999), it
is almost impossible to identify with certainty eachindividual captured outside the native geographic
range at the subspecies level. Therefore, we use only
the species name here and defer the problem regarding
the subspecific allocation of each individual or putative feral population to future investigations.
As is mentioned above, the natural distribution ofthis terrestrial species in Japan is confined to Ishigakijima and Iriomotejima of the Yaeyama Group. However, it has occasionally been brought to Okinawajima
since as early as the 19th Century (Fritze, 1894).
Recently, in particular, turtles of putative Taiwaneseorigins have been sold in pet shops on this island (e.g.,
Masuno et al., 1998), and one may sometimes discover individuals in the field that have been released,or have escaped from captivity.
Ota (1999) indeed referred to a few records of C.
flavomarginata, as well as its putative hybrids with anative turtle Geoemyda japonica, on Okinawajima
between the early 1980s and the early 1990s. However,
he deferred a conclusion regarding its establishment
on Okinawajima to future studies. In 1996, 1999 and2001, a total of five turtles were captured in the southern part of Okinawajima (H. Ota, unpublished data),
but all were found in the urban areas of Naha City andUrasoe City and thus are considered as escaped orreleased individuals rather than representatives of aferal self-breeding population. Therefore, establishment of this turtle on Okinawajima still remains to becarefully examined.
Recently, C. flavomarginata has also been recorded on Kumejima of the Okinawa Group (from the
middle 1990s to the early 2000s: Ota & Hamaguchi,
2003), Miyakojima of the Miyako Group (in 2003:
Oka, 2004; Shiroma & Ota, 2004), and Kuroshima of
the Yaeyama Group (in 2001 and 2003: H. Ota, unpublished data; N. Kamezaki, private communication;
information from three anonymous inhabitants).
Establishment of a feral population remains uncertain
for Kumejima, but is highly likely for the latter two
islands, considering the number of individuals
observed in a single year (five in 2003: Miyakojima)
and information from inhabitants (arguing that the
turtle is not so rare: Kuroshima).
Emergence of putative hybrids between C.
flavomarginata and G. japonica (Ota, 1999) predicts
an occurrence of introgression into the latter species
once C. flavomarginata has successfully colonizedFeral Populations of Amphibians and Reptiles in the Ryukyu Archipelago

Okinawajima or Kumejima where G. japonica occurs
as the only native turtle species (Ota & Hamaguchi,
2003). The impact of this omnivorous terrestrial turtle
on the native ecosystem of an island to which it has
successfully colonized remains to be studied.
2.2.3 Mauremys mutica
As in the case of C. flavomarginata, two subspecies are recognized of this species, M. m. mutica from
Taiwan and the eastern continent, and M. m. kamifrom the Yaeyama Group (Yasukawa et al., 1996). As
in the case of C. flavomarginata, however, the rangeof morphological variation more or less overlaps
between these two subspecies (Yasukawa et al., 1996),
making it occasionally difficult to identify with certainty each individual captured outside the native geographic range at the subspecies level. Therefore, weonly use the species name here and defer the problem
regarding the subspecific allocation of each individual
or putative feral population to future investigations.
As is mentioned above, in Japan the original rangeof M. mutica was supposedly confined to Ishigakijima,
Iriomotejima, and Yonagunijima of the Yaeyama
Group (Yasukawa et al., 1996). On the basis of dataand information obtained by the end of 1995, Ota(1999) recognized feral populations of this species on
two islands of the Okinawa Group and one island of
the Miyako Group (see Appendix). He also referred torecords of M. mutica from two other islands of the
Okinawa Group (Sesokojima and Zamamijima), butwith no information regarding its population status on
these islands (Ota, 1999).
Since then, seven individuals, including two hatchlings, have been found on Sesokojima (in 1998-1999:
H. Ota unpublished data), and this suggests establishment of a feral population of M. mutica on this island.
This turtle has also been found on Iheyajima (in 2002:
Honda et al., 2002; H. Ota, unpublished observation),
Tonakijima (in 1998: H. Ota, unpublished observation), Kumejima (some time between the middle
1990s and the early 2000s: Ota & Hamaguchi, 2003)of the Okinawa Group, Kurimajima of the Miyako
Group (in 2003: Shiroma & Ota, 2004), andHaterumajima of the Yaeyama Group (in 1999 and2004: K. Kinjo, private communication; Ota et al.,2004). Furthermore, Osawa and Osawa (1997), in
their photographic guide to the nature of Minamidaitojima, mentioned the presence of turtles, most likely
representing this species, on this island. On Haterumajima, M. mutica occurs in high densities in somecanals, and this obviously reflects establishment of abreeding colony on this island (Ota et al., 2004). The
status of this species on the other islands remains to bestudied.
Shiroma and Ota (2004) were concerned about theeffect of this omnivorous turtle upon the endemic,
highly endangered freshwater crab (Geothelphusa
miyakoensis) on Miyakojima. However, there arealmost no actual data on the impact of M. mutica on
the indigenous species of islands to which it has been137

introduced.
2.2.4 Trachemys scripta elegans
On the basis of data and information obtained by
the end of 1995, Ota (1999) recognized a feral population of this North American turtle on Okinawajima of
the Okinawa Group. He also referred to records of T. s.
elegans from Amamioshima of the Amami Group andtwo islands of the Yaeyama Group, but deferred a conclusion regarding its status on these three islands tofuture studies due to insufficient data available at that
date. Lever (2003) surmised that this turtle had already established a feral population on Amamioshima,
but without showing any additional data.
Since then, this turtle has been recorded from
Iejima (in 2002: Toyama, 2002) and Kumejima of the
Okinawa Group (some time between the mid-1990s
and early 2000s: Ota & Hamaguchi, 2003),
Minamidaitojima of the Daito Group (some time
between the late 1980s and the mid-1990s: Osawa &Osawa, 1997; Okudo, 2000), and Miyakojima of the
Miyako Group (in 2003: Oka, 2004; Shiroma & Ota,
2004). On Kumejima, Minamidaitojima, and Miyakojima, more than ten individuals were seen within small
areas (Ota & Hamaguchi, 2003; H. Ota, unpublishedobservations), and it is almost certain that this turtle
has already established feral populations on theseislands. Its population status on Iejima and Kumejima
remains to be studied.
Masuno et al. (1998) reported that in Okinawa Prefecture juvenile T. s. elegans were sold in large numbers at all pet shops they visited. It is thus likely thatthe putative feral populations in this region (seeabove) have originated from such pet individuals
through accidental escapes or intentional releases as inmany other regions of various countries (Lever, 2003).
Various types of impacts on the indigenous biodiversity have been shown for feral populations of T. s.
elegans outside Japan (see Lever (2003), and Cadi &Joly (2004), for a review). However, no pertinent data
are available yet for the Ryukyu populations.
2.2.5 Pelodiscus sinensis
On the basis of data and information obtained by
the end of 1995, Ota (1999) recognized feral populations of this soft-shelled turtle from two islands of the
Okinawa Group and two islands of the Yaeyama
Group (see Appendix). Since then, data have beencollected in large quantities on the origin and distribution of P. sinensis within the Ryukyu Archipelago(Sato et al., 1997; Sato & Ota, 1999). Analyses andsyntheses of these data indicate that: (1) breeding
populations of this turtle also occur on Kikaijima,
Amamioshima and Tokunoshima of the Amami Group,
Iheyajima of the Okinawa Group, Minamidaitojima of
the Daito Group, and Iriomotejima of the Yaeyama
Group; (2) none of these or other Ryukyu populations
are native; (3) sources of introductions differ between
the islands of Okinawa Prefecture (Okinawa, Daito,
and Yaeyama Groups) and those of Kagoshima Pre-

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fecture (Amami Group) (Taiwan for the former andthe main islands of Japan for the latter); and (4) introductions were carried out chiefly or exclusively forthe purpose of farming this turtle for food (Sato et al.,
1997; Sato & Ota, 1999). The result of questionnaire
surveys of inhabitants also suggests that most such
introductions were carried out during a period fromthe early 1950s to the late 1970s (Sato et al., 1997;
Sato & Ota, 1999). It is obvious from information
given in old newspapers, however, that on Okinawajima and Ishigakijima some individuals were released
earlier than 1909 and 1931, respectively (Toyama,
1998).
Besides those islands, Kitadaitojima of the Daito
Group (S. Shokita, private communication), andKohamajima (Tanaka, 2004) and Haterumajima of the
Yaeyama Group (Ota et al., 2004) may also have feral
populations of P. sinensis. Further surveys on theseislands are thus much desired.
Because of the putative predatory nature of P.
sinensis, impact of its feral populations upon theindigenous freshwater fauna is of much concern(Lever, 2003). Nevertheless, no concrete data are
available regarding the feeding habits of this species
in the field. Further studies are thus strongly desired
regarding this aspect of feral populations on eachcolonized island.
2.2.6 Hemidactylus frenatus
Ota (1999) did not refer to the Ryukyu populationsof Hemidactylus frenatus as nonnative. However, several lines of evidence, such as the geographic genetic
pattern (Moritz et al., 1993), temporary changes inknown ranges (Hunsaker, 1966; Case et al., 1994),
and the absence of populations on many uninhabitedislands (see Kohno & Ota (1991) and Ota et al. (1993)
for examples within the Ryukyu Archipelago),strongly suggest that the original range of H. frenatus
was confined to South and Southeast Asia and that its
current populations on most Oceanian and East Asianislands including those of the Ryukyus have derived
from recent human-associated dispersals (Lever,2003). Because Stejneger (1907) referred to a specimen collected on Okinawajima in 1854, however, it isalmost certain that H. frenatus had already colonized
to the Ryukyu Archipelago by the middle of the 19th
Century.
In the Ryukyu Archipelago, the distributional
range of H. frenatus had long been confined to islands
south of Amamioshima and adjacent islets (Ota, 1989).
Recently, however, this species has started to appear
on Kikaijima (in 1991: Ma. Toda, unpublished observation) and Amamioshima (in 2000: Kuze & Ota,
2001) of the Amami Group. Because more than ten
individuals were observed in Nase on Amamioshima
and Wan on Kikaijima in 2004 (H. Ota, unpublishedobservations), it is obvious that this species has
already established breeding colonies on these islands.
Considering that Tokunoshima, an island approximately 50 km southeast of Amamioshima, had long

been the northernmost range extremity of H. frenatus(Ota, 1989), such abrupt range extensions during thelast decade are puzzling. As of the late 1980s, several
authors suspected that low winter temperatures had
been playing a crucial role in preventing northward
colonization of this originally tropical human commensal (see Ota (1989) and papers cited therein). If
this is the case, then, such northeastward colonizationof this species may be attributable to the recent rise inglobal air temperatures (e.g., Domoto et al., 2000).
This needs substantial verification by examining therelationships between annual fluctuations of low winter temperatures and population sizes of H. frenatus on
these islands, and also by comparing low temperature
tolerance between H. frenatus and other gekkonid species showing greater northward extensions of ranges.
In the Ryukyu Archipelago, colonization by thisgecko may have been causing declines in a few indigenous geckos through competition (e.g., Ota, 1989),
as in some Pacific islands which H. frenatus has also
recently colonized (Case et al., 1994). Future studies
are needed to verify this possibility.
2.2.7 Hemiphyllodactylus typus typus
On the basis of data obtained by the end of 1995,
Ota (1999) assumed successful colonization of thisgecko on Miyakojima of the Miyako Group and
Iriomotejima of the Yaeyama Group. Since then,Hemiphyllodactylus typus typus has also been recorded on Taramajima of the Miyako Group (in 1998:
Kikukawa & Toda, 1998), and Ishigakijima (in 1999:
Japan Wildlife Research Center, 2001) and Haterumajima (in 2004: Ota et al., 2004) of the Yaeyama Group.
However, because only one individual has been confirmed on each of these three islands, the status of thisgecko on these islands remains uncertain. In 2004,
Tanaka (2004) also observed an individual gecko,
most likely belonging to H. t. typus, on Kohamajimaof the Yaeyama Group. This record, however, needs
verification on the basis of voucher material.
Hemiphyllodactylus t. typus is broadly distributed
among the insular and coastal regions of Southeast
Asia and Oceania (Bauer & Henle, 1994), and individuals recently found in the southern Ryukyus most
likely derive from recent accidental entries from one
or more localities within these regions (Ota, 1990;
Lever, 2003). Because all Ryukyu specimens hitherto
collected have been females (n=12: H. Ota, unpublished data), assemblages in this region are likely to beparthenogenetic, like those in the southern Pacificislands (Zug, 1991). The impact of colonization of H. t.
typus upon the indigenous biota remains to be studied.
2.2.8 Lepidodactylus lugubris
In the Ryukyu Archipelago, this parthenogenetic
gecko currently occurs in the southern half of the continental island chain (i.e., the Okinawa, Miyako, andYaeyama Groups) and the oceanic islands of the Daito
Group (Ota, 1999). Assemblages on the latter islands,
known since the middle 1930s, include a variety ofFeral Populations of Amphibians and Reptiles in the Ryukyu Archipelago

endemic clones and thus are obviously native to thisregion (Yamashiro et al., 2000). In contrast, assemblages in the Ryukyu Chain, first recorded in the early
1970s, consist solely of a single clone with a broad extralimital distribution and thus are considered to haveoriginated from recent accidental human transportation (Yamashiro et al., 2000; Ota et al., 2004).
On the basis of data and information available atthe end of 1995, Ota (1999) tentatively referred to L.
lugubris on two islands of the Okinawa Group, twoislands of the Miyako Group, and six islands of the
Yaeyama Group as already established (see Appendix).
Subsequent surveys have yielded data that indicate
establishment of successfully reproducing assemblages of this clonal gecko on Akajima (in 1996-2003:
H. Ota, unpublished data), Zamamijima (in 1996:
Yamashiro et al., 2000), Kubajima (in 2001 and 2002:
Ota & Masunaga, 2004) and Yakabijima (in 1999:
Yamashiro et al., 2000) of the Okinawa Group,
Ogamijima (in 1997: Nohina et al., 1998) and Taramajima (in 1998: Yamashiro et al., 2000) of the Miyako
Group, and Kohamajima (in 2004: Tanaka, 2004) of
the Yaeyama Group. As in the case of H. t. typus, theimpact of colonization of L. lugubris on the indigenous biota remains to be studied.
2.2.9 Anolis carolinensis
Ota (1999) referred to a dense population of A.
carolinensis, which was found in a limited area (Shuri,
Naha City) in the southern part of Okinawajima in
1994 (also see Ota et al., 1995). This population may
have declined subsequently, because a survey carriedout at this site in 2004 yielded no individual lizards atall (H. Ota, unpublished data). However, this assumption needs careful verification.
In August 2001, three adult males were collected(KUZ R52571-52573) and three other males wereobserved at Kanagusuku (Naha City), a site approximately 6 km southwest of Shuri (H. Ota, A. Kikukawa,& Ma. Toda, unpublished data). The population statusaround Oroku also needs careful assessment on thebasis of additional data. Serious efforts at eradication
should be made if the resultant data indicate the presence of a breeding population.
The impact of colonization of A. carolinensis on
the indigenous biota of Okinawajima remains to bestudied. Nevertheless, serious negative effects uponsome indigenous lizards through competition andupon some insects through predation are highly likely,
judging from the situation in the Ogasawara Archipelago, where feral A. carolinensis populations occur inhigh densities (e.g., Suzuki & Nagoshi, 1999; Karube& Suda, 2004; Takakuwa & Suda, 2004).
2.2.10 Ramphotyphlops braminus
In the Ryukyu Archipelago, this parthenogenetic
blind snake has long been known from many islands
south of Suwanosejima of the Tokara Group (Ota etal., 1991; Hikida et al., 1992). In 2000, however, several individuals were found in a limited area of139

Tanegashima of the Osumi Group (Kazuki, 2001).
This strongly suggests that the species was recently
introduced to this island and has already started reproduction successfully. The impact of colonization of R.
braminus upon the indigenous biota of Tanegashimaremains to be studied. Also, the origin of current assemblages of this skillful colonizer (Lever, 2003) onother Ryukyu islands needs careful investigation.
2.2.11 Elaphe quadrivirgata
This snake, originally endemic to the main islandsof Japan and a few adjacent islets, has a feral population on Kuchinoshima of the Tokara Group. Nagai(1938) stated that this population is derived from an
artificial introduction from Kuchinoerabujima of theOsumi Group (also see Hikida et al., 1992). The impact of its colonization upon the native Kuchinoshimabiota remains to be studied.
2.2.12 Elaphe taeniuraThis polytypic species originally occured in the
Miyako and Yaeyama Groups, Taiwan, the eastern andsoutheastern continent, and some Southeast Asianislands (Schultz, 1996). On the basis of data available
at the end of 1995, Ota (1999) surmised that there isan established feral population of E. taeniura in thesouth-central part of Okinawajima, and that it has
most likely been derived from individuals artificiallyintroduced from Taiwan or the eastern continent. In
2003, two individuals were found from the northern
forested part of Okinawajima (T. Otani, personal communication). The impact through predation of thissnake on the indigenous mammals and birds, mostly
concentrated in this part of the island (Ito et al., 2000),is of great conservation concern (Murakami &Washitani, 2002).
The name of the endemic Taiwanese subspecies, E.
t. friesi, is usually applied to the feral population of
Okinawajima (e.g., Otani, 1993, 1998; Murakami &Washitani, 2003). Indeed, specimens from Okinawajima are morphologically distinct from several othersubspecies, including E. t. schmackeri, in the southern
Ryukyus (H. Ota, unpublished data). However, it still
remains unclear as to whether they can be identified as
the Taiwanese or eastern continental subspecies (i.e.,
E. t. taeniura or E. t. mocquardi: H. Ota unpublisheddata). We thus refer to the Okinawajima population
simply as E. taeniura here.
Utiger et al. (2002) described a new genus,
Orthriophis, on the basis of molecular phylogenetic
analysis of the genus Elaphe sensu lato, to accommodate E. taeniura and a few other species. However,
because the bootstrap support to the validity of thisnew genus is not necessarily sufficient, we here take aconservative stance on this matter.
2.2.13 Protobothrops elegans
This species, as well as the next venomous snake,
was previously referred to as a member of Trimeresurus, a genus actually consisting almost exclusively of

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H. OTA et al.

Southeast Asian arboreal species (e.g., Malhotra &Thorpe, 2004). Although it was originally endemic to
the Yaeyama Group, P. elegans currently also occurs
in high densities around Itoman City in the southernpart of Okinawajima (e.g., Nishimura & Akamine,2002). The impact of colonization of this species upon
the indigenous biota of Okinawajima remains to bestudied.
2.2.14 Protobothrops mucrosquamatus
This venomous snake, originally distributed in
Taiwan and the southeastern continent, has recently
been recorded in Nago City in the north-central part of
Okinawajima (first in 1995: Akamine & Nishimura,1998). It is highly likely that there is a dense population of this species around Bi-mata in the city at present (e.g., Terada, 2003). Impact of colonization of P.
mucrosquamatus upon the indigenous biota of
Okinawajima remains to be studied.
2.3 Other species possibly having feral populations in the Ryukyu Archipelago
2.3.1 Geoemyda japonica
The known range of this turtle has long been confined to Okinawajima, Tokashikijima, and Kumejimaof the Okinawa Group. Recently, however, a few individuals were found on Gerumajima of the Okinawa
Group (Ota & Hamaguchi, 2003; Watanabe, in press).
According to several inhabitants, this turtle, though
not originally occurring on Gerumajima, has beenoccasionally observed on this island during the lastfew decades (H. Ota, unpublished information). Thus
its occurrence on Gerumajima as a feral population ishighly likely, although this needs future verification.
2.3.2 Gehyra mutilata
Lever (2003) surmised that the Ryukyu populations of this species were of exotic origins. He statedthat Toyama (1984) recorded G. mutilata as a recent
arrival on Okinawajima. Actually, however, there is nosuch statement in this report. Indeed, the species wasrecorded in the Ryukyu Archipelago earlier than the
beginning of the 20th Century (Stejneger, 1907). We,
therefore, tentatively retain this species as native to
the Ryukyu Archipelago, and defer the problem tofuture studies.

Acknowledgements
We thank Yoshinori Chigira, Kazumitsu Kinjo,
Maiko Okawa, Yushi and Keiko Osawa, Tsutomu
Otani, Fumiyasu Sato, Tsunehiro Shiroma, Shigemitsu
Shokita, Kenji Toji, Tsutomu Toji, Masanao Toyama,
and many other inhabitants of the Ryukyu islands for
pertinent information. Special thanks are due Masako
Izawa for critically reviewing the manuscript. We arealso much indebted to Tsutomu Hikida, Masafumi
Matsui, and Masatoshi Takakuwa for literature.
Data collecting was partially supported by Grants-

in-Aid from the Ministry of Environment, Japan (as aGlobal Environment Research Programme; Project
Leader: Koichi Goka), Research Institute for
Humanity and Nature (Iriomote Project; Project
Leader: Tokushiro Takaso), and Monbu-kagaku-sho (=the Ministry of Education, Sports, Culture, Science,
and Technology, Japan) (B-14380277: Project Leader:
Masako Izawa). Manuscript preparation was also
partially supported by a Grant-in-Aid to the 21stCentury COE Project at the University of the Ryukyus
from Monbu-kagaku-sho.
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(Received 25 December 2004, Accepted 31 December 2004)

Feral Populations of Amphibians and Reptiles in the Ryukyu Archipelago
Appendix.

143

List of islands of the Ryukyu Archipelago, where occurrence of nonnative amphibian or reptile population is assumed.
TG, Tanegashima. KN, Kuchinoshima. SW, Suwanosejima. AM, Amamioshima. KK, Kikaijima. YR, Yorojima, TK.
Tokunoshima. IH, Iheyajima. IZ, Izenajima. IE, Iejima. OK, Okinawajima. MN, Minnajima. SS, Sesokojima. YB,
Yabuchijima. YG, Yagajijima. IK, Ikeijima. TS, Tokashiki. AK, Akajima. ZM, Zamamijima. KM, Kumejima. MD,
Minamidaiotojima. KD, Kitadaitojima. MY, Miyakojima. KU, Kurimajima. OG, Ogamijima. IB, Irabujima. TR,
Taramajima. IG, Ishigakijima. TT, Taketomijima. KR, Kuroshima. KH, Kohamajima. HT, Haterumajima. HM,
Hatomajima. IR, Iriomotejima. YN, Yonagunijima.

Species
Rana catesbeiana

Osumi Gr.

Tokara Gr.

Amami Gr.
YR*, TK*

Fejervarya sp.
Polypedates
leucomystax
Microhyla ornata
Bufo gargarizans
miyakonis
B. marinus
Chinemys reevesii
Cuora
flavomariginata
Mauremys mutica

Daito Gr.

Miyako Gr.

Yaeyama Gr.
IG*#, KH*

MD*, KD*TR*
MY, IB, KU,
TRKR*, YN**

TR*

KR*

OK*, SS*, YB*,
YG*, IK*, IH,
IZ, IE, TN
SW*
OK*#

MD*, KD*
MD*, KD*

IG*, HMOK

Trachemys scripta
elegans
Pelodiscus sinensis
Hemidactylus
frenatus
Hemiphyllodactylus
typus typus
Lepidodactylus
lugubris
Anolis carolinensis
Ramphotyphlops
TGbraminus
Elaphequadrivirgata
E. taeniuraProtobothrops
elegans
P. mucrosquamatus

Okinawa Gr.
IH*, IZ*, IE*,
OK*, TS*#,
TN*#, KM*, SS

MY

KR
HT

OK*, AK*, SS,
ZM
OK*, KM

MD?

MY*

MD

MY

AM, KK, TK

OK*, KM*, IH

MD, KD?

Most inhabitedislands**

All inhabitedislands**

All inhabitedislands**

IG*, YN*,
IR, HR?
All inhabited All inhabitedislands**
islands**
MY*
IR*

OK*, MN* AK,
ZM, KB, YK

MY*, KU*,
OG , TR

OK*
KN
OK*
OK*
OK

* Given in Ota (1999).
# A population, though once established, seems to have disappeared.
** Populations were already known but were not regarded as nonnative at Ota’s (1999) date. See text.

IG*, TT*,
KR*, IR*,
HR*, YN*,
KH