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Wildlife Damage Management Conferences --
Proceedings
Wildlife Damage Management, Internet Center for
1-1-2005
Challenges of Invasive Reptiles and Amphibians
William Pi
USDA, APHIS, Wildlife Services, National Wildlife Research Center, Hilo, HI, USA
Daniel Vice
USDA, APHIS, Wildlife Services, Barrigada Heights, Guam, USA
Mike Pitzler
USDA, APHIS, Wildlife Services, Honolulu, HI, USA
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Pi, William; Vice, Daniel; and Pitzler, Mike, "Challenges of Invasive Reptiles and Amphibians" (2005). Wildlife Damage Management
Conferences -- Proceedings. Paper 84.
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112
CHALLENGES OF INVASIVE REPTILES AND AMPHIBIANS
WILLIAM C. PITT, USDA, APHIS, Wildlife Services, National Wildlife Research Center, Hilo,
HI, USA
DANIEL S. VICE, USDA, APHIS, Wildlife Services, Barrigada Heights, Guam, USA
MIKE E. PITZLER, USDA, APHIS, Wildlife Services, Honolulu, HI, USA
Abstract: Although worldwide distributions of many amphibians and reptiles are declining, a
handful of species are spreading rapidly throughout tropical regions of the world. The species
that have the greatest effect tend to be generalist feeders, have high reproductive rates, attain
large population sizes, and often due to their behavior and or small size, are easily transported or
are difficult to detect. The most notable of these species include the coqui frog, cane toad,
bullfrog, brown tree snake, and Burmese pythons. The effect of a few individuals typically is
small but the combined effect of large populations can be devastating to ecological communities
and agriculture. Currently, there are few methods available to effectively remove established
populations. However, invasive species management capabilities are developing, with more
effective methods in detecting incipient populations, improved control methods, more stringent
restrictions on movement of nonnative animals, and increased public support.
Key words: amphibians, brown tree snake, bullfrog, Burmese pythons, cane toad, coqui frog,
invasive species, reptiles
Proceedings of the 11
th
Wildlife Damage
Management Conference. (D.L. Nolte, K.A.
Fagerstone, Eds). 2005
INTRODUCTION
In the last 20 years, worldwide declines of
many amphibian and reptiles have been well
documented. At the same time, a growing
number of species have invaded new
habitats and have reached population levels
that have had negative consequences on
native flora and fauna, agriculture, and local
economies (Mooney and Hobbs 2000). Five
of the 24 vertebrate species listed as the
worst invasive species are amphibians and
reptiles (Lowe et al. 2004). Invasive
amphibians and reptiles generally have a
high reproductive rate, which facilitates
rapid population growth and recovery from
stochastic events. They have generalized
diets that effectively utilize locally abundant
resources. Typically, successful invaders are
small or secretive, which allows undetected
movement in transportation networks.
These cryptic behaviors also allow the
development of incipient populations that
are difficult to detect until the animal is well
established. Species that exhibit all of these
attributes tend to be most successful at
colonizing new environments .
The probability of a successful
invasion is also dependent on the qualities of
the ecosystem invaded (Simberloff and Von
Holle 1999). Beyond a suitable climate and
habitat, ecosystems with a limited
assemblage of resident species have fewer
potential competitors and predators and,
therefore, enhanced probability of successful
colonization. Lastly, as the frequency of
invasion events by a species increases, the
likelihood that the species will successfully
establish increases. Insular areas are
113
generally more susceptible than mainland
areas, as islands support few predators or
competitors, often receive heavy air and sea
traffic, and typically provide a favorable
climate for many potentially invasive
species (Elton 1958, Simberloff 1995).
Currently, 33 non-native amphibians and
reptiles have been established in Hawaii and
more species continue to arrive (M.
Wilkinson, Hawaii Department of Land and
Natural Resources, personal
communication). For example, six snake
species have been intercepted in
transportation networks in the Pacific and at
least six species of frogs have established
populations in Guam in the past three years
(D. Vice, personal communication). While
the mechanism for arrival differs among
locales and species, the rapid and expanding
colonization of invasive reptiles and
amphibians is affecting ecological and
economic systems worldwide.
The pathways that transport invasive
species are varied and likely increasing.
Rapid increases in global transportation
networks move people and commodities to
previously remote destinations, increasing
the homogeneity of global floral and faunal
communities (Mack et al. 2000). Generally,
species are either accidentally or
intentionally transported. Accidental
movements include stowaways in air and sea
cargo, shipping containers that holds cargo,
or vessels that move people and
commodities (e.g., brown treesnake),
hitchhikers on agricultural products (e.g.,
coqui frogs, geckos, blind snakes) and pet
escapes (e.g., pythons and Jackson
chameleons). Intentional releases include
those that were intended to provide food for
people (e.g., bullfrogs and turtles), to
combat other species (e.g., cane toads and
poison dart frogs), or for aesthetic reasons
(e.g., veiled chameleons). Although many
intentional releases are altruistic in intent,
some are for insidious or financial reasons.
Species smuggled and released for the pet
trade are increasing threats, especially in
tropical environments and difficult to
prevent as border security measures and the
realignment of customs inspections are not
focused on invasive species.
HIGHLIGHTED SPECIES
Several species have become widely
publicized for their overall effect as invasive
species or as successful invaders in multiple
regions. To understand the effects of
invasive amphibians and reptiles and
potential problems with control efforts, we
provide a brief summary of several
noteworthy species. Further, we provide a
brief discussion regarding the social,
biological, and political complexity of the
invasive species issue.
Coqui Frogs
The coqui frog (Eleutherodactylus
coqui) was introduced into Hawaii during
1988-1995, likely from infested plant
shipments from Puerto Rico (Kraus et al.
1999). Sizeable populations are now found
on the islands of Hawaii, Maui, Oahu, and
Kauai. The super-abundant terrestrial frog
threatens Hawaii’s multi-million dollar
floriculture, nursery, real estate, and tourist
industries, as well as its unique ecological
systems (Beard and Pitt 2005). Effects from
the coqui are predominantly associated with
the frog’s piercing call (80-90 dBA at 0.5 m)
and the extremely high population densities
that have exceeded 50,000 individuals ha in
Hawaii. Beyond the noise nuisance, the
loud nighttime choruses of male frogs has
affected real estate values, as people desire
coqui free property (A. Hara, University of
Hawaii, unpubl. data). The floriculture
industry may also be affected through the
refusal of export shipments, reduced sales,
and increased costs associated with control
and quarantine efforts. Further, the frogs
may affect native insect populations, forest
114
nutrients, compete with native birds and
bats, and alter ecosystem processes (Beard
and Pitt 2005). Due to the high densities of
frogs and their present range, few options
exist for control of wild populations.
Mechanical controls include hand capturing,
habitat alteration, and trapping. These
methods have limited effectiveness, as the
logistic constraints in applying across a
large, complex environment with heavy frog
populations preclude large-scale application.
Some success has been documented using
hot water treatments for quarantine efforts in
ornamental plant shipments (M. Wilkinson,
pers. comm.). Biological control or the
release of organisms to combat the frog
likely will have little success and could have
many unintended consequences.
Unfortunately, disease organisms have a low
potential for controlling coqui frogs in
Hawaii, primarily because viruses and
diseases are most effective when applied to
small populations of species with low
reproductive capacity (Brauer and Castillo-
Chavez 2001, Daszak et al. 2003). In large
populations, diseases may induce temporary
population declines, but surviving
individuals may develop resistance,
resulting in population levels similar to
those pre-treatment. As most of the major
frog diseases infect tadpole stages (Daszak
et al. 2003), coqui, which develop directly
into tiny frogs inside terrestrially-deposited
eggs, are less likely to be affected by disease
organisms. Although many frogs are quite
susceptible to a variety of chemicals, the
terrestrial coqui frog has been unaffected by
a wide range of potential pesticides.
Currently, only citric acid and hydrated lime
have proven to be effective and registered
for use to combat the frogs (Pitt and Sin
2004a, Pitt and Doratt 2005). Although
these chemicals are effective if sprayed
directly on the frogs, there are limitations
with these products, including varying
efficacy affected by weather conditions,
potential phytotoxicity to plants, high costs
associated with repetitive spraying of large
areas, access to remote or private land, and
other factors (Pitt and Sin 2004b).
Burmese Pythons
Burmese pythons (Python molurus
bivittatus) became established in Everglades
National Park during the 1990s as the result
of unwanted or accidentally released pets (S.
Snow, National Park Service, pers. comm.).
Burmese pythons, native to Southeast Asia,
are large snakes (>7 m) with high
reproductive rates and are common pets in
the United States (Pough et al. 1998).
Pythons may compete with native snake
species, prey on many native mammals and
birds, transmit disease to native reptiles, and
disturb visitors due to their large size. The
number of snakes removed has quickly
increased in recent years and may represent
a rapidly increasing population (S. Snow,
unpubl. data.). Sources of mortality for the
snakes in the Park include motor vehicles,
mowing equipment, fire, and possibly
alligators (S. Snow, unpubl. data). Currently,
management actions center on direct control
and education efforts to prevent further
introductions. Control techniques include
trapping, hand capture, and early detection
using dogs.
Bullfrogs
Bullfrogs (Rana catesbeiana) from
the eastern United States were widely
introduced from 1900-1940 into many
western states, including Hawaii, as a food
resource. Bullfrogs are responsible for
significant ecological effects and have been
difficult to control as they are highly mobile,
exhibit generalized eating habits, and have
high reproductive capacity (Moyle 1973).
Bullfrogs may cause the extirpation of other
species due to intense predation and
competition (Kats and Ferrer 2003), and
may be a primary predator of several
115
federally endangered waterbirds in Hawaii.
Management of bullfrog populations is
difficult, in part due to commingling with
native species in aquatic habitats. Adult
frogs are removed by trapping or hand
captures and tadpoles are destroyed by
draining ponds or chemical treatment with
limited success. Fencing may also be used
to limit frog movements away from infested
habitats.
Cane Toads
Giant neotropical (Bufo marinus) or
cane toads were widely introduced from
Central America into sugar cane producing
regions worldwide to control beetles causing
damage to crops (McKeown 1978).
However, the effort had very limited
success, as the beetles could climb into the
vegetation to escape foraging toads. Cane
toads may compete with native species for
food, compete with native amphibians for
breeding sites, and prey on a variety of
invertebrate and vertebrate species (McCoid
1995, Catling et al. 1999, Williamson 1999,
Boland 2004). Further, native species
preying on cane toads may be poisoned by
the toad’s parotoid glands (McCoid 1995).
The frogs also may be a nuisance when large
numbers congregate for breeding in ponds or
water features and may foul water sources.
Australia has been aggressively pursuing
control options but has had little success in
developing effective methods (Luntz 1998).
Currently, the only effective strategies are
pond drying, hand capture, and trapping.
Brown Treesnakes
Brown treesnakes (Boiga irregularis;
BTS) were accidentally introduced into
Guam shortly after World War II from their
native range in Australia and Papua New
Guinea. The slender arboreal snakes average
approximately 1 m in length, with large
individuals capable of exceeding 2.5 m.
They have reached extremely high
population levels (> 20 per acre) on Guam,
in part because of abundant food and the
lack of predators and ecological competitors.
The extreme densities of BTS have resulted
in the extirpation of most of Guam’s native
forest birds (9 of 11), reductions in native
lizard populations, and extirpation of two of
the three native bats (Savidge 1988, Rodda
and Fritts 1992, Vice et al. 2005b). Beyond
the severe ecological effects, brown
treesnakes threaten human health and safety,
agriculture, poultry production, and pets..
The snakes are poisonous and may cause
trauma to small children, with numerous
bites treated by medical facilities annually
(Fritts et al. 1994). The largest economic
impact from the snakes is the disruption of
power systems. The aboreal snake
frequently climbs utility poles, power lines,
and other structures, searching for birds and
lizards. Snakes occasionally disrupt these
systems when they cross from grounded to
live structures, causing an estimated 1.4
million (U.S. dollars) in damages from
power outages (Fritts et al. 1999). The
cryptic, nocturnal snake is especially adept
at stowing in cargo and dispersing off Guam
via commercial and military traffic (Vice
and Vice 2004), creating substantial risk to
surrounding islands. A variety of methods
are employed to control snakes and restrict
their access to aircraft and cargo leaving the
islands, including hand capture off fences
(Engeman and Vice 2001), trapping (Vice et
al. 2005a), and inspection of outbound cargo
using detector dogs (Vice and Engeman
2000, Vice and Pitzler 2002) . Other
developing and potential methods include
the use of oral toxicants, repellents,
reproductive inhibition, and barriers.
Curious Skink
Often the effects of invasive species
are not predictable, and the combined effects
of two or more invasive species may result
in synergistic effects that exceed the sum of
116
the individual species effects. Such is the
case of the curious skink (Carlia
ailanpalai), skink, a small terrestrial lizard
that was introduced to Guam in the 1960s
(Zug 2004). This lizard has reached
extremely high population densities,
(approaching 10,000 acre in snake-free
areas) on Guam both in forested habitats and
near human habitation (Campbell 1996).
Due to sheer number of lizards, they may
compete with native lizards for food and
physically displace other native terrestrial
skinks through territorial interactions.
However, this is only a small part of the
overall effect on Guam. The high
population levels of the lizards have
exacerbated other problems, as the skink
serves as the primary food source supporting
the abundant BTS population on island. The
abundance of skinks in close proximity to
human habitation brings snakes into contact
with cargo facilities, power generation and
distribution stations, and agricultural
production, increasing the risk of snake-
caused damages associated with these
activities. The invasive skink has now
colonized the remainder of the populated
Northern Mariana Islands, and may increase
the probability of successful colonization by
the BTS, as the skink will provide an
abundant ectothermic food source for
juvenile snakes, should they reach the
currently snake-free islands. Further, the
skink population has facilitated growth in
Guam’s population of native yellow bittern
(Ixobrychus sinensis). Increasing bittern
populations near Guam’s airport has created
an aviation safety risk, as bitterns frequently
forage for skinks in the manicured turf
around the airfield and are subsequently
struck by aircraft (Vice and Pitzler 1999).
Thus, a small invasive species with few
predictable effects may cause a myriad of
significant emergent effects.
PRIORITIES OF INVASIVE SPECIES
The priorities of invasive species
management are generally divided between
prevention and control. Prior to
establishment, research and funding should
go to prevention and early detection to
decrease the potential for species becoming
a problem. To increase the effectiveness of
limited funding, a risk analysis should be
performed to promote awareness of species
that could cause significant effects. Further,
coordination and cooperation among state
and local agencies decreases the potential
for duplicated efforts and increases the
response efforts for incipient species. After
a species has become established, research
and funding is shifted to documenting
effects of the species on ecological services,
agriculture, and local economies.
Development of control strategies and
public awareness area are priorities after
establishment to control the effects of the
new species.
It is widely accepted that prevention
is the preferred means of dealing with
invasive species, as post-colonization
eradication efforts require massive funding
and resource commitments. Additionally,
complete eradication of vertebrate species
has rarely been accomplished in large
landscapes. Unfortunately, the line that
separates the priorities before and after
establishment dictates the amount of funding
available and the cost of the eradication
effort. Prior to species establishment, the
cost of controlling a species is low and the
probability of success is high. However, the
amount of funding and public interest in
dealing with the potential problem is
generally low at this time. Federal funding
for research and interdiction efforts prior to
species establishment is typically not a part
of congressional funding. Funding for
research and interdiction efforts is usually
only secured with public support and
congressional backing. After the species is
117
established, funding typically becomes more
available and public interest in dealing
increases. Conversely, the costs of control
and/or eradication efforts sky rocket and the
probability of successful eradication drops
after a species is established. This same
scenario has been repeated in many areas
with many species. A recent example is the
above-mentioned case of the coqui frog in
Hawaii. Although the species became
established by the early 1990s in parts of
some islands, no funding was available,
even though the potential to eradicate was
still high. The primary public opinion was
that this was not a major problem and there
were likely to be few negative consequences
associated with this introduction. Ten years
later, public opinion is extremely supportive
of dealing with the issue and several studies
have documented the effects of the frogs on
ecological communities, real estate,
agriculture, and human health (Kraus and
Campbell 2002). To highlight this change, in
March 2004, the Mayor on the island of
Hawaii declared a state of emergency in
dealing with the coqui situation.
Unfortunately, this response occurred once
the frog had populated massive tracts of
habitat on the island, rendering eradication
unlikely.
In conclusion, invasive amphibians
and reptiles are an increasing worldwide
problem, causing a diverse array of
problems that cannot be easily predicted.
Invasive reptiles and amphibians may cause
more significant problems on island systems
than mainland areas. The number of new
introductions is likely to continue escalating,
as many pathways of invasion are not
subject to stringent quarantine and/or
control. Currently, there are few effective
options in controlling established invasive
reptile and amphibian species, and the cost
for control efforts is often extreme once a
species becomes widespread. Although
politically challenging, the most cost
effective approach to invasive species
management is to secure funding for
research and interdiction efforts to prevent
establishment.
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