Leaf Litter 3.1

Leaf Litter
The Magazine of Tree Walkers International and Amphibian Conservation

EDITOR
Ron Skylstad
ASSISTANT EDITORS
Lee Hancock
Jason Konopinski
Ed Kowalski
PHOTO EDITOR
Tim Paine
LAYOUT
Elizabeth Brock
TWI DIRECTOR
Ron Skylstad
TWI PROGRAMS DIRECTOR
Brent L. Brock
MISSION STATEMENT
Tree Walkers International supports the
protection, conservation, and restoration of wild
amphibian populations through hands-on action
both locally and internationally.
We foster personal relationships between
people and nature by providing opportunities for
citizens of all ages to become directly involved in
global amphibian conservation.
Through this involvement, our volunteers
become part of a growing and passionate
advocacy for the protection and restoration
of wild amphibian populations and the
environments on which they depend.
Leaf Litter
VOLUME 3 | ISSUE 1
1 FROM THE EDITOR
FEATURES
4 Ospring Begging, Larval Relocation
and Parental Care in Dendrobates
auratus
6 Te Use of a Modied Shipping
Container as an Amphibian Laboratory
at the Atlanta Botanical Garden
10 A Decade of Amphibian Conservation
Action in Panama: From Golden Frogs
to Marsupials
19 Atelopus Zeteki
26 Further Notes on Dendrobates auratus:
Te Variability of the Panamanian
Green and Black Poison Dart Frog
40 Confessions of a Pumilio Chaser
54 Te Oophaga pumilio of Escudo de
Veraguas
COVER Cochranella albomaculata
photo by Tim Paine
from the editor | 1
from the editor
The country of Panama is a convergence of many things.
The most recent theories about its history postulate that it was formed by a combination of
both colliding continental plates and volcanic activity, which created various oceanic ridges
and islands. Over time these islands and ridges expanded, shifted and coagulated, ultimately
creating an isthmus of land that connected the land masses of North and South America.
Plants and animals from each continent made their way across this ecologically developing
bridge, intermingling and settling into new niches and populations as they went.
Although Panama may not be the most biodiverse country on the planet, it is one of the most
uniquely biodiverse places on earth as a profusion of species and environments coexist in
a relatively small piece of land. In recent years Panama has been known for its amphibian
crisis and spread of the chytrid fungus, which has traveled wave-like through the country. In
response, various attempts have been made to save and conserve its numerous vulnerable
species of amphibians.
This issue of Leaf Litter is a celebration of this unique country, showcasing some of its
amphibians and the conservation efforts that have sought to rescue these amazing creatures.
Ron Skylstad
EDITOR-IN-CHIEF
photo by Dant Fenolio
2 | Leaf Litter :: Vol 3 Issue 1
El Valle
Penonome
Bocas del Toro
David
Boquete
El Cop
Calobre
Santa Fe
San Felix R
io
S
a
n
ta
M
a
ria
Caldera
Escudo de Veraguas
Volcn Bar
!"#"$"
$"! &' ()* +&,#(-. &'
Chiriqui Grande
Map of Panama | 3
Panama City
Taboga Island
La Chorrera
Portobelo
Capira
Altos de Campana
National Park
Kuna Yala (San Blas)
Darien
IN THE UPPER MANOA VALLEY
on the island of Oahu, Hawaii, exists
an introduced population of the green
and black dart-poison frog (Dendro-
bates auratus). This population exists
as the result of an early experiment for
the biocontrol of mosquitoes in which
approximately 200 individuals from
Taboga Island, Panama were released
in the valley. In the ~75 years since its
establishment, the population appears
to be contained within the Manoa Val-
ley (a separate population does exist
on the other side of the island in the
Waihole Valley, however there is some
debate as to whether this was part of
the initial introduction or occurred
later).
Part of our research group (CG Farmer
laboratory at the University of Utah)
wus IorLunuLe enougI Lo conducL heId
studies on this population. These
studies revealed details of parental care
behaviours within this species previ-
ously not observed which I would like
Lo brIey summurIse In LIIs urLIcIe. n
order to fully appreciate these observa-
tions, however, a short introduction to
parental care behaviours is required.
Parental care behaviours, loosely de-
hned us uny posL-ovIposILIonuI InvesL-
ment by a parent, play critical roles in
the life histories of numerous animal
species. These behaviours are pro-
IoundIy InuenLIuI IuceLs oI un unImuI`s
life history, yet in many species these
behaviours have yet to be studied in
great detail.
Within the Anura, these behaviours
are widespread, having apparently
convergently evolved numerous times.
The most common form of parental
care within this group is clutch atten-
dance, the behaviours of which range
in investment from simple clutch atten-
dance to maternal provisioning of
larvae in the form of feeder eggs. The
subfamily Dendrobatinae contains sev-
eral parental care behaviours, the most
basal of which is clutch attendance by
the male followed by larval deposition
into bodies of water. This is the pattern
of behaviour currently described within
D. auratus.
DurIng our heId sLudIes uL (und
around) the Harold B. Lyon Arboretum
on Oahu, we observed several larval
depositions by male D. auratus into
the axils of a large bromeliad (Vriesea
imperialis). Upon arrival of a tadpole-
carrying male D. auratus to a water-
hIIed uxII, one oI LIree evenLs occurred:
1) the male (with tadpole) complete-
ly submerged into the axil but the
tadpole did not detach and the male
continued to another axil where the
tadpole was eventually deposited;
2) the male detached the tadpole
manually with the use of his rear
Ieg, IckIng LIe LudpoIe oII wIIIe
submerged in the axil;
3) the tadpole detached in the pool,
but later re-attached to the back
of the male and was relocated to a
different axil following a distinct
begging behaviour.
It is the third event that we would like
to discuss in this note, as we feel it
demonstrates complex and innately
fascinating behaviours not typically at-
tributed to this species. These behav-
iours occurred in approximately 15%
of the deposition events and relocation
occurred between 5 minutes after ini-
tial deposition up to 48 hours.
The begging behaviour mentioned
above involved the tadpole stiffening
its tail, rapidly vibrating it for a few
Offspring Begging, Larval Relocation and Parental Care
in Dendrobates auratus
BY BENJAMIN K. CHAN
PHOTOGRAPHS BY THE AUTHOR
University of Utah Department of Biology
357 S 1400 E, Salt Lake City, UT 84112
Ofspring Begging, Larval Relocation and Parental Care in Dendrobates auratus | 5
seconds, and then swimming erratically
around the male, especially under
the males throat and around his rear,
until the tadpole was able to align with
the backside of the adult and re-attach.
This larval begging behaviour differed
from typical swimming patterns ob-
served when tadpoles were manually
disturbed, frightened, or under attack
Irom conspecIhcs. urLIermore, In
each case the frog that approached was
the same individual that had initially
deposited the tadpole (determined
by distinctive colour patterns), and
these behaviours did not occur when
other frogs (of the same species) were
observed in or near the axil. A similar
swimming pattern has been observed
in oophagous tadpoles of Oophaga
pumilio in response to maternal visita-
tion (Brust, 1993). Described as beg-
ging, this behaviour occurs immedi-
ately before a nutritive egg is laid.
Examination of one rejected axil after
the male left with the tadpole revealed
a larger tadpole in the base, but this
was not true of all axils rejected by
tadpoles. In two relocation events, the
male left the pool but returned one day
later. Upon return of the male, the tad-
pole performed the begging behaviour
LIree LImes. TIe muIe LIen ouLed InLo
the pool, and the tadpole reattached to
his back. The tadpole was then depos-
ited in a neighboring axil within the
same bromeliad.
The use of care-eliciting signals by D.
auratus larvae represents a more com-
plex parent-offspring relationship than
previously described. Our observations
suggest that both parent and offspring
have some degree of choice regarding
the site of larval development. These
behaviours arent without costs, how-
ever, and the reasons for this increased
male investment are unknown. We
have conducted laboratory studies that
suggest acute changes in phytotelm
volume may contribute to higher inci-
dences of larval relocation.
Parental investment theory predicts
that the degree of parental care will
depend on offspring value, costs of care
beIuvIours, us weII us oIIsprIng benehLs
associated with these behaviours.
Competition between offspring may
have detrimental effects on offspring
value (particularly in nutrient poor
microhabitats) and, therefore, we
would expect competition-avoiding
mechanisms to evolve in these environ-
ments. Several species of amphibian
utilize nutrient-poor microhabitats for
the development of their offspring and
it is likely that many similar behaviours
have evolved across taxa.
_________________
Works Cited
Brust, D.G. 1993. Maternal brood care by
Dendrobates pumilio a frog that feeds
its young. Journal of Herpetology 27:96-98
AMPHIBIAN DECLINE NOW
ENDANGERS at least one third of
the worlds 6,000 plus amphibian
species
1
. Time is short and amphibian
conservation efforts are in dire need
of additional training and capacity
building programs
2
. The Atlanta
Botanical Garden (the garden)
has been committed to amphibian
conservuLIon eIIorLs Ior over hILeen
years and has enacted in situ and ex
sILu cupLIve breedIng und heId reseurcI
programs across eight countries. The
garden is the only one of its kind
with an amphibian conservation
program that employs at least three
amphibian specialists at any given
time, including a Ph.D. level amphibian
conservation scientist. Program efforts
include on site captive amphibian
management and reproduction,
training of amphibian husbandry and
breeding techniques in the United
States and in Latin America, ex situ
collaborations in the U.S. and abroad,
and in situ research and conservation
activities. Facilities include a dedicated
quarantine room, a live food culturing
room, a series of portable ponds for
rearing endangered ranid tadpoles,
and two on-site amphibian laboratories
that accommodate captive groups of
highly endangered amphibian species.
This article will focus on one of the
on-site amphibian laboratories, our
amphibian pod.
In 2007 the garden had a fundraising
event to support their amphibian
conservation program. During
the lecture, a point was made that
an Australian biologist, Dr. Gerry
Marantelli, had pioneered the use
of shipping containers as mobile
amphibian laboratories in Australia.
Gerry and his crew were successfully
breeding endangered Australian frogs
wILIIn LIe conhnes oI LIese mobIIe
labs. To our great surprise and delight,
several of our supporters decided to
fund a repurposed shipping container
turned amphibian breeding laboratory
(a pod) here at the garden. The
opportunity would provide experience
in establishing an on-site lab and be of
benehL In deveIopIng LIe experIence
necessary to set up future facilities
off site. Further, a collection of
endangered Panamanian amphibians
would be moved into the pod, allowing
the collection more space.
Without getting too detailed regarding
all of the parts that went into making
our pod functional, we would like to
cover some of the basics. A shipping
container was purchased from
Container Tech, a local business in
Atlanta, which made the necessary
modIhcuLIons Lo ouLhL LIe conLuIner
as an amphibian lab. Our shipping
container is 40 feet in length, 8 feet
wide, and 8 feet tall. Container Tech
added two windows, two doors, and
subdivided the inside of the container
into two rooms of 30x8x8 feet and
1ox8x8 IeeL. TIe hrsL room serves
as a biosecurity area where shoes
can be changed and hands washed.
The larger room accommodates over
80 terrariums. The terrariums are
ZooMed brand glass enclosures that are
situated on epoxy coated metal racks
and plumbed with misting systems on
timers. The misting system currently
provides 6 misting events a day but
can be adjusted to accommodate dry/
weL seuson cycIIng. WuLer Is hrsL
rouLed LIrougI curbon hILers, mIsLed
into our terrariums, and then leaves
LIem LIrougI oor druIns. uIse
bottoms made of egg crating keep
the substrate drained. Wastewater
is plumbed into water tubes. These
water storage devices were designed by
Advanced Coastal Technologies and are
used In un eIIorL Lo be eIhcIenL wILI our
water since we are in a region affected
by drought. Wastewater is treated with
bleach and can be reused where needed
around the garden.
The collection of amphibians in the
gardens pod includes species of critical
conservation need in response to the
spread of amphibian chytrid fungus,
BY DANT FENOLIO, ROBERT HILL, JULIA KAYLOCK, AND JENNY CRUSE SANDERS
PHOTOGRAPHS BY DANT FENOLIO
Atlanta Botanical Garden, Department of Conservation Research
1345 Piedmont Ave NE, Atlanta, GA 30309
The Atlanta Botanical Gardens frog pod is a 40 foot long repurposed shipping container
modifed to include: two internal rooms, two windows, two doors, and internal environmental
controls.
he Use cI c Mcd|hed Sh|pp|ng CcnIc|ner cs cn
Amphibian Laboratory at the Atlanta Botanical Garden
The Use of a Modifed Shipping Container as an Amphibian Laboratory at the Atlanta Botanical Garden | 7
Batrachochytrium dendrobatidis (Bd),
through Panama. In 2005, Bd was
headed eastward through Panama.
Research biologists had documented
what happened after Bd had arrived
further northwest. The scene at
localities west of El Valle had been grim
and most amphibians disappeared
shortly after the fungus arrived. Ahead
of the arrival of Bd at El Valle, a team
of researchers, in collaboration with
the Panamanian government, collected
small groups of frogs to form captive
assurance colonies. The garden and
Zoo Atlanta acquired collections of
these frogs from El Valle. The frogs
are now a key component of the
live collection at the garden. Our
umpIIbIun Leum Is Iocused on rehnIng
captive care and reproduction methods
for 13 Panamanian species. Colonies of
these same species exist at the captive
amphibian breeding center at El Valle,
Panama, known as EVACC. It should
be pointed out that EVACC is a facility
collaborating with and sponsored
by a wide variety of institutions,
spearheaded by the Houston Zoo.
The garden has a strong collaboration
with EVACC and the Houston Zoo
in capacity building and information
exchange for captive husbandry of
species in our conservation collections.
The garden also sends one of its
amphibian specialists down annually to
assist at the EVACC facility.
In the wild, the future of some
endangered Panamanian species is
uncertain. Several of the frogs that
have disappeared entirely from El Valle
are in the collections of EVACC and the
garden. For example, the Rusty Robber
Frog, Strabomantis bufoniformis,
and Bobs Robber Frog, Craugastor
punctariolus, are of conservation
concern in the wild. In captivity, eggs
have been produced but both species
have yet to yield an F1 generation. One
species at the garden has not been
bred, the Evergreen Robber Frog,
Craugastor gollmeri. Other species
Iuve been bred buL dIIhcuILIes Iuve
been encountered with rearing the
young. For example, the Fort Randolph
Robber Frog, Pristimantis gaigei, has
been bred at the garden; however,
the eggs and young have proven
dIIhcuIL Lo munuge. AnoLIer exumpIe
involves the Horned Marsupial Frog,
Gastrotheca cornuta, which was bred
Ior LIe hrsL LIme In cupLIvILy uL LIe
garden. In the past, the young had
proven dIIhcuIL Lo reur beyond sIx
or eight months of age. However, in
2008 we successfully bred the Horned
Marsupial Frog and 7 young frogs are
still alive and growing. The success
has come with the additional space the
frogs have in the pod and a change in
vitamin supplementation. Another
challenge has been Rabbs Fringe Limb
Tree Frog, Ecnomiohyla rabborum,
a recently described species. The
hrsL recorded cupLIve breedIng oI LIe
species took place at the garden but we
have much to learn about the captive
management of these endangered
frogs.
A number of species in the Panamanian
The fnished amphibian room in the Atlanta Botanical Gardens frog pod holds over 80 terrariums and has rack space to accommodate tadpoles
and juvenile frogs.
collection have done well in captivity.
The Thorny Crowned Frog, Anotheca
spinosa, has been bred at the garden
quite a few times. Adult and subadult
hrsL generuLIon Irogs serve us LungIbIe
evidence of the success. Several
other species have been bred with
regularity including the Lemur Leaf
Frog, Hylomantis lemur, Pratts Rocket
Frog, Colostethus pratti, and the
White Spotted Glass Frog, Cochranella
albomaculata. Particularly in the
case of the Lemur Leaf Frog, dozens of
captive offspring have been distributed
to zoos and aquariums for education
and conservation exhibits.
The gardens amphibian conservation
team now focuses on long-term
nutritional requirements with their
captive Panamanian collection.
Our program works closely with
veterinarian and amphibian specialist
Dr. Brad Wilson. This collaboration
involves a series of controlled
experImenLs desIgned Lo cIurIIy specIhc
vILumIn und mIneruI dehcIencIes In
second and third generation captive
amphibians. The goal is to improve
mineral supplementation regimes,
aiming to eliminate reproductive
output declines in captive colonies
of amphibians. We are also
researching new approaches to captive
reproduction of species that have
not been bred frequently within our
collection.
The value of the gardens collection
housed in our amphibian pod is
severuI IoId: hrsL und IoremosL, our
exchange of information with the
EVACC facility in Panama is critically
important. The EVACC facility has
done exceptionally well with the
management and captive reproduction
of their populations of frogs; anything
the garden can contribute in care and
captive reproduction methodology will
only help to improve that outcome.
Further, the only frogs that might
some day be released into the wild
would be captive produced offspring
from EVACC (owing to the risk of
disease spread in sending captive
offspring back to Panama from foreign
facilities). Second, the assurance
colonies at the garden represent
valuable genetic material. Particularly
with species disappearing in the wild,
these collections carry increasing
importance. Additionally, captive
offspring produced at the gardens
facility have been distributed to over
25 zoos and aquariums around the
world. These individuals improve
public awareness of the amphibian
decline issue, encourage public
education of conservation issues facing
amphibians, and focus attention on
the great work EVACC has done. The
garden has supported researchers
investigating emergent infectious
disease by providing captive offspring
to populate laboratory studies. Captive
produced offspring provide a unique
resource Ior quuIIhed ucudemIc sLudy
und sIgnIhcunLIy expund LIe kInds oI
In the wild Strabomantis bufoniformis includes freshwater crabs in its diet. This is an imperiled
species and much focus has been put on breeding the individuals at the Atlanta Botanical Gar-
den. All egg clutches deposited to date have been infertile.
Pristimantis gaigei is a species that is being maintained and has been successfully bred in the
Atlanta Botanical Gardens frog pod.
The Use of a Modifed Shipping Container as an Amphibian Laboratory at the Atlanta Botanical Garden | 9
questions epidemiologists can ask in
the lab. Finally, the garden provides
internship opportunities, particularly
for collaborations with foreign
institutions looking to establish their
own captive breeding facilities. For
example, the lead veterinarian from
the National Zoo in Santiago, Chile,
interned at the garden in 2008. The
reason stems from a collaboration in
Santiago to develop a captive breeding
facility for endangered Darwins
Frogs (Rhinoderma). The internship
opportunities are perhaps the best
chance the garden has to transfer
information to other institutions
looking to combat amphibian
decline. Finally, the Panamanian
collection housed in our amphibian
pod allows the gardens amphibian
crew the experience of maintaining an
amphibian collection in a pod. The
lessons learned through our collection
management can be transferred to
future pod implementation programs
that we have in the works now.
With thousands of species facing
serious conservation threats including
habitat loss and infectious disease, the
orgunIzuLIons dedIcuLed Lo hgILIng
the causes of amphibian decline must
work together to address pressing
issues. As a well-established program
in the amphibian conservation
community, the garden plans on
facilitating the continued development
of collaborations with institutions such
as the National Zoo in Santiago, Chile,
and EVACC. The philosophy behind
the gardens amphibian conservation
program hinges on capacity building
with organizations and institutions
interested in combating amphibian
decline. Capacity building efforts,
particularly in developing countries
where funds and expertise may be in
short supply, are in dire need to ensure
longevity of successful conservation
programs. These activities include
establishment of captive breeding
facilities, staff training, and long-term
technical support. One important tool
is to develop methods for implementing
repurposed shipping containers, or
pods, such that pods can be used
to conserve rare amphibians in the
countries where they are declining. In
conjunction with training and support
programs, pods can be delivered to a
remote locality and be fully functional
within a short period of time.
ACKNOWLEDGEMENTS
ConLuIner TecI modIhed LIe sIIppIng
container that is now our frog pod
and was a huge help in working
with us through all of the twists and
Lurns us we InsLuIIed our hrsL Irog
pod. This experience has provided
us with the platform from which we
have begun to install new facilities.
Coastal Technologies Inc. produced
the water pillows that we use to store
and treat the waste water from the frog
pod. Their innovation and assistance
has helped us with the frog pod and
elsewhere in the Garden with water
conservation via capture and storage
of rainwater and waste water, for
later use around the grounds. Joe
Mendelson, Ron Gagliardo, Heidi Ross,
und Edgurdo GrIIhLI were InvoIved
with the collection and import of
Panamanian frogs that came to the US
and we appreciate their tremendous
efforts during that project; following
suit, we thank ANAM for permits and
logistical assistance to said individuals
while working in Panama. We thank
Mary Pat Metheson, Diana Champ
Davis, Fonaine Huey, Claire Simmons,
Bill Simmons, and Cindy Jeness for
their unwavering support of the frog
pod project. ZooMed has been a
partner with the Atlanta Botanical
Garden in amphibian conservation,
and the frog pod is no exception.
ZooMed terrariums and equipment
are used throughout the facility. Gerry
MurunLeIIy deveIoped und rehned
the concept of using repurposed
shipping containers as frog breeding
facilities. We appreciate his leadership,
creativity, and dedication to amphibian
conservation. We are thankful for
help during installation of the frog pod
from Jeremy Hodges, Jason Cameron,
Christy Jellets, Marcela Tirado, Ralph
Van Streels, and Tim Herman.
____________________
Works Cited
1 Stuart, S.N., J.S. Chanson, N.A. Cox, B.E.
Young, A.S.L. Rodrigues, D.L. Fischman,
and R.W. Waller. 2004. Status and trends
of amphibian declines and extinctions
worldwide. Science Express Reports
online, 14 October, 2004 (DOI: 10.1126/
science.1103538).
2 Gascon, C., J.P. Collins, R.D. Moore, D.R.
Church, J.E. McKay, and J.P. Mendelson III
(Eds.). 2005. Amphibian Conservation
Action Plan. Proceedings of the IUCN/SSC
Amphibian Conservation Summit 2005.
Cochranella albomaculata is a species that Atlanta Botanical Garden has just recently bred and
reared successfully in the frog pod.
Introduction
As the mystery surrounding the causes
of global amphibian declines moves
cIoser Lo beIng soIved, we hnd LIere
exists not one single culprit, but rather
a perfect storm of threats. These
include habitat loss, pollution, over-
collecting, introduced species, global
climate change and disease, all of which
contribute individually to the depletion
of amphibians from our planet (Berger
et al., 1998, Collins and Storfer, 2003,
and Gascon, et al., 2007). So at this
point we recognize this is happening
right before our eyes and that in a
matter of a decade, hundreds of species
may go extinct. All of these causes are
anthropogenic, resulting from careless
stewardship of the earth by our ever-
increasing human population. With
this generation about to witness what
could be the largest extinction event in
the history of humanity, should it not
also be the one that is challenged with
doing something about it?
With conventional
approaches to species
conservation through
habitat protection and trade
regulation being ineffective
against disease and climate
change, we are forced to
consider other options for
saving amphibians. As the
scope of the crisis becomes
clearer (Stuart et al.,
2004) and the threat of the
emerging infectious disease
amphibian chytridiomycosis
(Bd) continues to hang like
a black cloud over their
heads, conservation leaders
have recognized not only
the value of managed ex
situ breeding programs
in preventing further
extinctions, but also that
we must take action quickly
(Mendelson et al., 2006).
One part of the IUCN
Amphibian Conservation Action Plan
(ACAP, Gascon et al., zoo;) specIhcuIIy
calls for captive breeding and research
programs for threatened amphibians
to insure their survival while threats in
nature are mitigated. These programs
are implemented through global
partnerships facilitated through the
Amphibian Ark (www.amphibianark.
org, Zippel and Mendelson, 2008);
however, it is important to remember
that the use of captive breeding
programs as a conservation tool is
not new. Summaries can be found in
Zippel et al. (2006) and Gagliardo et
al. (2008). Here we report on what
comprises more than a decade of ex
situ conservation work in Panama,
including a summary of projects both
in and out of country, their strengths,
limitations, and experiences that will
assist in developing future projects.
Why Panama?
The disappearances and declines in
amphibian fauna in Central and South
America have been well documented
for decades (Pounds et al., 1997).
Survey work was particularly intensive
and productive in Costa Rica and
Panama, where Dr. Karen Lips and
her students diligently documented
amphibian declines for many years
(Lips, 2005). From these efforts a
pattern emerged as Bd moved in a
wave-like front from Costa Rica into
central Panama where amphibian
chytridiomycosis has now been
clearly linked with mass die-offs
of amphibian species from several
protected areas (Lips et al., 2006).
Panama has much at stake in relation
Lo umpIIbIun exLIncLIons: LIere ure
just over 200 species described from
this tiny country, 38 of which are
endemic. Currently, over 40 species are
considered Endangered (22 Critically
A Decade of Amphibian Conservation Action in Panama:
From Golden Frogs to Marsupials
BY PAUL CRUMP, RON GAGLIARDO, EDGARDO GRIFFITH, HEIDI GROSS, JOSEPH R. MENDELSON III, AND KEVIN ZIPPEL
The Horned Marsupial Frog, Gastrotheca cornuta, is one of the highest priorites for safeguarding in ex
situ programs. Captive born ofspring have proven difcult to rear due to what appears to be nutritional
defciencies. Photo by Brad Wilson, courtesy Atlanta Botanical Garden
A Decade of Amphibian Conservation Action in Panama | 11
Endangered) and at risk of becoming
extinct in the wild.
Project Golden Frog
Proactive response to amphibian
declines in Panama began in 1999
with Project Golden Frog (PGF, www.
projectgoldenfrog.org) as scientists
feared the loss of the culturally
sIgnIhcunL PunumunIun goIden Irog,
Atelopus zeteki. This spectacular
species is not only one of the worlds
most recognizable frogs but also one
of the most critically endangered
(Zippel, 2002). PGF became a multi-
institutional, international initiative
that has included primary partners
in Panama such as the Universidad
and Crculo Herpetolgico de Panama
along with many US-based zoos
(including the Cleveland, Colombus,
Denver, Detroit, and Maryland Zoos),
universities (Lee University and
Messiah College) and the Smithsonian
Tropical Research Institute. In 1999,
Bd was still a good distance (100 km
to the west) from the habitat of golden
frogs. However, as Lips and others
had witnessed the disappearance of
other harlequin frogs (e.g. Atelopus
senex, Atelopus chiriquiensis, Atelopus
varius and the Atelopus sp. from
Monte Verde) from higher elevation
sites in Costa Rica and Panama, the
writing was on the wall. The decision
was made to remove a small number
of golden frogs for safeguarding. PGF
obtained proper permits from the
Panamanian authorities (Autoridad
Nacional del Ambiente, ANAM) and
initiated the project focusing on work
in streams in central Panama in 2001.
From 2001 to 2005, PGF surveyed
transects in several streams, gathering
data on water quality as well as
surveying for frogs, tadpoles and egg
masses. Molecular data derived from
tissues of frogs from different sites
revealed that the locally well-known
goIden Irog wus ucLuuIIy Lwo specIes:
Atelopus zeteki and a golden morph
of A. varius (Zippel et al., 2007). PGF
gained permission to export animals
for an ex situ breeding program and
conducLed LIe hrsL exporL In zoo1.
Between 2001 and 2005, PGF collected
111 animals, including 26 amplectant
pairs and 59 newly metamorphosed
froglets. Because of the proximity of Bd
to golden frog habitat, the last group
of animals exported in May of 2005
received a prophylactic treatment
for Bd In heId prIor Lo exporL. TIese
IndIvIduuIs were hrsL LrunsporLed
to zoos in Detroit, Baltimore, and
Cleveland for initial quarantine and
maintenance. Regulations set in
place by the United States Fish and
Wildlife Service require that the golden
frogs be maintained only at facilities
accredited by the Association of Zoos
and Aquariums (AZA). Forty-one of
the wild-caught animals are currently
represented with over 1,300 animals in
the captive-bred population at nearly
50 AZA institutions.
During the course of this project, PGF
has effectively prevented the extinction
of both species of Panamanian golden
frogs. With Bd having moved through
all known golden frog habitats and
populations secured in AZA institutions
and in Panama (see section on
EVACC), PGF has chosen to broaden
its research, outreach, and ex situ
programs to include other Critically
Endangered harlequin frogs through
an initiative called the Atelopus
Conservation Trust (see www.
ranadorada.org/act).
Amphibian Research and
Conservation Coalition
(ARCC)
Thought to travel on the feet of
animals, insects, and even humans, Bd
was predicted to work its way down
the Isthmus of Panama toward the
mugnIhcenL umpIIbIun IoLspoL In EI
Cop. Working within the boundaries
of the protected Parque Nacional
General de Division Omar Torrijos
Herrera near El Cop, Lips and her
team had been surveying transects
Ior over q yeurs beIore hndIng LIe
hrsL suspecLed cuse oI umpIIbIun
chytridiomycosis in September of 2004
(Lips et al., 2006). Within 3 months
there was an 80% decline in overall
numbers of amphibians and a 50%
decline in species richness.
The scale of this mass die-off at El Cop
was so disturbing it moved researchers,
who had for years only studied and
documented these declines, to call on
coIIeugues ouLsIde LIeIr heIds Lo do
something about it. Lips summoned
experts from the zoological, academic,
und InLernuLIonuI dIpIomucy heIds Lo
a meeting in Atlanta in late 2004 to
report the sad news from El Cop and
brainstorm a plan of action. Realizing
that the man-made boundaries of this
Atelopus zeteki , the Panamanian Golden Frog, is now reproducing in range country thanks to
the eforts of PGF, ARCC and EVACC programs.
national park clearly did not prevent
the arrival of Bd and its devastating
eIIecLs, LIe sImpIe quesLIon wus posed:
What can we do? There were still
frogs in El Cop, but they were circling
the drain. Scientists predicted that
the area around El Valle de Anton (El
Valle), possibly the last upland area
of reasonably undisturbed fauna and
habitat of the Talamancan highland
region of eastern Costa Rica and
western Panama, would be the next
stop for Bd as it moved southward.
Thus began the rhetoric of emergency
response, salvage, and rescues
in regard to these sites. Endemic
species were at risk of extinction and
the challenge to stop it was daunting.
This meeting resulted in the formation
of the Amphibian Research and
Conservation Coalition (ARCC), a
temporary acronym to describe a
pilot operation set out by the Atlanta
Botanical Garden and Zoo Atlanta to
try to save not just one or two species,
a la PGF, but dozens via a triage
operation set up in temporary quarters
at El Cop (rescue and salvage) and
El Valle (pre-emptive collection). At
this time, as during the time of PGFs
operations in Panama, there was no
place to set up these types of operations
in-country. With full consent of the
Panamanian government, specimens
were exported to facilities in Atlanta for
safekeeping.
The ARCC project utilized the
bureaucratic, logistical, and personal
relationships set up by its predecessor,
PGF. Employing Panamanian
biologists, ARCC also obtained
the support and directive from the
Panamanian government needed
to collect and remove amphibians
from harms way. This was a pilot
project to investigate the feasibility
and challenges of conducting a multi-
specIes rescue quIckIy und eIhcIenLIy.
The project ran from 1 June to 1
September 2005 and utilized the skills
of 6 staff and dozens of volunteers
from around the world. The staff
was responsible for arranging all
IogIsLIcs, heIdwork, und LIe IormIdubIe
task of diligent care of captive
amphibians in makeshift facilities.
These facilities were primarily rented
houses or, in the case in El Valle, a
duplex provided at no cost by the
Hotel Campestre. Strict quarantine,
treatment, and maintenance protocols
were established to help maintain
the best possible hygiene under the
circumstances. Individual latex gloves
were used to handle each individual
and reduce the chances of cross-
contamination. Enclosures were
disinfected every other day using a
10% household bleach solution. Only
non-bleached paper towels were
used for routine cleaning protocols.
Each animal was swabbed for Bd
testing using the polymerase chain
reaction (PCR) method according
to Annis et al. (2004) and carefully
cataloged with collection data including
date of collection, gender, and the
general physical appearance of each
animal. Water used for amphibians
was simply rainwater collected in
95-liter containers without additional
dIsInIecLIon or hILruLIon. Hund-
collected and trapped food items were
used to maintain amphibians during
their in-country stay before being
exported.
At El Cop, transects previously
established by the Lips group were
monitored. These transects produced
very low numbers (60 total) of species
and individuals during the course
of the project and thus all were
collected, swabbed for subsequent
PCR testing to detect Bd, and treated
with Itraconazole (Supplied by PCCA,
Inc., Houston, Texas) according to
the method of Nichols & Larimande
(2000). Each frog was swabbed daily
for Bd during the 11-day treatment
regime and all PCR analyses were
graciously done by M. Poore and
M. Levy at North Carolina State
University, USA.
At El Valle, staff attempted to collect
up to 40 individuals (20 males/20
females) of species that were prioritized
using a prototype ranking system
developed by Smithsonian Institute
Scientist Dr. Roberto Ibez. This
prioritization of species was based on
a combination of factors, including
the degree of endemism, threat status,
and range. Taxonomy used was that
of the online reference Amphibian
Species of the World (Frost, 2007). A
major logistical challenge in El Valle
was holding many individuals from
multiple taxa with small numbers
of staff. Periodically, collection was
suspended as the routine care of
previously collected animals needed
to be addressed. At three intervals
between June and September
2005, all specimens were exported
with the permission of ANAM to
Atlanta Botanical Garden where they
underwent a 30-day quarantine.
During this time, they were monitored
and treated (if necessary) for internal
Frogs were swabbed for later Bd screening in El Cope. Photo by Ron Holt, courtesy Atlanta Botanical
Garden.
parasites and further acclimated to
new surroundings. Post-quarantine,
animals were distributed to facilities
at the Atlanta Botanical Garden, Zoo
Atlanta, and the Henry Vilas Zoo. Over
time, animals were distributed to other
institutions with the facilities, staff, and
commitment to maintain these species
and participate in the project.
ARCC collected only about
60 individuals from the
El Cop site. The taxa
in greatest numbers
included centrolenid
and leptodactylid frogs.
Findings in El Valle were
much more productive
since Bd had not yet
arrived. Hundreds of
individuals from a total
of nearly 30 species were
collected. Once the goal
of 20 of each sex was
achieved, duplicates were
released. In captivity,
some species such as
Hyloscirtus palmeri and
Hemiphractus fasciatus
proved exLremeIy dIIhcuIL
to maintain. Several
species did reproduce,
however, including
Hylomantis lemur,
Colostethus pratti,
Eleutherodactylus gaigae,
Gastrotheca cornuta,
Anotheca spinosa,
Oophaga vicentei, and
Ranitomeya minuta. In
some cases, these breeding
events (some taking
place within temporary
ARCC triage or EVACC
quarantine facilities)
represenL LIe hrsL In
captivity. Environmental
factors, such as extreme
buromeLrIc ucLuuLIons,
local food items or even
regular disturbance from
enclosure cleaning, were
possible triggers. In Hylomantis lemur,
hundreds of F1 offspring produced in
Atlanta have been dispersed to over 15
facilities in three countries, increasing
the ex situ safety net for this particular
species. Offspring of the rocket frog,
Colostethus pratti, have also been
distributed. Other taxa such as the
marsupial frog Gastrotheca cornuta,
despite several successful breeding
evenLs, Iuve proved dIIhcuIL In Lerms
of raising juveniles to adulthood due to
nutritional issues. This indicates that
there is still much to learn about the
captive care of many species.
El Valle Amphibian
Conservation Center (EVACC)
The drastic declines observed at
El Cop and predicted in El Valle
instigated the ARCC project where
amphibians were exported due to
lack of facilities. However, following
very closely behind was the concept
of performing this work in Panama.
Working in partnership with the
El Nispero Zoo in El Valle, the
Houston Zoo spearheaded the design,
installation, and implementation of
LIe hrsL In-counLry IucIIILy In Punumu.
This facility, the El Valle
Amphibian Conservation
Center (EVACC) was
created to provide an in-
country ex situ facility to
house assurance colonies
of critically endangered,
regional endemics, and
oLIer specIhcuIIy seIecLed
species that could be
maintained long-term
until pertinent threats
could be mitigated and
the species could be
reintroduced. This idea
could not have come
sooner as in April 2006,
LIe hrsL posILIve Bd test
and accompanying sick
and dying frogs were
found in El Valle!
Groundbreaking of
EVACC occurred in August
2005 at a site on the
grounds of the El Nispero
Zoo, but it would be nearly
18 months before it was
ready for animals. With
this, the Hotel Campestre
increased their already
sIgnIhcunL conLrIbuLIons
to amphibian conservation
by offering essentially an
entire wing devoted to
quarantining hundreds of
amphibians, raising their
food, and housing literally
hundreds of volunteers
from zoos, aquariums and
other institutions around
the world who came to
help in this massive triage
effort. In May 2006,
ANAM granted collection permits
and, using a ranking system similar to
that of the ARCC project, amphibians
were collected and maintained in
IndehnILe quurunLIne uL LIe CumpesLre
facility. As in the ARCC project, all
animals were carefully cataloged
Hylomantis lemur, the Lemur Leaf Frog, has been reproduced in both ARCC
and EVACC programs and many ofspring distributed to institutions for
safeguarding. Photo by Ron Holt, courtesy Atlanta Botanical Garden
and underwent prophylactic treatment for Bd as they
entered the program. Strict quarantine protocols were
implemented including dedicated, disinfected footwear
and other equipment that was not transferred between
rooms. Disposable, powder-free, vinyl gloves were used
for each individual enclosure. All discarded gloves,
refuse, and other materials were disinfected with 10%
sodium hypochlorite (bleach) before being discarded
to prevent any spread of pathogens from the collection.
AII oor surIuces In und ouLsIde oI LIe rooms und LIe
immediate areas were also disinfected daily with 10%
sodium hypochlorite. Individuals were fed daily or
every other day depending on species, and enclosures
and substrates were cleaned and disinfected with 10%
sodium hypochlorite on the days between feedings. This
allowed better fecal monitoring and removal of uneaten
food items. Water used for the amphibians was potable
tap water run through a 0.5 m curbon bIock hILer
to remove any chlorine, pesticides, or other harmful
subsLunces. InuIIy, In MurcI oI zoo;, LIe hrsL unImuIs
were moved into the EVACC facility!
Above: The EVACC educational exhibit area will help visitors to
the El Nispero zoo learn more about the amphibian crisis and local
response. Photo by Edgardo Grifth, courtesy Houston Zoo.
Left: Visitors to the El Nispero Zoo learn more about the eforts
behind the glass of the El Valle Amphibian Conservation Center.
Photo by Brad Wilson.
Being in country, the EVACC facility
would allow Panamanians to produce
their own golden frogs, safeguard
critically endangered amphibians
(without the major logistical issues
of exporting them), and educate
visitors to the El Valle region. Golden
frogs, highly revered for good luck
and posterity in Panamanian culture,
are important to tourist destinations
throughout the country. However,
between habitat loss, over-collecting
and Bd, their existence in the wild
has become seriously threatened.
Supplying captive-born offspring
for educational displays would help
decrease the pressure of wild collecting.
EVACC also acts a repository for
critically endangered amphibians from
the region, where these species can be
maintained long-term while threats are
addressed and, hopefully, species can
be reintroduced.
For both ARCC and EVACC, access to
amphibian food items was one of the
major challenges. In an emergency
response situation, time does not
always allow for proper setup of
adequate food colonies prior to starting
the project. Some food items such as
IruIL Ies (Drosophila melanogaster)
and springtails (Collembola sp.) were
cultured during the course of ARCC
and also EVACC; however, collecting
insects by hand, net or light traps
remained a part of the daily routine.
Providing adequate quantities of
the appropriate food proved to be a
challenge and careful record keeping
regarding the eating habits of the
different taxa allowed staff to modify
diets over time as the animals adapted
to different food items. Fortunately, an
in-country source of domestic crickets
(Acheta domestica) provided stock to
initiate permanent colonies at EVACC
and, more recently, methods to culture
katydids, springtails and roaches have
been perfected to offer more diversity
in diet and nutrition.
Like its predecessors, the EVACC
has seen success in small steps,
with the primary measure being to
keep some of these species alive.
Certain species, such as Hyloscirtus
palmeri and H. colymba have been
impossible to maintain in captivity
for any length of time both in and
out of Panama. Many taxa are doing
well, including Atelopus zeteki, A.
varius, Anotheca spinosa, various
Pristimantis species, Gastrotheca
cornuta, Hemiphractus fasciatus,
Hylomantis lemur and others. We
must keep in mind, however, that
the simple act of maintaining or
breeding threatened taxa in captivity
is not in itself conservation. The next
challenge is being able to produce
multiple generations that are carefully
managed genetically for potential
reintroduction. Many problems
remain to be solved, including those
uLLrIbuLed Lo nuLrILIonuI dehcIencIes
and veterinary issues as these new
generations are produced. Anotheca
spinosa, Pristimantis diastema,
Gastrotheca cornuta, Hemiphractus
fasciatus, and Hylomantis lemur have
been successfully reproduced so far
and dozens of captive-bred amphibians
exist as a result. Unfortunately, some
of the most critically endangered
Luxu Iuve proven dIIhcuIL boLI uL LIe
EVACC or Atlanta facilities and it
may turn out to be too late for those
species. Owing to these challenges,
work with surrogate taxa or hormone-
induced breeding techniques are
being employed as alternatives. All
amphibians that have died have
been submitted to Dr. Allan Pessier
at the San Diego Zoological Society
for comprehensive necropsy. The
mujor probIems IdenLIhed LIrougI
necropsy results are nutritional issues
and lungworm infections. The most
important success for EVACC is that,
thanks to many institutions and
individuals, there now exists a facility
in Panama where trained staff care
for and manage critically endangered
species that have much greater chances
of making their way back into their
natural habitats. More information
can be found at www.houstanzoo.org/
amphibians.
If we were able to do it all
again.
or LIe hrsL seven yeurs oI ucLIve
amphibian ex situ work in Panama,
there existed no ex situ facilities
in country and, therefore, animals
had to be exported to other places
for safeguarding. The price of doing
nothing was the potential loss of
species plus the associated real costs
for the projects. Doing nothing was
not an option. While PGF and ARCC
brought animals out of country for
Oophaga vicentei, a spectacular poison frog from Panama. Photo by Brad Wilson, courtesy Atlanta
Botanical Garden
safeguarding, EVACC sought to develop
a complementary program in Panama.
All three projects had successes and
limitations. These activities were
invaluable learning tools, and by
identifying aspects that work or do
not work, we gained a great deal of
knowledge and experience in several
key ureus:
1. Biosecurity
The risks involved in moving animals
and the pathogens they may harbor
across geographical barriers and the
potential of them exposing native
populations to new diseases are
tremendous. Having proper dedicated
space, staff, and protocol is crucial
in maintaining hygiene within the
collection and keeping any potential
pathogens contained. While testing
for Bd and other diseases is becoming
more common within institutional
and private collections, what about the
diseases that are either yet unknown
to us or for which there is no test? Just
because we cant see it or test for it yet
does not mean we should not practice
careful hygiene. Keeping animals
within their range country, ideally
within range habitat, greatly reduces
the risks associated with keeping living
collections and the introduction of new
pathogens.
2. Politics
Obtaining the permits for collecting
and exporting animals is very time
consuming and expensive, requiring
multiple visits with authorities and
submitting clear and open proposals.
Making sure there is also a positive
public perception within the range
country is also importantmany view
the export of any natural resource from
their country with a sense of regret or
suspicion, regardless of what they are
told. Of course, internal regulations
at zoos and other institutions can add
their own twist to the situation. Again,
keeping as much work in-country as
possible will help circumvent these
types of problems. In addition, there
must be transparency. Range country
citizens may be initially skeptical
of these programs but later become
excited and supportive after seeing
LIese IucIIILIes und projecLs hrsL Iund.
TIe oIhcIuI pubIIc openIng oI EVACC
on April 18, 2008 was not only a reason
to celebrate the completion of the
wonderful educational wing of the
center, but more importantly it was
a great way to build awareness and
support within Panama.
3. Resources and funding
All resources including funding,
sLuIhng, und In-counLry IogIsLIcs
should be carefully considered prior
to beginning these types of projects.
Funding for ARCC was sporadic
at times, resulting in some minor
delays with aspects of the project. The
Top: . The Crowned Tree Frog, Anotheca spinosa. Photo by Brad Wilson, courtesy Atlanta Botanical
Garden
Above: A new species of gliding tree frog from the El Valle area. Ecnomiohyla rabborum was
described in 2008 and named in honor of the contributions of world-renowned conservationists
George and Mary Rabb. Photo by Brad Wilson, courtesy Atlanta Botanical Garden
rapid response required accelerated
LImeIInes, und IL wus dIIhcuIL Lo puL
all necessary funding in place prior to
beginning the project. There will always
be unseen or unexpected costs to deal
with, on top of what was planned.
HuvIng uII oI LIe Iunds, sLuIhng,
and other resources fully in place at
the initiation will certainly help the
project move ahead more smoothly.
Again, in-country programs have
severuI udvunLuges Irom u hnuncIuI
perspecLIve: muLerIuIs und Iubor Lo
construct facilities is generally less
expensive outside of the U.S., as could
also be the cost of hiring and training
staff. A great deal of materials were
purchased for ARCC and EVACC
and shipped in. We must remember
that each country is different and the
costs in Panama, for example, may
be different from other countries. If
the airfare, lodging and other travel
expenses were removed from the
budget of an ex situ project that was
carried out in country instead, the cost
suvIngs couId be sIgnIhcunL. ncIude
the difference in cost of labor between
the U.S. and Panamanian institutions
and there are even greater savings by
keeping these projects in country as
much as possible.
4. Capacity
There are over 1,200 zoos and
aquariums in the world and if each
one had the ability to take on ONE
amphibian species, then we would be
able to rescue all 500 species needing
rescue. This, of course, is not the case.
In fact, while some institutions are
taking active roles in partnerships
that safeguard species, many simply
do not have or will not reallocate the
resources (hnuncIuI, LecInIcuI, socIo-
political) to carry out these projects.
There are currently over 44 threatened
amphibian species being worked with
at some level in ex situ programs
worldwide (!), a far cry from what is
needed considering there are around
500 species so critically threatened that
we are likely to lose them without ex
situ work. Global capacity is clearly an
issue.
In regards to working in Panama, the
ARCC did not have the capacity to
carry out its activities beyond a certain
time frame. Although it is permanent,
cupucILy oI EVACC Is noL InhnILe so
plans must be made to accommodate
hundreds of offspring as the various
species reproduce. Potential ideas
and solutions for future facilities are
reviewed by Pfaff and Crump (2007).
With limited space and resources,
it is important to prioritize species
for ex situ work and insure that such
valuable resources are spent wisely.
The Smithsonian Tropical Research
Institute (STRI) hosted the three-day
Panamanian Amphibian Prioritization
Workshop from the 12th-14th of
November 2008. The workshop was
well attended with thirteen participants
from ten different organizations. A
total of 204 species were evaluated
using the prioritization tool developed
by Amphibian Ark. The workshop
purLIcIpunLs worked very eIhcIenLIy,
completing all evaluations after
two days. The third day was spent
discussing both implementation
of the workshop results, and the
production of a larger national
amphibian conservation action plan.
TIe purLIcIpunLs IdenLIhed Iour busIc
groupings of species based on their
conservation need and thus dictates
their ex situ role.

1. Species that are likely extinct (ARK
10%).
2. Species that are stable now but will
be exposed to threatening processes
soon (RESCUE 15%).
3. Species for which the conservation
status are completely unknown (NONE
32%).
4. Species that require no conservation
action (NONE 43%).
All the species that fall into the
LIIrd group were ugged Ior in situ
assessment work.
Both ARCC and EVACC had to contend
with other issues that are taken for
granted at U.S. institutions, such as
the ability to order in food items and
other supplies, as well as easy access
to veterinary consultations. In-country
regulations in Panama made bringing
in domestic crickets (a common
and useful food source for many
cupLIve umpIIbIuns) dIIhcuIL, und LIe
manpower needed to collect local food
items only increased as the collection
grew. Another detail that cost varying
amounts of time for PGF, ARCC and
EVACC was the lack of bilingual
volunteers and staff.
Conclusions
Amphibians are now facing more
challenges than ever, making it
IncreusIngIy dIIhcuIL Lo sLuve oII
numerous species extinctions and
protect valuable biodiversity. The
threats are not likely to subside soon,
and for many species time is of the
essence. With the collaboration of
zoological institutions, academic
researchers, private breeders and
others, an opportunity to make a
difference is within our grasp. As future
conservation stewards, members of
these groups can begin by using the
recommendations proposed by the
Amphibian Ark (www.amphibianark.
org); Zippel et al., 2006) and ACAP
(Moore and Church, 2008) as a guide.
Ex situ work, used solely as a stopgap
while threats are mitigated, will be
critical for safeguarding many species
on the brink and, ultimately, to their
long-term survival in nature. The work
must be carefully carried out with great
attention to planning, implementation,
and exit strategies along with a serious
commitment to tasks at hand.
Our decade of conservation work in
Panama has demonstrated several key
prIncIpIes Lo rupId response: IuvIng
clear goals for the project (including
an exit strategy), proper infrastructure
(from administration and funding to
facilities and personnel), and trained
and committed staff to carry out the
planned activities to their logical end.
With careful planning, working within
the range country when possible, and
implementing appropriate biosecurity
protocols, ex situ operations may help
us in safeguarding many critically
endangered amphibians.
__________________
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K., Marantelli, G. & Parkes, H.
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& Parra-Olea, G. (2005): Amphibian
population declines in Latin
America: A synthesis. Biotropica 37:
222226.
Lips, K. R., Brem, F., Brenes, R., Reeve, J. D.,
Alford, R. A., Voyles, J., Carey, C., Livo,
L., Pessier, A. P. & Collins, J. P. (2006):
Emerging infectious disease and the
loss of biodiversity in a Neotropical
amphibian community. Proceedings
of the National Academy of Sciences
103: 3165-3170.
Moore, R.D. and Chruch, D. R. (2008).
Implementing the Amphibian
Conservation Action Plan.
International Zoo Yearbook 42(1):15-
23.
Nichols, D. K. & Lamirande, E. W.
(2000): Treatment of cutaneous
chytridiomycosis in blue-and-yellow
poison dart frogs (Dendrobates
tinctorius). Getting the jump! on
amphibian disease: 51. Moore, K. &
Speare, R. (Eds). Cairns: Rainforest
CRC.
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Johnson, K. (2008). Amphibian
Ark and the 2008 Year of the Frog
Campaign. International Zoo
Yearbook 42:(1):2429
Pfaf, S. and Crump, P. (2007) Chapter
6: Building ex situ facilities within
range countries. Amphibian
Conservation Resource Manual: 31
41. Grow, S. and Poole, V. A. (Eds).
Association of Zoos and Aquariums,
Silver Spring, USA. http://www.
aza.org/ConScience/Documents/
Amphibian_Resource_Manual.pdf
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& Gorman, G. C. (1997): Test of null
models for amphibian declines on
a tropical mountain. Conservation
Biology 11:13071322.
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Panamanian Golden Frog: Proyecto
Rana Dorada. Herpetological Review
33: 11-12.
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CBSG/WAZA amphibian ex situ
conservation planning workshop fnal
report. Apple Valley, MN: IUCN/SSC
Conservation Breeding Specialist
Group.
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Richards, C. L., Jaramillo A., C. A. &
Grifth, E. J. (2007): Implicaciones en
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de Panam, asociadas con su
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3: 2939.
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The Amphibian Extinction Crisis: A
Call to Action. Herpetol. Rev. 33:23-29.
__________________
ASSOCIATIONS
Paul Crump
Houston Zoo, 1513 MacGregor Way
Houston, TX 77030
USA
Ron Gagliardo
Amphibian Ark, Zoo Atlanta
800 Cherokee Ave
Atlanta, GA 30315
USA
Edgardo Grifth
El Valle Amphibian Conservation Center
El Valle de Anton, Panama
Heidi Gross
El Valle Amphibian Conservation Center
El Valle de Anton, Panama
Joseph R. Mendelson III
Zoo Atlanta
800 Cherokee Ave
Atlanta, GA 30315
USA
Kevin Zippel
Amphibian Ark/CBSG
12101 Johny Cake Ridge Rd
Apple Valley, MN 22124
USA
Atelopus zeteki | 19
In March of 2008, I took a
LrIp Lo Punumu wILI LIree IrIends: Lwo
Dutch and one Swede. I had already
traveled with my two Dutch friends to
Costa Rica in 2006, but this was the
hrsL LIme my SwedIsI IrIend wouId see
poison dart frogs in their natural habi-
tat.

The primary reason we travelled to
Panama was to see the many color vari-
ations and morphs of Oophaga pumilio
and Dendrobates auratus in the wild.
We also wanted to observe various
types of birds, such as the well-known
but secretive Quetzal. We were also
IopIng Lo hnd LIe purporLedIy exLIncL
species Atelopus zeteki.
n zoo6, uILer u BBC crew hnIsIed
hImIng LIIs specIes In LIe wIId, bIoIo-
gists from the Smithsonian Institute (as
well as other institutions) collected the
Iew survIvIng unImuIs LIey couId hnd
in the area in order to maintain them in
captivity. A number of zoos (including
institutions in Panama and the United
States) placed them into captive breed-
ing programs, many of which have
succeeded. Since this extraction, from
what Ive been told by several sources,
biologists from the Smithsonian
Institute have unsuccessfully attempt-
ed to locate more. Because of this,
Atelopus zeteki was declared extinct in
the wild in 2007. I was also informed
that the last living animals in the area
were most likely a few toads held at
Hotel Campestre in El Valle de Antn,
Panama, where they are kept in an out-
door vivarium beneath the roof next to
LIe recepLIon oIhce.

We began planning our search for A.
zeteki after arriving in Bocas del Toro.
We talked with people who knew
where Atelopus could previously be
found in the area between El Valle and
Campaa, but were told that chytrid
1

had devastated amphibian populations
in this area over the last few years. One
of the people we met knew about a se-
cret project for conserving A. zeteki in
the wild. Our hope was that we could
somehow get permission to visit this
projecL und, II we couIdn`L munuge, hnd
it on our own, could do so with the help
of local people. We had the name of a
person who knew where you could still,
uL IeusL LIe prevIous yeur, hnd ALeIopus
In LIe ureu. We Ioped Lo hnd IIm und
have him come with us.
Our Trip to El Valle
While in Cerro Punta, near Volcn
Bar in western Panama (also known
as Volcn de Chiriqui), we succeeded
BY DENNIS NILSSON
1 Batrachochytrium dendrobatidis is a fungus that, when contracted by an amphibian, causes the deadly chytridiomycosis. I will refer to the disease by its
shortened name chytrid throughout the rest of the article.
Atelopus zeteki
in reserving rooms at the Hotel
Campestre via telephone. This way we
would at least have a chance to see A.
zeteki alive, even if not in the wild. It
wus noL eusy Lo hnd vucunL rooms on
such short notice, and we where lucky
Lo hnd uny, especIuIIy sInce none oI us
spoke Spanish.
The next day we left the highland for-
est of Bar and made our way to El
Valle by way of David, Santiago and
Penonome. Although Cerro Punta had
been comfortably chilly, the tempera-
ture in the lowlands was between 30-
35C (86-95F). We were glad that our
car, a Toyota Land Cruiser Prado, had
air conditioning.
Many of the areas we passed through
seemed very dry, even desert-like, mak-
ing it unbelievable that these wonder-
ful toads can exist here. However, its
the many rivers and streams in the
region that make it possible for them
to survive. Once we reached Antn, we
turned off the main road and headed
up toward El Valle.
After seven long hours of driving we
arrived at El Valle. We were tired,
even though the three of us had taken
turns driving.
We immediately
checked into the
Hotel Campestre
and unloaded our
bags, then left
Lo hnd MurIo uL
the Serpentario,
a place where he
kept terrariums
with several kinds
of snakes on dis-
play. When we
arrived, however,
he was nowhere
to be found. We
did manage to talk
to his neighbour,
a very drunk man
(bottle in-hand)
wIo we hnuIIy un-
derstood to be tell-
ing us that Mario
could usually be
found there around
7 a.m.

While driving back through El Valle, we
stopped at a park you could visit for an
entrance fee. We talked with one of the
empIoyees LIere wIo conhrmed wIuL
we Iud Ieurd In Bocus: LIere wus, In
fact, a conservation project for A. zeteki
in the wild. We were also able to get
the names of places that werent on our
maps, including the name of the river
and area where the project was located.
Although this seemed to be good in-
formation, we were disheartened that
we couIdn`L hnd someone wIo couId
give us more details about the area and
projector perhaps even take us to the
project.
El Nispero
We decided to visit El Nispero, which
is a local zoo that had vivaria contain-
ing A. zeteki. We were suspicious that
these might be new animals that
were on display (I dont actually know
if these Atelopus came from the differ-
ent projects here and were provided by
biologists, or if they were animals they
had caught themselves. We had previ-
ously heard about the A. zeteki at El
Nispero, but were told they were very
old animals and were no longer alive,
so we were surprised to see these ones
on display).
In addition to the beautiful pair of
Atelopus, they also had many different
birds such as owls, parrots, and pea-
cocks (including albinos), as well as ta-
pirs, monkeys, jaguars and many more.
The unfortunate thing was that the
animals were in small cages and didnt
appear healthy, even though the per-
sonnel probably did their best for the
animalsI assume they just didnt have
the resources to provide better care and
enclosures for the animals.

There was also a large concrete cage
where the zoo had previously kept
A. zeteki, but it was no longer in use.
Perhaps this was because the risks
of animals being infected by chytrid
where a lot higher than in a regular
glass vivarium? I assume it was most
likely because it is easier to clean and
treat the animals in a glass enclosure if
they do happen to get infected.
Planning the Search
After observing the animals at El
NIspero, we IeIL Lo hnd u resLuurunL
where we could eat dinner and discuss
plans for the following day (the res-
taurant at the hotel wasnt available to
us because the hotel, except from the
two rooms we were renting, had been
Vegetation in the highland forests that surround Volcn Bar.
reserved for a French school). The food
we ate at the restaurant was delicious,
although it probably would have been a
lot better if we spoke Spanish and had
ended up with the food we thought we
were ordering.
We took out our maps and started
pIunnIng wIere we mIgIL hnd some
Atelopus, looking for distant places
where people didnt seem to live or per-
haps hadnt looked before. Along with
our plans, we also need a little bit of
luck, making sure we get started early
enough before the temperature got too
IIgI: In some pIuces In LIIs regIon IL
can get above 30C (86F), even if the
temperature in windy El Valle was only
around 24C (75F). After a while we
decided that this would be our reserve
plan if we werent able to contact Mario
und receIve IIs IeIp In hndIng LIe
project we were looking for.
We hnIsIed LIe meuI wILI u bununu
split and, after a couple of tries, suc-
ceeded in getting four cups of black cof-
fee, after which we made our way back
to the hotel to shower and get a couple
hours of sleep before it would be time
to get up again.

No Help
We rose with the sun the next morn-
ing around 6 a.m., eating breakfast
at the same restaurant we had eaten
dinner the night before. We wanted to
hnd MurIo us soon us possIbIe so we
couId hnd some ALeIopus beIore IL goL
too hot. So, after a very tasty omelette
and freshly made pineapple juice, we
went back to the Serpentarioonly to
hnd LIuL no one wus LIere. We wuILed
around for a half-hour, just to see if
someone might show up. The garden
there was very nice and there was a
small stream running through it. Small
green basilisks (Basalisca plumifrons)
were running aroundwe even man-
aged to see one run across the water
when it got scared, which was very in-
teresting to witness.
After the half-hour had passed we de-
cided to give up waiting and attempt
LIe seemIngIy IopeIess mIssIon Lo hnd
the amphibians ourselves. We had at
least some idea as to where to start
IookIng, und even II we couIdn`L hnd
any Atelopus, we were convinced that
it was going to be an interesting day
und LIuL we wouId no doubL hnd muny
other interesting animals.
A Search of Our Own
We took off in the car heading toward
our previously decided location. When
we got there (which took a while since
it was quite some distance away) we
couIdn`L even hnd LIe rIver. We usked
some kids in a village where it might
be, and they asked which river we
meant. Evidently there were several
rivers in the area, so we decided to visit
whichever one was closest.
The area was very dry and hot and the
vegetation wasnt what we where look-
ing for, so we followed the river up-
sLreum Lo hnd more suILubIe ALeIopus
habitat. We eventually found a more
suitable spot upstream and stopped to
watch some type of beautiful red bird.
At this location we found a tiny, dried-
out streambed that lead into the main
river channel. I only had one pair of dry
shoes left, and even though I hate wear-
ing rubber boots, I decided to put them
on anyway.
Even though it was the dry season and
the water level in the river was low, the
currenL wus sLIII owIng sLrong und
fast. The river was about 10-15 meters
wide with stones and gravel along its
edges. At some places the gravel almost
looked like sand. Large rocks and
boulders were everywhere, both in and
around the river. Along the shores were
larger, older trees mixed with palms.
At this location was some type of
grasshopper that called quite loudly,
and when multiple grasshoppers
started to call, it created an extremely
high and noisy chainsaw-like sound.
While my companions were walking
downstreamthey didnt have rubber
boots on and the shoreline was drier in
LIuL dIrecLIon- munuged Lo hnd one
of the grasshoppers. As I stood there
photographing it, changing lenses and
moving slowly so as not scare it, my
Swedish friend came back up to watch
what I was doing. Later on I discovered
he thought I was photographing the
beautiful Atelopus zeteki that was hop-
ping around just in front of me (I had
noticed that there was something mov-
ing around, but was so focused on tak-
ing pictures of the grasshopper, didnt
bother looking more closely at it).
It was a big, beautiful female. She was
mostly yellow with a few black spots
along her back (its primarily the males
that have the characteristic arrow-
shaped markings on their backs. They
are also quite a bit smaller in size than
the females). Its surprising how much
they look like skin and bones, but this
is exactly how they should look. We
didnt expect them to be as fast as they
The habitat of Atelopus zeteki.
ended up beIng: Louds normuIIy Lend Lo cruwI, buL A. zeteki
is very good at jumping. They werent at all afraid to dive
sLruIgIL InLo LIe coId, IusL-owIng wuLer oI LIe rIver, even
though you would think they would try to avoid it so as not
to get caught up and drowned by the swift current. When
they leap into the water, they go straight to the bottom like a
dry, yellow leafits almost impossible to distinguish them
with their yellow coloration and black markings. Their
coIoruLIon serves us boLI u perIecL cumouuge Ior uvoIdIng
discovery as well as a strong warning to predators of the
toxins contained in their skin. The temperature was surpris-
IngIy wurm: uround -C (q1-q) by LIe rIver und ;C
(99F) at the car.
IeL my IrIends Luke LIeIr pIcLures hrsL sInce wunLed Lo
make sure I got good photographs, and for that I would need
more space. I decided to leave the group for a little while to
see II couId hnd some more ALeIopus, desperuLeIy IopIng
my friends wouldnt lose the one we had found. I thought
that it would be nice to see two males meet and wave at
each other. Atelopus zeteki, as well as A. varius, has devel-
oped a sort of hand-waving signal as the noise of the streams
they inhabit make it almost impossible for their vocal calls to
be heard.
UnIorLunuLeIy, wusn`L ubIe Lo hnd uny more ALeIopus, even
though I was convinced there should have been more of
them. If it had been raining or the temperature was a little
cooler there might very well have been, because it seemed
like the perfect place with all the proper conditions for
Atelopus.
I eventually went back and took my photos, and afterward let
the beautiful A. zeteki crawl beneath some dry leaves. After
this I crossed the river, which wasnt easy, regardless of how
low the water seemed. It was deeper and more slippery than
I had thought it would be, but I managed to cross without
getting soaking wet. Once on the other side, I found quite a
few spiders sitting on the rocks in the sun. Some of them had
a grey-marbled coloration, looking exactly like part of the
rock.

We dIdn`L hnd uny more ALeIopus Iere, buL `m one Iundred
percent positive that this was the perfect habitat and condi-
tions for Atelopus zeteki and that there should had been
more present. Many of them could have been hiding because
it was so dry and hot. Perhaps the one we found was one
of the very last surviving A. zeteki in
this location? This was hard for us to
beIIeve: LIe dry surroundIngs Iud been
something we had dealt with in previ-
ous dendrobatid trips, and during the
dry season many frogs tend to seek
cover or congregate around rivers and
streams, which provides a much small-
er area for you to search. Our hope was
that other Atelopus in this area were
simply hiding under cooler cover.
No More Atelopus Found
We continued to make our way up-
stream where we found species of both
Colostethus and Eleutherodactylus.
There were many different animals in
LIIs ureu und pIenLy oI beuuLIIuI InsecLs:
buLLerIes oI uII coIours, grussIoppers,
us weII us dIIIerenL Lypes oI drugonIes.
After a couple more unsuccessful hours
oI seurcIIng, we decIded Lo hnuIIy gIve
up and head back to El Valle.
Male Atelopus zeteki live year-round
near the rivers, while females inhabit
the forest, only venturing to the riv-
ers Lo hnd u muLe und breed. We were
worrIed ubouL onIy hndIng LIIs one
female, but the heat and extremely dry
area made it very hard to know whether
or not she was the only individual left
inhabiting the area, or if others had
sought refuge. It could be that the fe-
male we found had been a juvenile in
2006-07 when all the other animals
disappeared, and had returned to the
river to mate. I hope that Im wrong,
and that A. zeteki can still be found
there.
Back to El Valle
When we got back to El Valle, we went
to the market to buy some souvenirs
before they closed for the day. I bought
some souvenirs of rana dorada (gold-
en frog in Spanish). We also talked
with some Americans that had moved
to the area and lived there throughout
the year. We showed them some of our
photos and they were very interested in
us telling them where the exact location
was we found the Atelopus, but we had
already decided to keep quiet about it
and talk only with the scientists that
work in the area and with this project.
After a very inexact and short descrip-
tion that they would never be able to
decipher, they told us that sometime in
LIe IoIIowIng weeks u hIm Leum Irom
Animal Planet was going to attempt to
hnd LIe goIden Irog uguIn. Ieurned
later that they did succeed, but I dont
know how many animals they found
or the exact spot in which they found
themif it was the same spot, then I
hope it remains a secret. I also heard
that there were many articles written
about this in the newspapers and that
it was blown up big in the media. Its
devastating for the last Atelopus zeteki,
but perhaps the authorities will open
their eyes and try to better conserve
this threatened species.
Top: One of the chainsaw-like grasshoppers found along the river.
Middle: Pristinmantis ridens
Bottom: Cranopsis melanchlora
After a shower and
change of clothes we
still hadnt had enough
from A. zeteki, so we
took the car and drove
to the area where we
thought the project
we had heard about
might be. We had to
ask about names that
werent on our maps,
and eventually we got
a good idea as to where
the project was.
To get there we had to
drive up an extremely
steep road. After an
unsuccessful attempt
to navigate the gravel
road over the moun-
tain we decided to give
up, mostly because we
were worried that we might damage
the rental car, but also because we were
content with what we had already seen
and experienced that day. The fact that
we were all hungry also helped make
the decision to head back into town.
More Atelopus Habitat
On our way back we stopped by one of
the small streams which, at least in an
area further downstream, is known to
formerly be habitat for A. zeteki. We
saw quite a few tadpoles in some of the
protected areas of the stream and man-
aged to catch some with a net, but it
wus dIIhcuIL Lo see exucLIy wIuL kInd oI
of tadpole they were. There were many
Iouses Iere, und hgurIng LIuL we were
most likely on someones private prop-
erty, we chose to go back to the car.
But right at that moment I saw some
sort of leafor tree frogleap into
LIe gruss. Ike u reex, swung my
camera onto my back and caught the
frog. I immediately regretted catch-
Ing IL: LIe Irog`s skIn wus more or Iess
falling off and some of its entrails were
hanging out of its side. I had never seen
anything like it. I was so shocked and
disgusted by it that I just let go. I didnt
take any photos of the frog, but think-
ing back, regret that I hadnt.

WIen we hnuIIy urrIved uL u resLuurunL
in El Valle, I washed my hands before
eating as I was still disgusted by the
malformed frog. After enjoying a very
nice meal we returned to our hotel,
where I noticed on one of the walls
a framed article with a photo of the
same species of frog I had caughtand
it was also loosing its skin. It looked
almost as horrible as the one I had
found. Unfortunately, the article was in
Spanish and I could only understand
small parts of it. (If someone read-
ing this knows what this condition is,
please contact me as I would appreciate
knowing more about it.)
The End of the Road
Back at Hotel Campestre we had a
few beers to celebrate our successful
day. Around us was an entire school of
French kids running around and play-
ing, teasing and making a lot of noise,
but it was fun. Our Panama trip had
only a day and a half left, which meant
we would drive to Panama City the next
day, where we would stay the night
beIore yIng buck Iome. DurIng LIuL
drIve we ended up IuvIng LIme Lo hnd
a couple more species of dendrobatids
along the way, but that is a different
story
Conclusion
Due to the risk of jeopardizing any part
of the science and conservation project
that is active for Atelopus zeteki in the
area around El Valle de Antn, I have
chosen to omit some details regarding
specIhc numes oI pIuces und peopIe.
The scientists that work with Atelopus
zeteki, both in situ and in the USA,
have received our information and my
photosit will hopefully serve as a bit
of help to keep the species alive in the
wild.
The biologists I talked to said they had
not seen any Atelopus zeteki in the area
we visited since the extraction project
in 2007.
There have since been new reports
about methods that could be used to
treat or mitigate the effects of chytrid
on amphibians in captivity, but as of
yet there still remains no safe way to
stop the fungus in the wild. The only
method currently seems to be catch-
ing wild amphibians and treating them
in captivity, then releasing them back
into the wildbut upon reintroduc-
tion, their chances of re-infection are
extremely high. There are new theories
regarding limited areas being treated
with certain types of bacteria that may
inhibit the growth of chytridiomycosis,
but we still have a long way to go before
we have something that is both effec-
tive and safe to use in the wild.
On the Trail of the Golden Frog | 25
Tose who have viewed at rst hand the steep, dark-green, forest-
covered slopes of the Cordillera de Talamanca-Chiriqui of Costa
Rica and Panama, with their ever changing aspect of sun and cloud,
moon and mist, bright blue sky and bright green mantle, driving rain
and boiling fog, come away with a feeling of overpowering awe and
mystery at the variety of nature and the magic of the human soul. It
is not surprising that the primitive peoples in this region also regarded
the mountains and their forests with mystical reverence, so near and
yet towering abruptly upwards to 4,000 meters from their lowland
valley habitations.
Among the Bribri, Cabecar, Boruca, Changina, and Chiriqui, when
the chicha has been drunk, the night grows late and dark, and the res
die down to burning embers, the wisest old man of the tribe tells his
engrossed listeners of a beautiful miraculous golden frog that dwells in
the forests of these mystical mountains. According to the legends, this
frog is ever so shy and retiring and can only be found after arduous
trails and patient search in the dark woods on fog shrouded slopes and
frigid peaks. However, the reward for the nder of this marvelous
creature is sublime. Anyone who spies the glittering brilliance of
the frog is at rst astounded by its beauty and overwhelmed with
the excitement and joy of discovery; almost simultaneously he may
experience great fear. Te story continues that any man who nds
the legendary frog nds happiness, and as long as he holds the frog
happiness will follow him everywhere. Te story tellers record many
men who have scaled the highest peaks and searched the darkest forests
for even a glimpse of the golden frog, but only a few ever see it. Fewer
still capture the cherished creature and hold him for a few moments,
and a very few are able to carry him with them for a longer period of
time.
Jay M. Savage
On the Trail of the Golden Frog: With Warszewicz and Gabb
in Central America
AN OFFER ON AN INTERNET WEBSITE READS:
FOR SALE - Panamanian Green and Black auratus. You
scroII down IurLIer und see unoLIer one: BIuck und BIue
auratus male wanted; will trade for female. Does this sound
familiar to you? I assume so. These are common terms when
it comes to selling or trading captive-bred auratus, but they
beur u greuL dunger: LIe mIxLure oI severuI IocuI Iorms LIuL
may appear similar, but actually originated in completely
different geographical regions! The result is the creation of
a genetic mix that has nothing to do with the aims of so-
called captive breeding programs for the conservation of
endangered species, but is rather, if I may be blunt, nothing
but genetic rubbish. Harsh words? Perhaps. But if a dart frog
hobbyist is serious about the frogs he is keeping (lets say,
for arguments sake, the black and blue one mentioned
above that needs a mate), the least he can do is deal with
LIeIr orIgIns und hnd ouL exucLIy wIuL IL Is LIuL dweIIs wILIIn
his vivaria. Unfortunately, the diligent frog keeper will
oILen reucI LIe boundurIes oI IIs eIIorLs IuIrIy quIckIy: no
one seems to be able to tell him for certain where his frogs
orIgInuIIy cume Irom. He cunnoL hnd uny soIId InIormuLIon
just mere speculation.

uckIIy, In IIs seurcI Ior LIe source oI IIs Irog, Ie hnds un
online morph guide. Within a few clicks he observes a photo
oI u specIhc coIor morpI LIuL Iooks very sImIIur Lo LIe one Ie
owns.weII, IL`s u bIL durker (perIups IL`s LIe usI LIuL mukes
the photo look too bright or the settings on his monitor),
(Further) Notes
on Dendrobates auratus
BY FRANK STEINMANN
UNLESS OTHERWISE NOTED
The Variability of the Panamanian Green and Black Poison Dart Frog
Dendrobates auratus is doubtless one of the most common poison dart frogs kept in
captivity. However, just keeping and caring for the species in captivity is not enough
if we choose to speak in terms of conservation. The following article is meant to portray
the importance of understanding what a local morph is and, furthermore, how
signicant the handling of local forms, designated by their geographical distribution,
is for the future of keeping and breeding this member of the dendrobatid family.
Authors Note: The title refers to an article written by Dunn in 1941 that dealt with the polymorphism of D. auratus (See Literature at end of article).
(Further) Notes On Dendrobates auratus | 27
but it has to be his aura-
tus! After all, its blue, and
thats what really counts.
Now, proudly informed,
he begins referring to
his frogs as being of the
Calobre morph, hailing
from the region of Calobre.
Our keepers desperate
and anonymous search
now comes to a glori-
ous end as he locates and
obtains another Calobre
auratus to pair up with his
previously solo frog. The
one thing he never knew,
however, is that there
ure more LIun hve dIIIer-
ent populations of blue
auratus described to date,
not to mention the ones
yet undiscovered. Whats
even more, some local
populations tend to have
green and blue individu-
als that can interbreed and
create forms that contain
every imagineable nuance of green,
blue, and everything in between. Given
this, imagine the authority of a single
photo used to depict an entire popula-
tion of frogs. It would be like using
Arnold Schwarzeneggers countenance
to represent Homo sapiens (a matter
on which I have no further comment).
We must realize that even the best
herpetological image or online morph
guide should be used carefully.

How then do we continue? Well, it
seems necessury Lo hrsL Iocus on
the term morph. What exactly is a
morph? And beyond that, what is a
color morph or a local morph? As far
as I am concerned, there is no valid
dehnILIon oI LIese Lerms us LIey reIuLe
to poison dart frogs. However, if you
were Lo seek ouL u dehnILIon, you
wouId hnd LIuL LIe rooL oI LIIs word
is based on the Greek and pertains to
a things form, shape, gestalt basi-
cally the appearance of something.
When describing the distributional
runge oI u specIhc Iorm oI Dendrobates
auratus, it is best to use the term local
morph or local form. For example,
there is a population of Dendrobates
auratus found along part of the Rio
Santa Maria in western Panama. As
there is no distinctive point of natural
occurrence such as a hill or village we
could use as a reference, it is best to use
the Rio Santa Maria for describing the
frogs found here. These frogs also tend
to have a varied colorationthere are
green and blue ones that mix and cre-
ate turquoise specimens. In the hobby,
the tendency is to then separate them
further into different color morphs of
the population (e.g. the blue morph of
the Rio Santa Maria population, etc.),
even though in the wild they freely
associate and breed with other color
forms within the population.
Geographical isolation of certain popu-
lations leads to the evolutionary devel-
opment of local forms (local morphs
or populations), and as we can see, it
is deceiving to make decisions based
on color morphs alone because there
can be several different colors within a
single population. It seems a bit more
accurate, then, to refer to the overall
local morphs or populations that might
contain different color forms.
To help keep things uniform within the
international hobby, I feel there should
be international communications and
agreements about the usage of these
terms. However, the quantum of
scientists in Europe and the U.S. cant
even seem to agree about the common
taxonomic reviews of the family Den-
drobatidae, so how will we be able to
agree on even more challenging terms
und cIussIhcuLIons? AnoLIer dIIhcuILy
is hobbyists that have a Humboldt,
Jr.
1
form of self-entitlement. There
are more new species and morphs
currently being found than the public
can pay attention to, and illegal self-
imports, narcissistic name creations,
and false assertions to protect newly
discovered populations or to keep away
competing smugglers result in nothing
more than false information and confu-
sion.
In 1994 German biologist Holger
Birkhahn et al. published an article in
the aquarist magazine DATZ, which
dealt with these same problems
(Birkhahn, H., Klpmann, V. & Was-
smunn, K. (1qqq): Zur VurIubIIILuL des
GoIdbuumsLeIgers, q; (q): ;o-;6).
1 Alexander von Humboldt was a 17th century naturalist and explorer whose botanical work became the Ioundation Ior the feld oI biogeography. He
traveled extensively throughout Latin America, describing species that had previously been unknown to Europeans.
Dendrobates auratus habitat in Soberania National Park, Panama.
To deal with them, Birkhahn suggested
implementing a cryptic morph code to
distinguish between different locales
of frogs. For example APZ3 was the
code for Auratus from Panama, fol-
lowed by the geographic region (here
Z for the German word zentral
or cenLruI), und hnuIIy u number
that describes the fact that the frogs
were the third local morph found in
that area. This type of code bears the
advantage of keeping sensitive data
away from people with dishonorable
intentions (such as smugglers) while
at the same time forming international
agreement over the terms of necessary
coding. Birkhahns model, however,
never gained ground. This could in
part have been due to the lack of com-
munication between people, as well
as the uncertainty about the usage of
these codes. For instance, it seems a
bit like the Illuminati when a circle of
chosen people encrypt their frogs with
the intension to protect wildlifeusing
such coded language might just be
making things more complicated than
they should be, and situations where
people are trading ACP2 for ACA1 can
seem just plain confusing. The logi-
cuI quesLIon LIen urIses: wIy noL IubeI
frogs after their actual geographic
source? I very much doubt that smug-
glers would eradicate entire popula-
tions of Dendrobates auratus, which
has never been that popular when it
comes to illegal imports, especially
when compared to species of Oophaga
or newly discovered thumbnails, like
the case of Ranitomeya benedicta. The
rapid loss of habitat, deforestation, as
well as the spread of Batrachochytrium
dendrobatidis (chytrid fungus) seem
to be the primary threats for several
populations of D. auratus.
There are currently thousands of
auratus being kept all over the globe
individual frogs being traded, named,
renamed, then shipped to other con-
tinents to be named and renamed yet
again. The result is selected bloodlines
that have been bred simply for their
color and phenotypic characteristics,
which then led to the creation of new
names. Over the last few years import-
ers Iuve been oodIng LIe murkeL wILI
Dendrobates auratus. Never before
seen color morphs (I use the term
color morph, because in most cases
there was no information regarding
LIe specIhc orIgIns oI LIese Irogs) Iuve
been made available and sold in the
United States, Canada, and beyond
to Europe. Many of these frogs were
labeled as farm
raised auratus
but where exactly
are these farms?
I doubt they are
actually in Pana-
ma, because if you
follow the CITES
documentation
for D. auratus,
there were no legal
exports of them during those years
from this country. So who collected the
parental frogs and where were they col-
lected? Background information on the
generations sold (F1, F2, etc.) would be
necessary and valuable information to
form a professional breeding stock in
terms of conservation. When contact-
ing the expert importers I have found
that you only gain factoids (no actual
answers) or stories that may or may not
be true. Why is that? Economic ways
by which to support conservation?
Competition? Ignorance?
For example, could someone explain
the origin of the Superblue morph of
D. auratus? This morph was originally
described as being a selected bloodline
of another morph, and then somewhere
along the line it became a supposed
wild-caught population or, in some
cases, it even became a long-bred and
seemingly long-lost morph that was
re-introduced to the hobby. If I browse
the offerings of several professional
breeders and importers (the only thing
An example of the amount of diversity that occurs within the
Birkhahn population of Dendrobates auratus. Without the proper
information regarding their origin, its easy to see how each frog could
be used to create new morphs in the hobby.
I would consider pro are their frog
prIces) cun hnd severuI morpIs oI
Dendrobates auratus, all of which look
sLunnIng, buL Iuve uII sorLs oI urLIhcIuI
names that provide no solid informa-
tion regarding their actual origins.
One of the best examples of this confu-
sion are the frogs that Holger Birkhahn
and Klaus Wassman discovered in the
early 1990s. Soon after their discovery
there were several, rather grey, imports
to Germany of these frogs, which
were numed uILer LIeIr dIscoverers:
Dendrobates auratus Birkhahn or
Wassman. The phenotypes and visual
characteristics of these frogs were then
used Lo creuLe more specIhc und cre-
ative descriptions Bronce/Bronze
or Birkhahn Bronce were common
terms used to describe the very same
frogs. This morph actually tends to
create several color forms within
LIe sume popuIuLIon: bIue, green, or
turquoise patterns on a bronze/cream
to brownish background. In addition
to this, these patterns may change as
the animal ages, such as a lightening
of the background color. After a while
people began to differentiate between
these variations and using names such
as Birkhahn Bronce Green, Birkhahn
Bronce Blue, and Birkhahn Bronce
Turquoise not paying attention to the
fact that all these animals were actually
one and the same morph and that these
differences were naturally occurring
variation from within the natural popu-
lation. To add to this chaos, someone
then created a division between the
Wassmann line and the Birkhahn
line, which is total nonsense because
they were all the same frogs! But the
sLone Iud uIreudy begun Lo roII: LIe
offspring of this truly beautiful morph
were traded and soon found their way
outside the borders of Germany and
into the Netherlands, Belgium and
even the United States. In each new
place all new keepers contributed to
the splintering of these frogs into new
forms and morphs. All of a sudden
we had Panama special, Bronce,
Highland Bronce and several other
descriptors. Some specimens tend to
have an extraordinarily fancy pattern
or color, which were separated out and
line bred with similar looking auratus,
creating even more color morphs. But
every single one of these separate
forms were descendents of the original
Birkhahn frogs. Nowadays it seems im-
possible to determine whether a speci-
men is a true Birkhahn or not, for too
many morphs were imported without
any reliable information about their
geographical origins and too many
were crossbred with Birkhahn-like
frogs. The multi-morph mix is irrevers-
ibly complete.
Another example is the El Cop
morpI. AL hrsL gIunce IL seems us
though many similar looking morphs
tend to populate the central Panamani-
an slopes and mountains that surround
the village of Cop. In actuality, the
national park (Parque General Divi-
sion Omar Torrijos Herrera) where the
hrsL bIg IIgIIund uuruLus oI LIuL regIon
were found has nothing to do with the
village of Cop, which is located just a
few miles from the park. If you explore
the surrounding area around Cop, fol-
lowing the Rio San Juan or Rio Toabre
in the Caribbean direction, what you
wIII hnd ure severuI dIIIerenL morpIs
A frog sold as an El Cop morph without any clear information as to its specifc origins. One
can only confdently know that it was collected somewhere in the general region of El Cop.
A specimen of the Super Blue morph of auratus...a morph whose origins seem to change with
the seasons.
that all look slightly similar but appear
to be their own separate populations
(the Birkhahn auratus are actually
from this region). One morph was
named Microspot by the person
who supposedly found it, but these
frogs are nothing but the descendents
of some metallic green and turquoise
large highland auratus that produced
offspring with smaller spots than their
parents. This is simply another exam-
ple of the irritating process of name-
genesis. The region of Cop, Penenom,
and further north and eastward is not
well explored when it comes to Dend-
robates auratus, but I am sure there
are several yet-to-be discovered popu-
lations here. To the extent that they are
actually distinct populations could be
verIhed by DNA-unuIysIs.u IucL LIuL
holds true for all forms of auratus.
The frogs that have been imported as
Cop could quite possibly be from
every place of that regionfrogs that
were formerly found in the Omar Tor-
rijos Herrera National Park might very
well have nothing in common with
frogs recently imported from this same
region. Specimens of Dendrobates
auratus within the national park seem
rare. According to local people they are
nearly extinct, most likely due to chy-
trid (whether or not this is a fact cannot
be upproved: Iong-Lerm unuIysIs wouId
be necessary to have variables such as
weather phenomena and wet and dry
seusons ure sImpIy InuencIng ucLIvILy
patterns of Dendrobates auratus in the
region).
Other victims of origin confusion
are the Campana and Kuna Yala
morphs. As their names suggest, the
frogs inhabit completely different geo-
graphic areas. However, in some cases
the two tend to have a very similar
physical appearance. Panamanian
species from the San Blas coast and
u Cumpunu (on LIe PucIhc cousL neur
the Canal Zone) have a D. histrionicus-
like dorsal pattern of many small,
light spots (Silverstone 1975). The
Campana morph is found on the
CenLruI PucIhc sIope neur LIe vIIIuge oI
Campana. The two villages of Campana
and Capira are very close to each other
and, as a matter of fact, the frogs dont
seem to care much about man-made
boundaries and spread out along the
Rio Capira and into Campana terri-
tory. Birkhahn wrote something about
another population of greenish aura-
tus close to the Campana morph and
a mixed population of the two found
along the boundaries of each morph.
Photos revealed a morph similar to the
so-cuIIed Cumouuge morpI. And
here we go yet again with self-made
Lerms In u scIenLIhc heId: KuIIuu und
Cream, Brown and Beige, Kahlua
and Cream Camothese are all names
for frogs that seem to have their origin
in the region of Campana and Capira.
But why are they given such obscure
Specimens from the many diverging populations of Dendrobates auratus found inhabiting the Campana region. The frogs in the lower two
images are examples of Camofage auratus.
and unprofessional names? When we
have inquired about the origin of the
Cumouuge uuruLus, LIe onIy unswer
a breeder shared with was something to
LIe eIIecL oI: TIe purenLuI Irogs were
picked on the other side from where the
brown and beige ones are from. When
we asked for more information, they
said they didnt have any more than
LIuL. MussIve numbers oI Cumouuge
auratus were imported and sold in
2006. Where did they come from? No-
body seems to know, and its a shame
that no hobbyist knows what sort of
genetic product he is keeping.
The Kuna Yala morph originates
in the eastern part of Panama, in the
Comarca San Blas near the natural
refuge Nusangandi. These frogs tend to
show a cream-white pattern on a dark
brown background, which may lighten
up as the frog matures. For years there
were frogs offered under the name
Kuna Yala in Germany. No one had
evidence as to where the frogs had
originally come from. Eventually, some
grey imports from the Campana region
came in, all of them looking exactly like
the so-called Kuna Yalas (the Cam-
pana morph is extremely variable and
some individuals show nearly white
patterns, which easily could be mis-
taken for Kuna Yala auratus). It is to
be assumed that pure Kuna Yalas are
extremely rare and most of the frogs
kept today are mixtures of Campana
and Kuna Yala, or specimens of just
Campanas. Apparently, the Kuna
YuIu morpI Is exLremeIy Iurd Lo hnd
Above: Auratus in the Campana region
were often found around boulders and rock
outcroppings like this one.
Below: A Kuna Yala auratus - a morph that is
difcult to fnd both in the wild and captivity.
in its natural habitatseveral people
uLLempLed un expedILIon Lo hnd evI-
dence of this frog, but they all returned
without success. However, a few years
ago a bird watcher accidentally found
some frogs in this region and managed
to take photos.
Another morph that leads to confu-
sion and the crossbreeding of different
forms is the blue and black morph. As
I stated in the beginning, there are sev-
eral blue morphs of auratus in Panama
that can look completely similar but are
in fact from very different populations
and regions. The most famous one, and
one of the most threatened, is a popula-
tion from the Calobre region. Its habi-
tat is found along the shoreline of the
nearby river, where only a few meters
of trees and shrubs are left to line the
riverbanks and function as habitat. The
conversion of the surrounding forest
into farmland has swallowed up all of
the picturesque forests, as well as this
populations biotope.
Marcus Bartelds from Tropical-
Experience.com followed the trail of
another blue auratus morph in 2007
and ended up near the town of Santiago
in central Panama, where he found a
population near the Rio San Pedro.
When we searched the very same spot
u yeur und IuII IuLer we couId noL hnd
a single frog. Was it because of the dry
season? The shoreline had been par-
tially cleared, so there could have been
several reasons.
The next population of blue specimens
is from the Caldera region of Panama.
Along the Rio Caldera, near the towns
of David and Boquete, you there are
several populations of Dendrobates
auratus that are blue in color, but also
Right top and middle: Frogs found along
the Rio Caldera...both specimens are from the
same population.
Right bottom: An auratus from the Calobre
region. Although it looks nearly identical to
the blue and black form of Rio Caldera aura-
tus pictured at the top, the two populations
are found over 100 miles apart.
Below: The banks of the Rio Caldera. The
remaining habitat for this population of frogs
consists of narrow bands of trees along the
shores of the river.
green or turquoise as result of natural
mixing and variation that occurs within
these populations. Now imagine using
only one photo as a reference guide for
these morphs. Its not possible.
Further east, near the town of Santa F,
lives another population of D. auratus
LIuL exempIIhes LIe pIenomenon oI
green and blue specimens within the
same population. This polymorphic
form can be found long the Rio Santa
Maria, at elevations around 100 to 500
m asl. Santa F itself sits within the
shadow of the divide between Atlantic
und PucIhc versunLs, und Is surrounded
by montane rainforests at heights be-
tween 1200 and 1550 m asl. Not quite
at the peak, but at heights around 800
und even up Lo 1,ooo m usI you cun hnd
one of the biggest Panamanian auratus
morphs, the Alto de Piedra morph.
With measurements up to 46 mm svl,
the females are giants by auratus stan-
dards. Evidently this morph is quite
rarewhen searching for it in January
zooq, we couId onIy hnd Lwo specI-
mens even though it was raining cats
and dogs and the conditions seemed
ideal (I cant help but wonder if their
low numbers are due to the cooler tem-
peratures at this altitude which provide
Top: A waterfall near Santa Fe in the
upland forests of the Altos de Piedra.
Above left: One of the largest forms of
auratus, the Altos de Piedra morph. It is
found at distinctly higher elevations than
most other populations of auratus.
Above center: A ventral view of an Alto
de Piedra auratus.
Above Right: A green form of D. auratus
from the Santa Maria population.
better conditions for chytrid to spread).
A very popular and common morph
is the Ancon Hill or Cerro Ancon
morph, alternatively the Canal Zone
morph. Both morphs seem to de-
scend from the same origin and, if you
look at a map of the surroundings of
Panama City, you will see that Ancon
Hill is the southernmost point of a
once connected band of forest that is
today divided in the Chagre National
Park, the Soberania National Park
and the National Interoceanic Park
due to the consequences of the build-
ing of the Panama Canal. This morph
populates the entire Canal Zone, and
popuIur IocuLIons Ior hndIng IL IncIude
Soberania National Park and Barro
Colorado Island. Ancon hill, however,
Is now IsoIuLed und serves us LIe hnuI
refuge for many animals of Panama
City. For me, this place is one of the
mosL IuscInuLIng spoLs In Punumu: sur-
rounded by the bustling activi-
ties of the Canal, the strange and
fascinating slum of El Chorrillo
and, as a contrast, Balboa (the
canal-administrational district
of Panama City), the domestic
airport, the terminal, and one of
Panamas largest malls, you can
hnd dozens oI specIes oI bIrds,
reptiles, small mammals, and
amphibians such as Dendro-
bates auratus living here. Rumors claim
that the hill has not yet been deforested
and developed because it is still under
U.S. ownership and contains a nuclear
fallout shelter from the times of the
Cold War. Regardless, it functions
today as a sort of informal national
monument and area of recreation, as
well as a viewpoint over Panama City.
The Ancon Hill auratus have a black,
dark brown, or brownish background
color and a pattern of often parallel
green, blue-green or greenish-yellowish
dots and spots. Some Individuals show
a dorsal stripe across their back. When
we last visited, we even found individu-
als with blue dots.
There is a high risk in mistaking the
Canal Zone morph and its brown
phenotype with another, similar look-
ing morph found even further west in
the province of Chiriqui. The San Felix
morph of Dendrobates auratus popu-
lates the surrounding forests of the Rio
San Felix and has the same body size as
the Ancon and Canal Zone auratus. The
background color is a light brown with
a gold-green or yellowish-green pattern
that often resembles the one that of the
Below left : An example of auratus found
inhabiting Ancon Hill. These specimens are
representative of frogs inhabiting the Canal
Zone region. Before the contstruction of the
Panama Canal these two populations were
part of a single, contiguous population.
Below right: A frog from the San Felix
population of D. auratus. Although similar
in appearance to the Ancon Hill/Canal Zone
morphs, it is from a completely seperate
population.
photo Thomas Ostrowski
Bottom: Ancon Hill (Cerro Ancon), an unlikely
habitat for Dendrobates auratus in the center
of Panama City.
Ancon Hill or Canal Zone aura-
tus. In fact, San Felix frogs have been
imported to the U.S. and Europe and
were crossbred with Ancon Hill aura-
tus because their owners never knew
that different populations existedthe
information was never provided by the
importers.
As you move west from Panama City
and leave the Canal Zone, you reach a
city called Arraijn. At a nearby river
you cun hnd uuruLus LIuL uIso uppeur
similar to the Canal Zone popula-
tion, but they tend to have more black
background color and their dots seem
to be more green-metallic and tend to
develop a more lined pattern.
Continuing west, near the
city of La Chorrera, you
cun hnd unoLIer morpI
of auratus. The waterfall
here, called El Chorro, was
once a scenic place to visit,
but has in recent years
been transformed into
a straightened and par-
tially cemented river and
gIven LIe uIr oI u sewuge
dump. Wastewater and
cIvIIIzuLIon`s gurbuge ouLs umongsL
submerged vegetation. Human debris
and refuse lines the shores, evidence of
LIe IusL oodwuLer. TIe puLrId smeII oI
waste and feces dominates the air. The
common form of auratus found here
tends to show a truly black background
color with greenish-bluish metallic
dots, commas or even slight reticulated
pattern. Some adults tend to show a to-
tal black colorization without showing
any pattern. Brownish individuals can
also be found within this population
but seem to occur more randomly. This
morph seems to not often be kept in
captivity, and there has been very little
information published on it. Referring
to Silverstone and his work about the
genus Dendrobates from 1975, speci-
mens of the Chorerra population are
being stored in the collection of the
University of Michigan Museum of
Zoology (UMMZ 95478).
A few miles offshore from Panama
CILy, on LIe PucIhc sIde und sILuuLed In
the Bay of Panama, are the islands of
Taboga and Taboguilla. The most com-
mon mistake concerning this morph of
auratus is referring to it as D. auratus
Tobago, which is incorrect because
Tobago is actually a small island in the
Caribbean that hosts no known mem-
ber of the dendrobatid family of frogs.
TIe hrsL ever LuxonomIcuIIy descrIbed
specimen of Dendrobates auratus
seems to have come from the Taboga
popuIuLIon. Ls hrsL descrIpLIon cun
be found in the Abstract of a report to
Lieut. James M. Gillis, U.S.N., upon
the Reptiles collected during the U.S.N.
Astronomical Expedition to Chili by
CIurIes GIrurd In 18q:
Phyllobates Auratus G. Tongue
narrow and elongated, free for
about the half or two-thirds of
its length. Anterior limbs, when
Above: The waterfall El Chorro, near the city of La Chorrera, where a nearly all-
black form of Dendrobates auratus can be found.
Left: An example of the La Chorrera morph of auratus.
stretched backwards, reaching
the vent with the tip of the longest
jner. Injerior surjcce oj thihs
granular. Color, above, metallic
golden; beneath, uniform bluish
broun. Collected on the Islcnd oj
Taboga, in the Bay of Panama, by
the late Prof. C.B. Adams, of Am-
herst College, Mass. (Phyllobates
auratus Girard, 1855 1854, Proc.
Acud. NuL. ScI. PIIIudeIpIIu, ;: zz6).
There are no records that describe the
whereabouts of this holotype, but it
seems to be USNM 10307 (National
Museum of Natural History, Washing-
ton DC). This is the same morph that
was introduced to some of the islands
of Hawaii to help aid in mosquito
control. In 1932, 206 specimens of
D. auratus from Taboga or Taboguilla
Islands, Panam, were released in the
upper Manoa Valley, Oahu, Hawaii;
their descendants form a breeding
population (Oliver & Shaw 1953; Ull-
mann 1976 in Silverstone 1975). The
origin of the creation of the German
trivial name of Dendrobates auratus
is quite interesting, as it is referred to
as Goldbaumsteiger, which, roughly
translated, means golden tree climb-
er. The term golden can be taken as
direct evidence for the appearance of
the gold-green pattern, not to mention
the fact that the Latin term aurum
means gold. It could in fact be incor-
Top: Dendrobates auratus habitat on the island of Taboga.
Above Left: A juvenile Taboga auratus. Is the coloration aposematic or cryptic? There is much room for debate...
Above Right: An example of the reticulated form found within populations of Taboga auratus.
rect to refer to Dendrobates auratus
as the Black and Green Poison Dart
Frog if the Taboga morph was in fact
the described holotype of the species
sInce LIIs descrIpLIon beLLer hLs LIe
morphs found in Costa Rica and some
parts of Panama as well as in Nicaragua
and Colombia. Perhaps a better choice
might have been Gold and Brown
Poison Dart Frog.
The population of Dendrobates aura-
tus on the island of Taboga is one of
the best to observe in the wild as the
frogs exhibit active patterns of behavior
throughout the entire day, even during
the drier periods of the year. As you
leave the main village on the coast of
the island, you will soon stumble across
LIe hrsL IndIvIduuIs dweIIIng umongsL
the leftovers of our civilization. Exer-
cising the motto out of sight, out of
mind, the locals dump their waste into
LIe IoresL, wIere you cun hnd Iundreds
of auratus hopping between rotting
hsIbones, empLy soIL drInk cuns, oId
canisters, clothes, and various other
items best left unmentioned. When last
visiting here, the olfactory peak came
in the form of a half-rotted cat carcass
amidst the still life of trash and buttress
roots, whose foul smell adumbrated
the already dingy scenario. The whole
scene is a shame, but who are we to
judge these people and their ways of
littering when we produce billion of
tons worth ourselves? As you leave the
signs of civilization behind, you soon
reach several small creeks that portray
idyllic scenic biotopes and contain hun-
dreds of Taboga auratus. It is interest-
ing that I have never once witnessed
an adult specimen of this morph that
had a black ground color, as it is often
described as havingonly young and
adolescent frogs seem to have this
color. As they mature, the frogs lighten
up und become brownIsI: some Lend
to show a darker brown, but they never
remain black. The color of the pat-
tern also changes over time and older
specimens show a fantastic metallic
goId-green. Very rureIy do you hnd
specimens with a bluish pattern. Since
this population of frogs is so easy to
identify, it is strange that they are not
more clearly recognized and are instead
labeled in vague terms such as auratus
from Panama or reticulated auratus.
Its true that the Taboga population
tends to show the reticulated pattern
relatively often, but the tendency for
it to have a gold-green pattern on a
brown background remains constant in
all specimens of this morph.
A fairly recent morph of Dendrobates
auratus was introduced to the hobby in
2007 and is found in an area surround-
ing a large lake east of Panama City.
Those who found the population didnt
want to divulge the actual locality, so
the frog was once again labeled with an
urLIhcIuI nume, MebuIo, wIIcI con-
sists of the names of those who found
it. Thomas Ostrowski of Dendrobase.
de replaced this name on his website
with Punta Celeste because of the
sky blue spots displayed on the frog
in the photograph (Ill let you wrestle
over the sense and nonsense of such
cryptic name creations but you should
be concerned about the fact that Mr.
Ostrowski was not allowed to present
LIe exucL hndIng pIuce oI LIe Irog due
Lo u promIse Ie mude Lo LIe hnders oI
the morph). This morph has a dark to
The Mebalo morph, a name that is more indicative of the people who found it than of the
location in which it was discovered.
Diferent Slopes,
Diferent Climates
All populations of Dendrobates
auratus found on the Pacifc slope
have to exist in a completely
diferent climate than their
Caribbean relatives. This diference
in climate is caused by the
Cordillera Central, the mountain
range that extends from east to
west and seperates Panama into
two diferent climate zones. The
majority of the rain clouds that
come in of the Caribbean lose
their cargo as they ascend the
Atlantic slopes, so even in the
drier season on this side you will
have a more humid climate. This
allows the Atlantic/Caribbean
populations of D. auratus a
diferent activity pattern than the
Pacifc populations, who tend to
hide during the dry season. You
can often search for hours and
not fnd a single frog, then turn
one stone or lift one dry palm leaf
and seemingly fnd all the auratus
in the area! This may also be the
explanation behind much of the
shyness in some auratus morphs:
they are simply accustomed to a
somewhat hidden lifestyle.
light brown background color accented
by a pattern of lines and dots or just a
few broken lines that create the ap-
pearance of a dotted pattern. The color
of the pattern is extremely variable,
consisting of anything from blue, green,
greenish-turquoise to several shades of
light blue. Altogether it shares the com-
mon morpIoIogIc scrIpL oI LIe pucIhc
forms of Dendrobates auratus which
are naturally smaller and more con-
hned LIun LIeIr CurIbbeun reIuLIves.
An exception is a morph whose distri-
bution spreads from the area around
the city of Colon, along the Caribbean
coastline, and clear into the surround-
ing forests of the village Portobelo.
TIere Is no oIhcIuI Lerm Ior LIIs
morph-- sometimes it is referred to as
Colon, but in general it is one those
amorphous Greens and Blacks. But
even here we have the phenomenon of
polymorphism, for the frogs can also
show a bluish or turquoise pattern on
a black or very dark brown body. All
of these color forms seem to show the
typical dorsal horseshoe mark on the
snout.
In discussing Green and Black au-
ratus we should understand that this
term is a very uncertain one. Although
it is used to describe the widespread
morph of auratus that is found along
the Caribbean coastline from Nicaragua
to Panama, even these populations dif-
fer genetically and have their own local
morphs. The popularity and abundance
of captive individuals of this form may
have also contributed to the creation of
its common name. In Panama we have
found this morph along the western
Caribbean coast, including the Bocas
del Toro area. It does not inhabit the
islands of the Bocas del Toro archi-
pelago, but is found on the peninsula,
which is divided into several parts such
as Cerro Brujo, Darkland, Gau-
chero, Buena Esperanza and so on.
It is to be expected that this morph is
genetically very close to the populations
found along the Caribbean Coast of
Costa Rica.
The last morph I will discuss in this
article is also one of the most spectacu-
lar, for it is completely distinct from
the typical color and pattern of Den-
drobates auratus. When introduced
into the dart frog hobby in 2006 there
were uLLerunces oI IybrId, urLIhcIuI,
and not to be taken seriouslybut
the call, behavior, and captive breeding
seemed to allude to the fact that we had
to consider it a new morph of Dend-
robates auratus: LIe goIden morpI.
This is a solid yellow or gold-yellow
morph without any major pattern at
all, except for the ventral view, which
offers some rather unspectacular dots
und IInes In u pInk or esI-coIored
tone. Some individuals carry these dots
on LIeIr unks us juvenIIes, buL seem Lo
lose them as they age. It is still a secret
where these frogs are found, but the
fact is that they were imported illegally.
Their discoverers never mentioned a
locality other than the Darien Region,
an area that still holds the reputation
of being a danger zone with guerillas,
drug lords, and militia armed to the
teeth. What better way to establish a
nice business than to create rumors
and sell smuggled frogs of which you
own the exclusive rights, and purport-
edly come from an area in which no
one would ever attempt to visit in order
The yellow or gold morph of Dendrobates auratus. Whether or not this is a selectively bred strain of frog or an actual population in the wild is a
subject of much controversy.
to verify their origins? And all of this, of
course, is done using the term conser-
vation, because what would happen if
the locality of these frogs became public
and everyone went out and collected
some? Regardless of its origins and the
motivation behind its introduction into
the hobby, this morph is being bred
well in captivity and seems to be divid-
Ing IobbyIsLs InLo Lwo groups: LIe ones
that admire it and the ones that meet
the frog with distrust as its appearance
seems way too exceptional.
There are plenty more morphs of
Panamanian Dendrobates auratus to
discuss, but to prevent total confusion
I will stop here it seems like more
than enough to deal simply with these
most popular forms. Also, it needs to
be understood that this article should
not be used as a morph guide or photo
guIde, buL ruLIer LIe exucL opposILe: Lo
show that it is not possible to identify a
certain morph simply by viewing a pho-
tograph or two! Instead of asking how
many different frogs we can keep and
breed, we should focus on quality man-
agement concerning the genetic integri-
ty and geographic locations of the frogs
we keep. As with any revolution, the
hrsL sLep begIns wILI LIe Inner revoIu-
tion. So-called breeders of dart frogs
are often doing nothing but augment-
ing the animals without any sense of
keeping local morphs separated, having
little to no regard for the true origins of
their animals. This responsibility falls
on the consumer choices of the hobby-
ist and their insistence on asking for
the locality of the Dendrobates auratus
specimens they purchase. If people
start to take this information seriously,
there might be a change in practice
of importers and breeders who sell
hundreds of pseudo-morphs, often with
incorrect and arbitrarily chosen names.
This might sound complicated or too
dIIhcuIL Lo cIunge, buL personuIIy IeeI
that this way of keeping and breeding
poison dart frogs (e.g. dealing with spe-
cIhc IocuIe und coIIecLIon InIormuLIon)
is one of the only ways we can begin to
be taken seriously when we speak of
conservation and the hobby.
As the world gets smaller and smaller,
LIe hnuI undIscovered reIugees oI
plants and animals seem to be fewer
and fewer. There is reason to hope that,
by changing the way we view the un-
known, there might be a change in how
we understand it.
_________________________
Works Cited:
Birkhahn, H., Klpmann, V. & Wassmann,
K. (1994): Zur Variabilitt des
Goldbaumsteigers.
Datz Dt. Aquar.Terrar.Zeitschr. 47 (9):
570-576
Cites (2005): Convention on International
Trade of Endangered Species of Wild
Fauna and Flora. Used online version in
march 2009 under http://www.Cites.org.
Dunn, E.R. (1941): Notes on dendrobates
auratus - Copeia, Vol. 1941, No. 2 (Jul. 8,
1941), Published by: American Society of
Ichthyologists and Herpetologists, 88-93.
Girard, C. (1855): Abstract of a report to
Lieut. J.M. Gillis, U.S.N., upon the reptiles
collected during
U.S.N. astronomical expedition to Chili. -
Proc.Acad.Nat.Sci.Phil 7: 226.
Oliver, J. A. & C. E. Shaw (1953). The
amphibians and reptiles of the Hawaiian
Islands. Zoologica (New York) 38(5):65-95.
Silverstone, P. A. (1975): A revision of
the poison-arrow frogs of the genus
Dendrobates Wagler. Sci. Bull. Nat. Hist.
Mus. 21: 1-55.
Ullman, H. (1967): A frog comes to Hawaii.
Natur. Hist. N.Y. 76(5):3637.
The Nominat Mistake
An error that often occurs is the use of the term nominat when it comes
to the description of local morphs of Dendrobates auratus. This is primar-
ily because there is no taxonomic classifcation within the context of
dendrobatids that would necessitate the usage of the term: only species
divided into subspecies are given a nominate form. The form that is
used to create the frst description is the so called nominate form and is
recognized by a scientifc name that includes two secondary names. As
there is no Dendrobates auratus auratus and there are no subspecies, the
term nominat cannot be correctly used in regard to this species or any
of its forms. A more acceptable phrase would be the local morph that
most resembles the holotype. Since the Taboga morph is the holotype
for Dendrobates auratus, it would be this morph we are referring to, not
the Green and Black morph that is often cited simply because of its
abundance in the captive hobby.
BY STEVE WALDRON
Confessions of a
Pumilio Chaser
Confessions of a Pumilio Chaser | 41
The Frog Gardens of
Bocas del Toro
IrsL, LIe conIessIon: my ucquuInLunce wILI Ooophaga
pumilio (and its highly evolved parental care behavior,
extreme color morph variation, and all around doe-eyed
cuteness) became a fatal attractionor at least one that lead
me to strange, obscure places like a Guaymi Indian settle-
ment on a tiny Caribbean island in western Panamas Bocas
del Toro region. Causing me to act out in strange, irratio-
nal behaviors, such as knocking on the door of a complete
strangers home who in addition to having to deal with the
shock of a large American appearing at their front door, was
suddenly forced to puzzle over questions such as
"Ixcose me, bot do yoo know where to nd the
tiny red frogs?
all executed in really badly broken textbook high school
Spanish. Im still amazed at how kindly these people re-
sponded to me (only once did I have to fend off a dog attack).
In many cases they knew that the tiny red frogs really
existed and werent an image from an insane mans halluci-
nation and they would sometimes even offer to lead me right
to where the tiny red frogs lived. Invariably, when I would
take up their offer to go frog hunting, they didnt lead me on
a multi-day hike into the deep, dark jungle, braving hordes
of biting insects, venomous serpents or treacherous tribes-
man (as you might expect from watching too many television
nature documentaries). Rather, these pumilio hunting expe-
ditions were no more than a simple stroll a few paces behind
my Panamanian friends homes and into their gardens.
Because Oophaga pumilio is not a frog of the rainforest.
It is a frog of man.
Of course, 9,000 or so years ago, before the arrival of
humans to Central America, the prehistoric proto-pumilio
was likely found only in the mythological, primeval, virgin
rainforests, perhaps near the forest edge of jungle streams.
They can still be found in the rainforests of such places like
the La Selva Biological Station in north central Costa Rica
where Joel Heinen studied this species and other members
PROLOGUE!
Confessions of a Pumilio Chaser is an ephemeral love song to a
singular kind of frog, Oophaga pumilio, typed out in one fnger-blis-
tering evening in a turn-of-the-20th-century bohemian San Fran-
cisco fat. It is based on a series of pumilio-watching trips I had
taken to Panamas Bocas del Toro Archipelago between 1998 and
2001 and was writen at a time when frogging was still a pursuit
for globetroting frog smugglers, going-for-broke frog collectors
and free-thinking herpetocultural experimenters. It was a golden
age known only to approximately twelve regulars on an Internet
listserve called Frognet.orgeccentric rebels, all of them, and
one-of-a-kind raconteurs. I printed out thirty hard copies of
Confessions... on my workplaces inkjet printer (complete with
color photos), hand-collated, hand-folded and hand-stapled. I
gave away twelve copies at a frogger event at a Holiday Inn in
suburban Seatle and sold none of themthe rest have dis-
solved into the ether and the compost heap. But now, Confes-
sions... has been resurrected, fnding a new home here within
the glossy pages of Leaf Liter Magazine. So cozy up in your
favorite reading chair, put on your favorite Panamanian reg-
gaeton CD, dim the lights, and take a vicarious journey with
me. In the pages of these confessions we will venture to the
land where O. pumilio hang from the rainforest vegetation like
psychedelically colored slime fairies and a bacterial infection
looms around every step of the fip-fop
of the leaf litter herpetofauna community in La Selvas mixed
habitat mosaic of rainforest and former agricultural plots
such as abandoned cacao (i.e. cocoa) groves (Heinen, 1992).
Curiously, Heinen found that not only is O. pumilio one of
the most common leaf litter vertebrates of rainforest and old
cacao stands (contributing to more than half the biomass of
species sampled in the cacao plots), but also found to be over
3X more abundant in cacao sites that had been abandoned
for 20 years than primary rainforest samples. Even more
impressive, cacao stands that had been abandoned for only
5 years held 6X as many pumilio as the rainforest leaf litter
microhabitat! A rainforest frog that seems to not only adapt
to human disturbances of the original vegetation but also
LIrIves us IL never Ius beIore? How couId LIIs be? To hnd
the answer, we must return to my human acquaintances of
Caribbean lower Central America and understand a bit about
their dinner menus.
CenLruI AmerIcu Ius gone LIrougI muny sIIILs In oruI
and faunal changes in its relatively short appearance on
the geological scene. Indeed, as a land bridge between the
North and South American continents, the Central American
isthmus, the interchange of biotas must seem like a busy
two-lane highway against the geologic time frame. Giant
evolutionary oddballs like ground sloths, anteaters, armadil-
los, herbivorous rhino-like toxodonts, predacious 4-meter
tall carnivorous Phorusrhachid birds (a.k.a. Terror Birds)
migrated north while cats, deer, mice and mastodonts
migrated south. Some theories suggest that the arrival of
humans from Eurasia and their hunting practices may have
caused the disappearance of many large grazing animals and
their associated predators from the Americas including all
of Central Americas gomphotheres (ancient mastodont-like
creatures), glyptodonts (giant armadillos), and megalony-
chids (giant ground sloths). This may be so, though Im not
entirely convinced of this theory. I am convinced, however,
that when in a foreign land and cultural differences make
hndIng conversuLIon LopIcs oI common InLeresL dIIhcuIL, one
should talk about food and much will be revealed
After a successful afternoon of pumilio chasing, my hosts
and I would sit around drinking jugo de pina (pineapple
juice), cooling off and trying to muddle through a conversa-
tion in my broken Spanish.
Have you ever eaten a jaguar?
INDIAN: No, I havent. But last month some Guaymi on
Isla Popa killed one that had swam over from the mainland
and ate it. The hide and teeth were hung up at the petrol
shop for everyone to see.
Pumilio Chaser: Wowthats terrible! El Tigre is a rare
animal and not many still exist.
Indian: Yesbut meat is hard to come by.
Pumilio Chaser: What else do people hunt for meat?
Indian: Oh, peccary, tapir, monkey, conejo
Pumilio Chaser: Conejo pinto (paca) or conejo (rabbit)?
Indian: Both...and armadillo, squir-
rel, bushrat, all types of birds, boa
Pumilio Chaser: Boas?
Indian: Yes, they are very tasty,
sweet meat, but many bones
It was starting to sound like a detailed
rundown of the evolutionary tree or
u specIes IIsL Irom u heId guIde Lo LIe
region
Indian: And turtle (a.k.a. Green Sea
Turtle, a.k.a. Bocas Beef), all types
oI hsI, sLreum sIrImp, snuIIs, Iund
crab, octopus, lizards
Pumilio Chaser: What kinds of
lizards?
Indian: U-grra (Helmeted
Iguana, Corytophanes cristatus) and jilida (Anolis
spp.) ure LIe eusIesL Lo hnd.
Pumilio Chaser: Do people eat frogs?
Indian: Oh yes, ogungo (medium-sized Eleutherodacty-
lus) and no-lu (tiny, thumbnail-sized Eleutherodactlyus).
Pumilio Chaser: And rcnc juroscente (Dendrobates
auratus) and ranitas rojas (Oophaga pumilio)?
Indian:
No, I dont think anyone around here eats
little red frogs. Do you eat them?
OI course, IL cun be Iurd Lo hnd u ruLIonuI expIunuLIon Ior
why one has traveled 2,000 miles to chase tiny red frogs
especially to those whose daily chasing after animal protein
Ior LIe dInner LubIe oILen goes unsuLIshed.
TIe ruruI peopIe oI WesLern Punumu hnd muny oI LIe ed-
ible small wild animals in the forest at the edge of their
current gardens or in feral regrowth plots of gardens past
(rusLrojos). n LIese gurdens und rusLrojos we hnd u key Lo
the riddle of O. pumilios enhanced abundance outside of the
rainforestNeotropical gardens not only sustain people but
also nurture poison frogs as well.
In the Guaymi and Bocatoreno garden patches of the aroids
Oto and Krunchi (otherwise known as Taro or Xanthosoma),
Oophaga pumilio hnds u nursery In LIe wuLer-hIIed uxIIs Ior
rearing its tadpoles. In banana, plantain and heliconia thick-
ets more tadpole deposition sites are found in those plants
axils. The strap-leafed terrestrial bromeliad, Aechmea
magdalenae, provides a spiny fortress for poison frogs to
hide and raise their offspring.
Frog chasers must be mindful of these
skin-shredding leaf edges when cornering their quarry. The
GuuymI uIso weuve LIe sLrong hbers Irom LIIs wIId pIneuppIe
into beautiful, durable carrying bags that I have seen carry
everything from babies to chainsaws. Discarded coco-
nut husks provide shelter and platforms for the territorial
vocalizations of wild poison frogs just as they do for captive
frogs. The abandoned canopy-shaded cacao groves planted
for export to satisfy the global demand for chocolatenow
lying fallow due to a price drop on the international market
and the ravages of an invasive fungusprovide thick leaf lit-
ter and an associated leaf litter invertebrate abundance that
feeds more poison frogs than does the rainforest. Threats
to the rainforest and the assaults on its beautiful and mys-
terious biodiversity by human overpopulation, industrial
resource extraction, and monoculture agricultural practices
are as intense in Bocas del Toro as anywhere in the trop-
ics. But it gives me great satisfaction to know that as long
as the rural people of Panama, Costa Rica, and Nicaragua
continue to garden in their traditional fashion of untidy plots
with wild and diverse indigenous crops, there will continue
to be a place for Oophaga pumilioand in abundance! So
abundant, in fact, that all you have to do is relax with some
friends, a cup of jugo de pina, and chase little red frogs until
the rains start.
Journal Excerpts from a
Pumilio Chasing Trip to
Bocas del Toro, Panama
2001
June 6, 2001
Today, the jet lag wore off and the naturalist awoke. Santi
took us to the easternmost (mas lejos) of the two Cayos
Zapatillas. A small island, I could walk its perimeter in an
hourand did. The forest and its inhabitants were refresh-
ingly unclutteredeasy traveling underneath coco palms
overgrown with giant Aechmea bromeliads (the middle-sized
plants were red spotted). No frogs. Perhaps they are there.
I understand they are green pumilio.
Perhaps they ARE there, up in the those green
pools of sky water waiting for the rains.
7 de Junio, 2001
Today I followed the black hawk again. Down Punta Vieja,
3 km or so on the Atlantic side, along the beach and palm
IoresL und Iound wIuL wus IookIng Ior: poIson Irogs, runILus
rojas! This race of Oophaga pumilio was uniquesmall,
ubouL LIe sIze oI LIe Cuyo Nuncy Iorm: red body, wIILe un-
derneath and brown legs w/ spots. Some were heavily black-
dotted and the dorsumfantastic! Pura rana vida!
They were fairly abundant in someones rastrojo and no-
where else to be found. Lots of the Guaymi bag-bromeliad
for nurseries. Before dinner we watched the incredible
display of phosphorescent worms in the lagoon. No stars in
the sky, but celestial illumination coming from the glowing
worm worIds oI EI Mur CurIbe. TIIs udjecLIve overused:
fantastic!
6/8/01
These days pass as in a dream. I wake throughout the night
to the thunder and lightning from the mainland and the
whine of swamp mosquito born from behind the beach. By
day I land on a beach of a new islandgood suerte foretold
by the omen of a Black Hawks* scolding. Hes angry I will
steal his big blue-clawed land crabs. He keeps watch over
me
*Eter time I'te come ccross c pumilio populction on the islcnds,
I'te hecrd the ccll oj the Common lccl Hcul (Buteogallus an-
thracinus) jrst. I'm not sure uh cnd uill hczcrd no uess. When
you hear the Black Hawk, stop and listen to what it is saying.
Isla Solarte, June 9, 2001
Toduy, uL hrsL, dId noL see LIe GoIden PumIIIo. Ieurd
their cohorts, the bird-voiced Colostethus of the hill and
the twittering Phyllobates hidden in the swamp. With time
I tuned into the SNAP! of dancing male Gold-collared
Manakin in the sapling thickets. And lured their females
away with my pish-ing alarm call imitations and their
nervous anticipation of some robber at their nest. The male
manakins have no such worries and continue the dance.
A lone Oropendola gurgled high in a treewhy is he here
alone? Why am I?
Igs IeII IIke munnu. ungus und IruIL Ies LIougIL so uL
IeusL. TIose sLrunge yeIIow owers IILLered LIe ground und
my gIunL hg Lree sLIII IeId LIe munukIn nesL Irom IusL yeur.
A Rufous-tailed Hummingbird buzzed my blue shirt. Giant
ponerine ants with fuzzy creamy white abdomens made their
way steadily and curiously, only traveling along rotted palm
petioles. Round, purely red tortoise beetles. A palm boot
hIIed wILI wuLer und dendrobuLId LudpoIes. And LIen. suw
LIe hrsL puIr oI GoIden pumIIIo.
And then Amen! I heard the Golden
pumilio chorus:
EP! - EP! EP!! EP! ! EP!
- u sLeudy, rupId Irequency hIIed LIe uIr. uL IemuIes re-
sponded to the call, thin ones ate while the males performed
their hurky jerky love dance. I was there to see this, today, el
nueve de Junio, 2001.
12 de Junio, 2001
I write this after four days of not writing. We have been
here, at 500 meters or so at Wilburto Martinezs Willie Mazu
RuncIo EcoIogIcuI. My hrsL uILernoonJnIgIL excILemenL:
meeting Ornel and his family, a naturalist of the highest
caliber, a peon (as he refers to himself) and a Chiriqueno.
Viewed his extensive local orchid collection (most with seed
pods) und IeurIng LIe possIbIIILIes Ior hndIng Irogs. WIen
returning from his home (hard rain, some hard coffee and
some suave chicha), I found a coral snake eating a caecilian
on the road. When the luz was lit in the rancho, 1000s and
1000s of insectos came out of the surrounding forest and a
large bat and a green leaf mantid and the biggest Bufo mari-
nus Ive ever seen came to feast on them. El sapo sabe.
The Toad knows. Hilma, the cook, would say later. But
hrsL sIe wouId cook me up u deIIcIous deep-IrIed hsI dInner
that would nearly kill me with food poisoning and worse,
keep me from one day and two nights of the best frogging
conditions during my mainland trip.
wretching while frogs were croaking,
bummed out and trembling in my tent.
TIe nexL duy we Iound z Lypes oI muInIund pumIIIo: LIe hrsL
in an old cacao grove/lowland forest remnant that will be
u rIce pIunLuLIon nexL yeur. TIe second: u gIunL oId growLI
tree growing along the highway near the Super de Chino
gus sLuLIon und mInI-murkeL. TIe Iormer, LIe cumouuged
quiet metallic variable-patterned pumilio. The latter, the
beautiful green tree pumilio. It rained hard and we had no
more time to frog. Ornel knew of a place where there were
pure yellow pumiliocomo oro. Like gold. We had to get
back to the rancho. We met a Peace Corps worker with an
unknown infection and got on the bus. A temporale wind
raged up the highway for like 20 minutes. The road was
littered with fallen trees, some snapping at right angles. A
cur wus Ipped. A vIIIuge`s uqueducL und wuLer pIpeIIne wus
broken by a tree. Banana plants tossed akimbo from the
gardens. A strong wind had come up from the ocean, built
strength on its way up to the hills and thrashed the roadside
communities, in only twenty minutes totally trashed the
roadside and all it supported. Desperation.
A Short List of Some
Known Oophaga
pumilio Color Morphs
in the Bocas del Toro
Archipeligo, Panama
Isla Bastimentos
Variable. There are at least two population centers on the
island. The population on the north end of the island (and
most familiar to frog enthusiast) are typically red with black
polka dots, lines and swirls on the body. Legs are often
white with brownish spots. These are bold and poisonous
frogs. There is also a form that is considerably more scarce
intermixed with this population, yellowish white replacing
the red. Variations between these two extremes of the Basti-
mentos pumilio spectrum are seen. No two frogs are alike.
On the southeast of the island an unusual small red form is
found. Red body with grayish-green legs occasionally spot-
ted in black. They are quite shy and not as variable as the
northno whitish yellow frogs were seen in this population
after a thorough survey.
Isla Solarte / Cayo Nancy
A tiny, pure tangerine orange frog,
often with white hands and feet. My
favorite morph of the islands. En-
countering these frogs in the wild is
a pure delightafter a morning rain,
hire a boatman from Bocas and have
him drop you off at Nancy Cay. As you
enLer LIe IoresL you`II hnd LIese LIny
glowing golden froglets hopping every-
where, almost like little forest fairies.
Look for them in the palm swamp and
the old United Fruit Companys grave-
yard (mostly malaria victimsbring
mosquito repellant).
Isla Colon
I have not explored this island nearly as
much as Id like to. Its the most heav-
ily populated island in the archipelago.
I found a small population of beautiful
green frogs with black-spotted bodes, yellow legs with brown
splotches. They were extremely wary and not at all easy to
chase. Mostly found in the trees and in large heliconia thick-
ets. An almost truly arboreal pumilio!
Loma Partida
If you dare risk the journey to Witch Hill (Cerro Brujo) you
might encounter the Blue Pumilio! They are sort of shy frogs
and blend in well with their surroundings. However, once in
hand, they are beautifulcolored like a moonlit night.
And what about the morphs found at Isla Pastores (Shep-
herd Island) rumored to have a snakeskin green pattern? Or
the remote island of Escudo de Veraguas where tiny pumilio
are found in red, white, and blue? This island also hosts an
incredibly dense population of an endemic salamander (Boli-
toglossa sp.). And Isla Popa? And Isla San Cristobal? And
the Mainland? And
WIerever you hnd u popuIuLIon oI pumIIIo IL wIII Iuve u
unIque cIurucLer uII ILs own. NoL jusL In LIe coIor or Irogs:
behavior, microhabitat preferences, and even subtleties in
vocalizations add to the rich diversity of Oophaga pumilio
in western Panama. Add to that the variations seen between
IndIvIduuI Irogs creuLes un uImosL InhnILe vurIeLy oI pumIIIo
to meetthere are truly no two pumilio alike. And the ad-
ventures youll have along the wayI encourage you to visit
these places and explore!
Midnight Interludes
One of the joys of frogging the Neotropics is that you can
chase frogs all day and all nightpumilio and other poison
frogs during the day and treefrogs and other strange creep-
ers during the nocturnal forays into the forest. Here are
a few journal entries written after night hiking through the
ruInIoresL:
4/28/95 Rara Avis 12:00 a.m.
RuIn, wuIkIng sLIcks, LIe uLLer oI buL wIngs uguInsL my
headlamp, foliage platforms alive with bizarre katydids, tink
Irogs, beeLIes und cockroucIes. TIe hsI sIeep, IuzIIy cruIsIng
the rivulets the iridescent shimmer of a male Rivulus kil-
IIhsI uguInsL my IeudIump. A spIder IIes In wuIL uL LIe edge
of a bromeliad leaf. Flash of thunder cracks, bringing in the
ruIn. TIe EIeuLIeroducLyIus songs, meLuIIIc und succIncL hII
the forest with the presence of life. Here, now, as always
the rain, the myriad of the living and me.
8 de Mayo, 2003
Rara Avis, Costa Rica, Rio Atelopus watershed
Lao by La Catarata
Here we are.
Tonight in this
Carib-bean
Rain-forest.
Along this Rio.
By this Water-fall
I dont have to ask
To know
what unites us
palm, fern, 10,000 types of leaf.
Brilliant-thighed forest frog on the trail.
Fan antennae beetle of the tree.
BuLs IoverIng, uLLerIng
My headlamp
illuminates an amber pool
where sleeping tadpoles swirl
and Rivulus hsI gIIde.
Raindrops drop droplets
on the cat-eyed snakes hunting eye.
Bocas del Toro
Agua de pipa para tomar
Hermosas play para caminar
Agua sal para banar
Bocas del Toro
Coconut juice for drinking
Beautiful beaches for walking
Saltwater for bathing
- lyrics from a Bocatoreno folk song
The Color of Love and
the Polka-Dot Poison
Frog, Oophaga pumilio,
Isla Bastimentos,
Panama
When someone asks why I would eschew the comforts of
home and modern convenience for the muck and biting
insects of the tropical forestwell, no one ever doesbut if
someone were to, I would have to say,
Where there are poison frogs,
there is Nirvana.
TIIs LrIp wus LurnIng ouL Lo be no excepLIon. My hrsL vIew oI
Isla Bastimentos and the rest of the Bocas del Toro archi-
pelago was an auspicious one. For hours, I had been hiking
along the spine of Panamas cloudforest continental divide;
LIe monLune IoresLs oI LIe ALIunLIc und PucIhc versunLs
spilling off on either side of the narrow trail. An enchanted
place, the cloud forest was alive with multi-hued colors of
mounLuIn Lunuger ocks, cIcudu sounds, pIercIng crIes oI
wIo knows? und LIe occusIonuI buLLery skIppIng LIrougI
the diffused light along the trail. Green epiphytes dripped
from tree branch to tree branch. The fog masses coming, go-
Ing und owIng. TIe cunopy IImbs oI gnurIed Lrees encrusLed
with a living blanket of moss, studded with seedlings of
unknown potential- orchids, bromeliads and countless other
epiphytes hung on every perch. I slipped, tripped and fell a
lot it, can be hazardous to hike and ologize in such rich
and slippery terrain.
I stumbled on and into one particularly memorable patch
of montane forest that was alive with the sounds of diurnal
frogsbut the familiar sounds of something like Oophaga
pumilio were coming from the trees. Craning my binocu-
Iurs upwurds und peerIng InLo bromeIIud puLcIes, hnuIIy
realized I was listening to the calls of an amazingly adapted
canopy poison frog, Oophaga arborea, and promptly fell
down a rather steep hill.
WIen hnuIIy sIId Iome, Iooked up und ucross u mud-
slide devoid of vegetation. The landslide revealed a window
through the dense foliage and an uninterrupted view of the
lowland coastal jungle, the Caribbean, and the Bocas del
Toro islands. The mountains were beautiful, but the nights
there were cold and rain incessant and it was time to hitch
hike down to las islas. The verdant cluster of rainforest is-
lands promised sunshine, spiny lobster, cold cerveza and, in
this unique case, polymorphic popula-
tions of Oophaga pumilio.
Ranging along the Caribbean
coast from northern Nicaragua
to western Panama, O. pumilio
is one of the more fascinating
frogs of the Central Ameri-
can isthmus. Hell, its got my
vote for the coolest frog in the
world!
In the northern part of its range, Ni-
caraguan and Costa Rican morphs of
O. pumilio are adorned in red-colored
bodies with blue legs. Traveling south
down the Atlantic lowlands of Costa
Rica, the color morphs of O. pumilio
grade from the blue jeans color pat-
tern of Nicaragua to frogs with less
blue in the leg (sometimes the blue
being replaced by black), increased
black spotting on the dorsum, and
neur LIe PunumuJCosLu RIcu border:
populations comprised of uniformly
red frogs. Pretty cool. However, it gets
even cooler, as we shall see. Something
extraordinary happened to the pumilio
color morphs of western Panama and
the stage was set for this marvel of
evolution in the archipelago of Bocas
del Toro.
The Bocas del Toro archipelago is
comprised of nine major islands, 59
smaller cays and countless mangrove
islets. Set in the Caribbean Sea, fringed
in coral reefs and covered in dense rain
forest, these islands are truly one of the
planets great paradisesand no greater
paradise exists for the pumilio chaser.
Most islands are home to some of
these frogs and nearly each one has a
uniquely colored population. Colors
runge LIe enLIre coIor specLrum: vIoIeL,
blue, green, orange, yellow, red, white
and patterns of spots, stripes, and color
combinations further add to this anuran
kaleidoscope. This remarkable case
of polymorphic variation (i.e. extreme
color differences between neighboring
populations) has long been admired by
scientists and only recently has been
studied in depth. Kyle Summers and
associates (Summers et al. 1997) have
put forth a hypothesis based on genetic
evidence and logic derived from the
theory of natural selection that the bewildering array of color
morphs of O. pumilio from the Bocas del Toro archipelago is
u producL oI pumIIIo Iove-LIe hnIcky LusLes oI cIoosy IemuIe
frogs. But we shall return to Dr. Summers and his evolution-
ary tale of frog romance later
The rains followed me to Isla Bastimentos. After two long
days of consistently inclement weather and wasted time
spent indoors, I was anxious to get into the forest and to
explore some of the other islands. I passed the time talking
with local boatmen, usually ending up in strange conversa-
tions, most much too strange to report here.
Boatman: What frog you seek, mon?
Pumilio Chaser: I seek the blue.
Boatman: I think I know this place. Where the blue frog
lives.
Pumilio Chaser: Great, man! When can we go?
Boatman: It is raining, mon.
Pumilio Chaser: UhI know, I have a rain jacket. You
can borrow it?
Boatman: No mon. The sea.
Pumilio Chaser: The sea?
Boatman: The sea, in a storm. It will take your life, mon.
It will take your life.
Even the seemingly idyllic Caribbean is no place for a leaky
dugout canoe, an overfed gringo and a 15 horsepower out-
board motor with a tendency to fail in the middle of a tropi-
cal storm
WIen LIe ruIns hnuIIy IIILed, so dId my spIrILs us Ieuded
ouL oI LIe vIIIuge Lo hnd LIe runILus rojus. Soon uILer IoI-
lowing the trail out of town, a grove of coconut palms was
reucIed und LIe hrsL Ioud cIIrps oI u muILILude oI muIe
poison frogs could be heard. It wasnt long before I realized
that love was in the air and, nearly everywhere I looked, a
red-bodied, white-bellied, black polka-dot-back Oophaga
pumilio could be found in the act of the reproductive pro-
cess. A veritable anuran orgyin broad daylight no less!
There were males with bulging vocal sacks calling out to
females from every coconut husk platform, females hopping
around and considering their mates, males wrestling and
grappling for territory.
It seems the days of steady rain and the sudden break in
weather had created the perfect conditions for courtship,
egg-laying, tadpole-carrying and tadpole-feeding. Whats
this tadpole feeding you ask? Well, perhaps you already
know but I will tell you anyway. Along with a few other
poison frogs found in lower Central America and the Choco
of Columbia and Ecuador, Oophaga pumilio is an egg feeder
and its tadpoles are obligate oophagous ovivores. Yeah, you
reud LIuL correcLIy: obIIguLe oopIugous ovIvores. mugIne
Sir David Attenborough and a BBC documentary production
relating the wonder in the following facts of O. pumilios life
IIsLory:
Besides performing the range of parental care tasks found
in other poison frogs (i.e. egg tending, tadpole carrying),
female O. pumilio somehow remember the spot of tadpole
deposILIon (e.g. LIe wuLer hIIed cIsLern oI u bromeIIud) und
amazingly, repeatedly return to lay unfertilized eggs for
their hungry offspring! A remarkable aspect of their natural
history and an incredible act of parental devotion. Which
reminds me of the good Dr. Summers
As previously mentioned, there are a lot of colourful pumilio
in Bocas del Toro, and Summers et al. set out to discover
Why? Well, it turns out that though wildly variable at the
morphological level, the Bocas del Toro populations of O.
pumilio show little detectable genetic variation amongst one
another. In fact, a similar level of genetic divergence (or lack
thereof) is found in island-bound Bocas del Toro populations
of other poison frog species, such as Phyllobates lugubris
and island Minyobates. Despite sharing the same evolution-
ary history and patches of forest as O. pumilio during their
6,000 year (?) stay on the islands, these other poison frogs
show little or no inter-population morphological differences.
A Phyllobates lugubris on Isla Bastimentos looks pretty
much like a Phyllobates lugubris on Cayo Solarte. Why
would natural selection act to produce a unique color form of
Oophaga pumilio on every island while keeping each Phyl-
lobates and Minyobates population relatively uniform? The
aforementioned differ from O. pumilio in that parental care
for these species is a much less involved process and ends
with the males deposition of the tadpoles to their rearing
sitesnot much of an energy expenditure on either parents
part. However, all that extra-unfertilized egg laying and
tadpole tending is quite energetically expensive for a female
Oophaga pumilio. Relative to her, the male is a negligent fa-
LIer: occusIonuI sLoppIng by Lo wuLer LIe eggs und, once LIe
tadpoles hatch out, he splits and is out looking for another
mate. Life is short, there are no eggs to be wastedshe must
choose her mate and his genetic contribution to her progeny
carefully.
Perhaps female O. pumilio are selecting their mates for
something subtle that may hint at good genes in those male
pumilio proteins that will convey health and good fortune to
her offspringthe intensity of his vocalizations, the stamina
of his back stroking during courtship process, or perhaps
something unique in his color and pattern. Whatever it is,
only the female O. pumilio knows for sure, but such intense
female scrutiny can provide a strong selection force. With
evolutionary time, sexual selection can produce elaborate
ornaments and bizarre male behavior such as the incredible
feathers and courtship dances of the New Guinea birds of
paradise, the antlers of stags, the wild colors of many male
cIcIIId hsIes. PerIups sexuuI seIecLIon couId even produce u
red frog with a white belly and a black polka-dot back?
I LIIs sLory seems unhnIsIed.IIke uII LIIngs on LIIs pIuneL,
its a work in progress. Evolution is an endlessly interesting
phenomenon sometimes producing results that can be clean-
ly laid out and rationalized with logical deductions based on
natural selection theory. But at other times it is perhaps best
Lo IIe down on LIe ruInIoresL oor, Irog`s eye IeveI, und suy Lo
yourseII:
Damn. Those are pretty frogs.
__________________
REFERENCES
Heinen, Joel T. (1992): Comparisons of the leaf litter herpeto-
fauna in abandoned cacao plantations and primary rain forest in
Costa Rica: some implications for faunal restoration. Biotropica
24:431-439.
Summers, K., Bermingham, E., Weigt., McCaferty, S., and Dahl-
strom, L., 1997. Phenotypic and Genetic Divergence in Three
Species of Dart-Poison Frogs with Contrasting Parental Behavior.
Journal of Heredity 88: 8-13.
Bocas del Toro Archipeligo | 53
/&+"0 1*2 (&-&
"-+)3!*234&
$"! &' ()*
BY JUSTIN YEAGER
UPON FINALIZING PLANS to spend part of the summer
in Panama, I began to brainstorm possible places I wanted to
explore while down there. On previous trips, years before, I
had explored a bit of the Bocas del Toro archipelago during
semana santa (the spring break for college students not from
Latin America) looking for Oophaga pumilio populations
and exploring the cloud forest of Boquete. More recently,
Erik Lindquist had generously invited me to his Atelopus
zeLekI sLudy sILe. WIIIe LIere, u BBC hIm crew vIsILed Lo hIm
un epIsode oI IIe In CoId BIood, IuIhIIIng u Iong-LIme dreum
of mine to see A. zeteki before they were extinct. For this
LrIp, Iowever, summer hnuncIuI und LIme resLrIcLIons IImILed
the scope of areas I could potentially visit. Regardless, I
really had my heart set on doing one special trip while down
there.
In doing a bit of reading online I realized there was an island
of extraordinary beauty and interesting natural history.
Escudo de Veraguas is known for being an island removed
from the rest of the Bocas del Toro archipelago, located
nearly 50 miles east of Bocas Town (on Isla Colon) and 20
miles off the mainland coast. Because of this, it has received
part of its fame for its unique endemics, including a species
of pygmy sloth, hummingbird and salamander. Adding to
this allure was a form of O. pumilio reported to be one the
most beautiful of the populations. I admittedly knew little
about this population of frogs before checking some online
sources, but upon seeing some photographs, I thought it
would be a fun trip. Luckily, several friends had previously
been there and I quickly learned the trip was very doable,
although expensive. Within a few weeks of proclaiming my
desire to visit Escudo, I heard that a fellow graduate student,
JP Lawrence of Michigan State, had planned on going there
and had room on his boat.
Upon arriving at the Smithsonian Tropical Research
Institute (STRI), where we would all be working for
the next weeks, we had a small meeting of poison frog
The Oophaga pumilio of Escudo de
Veraguas
The Oophaga pumilio of Escudo de Veraguas | 55
researchers over dinner (to call this meal dinner is quite an
undersLuLemenL: LIe smuII resLuurunL, cuIIed GuurI GuurI,
is owned by a young German/Spanish couple who served
us a decadent six-course meal complete with table service).
We planned the trip to take place just a few days from that
nIgIL, IeuvIng STR uL q:o AM. By uII uccounLs LIe LrIp Lo
the island can be rough even during the best of conditions, so
leaving early and having good weather were imperative. The
night before we left I made sure to charge my batteries, pack
my food, and get various things ready for our early morning
start. I was expecting our boat ride to the island to be similar
to the river traveling I had done in Peru, so I planned to wear
my rain jacket, pull the hood over my eyes, turn on my iPod,
and sleep for the 2.5-hour trip.
The next morning we awoke to a pitch-black room, quickly
gathered our things and walked to STRI. The morning air
was warm and the sky appeared clear. We boarded the boat
and arranged our gear. Veterans of Escudo had graciously
mentioned that we would want several extra life jackets to
pad our seats as the ride would likely be a rough one. As
we settled in I situated myself between two seats, acquiring
what I thought was an ideal position for taking a nap.
Within minutes we were on our way, gently bouncing with
the waves.
As we entered open water,
passing between the Aguacate
Peninsula and the islands, the
gentle bouncing of the boat got
harder and harder, but I was
intent on sleeping. Before long
it was evident that my concern,
previously focused on taking a
nap, would best be shifted to
stabilizing my body. The waves got
progressively larger and, for the
hrsL LIme In yeurs, Ioped LIuL my
childhood motion sickness would
not return.
If only I was so lucky.
The boat continued hitting the
swells, crashing down brutally after
each one. At times the force was
so hard I worried about breaking
a tooth as my upper and lower jaws
crunched together. It was about this
time that the weather turned sour.
Though still dark, I noticed there was
a mix of both of fresh and salt water hitting my facemy lips
relished the fresh taste of the rain.
Despite my best efforts to keep down my pineapple-and-
peur-juIce breukIusL, wus unsuccessIuI. DurIng my hrsL
bouL oI IeedIng LIe hsI usked LIe bouL drIver Lo sIow Ior
a moment so I wouldnt get hit by the waves. He cheerfully
obliged, if for no other reason than to chuckle at me. In
my head I couldnt help but start singing the theme song to
Gilligans Island as the sun began to rise, revealing the size
of the waves. During ensuing bouts of seasickness (seven to
be exact), waves hit me in the face as I my stomach emptied
itself, the force of which felt like being punched. At one
point I looked at my watch to discover we were only halfway
through the predicted two and a half hour trip. My optimism
sank.
Even the most miserable experiences eventually have a
light at the end of the tunnel. Mine came when the weather
cleared and we could make out the faint outline of an
island. And for a moment, just as we began to circle the
island, the rain stopped completely and the sun burst out
from behind the clouds, illuminating what appeared to be a
mystical paradise before us. A few moments later the rains
reLurned.buL my spIrILs couId noL Iuve been IIgIer: we Iud
hnuIIy mude IL Lo Escudo do Veruguus.
Once on the island we quickly gathered our things and took
sIeILer under u smuII hsIIng sIuck. TIere were u Iew IocuI
hsIermen us weII us u IumIIy wILI uII oI us, cuddIIng under
the shelter and trying to stay out of the rain. It continued to
pour for the next hour, eventually slowing to a drizzle. By
that point I couldnt contain my excitement to go look for
frogs. I ventured down the beach looking for any trails that
could take me into the forest. I entered through a grassy
area and was soon enclosed by trees. The muddy path I was
following quickly became deeper until I was wading through
almost three feet of water. Around twenty feet in I began to
hear what I swore was an O. pumilio call, only quicker and
higher pitched than the call I was accustomed to hearing
As with many populations of Oophaga pumilio, the population on Escudo de Veraguas demon-
strates high levels of variability.
The Oophaga pumilio of Escudo de Veraguas | 57
from the species. I continued down
LIe LruII, oILen munugIng Lo hnd LIe
deepest mud holes, until I heard calling
again. I was convinced that I was close
enougI Lo LIe source Lo hnd LIem und,
sure enough, on the back side of a large
tree covered in epiphytes, I found a pair
within a foot of one another, the male
calling intently to the female. They
were slightly smaller than I expected,
roughly the size of a Ranitomeya
reticulata, but just as beautiful. As
is often the case when hunting for
unImuIs, once we Iound LIe hrsL
individuals, more were soon to follow.
It soon became apparent that there was
a lot of variability in the coloration of
this population, just like many other O.
pumilio localities. The predominant
phenotype had a red splash that started
on the head and ended in a ring on the
lower back, fading into light blue. Other
IndIvIduuIs Iooked quILe dIIIerenL: some
were solid blue, others looked to even have a bit of green or
brown on them. The frogs were quite abundant, though I
attribute it partially to there being such a heavy rain prior to
us hunting for them. We ended up combing the majority of
our side of the island. Numerous individuals were spotted
on the hillsides amongst grasses, on trees, and around plants
such as banana or Heliconia. In our search we also turned
up several Phyllobates lugubris hopping through the leaf
litter.
Phyllobates lugubris
We continued to wind our way through the forest and, before
long, ended up in a marshy area full of crabs and anoles
jumping from branch to branch. I looked up from my frog
searching and saw a cove ahead with waves coming inwe
had unknowingly walked much of the island. It was then
that I was reminded of just how small this island actually is.
After sitting and soaking in the beautiful scenery, and resting
from the hiking and earlier bouts of seasickness, we turned
around and headed back to the shack.
As JP photographed frogs for his morphometrics project
we uII Look Lurns pIoLogrupIIng LIe Irogs Ie Iud hnIsIed
wILI. TIe numbers Ie requIred Ior IIs sLudy benehLed us
all as we were able to get a better idea of the variability of
the population as a whole, rather than just catching a few to
photograph. As we snapped away our boat driver reminded
us the weather was likely to worsen again and we should
think about wrapping up our photography. Until that point I
had repressed the thought of the return boat ride, preferring
to revel in the temporary bliss of being on a beautiful isolated
island.
As luck would have it, the return trip was much more
merciful on my stomach, though the waves continued to
be a source of considerable discomfort. Leaving Escudo
de Veraguas, we headed across open water for some time
beIore hnuIIy spoLLIng LIe muInIund uguIn. DurIng LIIs LIme
IL wus exLremeIy dIIhcuIL Lo Iocus on unyLIIng LIuL wouId
quell feelings of seasickness. The swells of the waves would
lift our boat up only to come crashing down yet again. One
could not help from groaning or grunting every few crashes,
though there was nothing the sea could throw at us that
could take away our experience of Escudo de Veraguas.
59
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ur. kevln WrlghL and Lhe Arlzona Lxouc Anlmal PosplLal are now oerlng 1Wl members
a 10 dlscounL on amphlblan veLerlnary servlces and LesLs.
ur. WrlghL ls co-auLhor/edlLor of Lhe book Ampblbloo MeJlcloe ooJ copuve nosbooJty
and was named Lhe 2009 norLh Amerlcan veLerlnary Conference's Lxouc Speaker of
Lhe year, as well as Lhe 2008 uvM of Lhe year.
1o lnqulre abouL prlces and servlces, slmply conLacL Lhe hosplLal and menuon LhaL you
are a 1Wl member.
www.azeah.com
Gray, Heather M., Kyle Summers and Roberto Ibez D. (2009) Kin discrimination in cannibalistic tadpoles of the
green poison frog, Dendrobates auratus (Anura, Dendrobatidae). PIyIIomedusu. 8 (1): q1-o.
DESCRIPTION: Tests on cannibalism in Dendrobates auratus tadpoles were conducted to see if there was a bias to-
wards consuming unrelated tadpoles over related tadpoles. In the tests, larger tadpoles showed a distinct bias towards
attacking and consuming related tadpoles over unrelated tadpoles when presented a choice. There were no preferences
in attacking and consuming related and unrelated tadpoles when the choice was a single tadpole, these tests demon-
strate that D. auratus tadpoles are indiscriminate predators.
Hagemann S.; Prhl H. (2007) Mitochondrial paraphyly in a polymorphic poison frog species (Dendrobati-
dae; D. pumilio). MoIecuIur PIyIogeneLIcs & EvoIuLIon q, z: ;qo-;q;.
DESCRIPTION: Mitochondrial analysis of Oophaga pumilio has resulted in data that indicates that there are poten-
tially three species currently recognized as one species.
Puper uvuIIubIe Iere: ILLp:JJbIo-nIcu.InIoJBIbIIoLecuJHugemunnzoo;MILocIondrIuI.pdI
Lock, F.S. (1895) On a supposed action of distilled water as such on certain animal organisms. J. Physiol.
18(q): 1q-1.
DESCRIPTION: A review and comparison of prior information on the supposed danger of distilled water on tadpoles
and tubifex worms. Data indicates that contaminants of metal salts residues from the distillation process and not the
distilled water is the causative agent in the reported deaths.
Pryor. G.S. & Bjorndal, K.A. (2005) Effects of the nematode Gyrinicola batrachiensis on development, gut
morphology, and fermentation in bullfrog tadpoles (Rana catesbeiana): a novel mutualism. J. Exp. Zool.
oA:;oq-;1z.
DESCRIPTION: Contrary to popular wisdom on parasitic infections in captive anurans, infections by pinworms may
ucLuuIIy be benehcIuI Lo LIe IosL. n buIIIrog (Rana catesbeiana) LudpoIes, InIecLIons by u specIhc nemuLode demon-
sLruLed LIuL LIe nemuLodes ucLuuIIy Increused dIgesLIve eIhcIency Ior LIe LudpoIes.
Seung Yun Lee & Richard P. Elinson (2008) Abnormalities of forelimb and pronephros in a direct developing
frog suggest o retinoic ocid depciencg. AppIIed HerpeLoIogy. (1): -q6.
DESCRIPTION: In a direct developing frog (Eleutherodactylus coqui), disrupting the metabolism of retinoic acid
results in developmental abnormalities in both embryos and metamorphs. This indicates that developmental problems
muy be u eIIecLIve meLIod Lo deLecL InsuIhcIenL dIeLury vILumIn A (us reLInoIJreLInyI puImILuLe) In cupLIve Irog popuIu-
tions.
Amphibian Related Papers and Publications
Adam Bilsing
Adam Hess
Adam Rees
Adrienne Classen
Alan Walters
Alex Nemec
Alexander McKelvy
American Frog Day
Andrew Attea
Andrew Grossi
Aric Anderson
Ash Barnes
Beth Sims
Brent Barrick
Brian Mark
Brian Sexton
Bruce Maclean
Chad Marcum
Charles Templeton
Christina Hanson
Christopher Archuleta
Christopher P. Desmond
Christopher Miller
Cindy Dicken
Cliff Miller
Crystal Huempfner
Dan Wright
David Bolt
David Jones
Debra Valentine
Derek Benson
Dolores Santucci
Domenic Valenti
Donn Edwards
Doug Florian
Douglas Peel
Dave & Erin McLay
Ed Connors
Ed Kowalski
Emily Lisborg
Eric Powell
Eric Malolepsy
Gary McCarthy
Gerald Pottern
Greg & Amanda Sihler
Iam Maguire
Jack Reese
Jake Mallory
Jamie Neel
Jan Roland
Jason Geitzenauer
Jason Stein
Jennifer Macke
Jenny Pramuk
Joe Burman
Joe Milmoe
Johan Mattsson
John Allendorf
Jonathan Garrett
Jordan Ondras
Joseph Mailhot
Joshua Kennison
J.P. Lawrence
Kim Klisiak
Kristina Zunker
Kyle Kopp
Lee Hancock
Lissette Marin
Logan Benedict
Lonnie Cornell
Marc Knox
Mark Truax
Mark Budde
Martin Gruber
Martin Haberkern
Matthew Mirabello
Michael Khadavi
Michael Wallitis
Mitchell Heller
Nathalie Blocry
Nathan Bussard
Nathaniel Wyatt
Northwest Frog Fest
Olivia Morris Mueller
Patti Granzin
Paul Rust
Peter Keane
Raymond Coderre
Richard Hartman
Richard Revis
Richard Terrell
Robert Nhan
Robert Ossiboff
Ron Gagliardo
Ryan Stramaglia
Salvatore Zimmitti
Sarah Ehmer
Sarah Smith
Scott Menigoz
Scott Sliwinski
Sean Higgins
Sean Pennington
Sean Wilcox
Sonny Siemiller
Steve Kline
Steven Leising
Thomas Manchester
Tim Paine
Timo Paasikunnas
Todd Kelley
Walter Merker
William Harris
William Heath
William Hunter
Yasmine Callaham
Ylli Mujaj
Your 2009 contributions have enabled Tree Walkers International to continue moving forward in our mission to support the
protection, conservation, and restoration of wild amphibian populations. Without your generosity and commitment, this
work would not be possible.
Our S|ncere hcnks
Photo by Tim Paine
Amphibians are in
trouble.
You can help.
TREE WALKERS
INTERNATIONAL
United to save amphibians.
www.treewalkers.org

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