Leaf Litter 3.1

Leaf LitterThe Magazine of Tree Walkers International and Amphibian Conservation

EDITORRon Skylstad

ASSISTANT EDITORSLee Hancock

Jason KonopinskiEd Kowalski

PHOTO EDITORTim Paine

LAYOUTElizabeth Brock

TWI DIRECTORRon Skylstad

TWI PROGRAMS DIRECTORBrent 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 LitterVOLUME 3 | ISSUE 1

1 FROM THE EDITOR

FEATURES4 O!spring Begging, Larval Relocation and Parental Care in Dendrobates auratus

6 "e Use of a Modified 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: "e Variability of the Panamanian Green and Black Poison Dart Frog

40 Confessions of a Pumilio Chaser

54 "e Oophaga pumilio of Escudo de Veraguas

COVER Cochranella albomaculataphoto 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 SkylstadEDITOR-IN-CHIEF

photo by Danté Fenolio

2 | Leaf Litter :: Vol 3 Issue 1

El VallePenonome

Bocas del Toro

David

BoqueteEl Copé

Calobre

Santa Fe

San Felix Rio Santa Maria

Caldera

Escudo de Veraguas

Volcán Barú

PANAMAMAP OF THE COUNTRY OF

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 VALLEYon the island of O’ahu, Hawai’i, existsan introduced population of the greenand black dart-poison frog (Dendro-bates auratus). This population existsas the result of an early experiment forthe biocontrol of mosquitoes in whichapproximately 200 individuals fromTaboga Island, Panama were releasedin the valley. In the ~75 years since it’sestablishment, the population appearsto be contained within the Manoa Val-ley (a separate population does existon the other side of the island in theWaihole Valley, however there is somedebate as to whether this was part ofthe initial introduction or occurredlater).

Part of our research group (CG Farmerlaboratory at the University of Utah)

studies on this population. Thesestudies revealed details of parental care

behaviours within this species previ-ously not observed which I would like

order to fully appreciate these observa-tions, however, a short introduction toparental care behaviours is required.Parental care behaviours, loosely de-

-ment by a parent, play critical roles inthe life histories of numerous animalspecies. These behaviours are pro-

life history, yet in many species thesebehaviours have yet to be studied ingreat detail.

Within the Anura, these behavioursare widespread, having apparentlyconvergently evolved numerous times.The most common form of parentalcare within this group is clutch atten-dance, the behaviours of which rangein investment from simple clutch atten-dance to maternal provisioning of

larvae in the form of ‘feeder eggs.’ Thesubfamily Dendrobatinae contains sev-eral parental care behaviours, the mostbasal of which is clutch attendance bythe male followed by larval depositioninto bodies of water. This is the patternof behaviour currently described withinD. auratus.

around) the Harold B. Lyon Arboretumon O’ahu, we observed several larvaldepositions by male D. auratus intothe axils of a large bromeliad (Vrieseaimperialis). Upon arrival of a tadpole-carrying male D. auratus to a water-

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 tadpolemanually with the use of his rear

submerged in the axil;

3) the tadpole detached in the pool,but later re-attached to the backof the male and was relocated to adifferent axil following a distinct“begging” behaviour.

It is the third event that we would liketo discuss in this note, as we feel itdemonstrates complex and innatelyfascinating behaviours not typically at-tributed to this species. These behav-iours occurred in approximately 15%of the deposition events and relocationoccurred between 5 minutes after ini-tial deposition up to 48 hours.

The “begging” behaviour mentionedabove involved the tadpole stiffeningits tail, rapidly vibrating it for a few

Offspring Begging, Larval Relocation and Parental Care

in Dendrobates auratusBY BENJAMIN K. CHANPHOTOGRAPHS BY THE AUTHOR

University of Utah Department of Biology357 S 1400 E, Salt Lake City, UT 84112

O!spring Begging, Larval Relocation and Parental Care in Dendrobates auratus | 5

seconds, and then swimming erraticallyaround the male, especially underthe male’s throat and around his rear,until the tadpole was able to align withthe backside of the adult and re-attach. This larval “begging” behaviour differedfrom typical swimming patterns ob-served when tadpoles were manuallydisturbed, frightened, or under attack

each case the frog that approached wasthe same individual that had initiallydeposited the tadpole (determinedby distinctive colour patterns), andthese behaviours did not occur whenother frogs (of the same species) wereobserved in or near the axil. A similarswimming pattern has been observedin oophagous tadpoles of Oophagapumilio 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 afterthe 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

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 thanpreviously described. Our observationssuggest that both parent and offspringhave some degree of choice regardingthe site of larval development. Thesebehaviours aren’t without costs, how-ever, and the reasons for this increasedmale investment are unknown. We have conducted laboratory studies thatsuggest acute changes in phytotelmvolume may contribute to higher inci-dences of larval relocation.

Parental investment theory predictsthat the degree of parental care willdepend on offspring value, costs of care

associated with these behaviours.

Competition between offspring mayhave detrimental effects on offspring‘value’ (particularly in nutrient poormicrohabitats) and, therefore, wewould expect competition-avoidingmechanisms to evolve in these environ-ments. Several species of amphibianutilize nutrient-poor microhabitats forthe development of their offspring andit is likely that many similar behaviourshave evolved across taxa.

_________________Works Cited

Brust, D.G. 1993. Maternal brood care byDendrobates pumilio – a frog that feedsits young. Journal of Herpetology 27:96-98


AMPHIBIAN DECLINE NOW ENDANGERS at least one third of the world’s 6,000 plus amphibian species1. Time is short and amphibian conservation efforts are in dire need of additional training and capacity building programs2. The Atlanta Botanical Garden (the garden) has been committed to amphibian

years and has enacted in situ and ex

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

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

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

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

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/

into our terrariums, and then leaves

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

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 garden’s pod includes species of critical conservation need in response to the spread of amphibian chytrid fungus,

BY DANTÉ FENOLIO, ROBERT HILL, JULIA KAYLOCK, AND JENNY CRUSE SANDERSPHOTOGRAPHS BY DANTÉ FENOLIO

Atlanta Botanical Garden, Department of Conservation Research1345 Piedmont Ave NE, Atlanta, GA 30309

The Atlanta Botanical Garden’s frog pod is a 40 foot long repurposed shipping container modi"ed to include: two internal rooms, two windows, two doors, and internal environmental controls.

Amphibian Laboratory at the Atlanta Botanical Garden

The Use of a Modi"ed 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

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 Bob’s 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

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

involves the Horned Marsupial Frog, Gastrotheca cornuta, which was bred

garden. In the past, the young had

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 Rabb’s Fringe Limb Tree Frog, Ecnomiohyla rabborum, a recently described species. The

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 "nished amphibian room in the Atlanta Botanical Garden’s 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

evidence of the success. Several other species have been bred with regularity including the Lemur Leaf Frog, Hylomantis lemur, Pratt’s 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 garden’s 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

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 garden’s collection housed in our amphibian pod is

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 garden’s 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

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 Garden’s frog pod.

The Use of a Modi"ed 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 Darwin’s 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 garden’s 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

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 garden’s 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

container that is now our frog pod and was a huge help in working with us through all of the twists and

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,

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

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

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.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

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 o!spring have proven di#cult to rear due to what appears to be nutritional de"ciencies. 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

Atelopus zeteki. This spectacular species is not only one of the world’s 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 Círculo Herpetológico 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

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

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

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

Copé. Working within the boundaries of the protected Parque Nacional General de Division Omar Torrijos Herrera near El Copé, Lips and her team had been surveying transects

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 Copé was so disturbing it moved researchers, who had for years only studied and documented these declines, to call on

something about it. Lips summoned experts from the zoological, academic,

a meeting in Atlanta in late 2004 to report the sad news from El Copé 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 e!orts of PGF, ARCC and EVACC programs.


national park clearly did not prevent the arrival of Bd and its devastating

“What can we do?” There were still frogs in El Copé, 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 Copé (rescue and salvage) and El Valle (pre-emptive collection). At this time, as during the time of PGF’s 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 harm’s way. This was a pilot project to investigate the feasibility and challenges of conducting a multi-

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

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

collected and trapped food items were used to maintain amphibians during their in-country stay before being exported.

At El Copé, 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 Ibáñez. 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 Copé 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

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)

captivity. Environmental factors, such as extreme

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

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 Copé 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

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

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,

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

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

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 o!spring 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.

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 to remove any chlorine, pesticides, or other harmful

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 Gri!th, courtesy Houston Zoo.

Left: Visitors to the El Nispero Zoo learn more about the e!orts “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

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

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

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

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….

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

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 can’t 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 important—many 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

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,

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

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.

and other resources fully in place at the initiation will certainly help the project move ahead more smoothly. Again, in-country programs have

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

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

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,

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

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.

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 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

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

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

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.


__________________REFERENCES

Annis, S., F. P. Dastoor, H. Ziel, P. Daszak, and J. E. Longcore. (2004). A DNA-based assay identi"es Batrachochytrium dendrobatidis in amphibians. Journal of Wildlife Diseases 40:420–428.

Berger, L., Speare, R., Daszak, P., Green, D., Cunningham, A., Goggin C., Slocombe, R. Ragan, M., Hyatt, A., McDonald, K., Hines, H., Lips, K., Marantelli, G. & Parkes, H. (1998). Chytridiomycosis causes amphibian mortality associated with population declines in the rain forests of Australia and Central America. Proceedings of the National Academy of Sciences 95: 9031–9036.

Collins, J. and Storfer, A. (2003). Global amphibian Declines: sorting the hypotheses. Diversity and Distributions 9:89-98.

Frost, Darrel R. (2007). Amphibian Species

of the World: an Online Reference. Version 5.1 (03 January 2009). Electronic Database accessible at http://research.amnh.org/herpetology/amphibia/index.php. American Museum of Natural History, New York, USA.

Gagliardo, R.W., Crump, P., Gri#th, E., Mendelson, J.R., Ross, H., and Zippel, K. (2008). The Principles of rapid response for amphibian conservation, using the programmes in Panama as an example. International Zoo Yearbook 42:125-135.

Gascon, C., Collins, J.P., Moore, R.D., Church, D.R., McKay, J.E., and Mendelson III, J.R. (eds.). (2007). Amphibian Conservation Action Plan. IUCN/SSC Amphibian Specialist Group. Gland, Switzerland and Cambridge, UK. 64pp.

IUCN (2008). 2008 IUCN Red List of Threatened Species. <www.iucnredlist.org>.

Downloaded on 03 January 2009.

Lips, K. R., Burrowes, P. A., Mendelson, J. R. & Parra-Olea, G. (2005): Amphibian population declines in Latin America: A synthesis. Biotropica 37: 222–226.

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.

Pavajeau, L., Zippel, K.C., Gibson, R., & Johnson, K. (2008). Amphibian Ark and the 2008 Year of the Frog Campaign. International Zoo Yearbook 42:(1):24–29

Pfa!, 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 Zoo’s and Aquariums, Silver Spring, USA. http://www.aza.org/ConScience/Documents/Amphibian_Resource_Manual.pdf

Pounds, J. A., Fogden, M. P., Savage, J. M. & Gorman, G. C. (1997): Test of null models for amphibian declines on a tropical mountain. Conservation Biology 11:1307–1322.

Zippel, K. C. (2002): Conserving the Panamanian Golden Frog: Proyecto Rana Dorada. Herpetological Review 33: 11-12.

Zippel, K., Lacy, R. & Byers, O. (Eds.) (2006): CBSG/WAZA amphibian ex situ conservation planning workshop "nal report. Apple Valley, MN: IUCN/SSC Conservation Breeding Specialist Group.

Zippel, K. C. , Ibáñez D., R., Lindquist, E. D., Richards, C. L., Jaramillo A., C. A. & Gri#th, E. J. (2007): Implicaciones en la conservación de las ranas doradas de Panamá, asociadas con su revisión taxonómica. Herpetotropicos 3: 29–39.

Zippel, K.C. and J.R. Mendelson III. 2008. The Amphibian Extinction Crisis: A Call to Action. Herpetol. Rev. 33:23-29.

__________________ASSOCIATIONS

Paul CrumpHouston Zoo, 1513 MacGregor WayHouston, TX 77030USA

Ron GagliardoAmphibian Ark, Zoo Atlanta800 Cherokee AveAtlanta, GA 30315 USA

Edgardo Gri!thEl Valle Amphibian Conservation CenterEl Valle de Anton, Panama

Heidi GrossEl Valle Amphibian Conservation CenterEl Valle de Anton, Panama

Joseph R. Mendelson IIIZoo Atlanta800 Cherokee AveAtlanta, GA 30315 USA

Kevin ZippelAmphibian Ark/CBSG12101 Johny Cake Ridge RdApple Valley, MN 22124USA

Atelopus zeteki | 19

In March of 2008, I took a

Dutch and one Swede. I had already traveled with my two Dutch friends to Costa Rica in 2006, but this was the

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

species Atelopus zeteki.

-gists from the Smithsonian Institute (as well as other institutions) collected the

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 I’ve 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 Antón, Panama, where they are kept in an out-door vivarium beneath the roof next to

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 Campaña, but were told that chytrid1 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

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,

have him come with us.

Our Trip to El Valle While in Cerro Punta, near Volcàn Barú in western Panama (also known as Volcàn de Chiriqui), we succeeded

BY DENNIS NILSSONPHOTOGRAPHS BY THE AUTHOR

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

such short notice, and we where lucky

spoke Spanish.

The next day we left the highland for-est of Barú 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 30-35°C (86-95°F). 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, it’s the many rivers and streams in the region that make it possible for them to survive. Once we reached Antón, 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

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)

-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

fact, a conservation project for A. zeteki in the wild. We were also able to get the names of places that weren’t 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

give us more details about the area and project…or perhaps even take us to the project.

El NisperoWe 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 don’t 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 didn’t appear healthy, even though the per-sonnel probably did their best for the animals—I assume they just didn’t 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

where we could eat dinner and discuss plans for the following day (the res-taurant at the hotel wasn’t available to us because the hotel, except from the two rooms we were renting, had been

Vegetation in the highland forests that surround Volcán Barú.


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

Atelopus, looking for distant places where people didn’t seem to live or per-haps hadn’t 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

can get above 30°C (86°F), even if the temperature in windy El Valle was only around 24°C (75°F). After a while we decided that this would be our reserve plan if we weren’t able to contact Mario

project we were looking for.

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 HelpWe 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

too hot. So, after a very tasty omelette and freshly made pineapple juice, we went back to the Serpentario…only to

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 around—we 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

the amphibians ourselves. We had at least some idea as to where to start

any Atelopus, we were convinced that it was going to be an interesting day

other interesting animals.

A Search of Our OwnWe 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

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 wasn’t what we where look-ing for, so we followed the river up-

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

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 downstream—they didn’t have rubber boots on and the shoreline was drier in

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, didn’t bother looking more closely at it).

It was a big, beautiful female. She was mostly yellow with a few black spots along her back (it’s 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). It’s surprising how much they look like skin and bones, but this is exactly how they should look. We didn’t expect them to be as fast as they

The habitat of Atelopus zeteki.


A. zeteki is very good at jumping. They weren’t at all afraid to dive

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 leaf—it’s almost impossible to distinguish them with their yellow coloration and black markings. Their

discovery as well as a strong warning to predators of the toxins contained in their skin. The temperature was surpris-

(99°F) at the car.

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

my friends wouldn’t 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.

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 wasn’t 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.

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

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 FoundWe continued to make our way up-stream where we found species of both Colostethus and Eleutherodactylus. There were many different animals in

After a couple more unsuccessful hours

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-

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 I’m wrong, and that A. zeteki can still be found there.

Back to El ValleWhen 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

Animal Planet was going to attempt to

later that they did succeed, but I don’t know how many animals they found or the exact spot in which they found them—if 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. It’s 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 ridensBottom: Cranopsis melanchlora


After a shower and change of clothes we still hadn’t 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 weren’t 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 HabitatOn 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

of tadpole they were. There were many

most likely on someone’s private prop-erty, we chose to go back to the car. But right at that moment I saw some sort of leaf—or tree frog—leap into

camera onto my back and caught the frog. I immediately regretted catch-

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 didn’t take any photos of the frog, but think-ing back, regret that I hadn’t.

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 caught…and 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 RoadBack 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

a couple more species of dendrobatids along the way, but that is a different story…

ConclusionDue 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 Antón, I have chosen to omit some details regarding

The scientists that work with Atelopus zeteki, both in situ and in the USA, have received our information and my photos—it 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 wild…but 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

“!ose who have viewed at first 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 fires 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 finder of this marvelous creature is sublime. Anyone who spies the glittering brilliance of the frog is at first astounded by its beauty and overwhelmed with the excitement and joy of discovery; almost simultaneously he may experience great fear. !e story continues that any man who finds the legendary frog finds happiness, and as long as he holds the frog happiness will follow him everywhere. !e 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. SavageOn 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

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

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 (let’s say,

for argument’s sake, the “black and blue” one mentioned above that needs a mate), the least he can do is deal with

his vivaria. Unfortunately, the diligent frog keeper will

one seems to be able to tell him for certain where his frogs

– just mere speculation.

online morph guide. Within a few clicks he observes a photo

the photo look too bright or the settings on his monitor),

(Further) Notes on Dendrobates auratus

BY FRANK STEINMANNPHOTOGRAPHS BY THE AUTHORUNLESS 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

is for the future of keeping and breeding this member of the dendrobatid family.

Author’s 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, it’s blue, and that’s 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 keeper’s 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

-ent populations of blue auratus described to date, not to mention the ones yet undiscovered. What’s 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 Schwarzenegger’s 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

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

to poison dart frogs. However, if you

is based on the Greek and pertains to a thing’s form, shape, “gestalt” – basi-cally the appearance of something. When describing the distributional

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 coloration—there 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. can’t 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

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., Külpmann, V. & Was-

1traveled 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”

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. Birkhahn’s 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 wildlife—using 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-

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-

Dendrobates auratus. Never before seen color morphs (I use the term color morph, because in most cases there was no information regarding

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, it’s easy to see how each frog could be used to create new “morphs” in the hobby.


I would consider “pro” are their frog

Dendrobates auratus, all of which look

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 1990’s. Soon after their discovery there were several, rather grey, imports to Germany of these frogs, which

Dendrobates auratus “Birkhahn” or “Wassman”. The phenotypes and visual characteristics of these frogs were then

-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

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

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 Copé”

though many similar looking morphs tend to populate the central Panamani-an slopes and mountains that surround the village of Copé. In actuality, the national park (Parque General Divi-sion Omar Torrijos Herrera) where the

were found has nothing to do with the village of Copé, which is located just a few miles from the park. If you explore the surrounding area around Copé, fol-lowing the Rio San Juan or Rio Toabre in the Caribbean direction, what you

A frog sold as an “El Copé” morph without any clear information as to its speci"c origins. One can only con"dently know that it was collected somewhere in the general region of El Copé.

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 Copè, Penenomé, 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

holds true for all forms of auratus.

The frogs that have been imported as “Copé” could quite possibly be from every place of that region—frogs 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 necessary to have variables such as weather phenomena and wet and dry

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

the Canal Zone) have a D. histrionicus-like dorsal pattern of many small, light spots (Silverstone 1975).” The “Campana” morph is found on the

Campana. The two villages of Campana and Capira are very close to each other and, as a matter of fact, the frogs don’t 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

here we go yet again with self-made

Cream”, “Brown and Beige,” ”Kahlua and Cream Camo”…these 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 “Camo$age” auratus.


and unprofessional names? When we have inquired about the origin of the

a breeder shared with was something to

picked on the other side from where the brown and beige ones are from.” When we asked for more information, they said they didn’t have any more than

auratus were imported and sold in 2006. Where did they come from? No-body seems to know, and it’s 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

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 di#cult to "nd both in the wild and captivity.


in its natural habitat—several people -

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 population’s 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

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. It’s not possible.

Further east, near the town of Santa Fé, lives another population of D. auratus

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 Fé itself sits within the shadow of the divide between Atlantic

by montane rainforests at heights be-tween 1200 and 1550 m asl. Not quite at the peak, but at heights around 800

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 rare—when searching for it in January

-mens even though it was raining cats and dogs and the conditions seemed ideal (I can’t 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

Soberania National Park and Barro Colorado Island. Ancon hill, however,

refuge for many animals of Panama City. For me, this place is one of the

-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 Panama’s largest malls, you can

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 existed—the information was never provided by the importers.

As you move west from Panama City and leave the Canal Zone, you reach a city called Arraiján. At a nearby river

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

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

dump. Wastewater and

submerged vegetation. Human debris and refuse lines the shores, evidence of

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

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.

specimen of Dendrobates auratus seems to have come from the Taboga

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

“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

granular. Color, above, metallic golden; beneath, uniform bluish

Taboga, in the Bay of Panama, by the late Prof. C.B. Adams, of Am-herst College, Mass”. (Phyllobates auratus Girard, 1855 “1854”, Proc.

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 Hawai’i to help aid in mosquito control. “In 1932, 206 specimens of D. auratus from Taboga or Taboguilla Islands, Panamá, 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

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

the leftovers of our civilization. Exer-cising the motto “out of sight, out of mind,” the locals dump their waste into

of auratus hopping between rotting

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 having—only young and adolescent frogs seem to have this color. As they mature, the frogs lighten

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

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.” It’s 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 didn’t want to divulge the actual locality, so the frog was once again labeled with an

-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 (I’ll 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

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.

Di!erent Slopes, Di!erent Climates

All populations of Dendrobates auratus found on the Paci"c slope have to exist in a completely di!erent climate than their Caribbean relatives. This di!erence in climate is caused by the Cordillera Central, the mountain range that extends from east to west and seperates Panama into two di!erent climate zones. The majority of the rain clouds that come in o! 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 di!erent activity pattern than the Paci"c populations, who tend to hide during the dry season. You can often search for hours and not "nd a single frog, then turn one stone or lift one dry palm leaf and seemingly "nd 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-

forms of Dendrobates auratus which are naturally smaller and more con-

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.

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

and “not to be taken seriously”…but the call, behavior, and captive breeding seemed to allude to the fact that we had to consider it a new morph of Dend-robates auratusThis 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

tone. Some individuals carry these dots

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-

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

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

-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

that this way of keeping and breeding poison dart frogs (e.g. dealing with spe-

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,

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., Külpmann, V. & Wassmann, K. (1994): Zur Variabilität 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 duringU.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):36–37.

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 classi"cation 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 "rst description is the so called “nominate form” and is recognized by a scienti"c 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 WALDRONPHOTOGRAPHS BY THE AUTHOR

Confessions of a

Pumilio Chaser

Confessions of a Pumilio Chaser | 41

The Frog Gardens of

Bocas del ToroOoophaga

pumilio (and its highly evolved parental care behavior, extreme color morph variation, and all around doe-eyed cuteness) became a fatal attraction…or at least one that lead me to strange, obscure places like a Guaymi Indian settle-ment on a tiny Caribbean island in western Panama’s Bocas del Toro region. Causing me to act out in strange, irratio-nal behaviors, such as knocking on the door of a complete stranger’s 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

tiny red frogs?”

all executed in really badly broken textbook high school Spanish. I’m 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 weren’t an image from an insane man’s 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 didn’t 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 !nger-blis-tering evening in a turn-of-the-20th-century bohemian San Fran-cisco "at. It is based on a series of pumilio-watching trips I had taken to Panama’s Bocas del Toro Archipelago between 1998 and 2001 and was wri#en at a time when frogging was still a pursuit for globetro#ing 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.org…eccentric rebels, all of them, and one-of-a-kind raconteurs. I printed out thirty hard copies of “Confessions...” on my workplace’s 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 Sea#le and sold none of them$the rest have dis-solved into the ether and the compost heap. But now, “Confes-sions...” has been resurrected, !nding a new home here within the glossy pages of Leaf Li#er 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 "ip-"op…


of the leaf litter herpetofauna community in La Selva’s 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

the answer, we must return to my human acquaintances of Caribbean lower Central America and understand a bit about their dinner menus.

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 America’s gomphotheres (ancient mastodont-like creatures), glyptodonts (giant armadillos), and megalony-chids (giant ground sloths). This may be so, though I’m not entirely convinced of this theory. I am convinced, however, that when in a foreign land and cultural differences make

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 haven’t. 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: “Wow…that’s terrible! El Tigre is a rare animal and not many still exist.”

Indian: “Yes…but 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

region…

Indian: “And turtle (a.k.a. Green Sea Turtle, a.k.a. “Bocas Beef”), all types

crab, octopus, lizards…”

Pumilio Chaser: “What kinds of lizards?”

Indian: “U-grra (Helmeted Iguana, Corytophanes cristatus) and jilida (Anolis spp.

Pumilio Chaser: “Do people eat frogs?”

Indian: “Oh yes, ogungo (medium-sized Eleutherodacty-lus) and no-lu (tiny, thumbnail-sized Eleutherodactlyus).”

Pumilio Chaser: “And (Dendrobates auratus) and ranitas rojas (Oophaga pumilio)?”

Indian:

“No, I don’t think anyone around here eats little red frogs. Do you eat them?”

why one has traveled 2,000 miles to chase tiny red frogs…especially to those whose daily chasing after animal protein

-ible” small wild animals in the forest at the edge of their current gardens or in feral regrowth plots of gardens past

the riddle of O. pumilio’s enhanced abundance outside of the

rainforest—Neotropical 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 pumiliorearing 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

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 chocolate—now lying fallow due to a price drop on the international market and the ravages of an invasive fungus—provide 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 pumilio…and 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

2001June 6, 2001Today, 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 hour…and did. The forest and its inhabitants were refresh-ingly uncluttered—easy 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, 2001Today I followed the black hawk again. Down Punta Vieja, 3 km or so on the Atlantic side, along the beach and palm

rojas! This race of Oophaga pumilio was unique—small, -

derneath and brown legs w/ spots. Some were heavily black-dotted and the dorsum—fantastic! Pura rana vida!

They were fairly abundant in someone’s 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

fantastic!

6/8/01These 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 island—good suerte foretold by the omen of a Black Hawk’s* scolding. He’s angry I will steal his big blue-clawed land crabs. He keeps watch over me…

*Buteogallus an-

thracinusyou hear the Black Hawk, stop and listen to what it is saying.

Isla Solarte, June 9, 2001

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 tree—why is he here alone? Why am I?

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

“And then — Amen!— I heard the Golden pumilio chorus:

“EP! - EP! – EP!! – EP! ! – EP!”

-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, 2001I write this after four days of not writing. We have been here, at 500 meters or so at Wilburto Martinez’s Willie Mazu

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

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, 1000’s and 1000’s of insectos came out of the surrounding forest and a large bat and a green leaf mantid and the biggest Bufo mari-nus I’ve ever seen came to feast on them. “El sapo sabe.” “The Toad knows.” Hilma, the cook, would say later. But

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.”

in an old cacao grove/lowland forest remnant that will be

tree growing along the highway near the “Super de Chino”

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 pumilio—“como 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

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, PanamaIsla BastimentosVariable. 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 north—no whitish yellow frogs were seen in this population after a thorough survey.


Isla Solarte / Cayo NancyA 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 delight—after a morning rain, hire a boatman from Bocas and have him drop you off at Nancy Cay. As you

glowing golden froglets hopping every-where, almost like little forest fairies. Look for them in the palm swamp and the old United Fruit Company’s grave-yard (mostly malaria victims—bring mosquito repellant).

Isla ColonI have not explored this island nearly as much as I’d like to. It’s 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 PartidaIf 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 beautiful…colored 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…

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

to meet…there are truly no two pumilio alike. And the ad-ventures you’ll have along the way…I encourage you to visit these places and explore!


Midnight InterludesOne of the joys of frogging the Neotropics is that you can chase frogs all day and all night—pumilio 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

4/28/95 Rara Avis 12:00 a.m.

headlamp, foliage platforms alive with bizarre katydids, tink

the rivulets – the iridescent shimmer of a male Rivulus kil-

of a bromeliad leaf. Flash of thunder cracks, bringing in the

the forest with the presence of life. Here, now, as always – the rain, the myriad of the living and me.

8 de Mayo, 2003Rara Avis, Costa Rica, Rio Atelopus watershed“Lao by La Catarata”Here we are.Tonight in thisCarib-beanRain-forest.Along this Rio.By this Water-fallI don’t have to askTo knowwhat unites us – palm, fern, 10,000 types of leaf.Brilliant-thighed forest frog on the trail.Fan antennae beetle of the tree.

My headlamp – illuminates an amber poolwhere sleeping tadpoles swirland RivulusRaindrops drop dropletson the cat-eyed snake’s hunting eye.

Bocas del ToroAgua de pipa para tomarHermosas play para caminarAgua sal para banar

Bocas del ToroCoconut juice for drinkingBeautiful beaches for walkingSaltwater for bathing

- lyrics from a Bocatoreno folk song


The Color of Love and

the Polka-Dot Poison

Frog, Oophaga pumilio,

Isla Bastimentos,

PanamaWhen someone asks why I would eschew the comforts of home and modern convenience for the muck and biting insects of the tropical forest…well, no one ever does…but if someone were to, I would have to say,

Where there are poison frogs, there is Nirvana.

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 Panama’s cloudforest continental divide;

spilling off on either side of the narrow trail. An enchanted place, the cloud forest was alive with multi-hued colors of

the diffused light along the trail. Green epiphytes dripped from tree branch to tree branch. The fog masses coming, go-

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 frogs…but the familiar sounds of something like Oophaga pumilio were coming from the trees. Craning my binocu-

realized I was listening to the calls of an amazingly adapted canopy poison frog, Oophaga arborea, and promptly fell down a rather steep hill.

-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, it’s 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

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 planet’s great paradises…and 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

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

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: “Uh…I 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…

-

lowing the trail out of town, a grove of coconut palms was

poison frogs could be heard. It wasn’t 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 orgy—in 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. What’s 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

Sir David Attenborough and a BBC documentary production relating the wonder in the following facts of O. pumilio’s life

“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

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 male’s deposition of the tadpoles to their rearing sites—not much of an energy expenditure on either parent’s 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-

tadpoles hatch out, he splits and is out looking for another mate. Life is short, there are no eggs to be wasted—she 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 offspring…the 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

red frog with a white belly and a black polka-dot back?

it’s 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

“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., McCa!erty, 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

BOCAS DEL TORO ARCHIPELIGO

MAP OF THE


BY JUSTIN YEAGER PHOTOGRAPHS BY THE AUTHOR

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

of mine to see A. zeteki before they were extinct. For this

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

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

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

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 face—my lips relished the fresh taste of the rain.

Despite my best efforts to keep down my pineapple-and-

a moment so I wouldn’t get hit by the waves. He cheerfully obliged, if for no other reason than to chuckle at me. In

my head I couldn’t help but start singing the theme song to Gilligan’s 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

Once on the island we quickly gathered our things and took

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 couldn’t 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

deepest mud holes, until I heard calling again. I was convinced that I was close

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

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

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 in—we 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

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

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

TREE WALKERS INTERNATIONAL PRESENTS

September 3-4, 2010

Lynnwood, WA

Microcosm is a two-day event that brings enthusiasts together through the common bond

of nurturing life within glass enclosures, from aquaria to vivaria, greenhouses and Wardian

cases. Just as species combine to form healthy, functioning ecosystems, Microcosm seeks

to form a coalitions of partners who apply their skills in cooperation to safeguard the

planet’s biodiversity.

For more information visit www.treewalkers.org


Gray, Heather M., Kyle Summers and Roberto Ibáñez D. (2009) Kin discrimination in cannibalistic tadpoles of the

green poison frog, Dendrobates auratus (Anura, Dendrobatidae)

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.; Pröhl H. (2007) Mitochondrial paraphyly in a polymorphic poison frog species (Dendrobati-

dae; D. pumilio)

DESCRIPTION: Mitochondrial analysis of Oophaga pumilio has resulted in data that indicates that there are poten-tially three species currently recognized as one species.

Lock, F.S. (1895) On a supposed action of distilled water as such on certain animal organisms. J. Physiol.

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.

DESCRIPTION: Contrary to popular wisdom on parasitic infections in captive anurans, infections by pinworms may Rana catesbeiana -

Seung Yun Lee & Richard P. Elinson (2008) Abnormalities of forelimb and pronephros in a direct developing

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

-tions.

Amphibian Related Papers and Publications

Adam BilsingAdam HessAdam ReesAdrienne ClassenAlan WaltersAlex NemecAlexander McKelvyAmerican Frog DayAndrew AtteaAndrew GrossiAric AndersonAsh BarnesBeth SimsBrent BarrickBrian MarkBrian SextonBruce MacleanChad MarcumCharles TempletonChristina HansonChristopher ArchuletaChristopher P. DesmondChristopher MillerCindy DickenCliff MillerCrystal HuempfnerDan WrightDavid BoltDavid JonesDebra ValentineDerek BensonDolores SantucciDomenic ValentiDonn EdwardsDoug FlorianDouglas PeelDave & Erin McLayEd ConnorsEd KowalskiEmily Lisborg

Eric PowellEric MalolepsyGary McCarthyGerald PotternGreg & Amanda SihlerIam MaguireJack ReeseJake MalloryJamie NeelJan RolandJason GeitzenauerJason SteinJennifer MackeJenny PramukJoe BurmanJoe MilmoeJohan MattssonJohn AllendorfJonathan GarrettJordan OndrasJoseph MailhotJoshua KennisonJ.P. LawrenceKim KlisiakKristina ZunkerKyle KoppLee HancockLissette MarinLogan BenedictLonnie CornellMarc KnoxMark TruaxMark BuddeMartin GruberMartin HaberkernMatthew MirabelloMichael KhadaviMichael WallitisMitchell HellerNathalie Blocry

Nathan BussardNathaniel WyattNorthwest Frog FestOlivia Morris MuellerPatti GranzinPaul RustPeter KeaneRaymond CoderreRichard HartmanRichard RevisRichard TerrellRobert NhanRobert OssiboffRon GagliardoRyan StramagliaSalvatore ZimmittiSarah EhmerSarah SmithScott MenigozScott SliwinskiSean HigginsSean PenningtonSean WilcoxSonny SiemillerSteve KlineSteven LeisingThomas ManchesterTim PaineTimo PaasikunnasTodd KelleyWalter MerkerWilliam HarrisWilliam HeathWilliam HunterYasmine CallahamYlli 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.

Photo by Tim Paine

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