Leaf Litter

The Magazine of Tree Walkers International and Amphibian Conservation

DARWIN S FROG 11
The Dar win s Frog
Cons er vat i on Ini t i at i ve
BY DANT F ENOL I O, MAURI CI O FABRY, ANDRES CHARRI ER,
MARCEL A TI RADO, MARTHA CRUMP, AND BI L L L AMAR
PHOTOGRAPHS BY DANT F ENOL I O
It was known as sapito vaquero or cowboy
frog by the Spaniards (Cei, 1962; Torres
and Castillo, 1973). Stranger than the
hcLIonuI unImuIs reporLed In LIe duy,
these amphibians have a bizarre physical
appearance to match their colorful name,
sporLIng u Iong, esIy, nose-IIke uppenduge.
Collectively referred to as Darwins Frogs,
the two species are in the genus Rhinoderma.
The Darwins Frog, Rhinoderma darwinii,
was formally described in 1841 (Dumril
and Bibron, 1841) and Chiles Darwin Frog,
R. rufum, was described in 1902 (Philippi,
1902). The former is found in Chile and
Argentina while the latter is a Chilean
endemic.
Darwins Frogs have a reproductive mode
that makes them noteworthy. Like many
anurans, male Rhinoderma darwinii
cuII Lo uLLrucL IemuIes on LIe IoresL oor.
However, the courtship behavior of the two
species can include direct kicks to the males
head and body by the female (Busse, 1989,
1991, 2002). Females deposit a clutch of
eggs on LIe dump IoresL oor und LIe muIe
fertilizes them. At that point, the female
leaves and her parental duties are done.
The male remains in the area of the clutch
as it develops. As the tadpoles hatch, the
male takes the developing larvae into his
mouth and manipulates them through an
opening (the vocal slit) below his tongue
and into his vocal sac. The tadpoles develop
inside the structure that is normally used
for vocalizing! After a developmental
period which can last from 45 to 60 days
and is temperature dependent, the male
spits out fully formed froglets. Male
Rhinoderma rufum spit out tadpoles in a
water source rather than accommodating
the larvae all the way through development.
Within the humid leaf litter of Chiles
temperate southern forests is a frog so unique that it captured
the attention of Charles Darwin when he visited the region.
12 LEAF LITTER VOL 4 ISSUE 1
The unique reproductive strategy of these frogs (vocal
sac brooding) has long been recognized and new
details about the behavior were added by 19th and
20th century investigators (e.g. De la Espada, 1872;
Harting, 1879; Howes, 1888; Barros, 1918; Wilhelm,
1qz;, 1qz; Puumer, 1q; Busse, 1q;o).
Unfortunately the Chiles Darwin Frog, R. rufum, has
not been seen since the 1970s (Busse and Werning,
2002; Crump, 2003). The decline of the Darwins Frog,
R. darwinii, is also documented and looks to be most
apparent in the northern portion of its limited range
(Busse, 2002; Busse and Werning, 2002; Crump,
2003; Crump and Veloso, 2005). No single cause
accounts for all amphibian declines (e.g., Blaustein,
1994; Pounds et al., 2006; Mendelson et al., 2006;
Collins and Crump, 2009). Likely contributing to the
Darwins Frog declines are problems of deforestation,
replacement of native forests with mono-cultured exotic
tree species, and non-sustainable land management
practices (Armesto et al., 1996; Aravena et al., 2002;
Cuevas and Cifuentes, 2009). Many of the frog
species that inhabit the dwindling southern temperate
humid forests of southwestern South America are
found nowhere else; there are minimally 37 anurans
unique to these forests in Chile, constituting no less
than 85% of the regional amphibian fauna including
three endemic genera and an endemic family to Chile
(Calyptocephalella, Insuetophrynus, Telmatobufo, and
the Calyptocephalellidae, respectively) (Daz-Pez et
al. 2008; Rabanal and Nuez, 2008; Vidal et al. 2008;
Fenolio et al., 2011). Many of them are now threatened.
Conservation actions will be required to ensure the
survival of these species, and useful activities include
the implementation of captive breeding and assurance
colonies.
Darwins Frogs have long been bred in captivity. The
eurIIesL recorded success LIuL we Iuve been ubIe Lo hnd
In LIe IILeruLure wus by Puumer (1q). AIso In LIe
early 1930s, Wilhelm removed tadpoles from a male
R. darwinii und urLIhcIuIIy reured LIem (WIIIeIm,
1qz). WIIIeIm wus uIso LIe hrsL Lo wILness und
report a male R. darwinii taking larvae into his mouth
(Wilhelm, 1932). More recently, Klaus Busse bred
a captive group in Germany multiple times (Busse,
1970, 1989, 1991, 2002). One of us (DBF) had a
captive group in the 1980s and 1990s and they bred for
R.darwiniihabitat.
DARWIN S FROG 13
several years. Recently, a captive breeding program
at the Universidad de Concepcin captively produced
R. darwinii and the program we detail here, at the
National Zoo of Chile, has groups of frogs breeding.
This is all great news. Having multiple programs in
Chile working with this endangered amphibian helps
to bolster the likelihood that they will survive into the
future. Collaboration between these groups is in the
best interest of these endangered amphibians.
The Darwins Frog Conservation Initiative is a
collaboration between the Atlanta Botanical Garden,
The National Zoo of Chile in Santiago and personnel
from the Center for Advanced Studies of Ecology and
Biodiversity at the Catholic University of Chile, from
Northern Arizona University at Flagstaff, and from
GreenTracks, Inc. The goals of the initiative focus on
elucidating the reasons behind the declines of Darwins
Frogs and other amphibians endemic to Chiles
temperate humid forests. Further, goals include
attempts to curb the declines using techniques such
as: (1) development of captive assurance colonies with
ex-situ breeding of endangered amphibians within
Chile, (2) monitoring of diseases in wild populations
to better inform policymakers and conservation
authorities in Chile, and (3) education of the general
public regarding problems that Chilean amphibians
face. Ultimately, we aim to provide information about
declining Chilean amphibians (including the spread of
emergent infectious amphibian disease) and to provide
conservation options. Another goal is to establish the
'DUZLQV )URJ DV D DJVKLS VSHFLHV WKXV EULQJLQJ
attention to the plight of all of south Chiles endangered
amphibians.
2QH RI RXU UVW SURMHFWV ZDV WR GHYHORS D FDSWLYH
breeding facility within Chile. Many projects involving
amphibian conservation have removed individuals of
the endangered species in question from the range
country for captive reproduction efforts elsewhere.
We feel that by aiding local conservation efforts,
complete with trained personnel and a modern
captive reproduction facility, the chances of long
term conservation success are greater. We hope the
program functions in perpetuity as long as Darwins
Frogs and other endemic amphibians of Chile require
conservation assistance. A grass roots approach
harnessing local enthusiasm and pride is an important
part of the equation. Additionally, by removing
individuals of an endangered species from the range
country, the frogs might be exposed to new pathogens
in the host country. This risk complicates the
14 LEAF LITTER VOL 4 ISSUE 1
prospects of returning captive bred individuals to the
range country for reintroduction should that become a
necessity. This is not a condemnation of past captive
reproduction efforts outside of host countries. Rather,
we feel that as the collective amphibian conservation
community has learned through its experiences,
range country breeding facilities, when possible, are
preferable. Cooperative projects involving the Atlanta
Botanical Garden have led to the same conclusion.
Our captive breeding facility is located on the grounds
of the National Zoo of Chile in Santiago (Parque
Metropolitano de Santiago, Chile). The building was
designed with one of its walls made of glass so that
the visiting public could see into the laboratory. We
wanted to use the facility for education and outreach
with the general public. To draw visitors to the facility,
we commissioned a large sculpture of a Darwins Frog
made by a Chilean artist and placed it to one side
of the lab. Additionally, signage with information
about Darwins Frogs and other threatened Chilean
amphibians educates visitors. To emphasize education
for the general public, we developed a bilingual website
featuring our project: www.savedarwinsfrogs.org.
We have also emphasized education for the personnel
running the facility. The lead veterinarian for the zoo,
Marcela Tirado, visited the Atlanta Botanical Garden
for captive amphibian husbandry training. She also
attended the Association of Zoos and Aquariums
captive amphibian management program.
The captive breeding facility at the National Zoo of Chile
is modeled after the amphibian breeding facilities at the
Atlanta Botanical Garden. Front opening terrariums
streamline maintenance and upkeep. ZooMed
Laboratories supported our project with discounted
rates on equipment and in serving as a staging ground
for a host of equipment purchased in the United States
before it was shipped to Chile. There are 32 18x18x18
inch (roughly 46x46x46cm) terraria in the facility to
accommodate the same number of breeding groups.
7KHJODVVWHUUDULDZHUHGULOOHGDQGWWHGZLWKEXONKHDG
WWLQJVWRLPSOHPHQWGUDLQDJHV\VWHPV(DFKXQLWKDV
a false bottom made of egg crating to keep fouled
water away from the inhabitants and to provide ample
drainage of the media. The terraria are supported
on lab racks. The lab racks are arranged so that the
visiting public can see them through the wall of glass.
DARWIN S FROG 15
Automated misting systems keep the humidity in
each enclosure at appropriate levels and dampen the
terraria a number of times daily (the number of misting
evenLs Is uILered Lo repIIcuLe heId condILIons dependIng
on the season). Fluorescent lighting illuminates
the enclosures. The building has an environmental
control system strong enough to keep the ambient
temperatures cool during the hottest of Santiagos
summer days. Floor drains facilitate cleaning. Should
the electricity fail, a self-starting generator has been
provided that can power the building for an extended
period through potential blackouts. A spare water
storage unit was also implemented on a hillside above
the breeding facility, should the water supply to the
zoo be interrupted. The 350-gallon (1325L) reserve
tank has enough water to run the facility for more than
four weeks. The water will gravity-feed to the breeding
IucIIILy In LIe ubsence oI eIecLrIcILy. TIe boLLom oor
of the captive breeding facility is an insect culturing
IucIIILy. CrIckeLs, coIIemboIuns, und our beeLIes ure
already in production. We are working to add bean
beeLIes, upIIds, und wIngIess IruIL Ies. AddILIonuI
good news Is LIuL In zo1o our IucIIILy survIved LIe hILI
strongest earthquake in recorded history. In particular,
the self-starting generator and backup water supply
worked uwIessIy.
Our hrsL IIve Irogs were udded In zo1o. We ure pIeused
to report that they are already breeding. The goal is
to maintain the collection of wild collected frogs by
locality. We would like to have as many localities
represented in our facility as space permits. Breeding
groups ure LIree Lo hve Irogs In sIze. CupLIve oIIsprIng
are accommodated individually in deli cups when
young, and then in plastic tubs on shelves.
Field work has involved visiting historic localities where
Darwins Frogs have been reported. It also includes
checking forests without reports of Rhinoderma that
have appropriate habitat. Since 2008, we have visited
dozens oI IocuIILIes IookIng Ior LIe Irogs. WIen we hnd
populations, skin swabs are collected. The swabs are
tested for the presence of amphibian chytrid fungus
(Bd) in the laboratory of M. Levy at North Carolina
State University, School of Veterinary Medicine. At the
end of our study, we will publish a report documenting
where we observed frogs and where we have detected
amphibian diseases.
VIsILs Lo heId sILes Iuve InvoIved u Iew IocuIILIes
accessible via a 2-wheel-drive vehicle. However,
4-wheel-drive vehicles are usually required. Hiking in
from the ends of logging roads and other non-developed
access ways is also routine. For a few remote localities,
renting horses and mules has been necessary. The pack
animals allow for access to distant forest fragments
wILI cumpIng equIpmenL und u heId crew Ior severuI
days. The trips to these remote localities have been
memorable. Uncut forest fragments offer a glimpse
into what southern Chile was once like.
Our project has combined three key components: (1) an
ex-situ captive breeding facility within Chile, complete
wILI LruIned personneI, (z) u heId progrum Lo IeIp us
better understand the causes for the declines of Darwins
Frogs, and (3) an education and outreach program to
help us communicate our message to the public. We
feel that projects with all of these components have a
better chance at producing viable conservation options
for declining amphibian species. We are currently
looking to expand our capacity at the National Zoo of
Chile to accommodate six to twelve additional species
of amphibians that have experienced marked declines
in recent years. Sadly, there is no shortage of candidate
species. Building from the platform of an educated
husbandry staff in Santiago, the experiences we have
had with Darwins Frogs, and the relationships we have
developed with wildlife authorities in the government
and throughout Chile, we are optimistic about our
chances. Nevertheless, the issues threatening the
amphibians of Chiles southern humid forests are not
going to be solved through these measures alone. Non-
sustainable forestry practices are eliminating critical
habitat. Alteration of habitat in other ways is also
contributing to the problems. Invasive species, like
trout, could be impacting local anurans by feeding on
uquuLIc Iurvue. CupLIve ussurunce coIonIes, heId work,
and educational outreach programs offer steps in the
right direction, but there is still much work to be done.
Charles Darwin referenced the origin of species as
the mystery of mysteries in the opening paragraph
of On the Origin of Species (Darwin, 1859). Over 150
years after his landmark publication, the mystery of
mysteries might not be the evolutionary origins of
Darwins Frogs, but rather their fate at the hands of
humans.
Acknowledgements: We are grateful to the following
Ior hnuncIuI supporL: TIe AssocIuLIon oI Zoos und
Aquariums and their Conservation Endowment Fund
(grant No. 08-809), The Shared Earth Foundation,
The Sophie Danforth Conservation Biology Fund,
The Chicago Board of Trade Endangered Species
Fund, Cisco Systems, and the George and Mary Rabb
Foundation. We would like to thank Klaus Busse for
his help throughout our project and with this article.
Klaus has more experience in keeping and breeding
Rhinoderma than anyone else so we are grateful for his
input. We would like to thank ZooMed Laboratories
for their generous assistance with this project. We
thank Robert Hill and Mark Mandica for their time
and suggestions as we developed the breeding center.
Osvaldo Cabezas, our amphibian keeper at the National
Zoo of Chile, has done a tremendous job in keeping our
animals happy and healthy.
16 LEAF LITTER VOL 4 ISSUE 1
Ltrnntunr Ctrb
Barros, R. 1918. Notas sobre el sapito de vaquero
(Rhinoderma darwinii). Revista Chilena de Historia
Natural 22: 7175.
Blaustein, A.R. 1994. &KLFNHQOLWWOHRU1HURVGGOH"
A perspective on declining amphibian populations.
Herpetologica 50: 8597.
Busse, K. 1970. Care of the young by male
Rhinoderma darwinii. Copeia 2: 395.
Busse, K.1989: =XP%UXWSHJHYHUKDOWHQGHV
Nasenfrosches Rhinoderma darwinii (Anura:
Rhinodermatidae). Tier u. Museum 1 (3): 59 63.
Busse, K.1991: Bemerkungen zum
)RUWSDQ]XQJVYHUKDOWHQXQG]XU=XFKWYRQ
Rhinoderma darwinii: Balz bis Eiablage.
Herpetofauna 13 (71): 11 21.
Busse, K. 2002 (2003): )RUWSDQ]XQJVELRORJLHYRQ
Rhinoderma darwinii (Anura: Rhinodermatidae)
und die stammesgeschichtliche und funktionelle
Verkettung der einzelnen Verhaltensablufe. Bonn.
zool. Beit.r 51 (1); 3-34.
Busse, K. and H. Werning. 2002. Another extinct
DPSKLELDQVSHFLHV"7KHIDWHRIWKH'DUZLQ)URJV.
ZGAP Mitteilungen 18(2) 2: 1618.
Cei, J.M. 1962. Batracios de Chile. Ediciones de la
Universidad de Chile. Santiago de Chile. 128pp.
Cei, J.M. 1980. Amphibians of Argentina. Italian
Journal of Zoology Monograph 2: 1609.
Collins, J.P. and M.L. Crump. 2009. Extinction
in Our Times: Global Amphibian Decline. Oxford
University Press.
Crump, M. 2003. Vocal sac-brooding frogs. From
Grzimeks Animal Life Encyclopedia, second ed.
Hutchins, M. ed. Gale Group.
Darwin, C. 1859. On the Origin of Species by Means
of Natural Selection, or the Preservation of Favoured
Races in the Struggle for Life. John Murray, London.
De La Espada, J. 1872. Sobre la reproducin de
Rhinoderma darwinii. Annales de la Sociedad de
Historia Natural de Madrid I: 139-151.
Dumril, A.M.C. and G. Bibron. 1841. Erptologie
Genrale ou Histoire Naturelle Complte des
Reptiles. Volume 8. Paris: Librarie Enclyclopedique
de Roret.
Goicoechea, O., O. Garrido, and B. Jorquera. 1986.
Evidence for a trophic paternal-relationship in the
frog Rhinoderma darwinii. Journal of Herpetology
20(2): 168178.
Harting. 1879. Rhinoderma darwini- Album der
Natur. Haarlem (Venesch Bijland) 78.
Howes, B.G. 1888. Notes on the gular brood-pouch
of Rhinoderma darwini. Proc Zool Soc 231-237.
Mendelson, J.R. III, E.D. Brodie Jr., J.H. Malone,
M.E. Acevedo, M.A. Baker, N.J. Smatresk, and
J.A. Campbell. 2004. Factors associated with the
catastrophic decline of a cloudforest frog fauna
in Guatemala. Revista de Biologia Tropical 52(4):
9911000.
Ottmar, W.G. 1927. La Rhinoderma darwinii D. y B.
Boletin de la sociedad de Biologia de Concepcin Ao
1, Nos. 1 and 2: 431.
Ottmar, W.G. 1932. Nuevas onservaciones acerca de
la neomelia de la Rhinoderma darwinii. Extraxto de
la Revista Chilena de Historia Natural XXXVI (1932):
166170.
Puumer, K. 1q. Observaciones biolgicas acerca
de la Rhinoderma darwini D & B. Revista Chilena de
Historia Natural 39: 28-30.
Philippi, R.A. 1902. Suplemento a los Batraquios
Chilenos Descritos en la Historia Fisica I Piltica de
Chile de don Claudio Gay. Santiago de Chile: Librera
Alemana de Jose Ivens.
Pounds, J. A., A.C. Carnaval, R. Puschendorf, C.F.B.
Haddad, and K.L. Masters. 2006. Responding
to amphibian loss. Science 314 (8 December
2006):15411542.
Rabanal F.E. and J. Nuez. 2008. $QELRVGH
los Bosque Templados de Chile. First Edition.
Universidad Austral de Chile, Valdivia , Chile. 206 pp.
Torres, D. and H. Castillo. 1973. Notas sobre
ODGLVWULEXFLyQJHRJUiFDGHOVDSLWRYDTXHUR
Rhinoderma darwini Dum y Bibr 1841 (Amphibia,
Anura, Dendrobatidae). Boletn del Museo Nacional
de Historia Natural (Chile) 203-204: 7-9.
DARWIN S FROG 17