Developing a Captive Breeding Protocol for Georgia`s Blind

Haideotriton wallacei. Almost nothing is known of the reproductive biology of this species nor has any VWXG\ FODUL¿HG SRSXODWLRQ HFRORJ\ EHKDYLRU JHQHWLF
similarities/differences between known populations, etc. In fact, the limits of its range are not even clearly GH¿QHGRZLQJWRWKHLQDFFHVVLELOLW\RIWKHKDELWDWZLWKLQ
which the salamander lives (the Floridan Aquifer). Because these salamanders inhabit groundwater, any study aiming to clarify their biology and ecology will be a challenge.
Developing a Captive Breeding Protocol for
Georgia’s Blind Salamander (Haideotriton
wallacei) at the Atlanta Botanical Garden
A n Up d ate o n T WI Gr ant No. 01
B Y D A N T É F E N O L I O, R O N A L D B O N E T T A N D M AT T H E W N I E M I L L E R
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adaptations in morphology, ecology, and behavior that often demonstrate unique and peculiar phenotypes. Salamanders have successfully exploited the extreme environments of subterranean waters that lack light and often have very low food and nutrient resources. Scientists are just beginning to unravel the complexities in biodiversity, adaptation, and population ecology of groundwater inhabiting salamanders (termed VW\JRELWLFVDODPDQGHUV,QWHQVL¿HGFDYHH[SORUDWLRQ
coupled with genetic analyses has produced a rapidly expanding list of caudates that are restricted to, and adapted to, subterranean habitats;; at least 11 North American species from four different lineages exist (Chippindale et al., 2000;; Wiens et al., 2003;; Bonett and Chippindale, 2004;; Niemiller et al., 2008) and more are on the way (Bonett et al., in prep.). One convergent characteristic in all but one or two of the North American species is paedomorphosis, whereby salamanders maintain a larval morphology and achieve reproductive maturity in this condition, all the while living in persistent bodies of water (i.e., groundwater). This strategy may be advantageous when aquatic environments are more productive, or terrestrial habitats are inhospitable or nonexistent (Wilbur and Collins, 1973;; Sprules, 1974;; Bruce, 1976;; Bonett and Chippindale, 2006;; Niemiller et al., 2008). However, very little is known about the life history strategies or population ecology of any subterranean salamander. We may lose some of the biodiversity inhabiting groundwater before we have a chance to understand the biology and ecology of these fascinating fauna. Human activities on the surface and within recharge zones of aquifers can degrade groundwater quality, threatening future human use of the resource as well as subsurface ecosystems that exist there (Crunkilton, 1982, 1984;; Tercafs, 1992;; Simon and Buikema, 1997;; Wood et al., 2002;; Graening and Brown, 2003). Agricultural pollution and industrial runoff have been documented to pose serious risks to groundwater quality (Crunkilton, 1982, 1984;; Gunn et al., 2000;; Culver et al., 2000;; Wood et al., 2002). In fact, the risk of groundwater contamination in the U.S. is greatest in agricultural areas where, ironically, an estimated 95% of local residents rely directly on the resource for their freshwater needs (USGS, 2005). Groundwater communities often include amphibians and invertebrates that are sensitive to contaminants in the environment, particularly fertilizers and pesticides (e.g., Cole and Casida, 1983;; Crunkilton, 1982, 1984;; Hecnar, 1995;; Sparling et al., 2001;; de Wijer et al., 2004;; Relyea 2005a, 2005b, 2005c). From a wildlife management perspective, groundwater contamination is a serious problem;; because so many groundwater species are found in a single aquifer system, one contamination event can pose an extinction threat. For example, in 1981 a liquid ammonia nitrate and urea fertilizer spill and subsequent contamination of Ozark groundwater (in Missouri), demonstrated the seriousness of such events. Roughly 21 km from the spill site, living and dead groundwater fauna ranging IURP EOLQG FDYH¿VK WR JURXQGZDWHU FUD\¿VK WR FDYH
salamander larvae washed out of a spring fed by the aquifer (Crunkilton, 1982, 1984). Because an aquifer is the entire available habitat for groundwater fauna, any change in water quality, and especially contamination of it, poisons the environment without anywhere for fauna to seek refuge.
Over-­harvest of groundwater also threatens the communities of organisms living in the habitat. There are some points to consider that demonstrate groundwater fauna as exceptionally vulnerable to human activities, including excessive water extraction. For example, subsurface habitats typically display decreased diversity in community complexity and reduced species abundance relative to above ground Perhaps the most enigmatic of the stygobitic ecosystems;; this means there are fewer species in salamanders is the Georgia Blind Salamander, subterranean habitats and fewer individuals than in HAIDE OTR I T O N WA LLA C E I 4 1
related species living in surface habitats (Holsinger, 1988). Processes that isolate subterranean populations of organisms, and evolutionary adaptation of those species to their environments, can produce extreme patterns of endemism (Barr and Holsinger, 1985;; Culver et al., 2000). For example, the combined ranges of over 50% of the described species and subspecies of groundwater dwelling fauna in the continental United States are estimated to constitute less than 1% of the total surface area of that region (Culver et al., 2000). Within the United States, subterranean fauna constitute more than 50% of the imperiled (G1–G2) species recorded in the Natural Heritage Program;; however, less than 4% have received federal protection (Culver et al, 2000). These conditions in subterranean species render them vulnerable to human activities. For example, it is believed that no fewer than 10 species of troglobites and stygobites have gone extinct owing to anthropogenic activities (Elliott, 2000).
Haideotriton wallacei
The situation for the Georgia Blind Salamander and other species that inhabit the Floridan Aquifer is no different. Despite the fact that the Floridan Aquifer has been designated as an at risk aquifer for fertilizer contamination by the United States Geological Survey (Nolan et al., 1998), no regular activities are in place by any agency or institution to monitor the populations of endemic and imperiled wildlife within the aquifer.
To begin to understand and monitor the populations of groundwater organisms inhabiting the Floridan Aquifer, a group of biologists has teamed up to study the biology and conservation status of the Georgia Blind Salamander and one of its fellow aquifer inhabitants, WKH 'RXJKHUW\ 3ODLQ &DYH &UD\¿VK (Cambarus cryptodytes). Both in-­situ and ex-­situ studies are planned. Further, Tree Walker International and a private donor have supported the work through a research grant to help establish a basic lab wherein live specimens are being maintained in captivity at the Atlanta Botanical Garden. Cambarus cryptodytes
4 2 L( $) /,7 7 ( 5 ‡ 9 2 / , 6 68 ( Using the TWI grant, a laboratory has been developed at the Atlanta Botanical Garden to study the reproductive biology and initiate a breeding colony for both the Georgia Blind Salamander and Dougherty Plain Cave &UD\¿VK7KHGHVLJQRIWKHV\VWHPVKDVEHHQPRGHOHG
after the systems used by the United States Fish and Wildlife Service to breed the Texas Blind Salamander (Eurycea rathbuni) and the Austin Blind Salamander (E. waterlooensis), which is bred in captivity by the City of Austin. Our system is composed of a series RI DTXDULD WKDW DUH SOXPEHG LQWR D FHQWUDO ¿OWUDWLRQ
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from one of the localities where the salamanders and FUD\¿VK KDYH EHHQ FROOHFWHG 7KH DSSURDFK LQFOXGHV
using local limestone so that any trace minerals that may be present in the limestone are also present in the water of the system. A chiller maintains the water temperature at 68°F, the groundwater temperature in the aquifer where these species are found. Capacity IRUJURXSVRIVDODPDQGHUVDQGFUD\¿VKRIHDFK
are present;; although much smaller groups have been collected to start the project. The system is maintained in complete darkness. Live specimens of both the Georgia Blind Salamander and the Dougherty Plain &DYH &UD\¿VK KDYH EHHQ FROOHFWHG IURP )ORULGD DQG
Georgia. These animals are settling into their captive conditions nicely. Importantly, this project would not be possible without the assistance of a number of cave divers, four of which have participated so far (Kelly Jessop, Ben Martinez, Mike Stine and Bonnie Stine).
The Georgia Blind Salamander, and all the inhabitants of the Floridan Aquifer, face daunting environmental challenges. As water quality deteriorates in this aquifer, the species that call it home are likely to disappear. Taking steps now to develop captive breeding protocols for some of these imperiled species will enable the rapid establishment of assurance colonies in the future, should they be necessary. Further, learning about the reproductive biology of the salamander DQGWKHFUD\¿VKZLOOH[SDQGXSRQZKDWOLWWOHZHNQRZ
of these “extremophiles.” Immediate goals of this project include developing a protocol for the captive PDQDJHPHQW RI WKH FUD\¿VK DQG WKH VDODPDQGHU DV
well as learning more about the reproductive output and behavior of both species. Future goals include learning more about the range of both aquifer inhabitants, learning how different populations are similar or different based on genetic analysis, studying the population ecology of these unique creatures, and determining if the salamander is suffering from emergent infectious amphibian diseases. The long-­
term conservation of imperiled groundwater fauna, starting with the Georgia Blind Salamander and the 'RXJKHUW\3ODLQ&DYH&UD\¿VKLVRXUNH\JRDO
HAIDE OTR I T O N WA LLA C E I 4 3
Acknowledgements: We would like to thank TWI and Lee Moran for funding assistance. We are indebted to several cave divers who have assisted us in acquisition of live animals (Kelly Kessop, Ben Martinez, Mike Stine, Bonnie Stine). We appreciate the time and assistance of the Georgia Department of Natural Resources (permit No. 29-­WBH-­11-­79) and the Florida Fish and Wildlife Conservation Commission (permit No. LSSC-­09-­0288). Valuable time and assistance has been provided from Atlanta Botanical Garden’s amphibian specialist, Robert Hill, and amphibian conservation coordinator, Mark Mandica. We are also thankful for the valuable time and suggestions regarding this project from W.W. Lamar and Dr. S. Opsahl.
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ďƌĞĞĚŝŶŐƚŚĞdĞdžĂƐůŝŶĚ^ĂůĂŵĂŶĚĞƌ͕Eurycea rathbuni͕ĂŶĚƚŚĞƉƌŽŐƌĂŵƌƵŶďLJƚŚĞĐŝƚLJŽĨƵƐƟŶ͕dĞdžĂƐ͕ĨŽƌƚŚĞ
ƵƐƟŶůŝŶĚ^ĂůĂŵĂŶĚĞƌ͕E. waterlooensis.
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