How Did the Mountain Chicken Frog Get to the Caribbean?

How Did the Mountain Chicken Frog Get to the Caribbean?

Vol. 6(11), pp. 754-764, November 2014 DOI: DOI: 10.5897/IJBC2011.075 Article Number: 10FEB8D48641 International Journal of Biodiversity ISSN 2141-243X Copyright © 2014 and Conservation Author(s) retain the copyright of this article http://www.academicjournals.org/IJBC Full Length Research Paper Conservation and historical biogeography: How did the mountain chicken frog get to the Caribbean? Jay D. King1* and Pamela C. Ashmore2 1Rare Species Conservatory Foundation, Loxahatchee, Florida, United States. 2Department of Sociology, Anthropology and Geography, University of Tennessee, Chattanooga, United States. Received 15 March, 2011; Accepted 3 November, 2014 Leptodactylus fallax, commonly known as the mountain chicken frog, is a large terrestrial frog currently found on two islands in the Caribbean. Habitat destruction, overhunting and disease outbreaks have contributed to declining population numbers. In order to identify appropriate conservation strategies, the historic geographic distribution of this frog must first be determined. Because no archeological evidence exists, this was accomplished by reviewing historical documents and inspecting museum collections. Inaccuracies in location and species names were identified in documents as well as in the mislabeling of museum specimens. Two means for natural immigration (dispersal and vicariance) and the artificial introduction by humans were considered. The authors concluded that the Amerindians transported L. fallax to eight islands throughout the Lesser Antilles as potential food resources as they colonized this area. The implication that 75% of the historical distribution is currently unoccupied by this species is considered in light of future reintroduction projects. Key words: Leptodactylus fallax, Amerindians, dispersal, vicariance. INTRODUCTION Wildlife biologists, zoo and aquarium personnel, and species that could benefit from this process is Leptodactylus conservationists are often asked to “save a species” fallax, commonly known as the mountain chicken frog sometimes in a specific location. But should they? Important (Figure 1). It is a large terrestrial frog (SNV 121 to 167.2 considerations need to be addressed before answering mm) (Heyer, 1979) with a unique form of reproduction this question. Is the location part of the historic geographic called obligatory oophagy (Gibson and Buley, 2004): a range? How and when did the species arrive? Is it an female lays eggs in a foam nest in which tadpole develop- endemic or an exotic? ment occurs while the female produces infertile eggs as a Historical biogeography is the study of the geographic source of nutrient for the young. distribution of an organism and how that distribution Currently this species is found on the islands of the occurred (Crisci et al., 2003). Understanding the past and Commonwealth of Dominica and Montserrat, but histori- present range of a species is necessary to appropriately cally may have been found throughout the Lesser Antilles, identify and implement viable conservation strategies. One the southeastern arm of the archipelago of the Caribbean *Corresponding author. E-mail: [email protected]. Tel: 314-664-2368. Fax: 314-644-0548. Author(s) agree that this article remain permanently open access under the terms of the Creative Commons Attribution License 4.0 International License King and Ashmore 755 Figure 1. The mountain chicken frog (L. fallax). Figure 2. Historic distribution of Leptodactylus fallax. Islands (Figure 2). Conservation organizations have create ancillary populations as a “safety net” to catastrophic expressed interest in reintroducing this species to several events: the population inhabiting the island of Montserrat islands in order to re-establish its former range and is threatened by volcanic activity and populations found on 756 Int. J. Biodivers. Conserv. this and the island of the Commonwealth of Dominica are delicate creatures, extremely susceptible to salt water, to desic- being decimated by the pathogenic chytrid fungus, cation, and to the heat of the sun, and since they do not possess Batrachochytrium dendrobatidis. However, it must be wings, it is difficult to imagine how they could cross sea barriers.” There are two possible modes of natural immigration: vicariance determined which islands actually composed the historical and dispersal. Vicariance is the process by which one population is range of L. fallax before reintroduction should be divided into two by the formation of a barrier, such as the rising of a considered. mountain range or the flooding of a plain. Dispersal is the process A number of factors contribute to the puzzle of recon- by which members of a population cross a pre-existing barrier, structing the historical distribution of L. fallax. First, there sequentially becoming an isolated population. The arrival of L. fallax on islands in the Caribbean was further assessed by investigating appears to be no archaeological evidence, such as skeletal known incidents of rafting and studies of ocean currents. In addition, remains. As a result, we are forced to rely entirely on the more unusual modes of dispersal were noted in the historical literature. historical literature and preserved specimens. Second, there have been inconsistencies in the taxonomic nomen- clature that has been assigned to this species (see RESULTS “Museum Specimen Search”). Third, preserved specimens have been mislabeled or lost, as will be documented The historical distribution of L. fallax by island in a below. And finally, several important historical references north to south distribution describing the geographic distribution of the frog are in French and mistranslations have been done. This reconstruction is based on accounts from the historical literature, inventories of museum collections, and in some cases a review of the specimens included in the collections. METHODOLOGY Literature search Jamaica A comprehensive literature search was conducted for the keywords “Leptodactylus” and “L. fallax”. All relevant literature was obtained The Jamaica Journal tells the story of a woman bringing and pertinent references found in this literature were also gathered. a pair of L. fallax to this island and the successful breeding Often these consisted of field notes from early explorers and many at two different sites (Proctor, 1973). This journal does not of these accounts were in French. The original French sources have a scientific focus and normally contains essays and were then translated and compared with earlier translations to poems, so it is likely that this was a fictional account. identify if any inaccuracies existed which are relevant to the geographic distribution of this frog species. The information in these sources Additionally, a West Indies herpetologist confirmed no past was used to piece together the historical biogeography of L. fallax. or present reports of this species in Jamaica (Personal communication – Jay King). It is unlikely that L. fallax occurred on this island. Museum specimen search An index to herpetology collections in the United States can be Dominican Republic found on the Internet (California Academy of Sciences, 2014). This source lists eight institutions holding specimens of L. fallax (Table Lescure (1979) states that there is a translation error in 1). The web page for each institution was reviewed for information the word Dominica in the book “Amphibiens vivants du regarding number of individual specimens, location and date collected. In two cases, data were not available on-line and monde (World living amphibians)”. It states that L. fallax institutional personnel were able to supply collection records. One was a species found in the Dominican Republique, instead institution, the University of Kansas, listed ten specimens from the of on the island of the Commonwealth of Dominica. With “Republica Dominicana” however there are no historical accounts of the exception of a listing of L. fallax specimens at the this species ever occurring in the Dominican Republic. University of Kansas from the Dominican Republic, no other Consequently, field notes were obtained, the collection was visited, accounts can be found. It was determined that these and the specimens examined. specimens were actually L. albilarbris (Personal obser- vation – Jay King). It is unlikely that L. fallax occurred on Biogeographical reconstruction this island. Attempting to reconstruct the historical distribution of L. fallax requires consideration of how this species first occurred on islands Puerto Rico in the Caribbean. Our own training and field experiences and a review of relevant literature on island immigration enabled us to Attempts were made in 1929 and 1932 to introduce L. determine possible means for the arrival of the frog on the islands. fallax to this island, but both failed. Barbour (1937) states, Groome (1970) appropriately describes the challenge: “In general, “The imported population which was taken while calling at Amphiba are absent on oceanic islands, for their eggs and tadpoles require fresh water and their skins are totally allergic to salt.” night from Dominica, may be males only, according to Similarly, Meyer (1953) identifies the improbability of a natural Major Chapman Grants.” Other than the brief unsuccessful immigration of the frogs to the islands: “Amphibians as a group are attempts to introduce this species, it is unlikely that this King and Ashmore 757 Table 1. Institutions with collections of preserved specimens of L. fallax. No. of Institution Collection

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