Parasites and Disease-causing Organisms Reported from Wisconsin Amphibians and Reptiles Bureau of Science Services Wisconsin Department of Natural Resources P.O. Box 7921 Madison, WI 53707-7921 Miscellaneous Publication PUB-SS-1145 2014 ______________________________________________________ Summary: This report compiles and summarizes published information on parasites and disease-causing organisms reported from Wisconsin amphibians and reptiles in an accessible general reference for use by wildlife health and conservation practitioners. The catalog consists of two primary tables that summarize the available literature. The first lists parasite species by taxonomic group and provides a host list for each parasite. The second provides a checklist of Wisconsin amphibians and reptiles along with their reported parasites. A short “Discussion” section presents observations and summary conclusions derived from a cursory analysis of the records contained in these tables. Investigators have looked at a wide range of parasites affecting Wisconsin amphibians and reptiles: at least six bacteria, three fungi, 15 protoctists, 46 trematodes, five monogeneans, four cestodes, 22 nematodes, two acanthocephalans, three leeches, one mollusk, and 11 flies. The parasites of amphibians have been treated more frequently than those of reptiles. No papers address the parasites of 21 Wisconsin species, demonstrating that there is much yet to be learned in our area. The effects of most parasite species on the health and reproduction of their hosts remain largely uninvestigated. This report provides a foundation for documenting the composition of the regional parasite fauna and can help biologists understand changes brought on by environmental variation and the introduction and spread of invasive species. Useful references are identified as starting points for future efforts. Cover Illustrations: Various parasitic organisms. Source: Arthur’s Free Clipart, www.arthursclipart.org. The Wisconsin Department of Natural Resources provides equal opportunity in its employment, programs, services, and functions under an Affirmative Action Plan. If you have any questions regarding this plan, please write to Equal Opportunity Office, Department of Interior, Washington, DC 20240. This publication is available in alternative format (large print, Braille, audio tape, etc.) upon request. Please call (608) 266-0531 for more information. Amphibian and Reptile Parasites Parasites and Disease-causing Organisms Reported from Wisconsin Amphibians and Reptiles Dreux J. Watermolen Bureau of Science Services Wisconsin Department of Natural Resources P.O. Box 7921 Madison, WI 53701-7921 December 2014 i Amphibian and Reptile Parasites ii Amphibian and Reptile Parasites Contents Introduction 1 Methods 2 Structure of the Catalog 3 Catalog of Parasites and Disease-causing Organisms 5 Table 1. Parasites and disease-causing organisms reported from 5 amphibians and reptiles in Wisconsin Table 2. Wisconsin amphibian and reptile species and their parasites 25 Discussion 37 Table 3. Amphibian and reptile species for which no published 38 parasite records were found Literature Cited 43 Appendix A 57 iii Amphibian and Reptile Parasites “They are, people will assure you, the hangnail on life’s little toe – insignificant uglies and annoying to boot.” “Yet such organisms are parts of that great and ponderous machine we call life.” - Robert Michael Pyle (1989) iv Amphibian and Reptile Parasites Introduction Parasites and disease-causing organisms are a growing concern among conservationists and wildlife health professionals. In particular, the potentially deleterious impacts of widespread infectious diseases on amphibian and reptile populations have garnered increasing attention (Green et al. 2002, Carey et al. 2003, Origgi 2006, Lee 2013, Duffus et al. 2014, Martel et al. 2014, Price et al. 2014). A report of the “largest mass die-off of amphibians ever recorded in the academic literature” and linked to ranavirus recently appeared in print (Wheelright et al. 2014), capturing the interest of both the scientific community and general public (Earl and Gray 2014, Whittle 2014). Further, Brenes et al. (2014) have shown that this pathogen can be transmitted between fish, amphibians, and reptiles, thus contributing to ranavirus’ persistence in the environment, potential spread, and broader impact. Chinchar and Waltzek (2014) have identified this organism as an emerging threat to aquaculture. The impact of chytridiomycosis on frog populations has also been reported widely (Skerratt et al. 2007, James et al. 2009, Vredenburg et al. 2010), and Martel et al. (2014) recently described Batrachochytrium salamandrivorans’ effect on salamander populations. Similarly, the U.S. Geological Survey’s National Wildlife Health Center has confirmed snake fungal disease in snakes in several eastern and Midwestern states, including Wisconsin, raising concerns that it could pose a threat to snakes similar to what white-nose syndrome does in bats (Miller 2013, Sleeman 2013). Many host-parasite relationships appear to be relatively innocuous, with some individual hosts harboring large numbers of microorganisms and worms of several different species.1 In some cases, however, parasites may present disease threats to the animals they infect or could infect, a particular concern for endangered species and small, isolated populations. For example, trematode parasites (Ribeiroia sp.) have been implicated as a cause of deformities and mortality, and possibly declines, in some frog populations (Johnson et al. 2002, Johnson and Sutherland 2003, Johnson et al. 2004). The nematode Serpinema microcephalus has been associated with pancreatitis in red-eared sliders (Trachemys scripta elegans) (Wieczorowski 1939, Hidalgo-Vila et al. 2011), raising concerns about the spread of this potentially invasive host species into new habitats. Pentastomes have been found to infect some rare and endangered species, with infections sometimes leading to death (Riley 1986). Myiasis (infestations by fly larvae that grow inside the host while feeding on its tissue) leading to death is believed to be more widespread in frogs than previously reported (Bolek and Janovy 2004, Wolff et al. 2013). In addition, the presence of some parasites may predispose their host to other ailments either by a reduction in general vitality resulting in increased susceptibility or through the introduction of other disease-causing organisms. Parasites and zoonotic diseases may also pose human health concerns. For example, Harris et al. (2009) have referred to reptile-associated Salmonella infections in humans as a significant public health problem. Furthermore, Eckner et al. (2011) have shown that lizards may serve as reservoirs of Borrelia burgdorferi, the causative agent of Lyme disease. Other potential pathogens carried by reptiles include the bacteria Aeromonas, Campylobacter, and Mycobacterium, and various zygomycetes and protozoans. The effects of parasites may interact in complex ways with environmental conditions and other stressors (e.g., see Johnson and Chase 2004, Johnson et al. 2007, Kerby and Storfer 2009, Hartson et al. 2011, and Kerby et al. 2011). The observed relationships between environmental contaminants and parasites have led to the occasional use of parasitological data in environmental monitoring (Overstreet 1997, Marcogliese and Giamberini 2013). 1 It is important to acknowledge that very few individual animals of any species are completely free of parasitic organisms. 1 Amphibian and Reptile Parasites Our ability to manage proactively for concerns such as these has been somewhat limited due to a lack of basic information on the parasites and disease-causing organisms affecting our state’s fauna. Knowledge of epidemiology and control techniques appears to be of growing importance, but the literature regarding the parasites of amphibians and reptiles remains widely scattered throughout a variety of sources, often crossing disciplines. Titles of many articles do not indicate that the parasites of Wisconsin species were studied (e.g., Goodchild and Dennis 1967); occasionally, the titles may even suggest otherwise (e.g., Bolek et al. 2009b). Sometimes, particularly in older references, authors have failed to indicate the source of their study organisms or the location where their work was performed. As a result, attempts to find information specific to Wisconsin species can pose significant challenges. Even with the availability of modern electronic indexing and abstracting services, presenting a basic host-parasite species list for the state has proven to be a difficult task. The objective of this report, therefore, is to compile and summarize the published information on parasites and disease-causing organisms of Wisconsin amphibians and reptiles in an accessible general reference for use by wildlife health and conservation practitioners. Similar comprehensive catalogs have been prepared for only one other state in our region: Michigan (Muzzall 2005). Together these “snapshots” provide a starting point for characterizing and quantifying the regional richness and diversity of parasite communities across the host species’ geographic ranges. Methods Parasite records were gleaned from a variety of sources including bibliographies focused on Wisconsin herpetology (Dlutkowski et al. 1987, Watermolen 1991, Watermolen 2013). I searched for Wisconsin records in standard references addressing the various parasite taxa (Bychowsky 1957, Petrochenko 1971, Yamaguti 1971, Schell 1985, Riley 1986, Levine 1988, Barta 1991, Anderson 1992, Walters et al. 2011), as well as references specific to those species parasitic in amphibians and reptiles (Reiber et al. 1940; Hughs, Baker and Dawson 1941a, 1941b, and 1942; Hughes, Higginbotham, and Clary 1941, 1942a, and 1942b; Nigrelli 1945; Bardsley and Harmsen 1973; Ernst and Ernst 1977, 1979, and 2006; Prudhoe and Bray 1982; Baker 1987; Dyer 1991; Telford 1995; McAlpine 1996; Kuzman et al. 2003; Duszynski et al. 2007; and Fajfer 2012). I also searched various electronic databases (Biological Abstracts, Zoological Record, Web of Science, etc.) and Google Scholar for the names of Wisconsin biologists and various combinations of taxonomic and subject terms to identify titles that might cover Wisconsin or involve Wisconsin specimens. I then reviewed the works cited in each source found for additional likely references. Search terms covered a wide range of parasite taxa. No attempt, however, was made to identify studies addressing viruses, and although several bacteria, fungi, and protoctists are included in the catalog, only limited efforts were made to investigate these groups. Greater focus was placed on helminths and leeches, taxa with which I am more familiar. No effort was made to investigate the freshwater mollusk (Simpsonaias ambigua) larvae that parasitize mudpuppies (and possibly other salamanders), although a record is included as a result of other work. Records of arthropod ectoparasites were noted when they were encountered, but as with mollusks were not a focus of the effort. I made no efforts to sift through the “gray” literature, but included agency reports when I stumbled upon them in the course of my other searching. Several academic theses are included, but I did not tabulate the records contained in these documents as those records 2 Amphibian and Reptile Parasites were generally published in other papers that are cited herein. I applied a similar approach to the abstracts that appear in the annual reports of the UW-Milwaukee Field Station. The result of these efforts is an extensive catalog of parasite records from the state published through the end of 2014, including several missed during previous compilations and reviews. In spite of best efforts, however, I have undoubtedly missed some references with Wisconsin information and will appreciate learning about additional records. Every attempt has been made to ensure the accuracy of the catalog entries and literature citations, but it is nonetheless conceivable that errors may have crept into the work. For these, I take responsibility. Future workers should check the original sources to avoid perpetuating any such errors. Structure of the Catalog The catalog consists of two tables that summarize the available literature. Table 1 (pages 5-24) lists parasite species by taxonomic group and provides a host list for each parasite. Parasites are listed to the lowest taxonomic unit possible and nomenclature appears here as it did in the original work, with only minimal attempts to address the past complexities of parasite taxonomy and the resulting synonyms. In cases where I was able to update parasite nomenclature, the species name as reported in the original source(s) usually follows the updated name in parentheses. It should be recognized, however, that review of the validity of the published names was well beyond the scope of current efforts. Parasite species are listed alphabetically by family and genera. The bewildering diversity of protoctists has led to continuous debate about organismal names and classification schemes. I have generally followed the classification presented by Margulis et al. (1990), with a few minor revisions based on information contained in the online Encyclopedia of Life resource 2. Parasite location within/on the host (anatomical habitat) is indicated in parentheses when this information has been included in the sources. Each entry includes a citation to the relevant literature source(s). Entries in Table 1 are presented in the following format: Parasite Species (Name in Original Source, if Updated by Me) Host Species (Anatomical Habitat) – Literature Citation(s) Since this is not a taxonomic work and for the sake of simplicity, I omitted the author and year of description associated with genus-species names for both parasites and hosts. These are generally available in the cited references. Table 2 (pages 25-36) provides a checklist of Wisconsin amphibians and reptiles along with their reported parasites. Amphibian and reptile nomenclature follows Casper and Anton (2013), with species names listed alphabetically by order. Although included in Table 1, nonnative host species reported from captive settings in the state are omitted from this table. County of occurrence is provided when this information has been included in the reference. As in Table 1, each entry is followed by a citation to the relevant literature source(s). 2 http://eol.org/ 3 Amphibian and Reptile Parasites Entries in Table 2 are presented in the following format: Host Species (Host Common Name) Taxa Group: Parasite Species (Geographic Location) – Literature Citation(s) In sum, Table 1 lists parasites and their hosts and Table 2 lists hosts and their parasites. A short “Discussion” section follows these tables and presents observations and summary conclusions derived from a cursory analysis of the records contained in the tables. Table 3 in this section (page 38) lists host species for which I was unable to find any published parasite records from Wisconsin. All references cited in the tables are listed together in the “Literature Cited” section that follows the discussion. Finally, Appendix A concludes the catalog by providing a list of references that specifically address parasites of Wisconsin amphibians and reptiles. Notes 4