Geographic Distribution: Oxyrhopus melanogenys orientalis

Igor J Roberto

Abstract. A new species of the gekkonid genus Cnemaspis is described from Pulau Pemanggil, Johor, West Malaysia on the basis of its unique combination of color pattern, scale characteristics, and snout–vent length. It resembles the insular endemic C. baueri from the adjacent islands of Aur and Dayang. This species is the only known reptile endemic to Pulau Pemanggil. Key Words. Cnemaspis; New Species; Pulau Pemanggil; Seribuat Archipelago; West Malaysia; Gekkonidae.

The agamid genus Harpesaurus is poorly understood, partially due to their scarcity in collections. Two taxa have been described from Borneo, originally assigned to two different genera: Harpesaurus [Hylagama] borneensis and Harpesaurus thescelorhinos. Today thescelorhinos is generally considered a junior synonym to borneensis. We here report on three additional specimens collected in Sarawak in 2005. While the two new females had some scale characters distinctly different from the two females previously known, we consider all known specimens from Borneo conspecific. Most notably, the two new females were soon to give live birth. This constitutes the first known example of live birth in agamid lizards from a tropical lowland area. We discuss why this unusual reproductive mode may have evolved in this particular taxon.

Lata Bukit Hijau is located within the Banjaran Bintang Ranges on the west coast of northern Peninsular Malaysia. The reptile fauna in this pristine area was intensively investigated from 2008 to 2011 on 10 consecutive visits. A total 37 species of reptiles from 31 genera and 10 families were recorded to inhabit this area. Out of this number, 17 species were lizards (13 genera and four families), 17 species were snakes (15 genera and four families) and three species were freshwater turtles (three genera and two families). These preliminary data increased the number of lizards, snakes and freshwater turtles reported from Banjaran Bintang from 31 to 41, 30 to 44 and three to five species, respectively.

Herpetological Review Volume 40, Number 3 — September 2009 2008 SSAR Ofﬁcers (2009) President BRIAN CROTHER Department of Biological Sciences Southeastern Louisiana University Hammond, Louisiana 70402, USA President-elect JOSEPH MENDLELSON, III Zoo Atlanta 800 Cherokee Avenue, SE Atlanta, Georgia 30315, USA Secretary MARION R. PREEST Joint Science Department The Claremont Colleges Claremont, California 91711, USA Treasurer KIRSTEN E. NICHOLSON Department of Biology, Brooks 217 Central Michigan University Mt. Pleasant, Michigan 48859, USA e-mail: kirsten.nicholson@cmich.edu Publications Secretary BRECK BARTHOLOMEW P.O. Box 58517 Salt Lake City, Utah 84158, USA e-mail: ssar@herplit.com Immediate Past President ROY MCDIARMID USGS Patuxent Wildlife Research Center Smithsonian Institution P.O. Box 37012 Washington, DC 20113-7012, USA Directors PAUL CHIPPINDALE (2010) TIFFANY DOAN (2010) TRAVIS LADUC (2010) STEPHEN RICHTER (2010) DAVID CUNDALL (2012) KEVIN DE QUEIROZ (2012) PATRICK GREGORY (2012) ANN PATERSON (2012) Trustee GEORGE R. PISANI University of Kansas, USA HERPETOLOGICAL REVIEW The Quarterly News-Journal of the Society for the Study of Amphibians and Reptiles Editor ROBERT W. HANSEN 16333 Deer Path Lane Clovis, California 93619-9735, USA HerpReview@gmail.com Associate Editors ROBERT E. ESPINOZA California State University, Northridge CHRISTOPHER A. PHILLIPS Illinois Natural History Survey DEANNA H. OLSON USDA Forestry Science Lab ROBERT N. REED USGS Fort Collins Science Center MICHAEL S. GRACE Florida Institute of Technology MARGARET S. GUNZBURGER Nokuse Plantation EMILY N. TAYLOR California Polytechnic State University GUNTHER KÖHLER Forschungsinstitut und Naturmuseum Senckenberg MEREDITH J. MAHONEY Illinois State Museum JESSE L. BRUNNER State University of New York at Syracuse Section Editors Book Reviews AARON M. BAUER Department of Biology Villanova University Villanova, Pennsylvania 19085, USA aaron.bauer@villanova.edu Current Research JOSHUA M. HALE Department of Sciences MuseumVictoria, GPO Box 666 Melbourne, Victoria 3001, Australia jhale@museum.vic.gov.au Current Research BEN LOWE Department of EEB University of Minnesota St Paul, Minnesota 55108, USA lowe0160@umn.edu Geographic Distribution ALAN M. RICHMOND Biology Department, Morrill IV South University of Massachusetts 611 North Pleasant Street Amherst, Massachusetts 01003-9297, USA alanr@bio.umass.edu Geographic Distribution INDRANEIL DAS Institute of Biodiversity & Environmental Conservation Universiti Malaysia Sarawak 94300, Kota Samarahan, Sarawak, Malaysia hamadryad2004@hotmail.com Geographic Distribution JERRY D. JOHNSON Department of Biological Sciences The University of Texas at El Paso El Paso, Texas 79968, USA jjohnson@utep.edu Geographic Distribution GUSTAVO J. SCROCCHI Instituto de Herpetología Fundación Miguel Lillo, Miguel Lillo 251 4000 Tucumán, Argentina soniak@unt.edu.ar Zoo View JAMES B. MURPHY Department of Herpetology National Zoological Park 3001 Connecticut Ave., NW Washington, D.C. 20008, USA jbmurphy2@juno.com Herpetological Husbandry BRAD LOCK Department of Herpetology Zoo Atlanta 800 Cherokee Ave., S.E. Atlanta, Georgia 30315, USA block@zooatlanta.org Natural History Notes CHARLES W. PAINTER New Mexico Dept. of Game & Fish P.O. Box 25112 Santa Fe, New Mexico 87504, USA charles.painter@state.nm.us Natural History Notes JAMES H. HARDING MSU Museum Michigan State University East Lansing, Michigan 48824, USA hardingj@msu.edu Natural History Notes JOHN D. WILLSON University of Georgia Savannah River Ecology Lab Drawer E Aiken, South Carolina 29802, USA willson@uga.edu Copy Editors BARBARA BANBURY RAUL DIAZ DANIEL PORTIK KYLE MILLER HESED Natural History Notes JACKSON D. SHEDD 533 Countryside Lane Chico, California 95973, USA Jackson.Shedd@gmail.com SSAR Editors Journal of Herpetology MATTHEW PARRIS, Editor Department of Biology University of Memphis Memphis, Tennessee 38152, USA Contributions to Herpetology KRAIG ADLER, Editor Department of Neurobiology & Behavior Cornell University Ithaca, New York 14853, USA Facsimile Reprints in Herpetology AARON M. BAUER, Editor Department of Biology Villanova University Villanova, Pennsylvania 19085, USA Herpetological Circulars JOHN J. MORIARTY, Editor 3261 Victoria Street Shoreview, Minnesota 55126, USA Catalogue of American Amphibians and Reptiles ANDREW H. PRICE, Editor Texas Natural History Collections The University of Texas at Austin Austin, Texas 78758-4445, USA SOCIETY FOR THE STUDY OF AMPHIBIANS AND REPTILES www.ssarherps.org The Society for the Study of Amphibians and Reptiles, the largest international herpetological society, is a not-for-proﬁt organization established to advance research, conservation, and education concerning amphibians and reptiles. Founded in 1958, SSAR is widely recognized today as having the most diverse society-sponsored program of services and publications for herpetologists. Membership is open to anyone with an interest in herpetology—professionals and serious amateurs alike—who wish to join with us to advance the goals of the Society. All members of the SSAR are entitled to vote by mail ballot for Society ofﬁcers, which allows overseas members to participate in determining the Society's activities; also, many international members attend the annual meetings and serve on editorial boards and committees. All members and institutions receive the Society’s primary technical publication, the Journal of Herpetology, and its newsjournal, Herpetological Review; both are published four times per year. Members also receive pre-publication discounts on other Society publications, which are advertised in Herpetological Review. To join SSAR or to renew your membership, please visit the secure online Allen Press website via this link: http://www.ssarherps.org/pages/membership.php Future Annual Meetings Herpetological Conservation JOSEPH C. MITCHELL, Editor Mitchell Ecological Research Services P.O. Box 5638 Gainesville, Florida 32627-5638, USA 2010 — Providence, Rhode Island, 7–12 July (with ASIH, HL) 2011 — Minneapolis, Minnesota, dates TBA (with ASIH, HL) 2012 — Vancouver, British Columbia, dates TBA (with ASIH, HL, WCH) About Our Cover: Gonocephalus robinsonii SSAR BUSINESS PHOTO BY HEOK HUI Of the 28 species of agamid lizards in Peninsular Malaysia and its associated archipelagos, only six occur in the genus Gonocephalus (Grismer 2008. Zootaxa 1860:28–34; Wood et al. 2009. Zootaxa 2012:28– 46). All Gonocephalus in Peninsular Malaysia are diurnal, arboreal species and most show signiﬁcant degrees of sexual dimorphism. Gonocephalus robinsonii is an arboreal, diurnal, upland species restricted to hill dipterocarp forests between 700 and 1700 m in elevation. This strange-looking lizard goes well with the mysterious lichen and moss-covered forests it inhabits. During the day, individuals are most commonly seen on the sides of both large and small tress, usually less than three meters above ground. Juveniles seem to be more active than adults and will ﬂee up the trunk of a tree at the slightest provocation, whereas adults generally tend to hold their position. At night, all age classes can be found sleeping on the branches of tree ferns as well as the vertical branches of smaller trees as high as 9 m above ground. Breeding generally coincides with the winter monsoons with hatchings appearing near the end of the season. Gonocephalus robinsonii occurs in at least two different isolated mountain ranges: the extensive Banjaran Titiwangsa in the west that runs nearly the entire length of the peninsula, and the much smaller Gunung Tahan in the east. The cover photo was taken by L. Lee Grismer, professor of biology at La Sierra University, Riverside, California. Grismer has been working intensively in Peninsular Malaysia since 2003 and in the last six years, his team has discovered and described over 25% of this region’s known lizard fauna, describing more species than had been identiﬁed during the previous century. They have also discovered several new species of frogs and snakes. Grismer’s latest book, The Lizards of Peninsular Malaysia, Singapore, and the Adjacent Archipelagos: Their Description, Distribution, and Natural History, is scheduled for publication in early 2010 and will be patterned after his earlier book on the herpetofauna of Baja California. The cover photo was recorded at Cameron Highlands, Pahang, with a Nikon D300 and a Sigma 14mm lens. The scene was metered to ambient light at ISO 200, and recorded using ﬁll ﬂash (Nikon SB20) set to -3EV at f22. According to Grismer, “What really made it work was after I began shooting, the clouds rolled in, surrounding the trees in mist, thus really adding a ‘cloud forest’ feel to the shot.” Journal of Herpetology Receives Award: Ranked Among Top 100 Journals in Biology and Medicine in the Past 100 Years In conjunction with the Special Libraries Association’s Centennial, the BioMedical & Life Sciences Division conducted a poll to identify the 100 most inﬂuential journals of Biology & Medicine worldwide over the last 100 years. The division was led in 2008 by Chair John Tebo, University of Cincinnati, and in 2009 by Chair Jean Crampon, University of Southern California. Three panels consisting of three experts each were recruited from the DBIO membership. Each panel composed a series of voter preference questions comparing journals in categories of biology or medicine that fell within its areas of expertise. The Journal of Herpetology was considered by the Natural History Panel and competed in the category of “Vertebrate Biology—Reptiles and Amphibians.” It surpassed some estimable competitors to make it into the top 100 journals. The other top journals recognized include Nature, Science, Ecology, Animal Behavior, and Proceedings of the Royal Society (Biology). SSAR was represented at a luncheon and awards celebration in Washington, DC on 16 June 2009 by Past-President, Roy McDiarmid, and Board Member, Kevin de Queiroz. SSAR is pleased to recognize the longstanding commitment SSAR President Brian Crother is ﬂanked by Matt Parris (Editor, Journal of Herpetology, left) and Kevin de Queiroz (SSAR Board Member, right), after sharing the Top 100 Journals award during the SSAR Board Meeting in Portland. Herpetological Review 40(3), 2009 257 258 Herpetological Review 40(3), 2009 to excellence by the editors of the Journal of Herpetology from its inception: Kraig Adler (1958–1963), James A. MacMahon (1964), Corson Jay Hirschfeld (1965–1967), J. P. Kennedy (1968–1979), Rodolfo Ruibal (1980–1989), Samuel S. Sweet (1990–1992), Richard A. Seigel (1993–2000), Brian K. Sullivan (2001–2005), Geoffrey R. Smith (2006–2008), Matthew J. Parris (2009–present). The award is also a credit to the associate editors, editorial board members, and hundreds of anonymous reviewers who have served over the 52-year existence of the journal. Consult the complete list of awardees at: The second award winner is Jeanine Refsnider. Her proposal deals with Painted Turtles (Chrysemys picta) and is designed to determine whether local adaptations to nest site choice are more genetically or environmentally driven (“Can maternal nestsite choice compensate for the effects of global climate change on reptiles with temperature-dependent sex determination? A common garden experiment using a model species”). She is working in Fred Janzen’s laboratory in the Department of Ecology, Evolution, and Organismal Biology at Iowa State University. http://units.sla.org/division/dbio/publications/resources/ dbio100.html NEWSNOTES 2008 Ofﬁcial Names List Now Available Online Free Panama Amphibian Rescue and Conservation Project The newest edition of Scientiﬁc and Standard English Names of Amphibians and Reptiles of North America North of Mexico, the ofﬁcial names list adopted by the American Society of Ichthyologists, The Herpetologists' League, and the Society for the Study of Amphibians and Reptiles, is now available as a free download from the SSAR website: Recognizing that Panama’s rich diversity of amphibians is an important natural treasure with signiﬁcant direct, cultural, biomedical, ecological and existence values that warrants protection from extinction, a group of organizations came together to respond to the crisis. Africam Safari Park (Mexico), Cheyenne Mountain Zoo (Colorado), Defenders of Wildlife (Washington DC), the Smithsonian National Zoological Park (Washington DC), the Smithsonian Tropical Research Institute (Panama), Zoo New England (Massachusetts) and Houston Zoo (Texas) have pooled their energy and resources, collectively pledging more than $750,000 in cash and in kind over the next three years to the Panama Amphibian Rescue and Conservation Project. The Project will consist of three distinct and complementary parts: 1) the ongoing operation of El Valle Amphibian Conservation Center (EVACC) in Western Panama, run by the Houston Zoo; 2) the amphibian chytrid cure research program to be initiated at the National Zoo in collaboration with Vanderbilt University; and 3) the construction and operation of the new Summit Park Amphibian Rescue Center in Panama. One “amphibian rescue pod” which is a biosecure, modiﬁed shipping container that will house the ﬁrst rescued species from Eastern Panama has already been established. Please ﬁnd out more about this project at: http://ssarherps.org/pages/HerpCommNames.php Kennedy Student Award 2009 The Kennedy Award Committee (Carl Anthony, Bill Lutterschmidt, Terry Schwaner, Lynnette Sievert, Chair) has completed its work for Volume 42 of the Journal of Herpetology. The Committee has selected “Preference for Local Mates in a Recently Diverged Population of the Lesser Earless Lizard (Holbrookia maculata) at White Sands” by Erica Bree Rosenblum (Vol. 42:572–583). The Kennedy Award carries with it a cash prize of US $200 or the winner’s selection of any SSAR publications valued at twice that amount. The committee invites all student members of the Society to submit their work to the Journal, and encourages members who supervise the work of students to draw this award to the attention of those students. http://www.amphibianrescue.org/ Dean E. Metter Memorial Award for 2009 There were 25 proposals submitted this year for the Dean E. Metter Memorial Award. The proposals were all of high quality; two stood out from the others. The committee requested permission from President Crother to make two awards this year and that request was granted. The ﬁrst award winner is Benjamin Jellen of Saint Louis University. He is a Ph.D. student in the Department of Biology working with Robert Aldridge. His proposal is entitled “Pre-and post-copulatory determinants of reproductive success in Missouri northern watersnakes (Nerodia sipedon).” The research attempts to show how these snakes communicate using pheromones and how this relates to male and female reproductive success. MEETINGS Meetings Calendar Meeting announcement information should be sent directly to the Editor (HerpReview@gmail.com) well in advance of the event. 2–4 October 2009—Symposium on Anolis Biology, Harvard University, Cambridge, Massachusetts, USA. Information: http:// anolis.oeb.harvard.edu. 17 October 2009—Texas Herpetological Society Fall Meeting and Lizard Symposium, Brazos Center, Bryan, Texas, USA. Information: http://www.thehibbitts.net/THS/. Herpetological Review 40(3), 2009 259 17–19 October 2009—Gaffney Chelonian Fossil Symposium, Drumheller, Alberta, Canada. Information: http://www.tyrrellmuseum.com/events/turtlesymp09.php 26–28 February 2010—35th Annual Desert Tortoise Council Symposium, Ontario, California, USA. Information: http://www. deserttortoise.org/symposia.html 22–24 April 2010—57th Annual Meeting, Southwestern Association of Naturalists, Llano River Field Station, Junction, Texas, USA. Information: http://www.biosurvey.ou.edu/swan/annualm10. html 7–12 July 2010—Joint Meeting of Ichthyologists and Herpetologists (ASIH / HL / SSAR), Providence, Rhode Island, USA. Information: http://www.dce.k-state.edu/conf/jointmeeting/ 21–24 July 2010—33rd Annual International Herpetological Symposium, Tucson, Arizona, USA. Information: http://www. kingsnake.com/ihs/ CURRENT RESEARCH The purpose of Current Research is to present brief summaries and citations for selected papers from journals other than those published by the American Society of Ichthyologists and Herpetologists, The Herpetologists’ League, and the Society for the Study of Amphibians and Reptiles. Limited space prohibits comprehensive coverage of the literature, but an effort will be made to cover a variety of taxa and topics. To ensure that the coverage is as broad and current as possible, authors are invited to send reprints to the Current Research section editors, Joshua Hale or Ben Lowe; postal and e-mail addresses may be found on the inside front cover. The current contents of various herpetological journals and other publications can be found at: http://www.herplit.com/contents. Impacts of Climate Change on Tropical Lizard Communities As tropical lizard taxa occurring in forested habitats are adapted to narrow temperature intervals and are functionally thermoconformers, slight elevations in mean temperature due to climate change may disproportionately affect them. To gain insight into this phenomenon, the authors ﬁrst accumulated body temperature, thermal maximum, minimum, and optimum, maximum latitude, habitat type (open or forested), thermoregulation method (basking or not), and phylogenetic relationship data for 66 species (determined in previous studies) and performed a meta-analysis in order to reveal correlations between the physiological traits and the ecological, behavioral, distributional, and phylogenetic information. Second, ﬁne-scale environmental and physiological data for diurnal Puerto Rican Sphaerodactylus and Anolis (four and two species, respectively) were used to determine the effect predicted climate change will have on lizards occupying different environments. The meta-analysis reveals that some tropical taxa indeed have low thermal maxima relative to the environment in general and are restricted to cool microclimates. Furthermore, the study of the Puerto Rican taxa reveals that these thermoconformers are already operating at temperatures in excess of their thermal 260 maxima and likely will be pushed beyond their thermal tolerance with predicted temperature increases. This study further substantiates that the effects of climate change will be multifarious and will impact every sector of the globe. HUEY, R. B., C. A. DEUTSCH, J. J. TEWKSBURY, L. J. VITT, P. E. HERTZ, H. J. ÁLVAREZ PÉREZ, AND T. GARLAND JR. 2009. Why tropical forest lizards are vulnerable to climate warming. Proceedings of the Royal Society B 276:1939–1948. Correspondence to: Raymond Huey, Department of Biology, University of Washington, PO Box 351800, Seattle, Washington 98195, USA; e-mail: hueyrb@u.washington.edu. Greater Species Richness in the Tropics: Extinction, Speciation, or Historical Biogeography? Traditionally, the elevated species richness in the tropics has been explained by higher rates of speciation or lower rates of extinction relative to temperate regions. However, recent research has revealed that in some systems, rates of speciation and extinction are comparable in the two regions and that the higher species richness in the tropics is a result of the lineages of interest having occupied these regions for a longer period of time. In this study the authors aimed to determine if this was the case with ranid frogs. In addition to presenting a new phylogeny of the family Ranidae, divergence times and ancestral geographic distributions were estimated using the software BEAST and Mesquite, respectively. A simple regression revealed a highly signiﬁcant correlation between regional diversity and cumulative time of lineage presence in a region. A regression of diversiﬁcation rate of clade against latitudinal midpoint revealed a weak positive correlation, with higher latitudes exhibiting a slightly higher diversiﬁcation rate, which further demonstrates that higher species richness in the tropics is not due to higher speciation rates or lower extinction rates in that region. This study reiterates the importance of taking historical biogeography into account when trying to explain patterns of biodiversity. WIENS, J. J., J. SUKUMARAN, R. A. PYRON, AND R. M. BROWN. 2009. Evolutionary and biogeographic origins of high tropical diversity in Old World frogs (Ranidae). Evolution 63:1217–1231. Correspondence to: John Wiens, Department of Ecology and Evolution, Stony Brook University, Stony Brook, New York 11794, USA; e-mail: wiensj@life.bio.sunysb.edu. Documenting Salamander Declines in the Neotropics Anuran declines have received much deserved attention; however, in some parts of the globe, salamander species are disappearing at an equally alarming rate. In this paper the authors compare recent and historical salamander abundance data from several sites in southern Mexico and Guatemala, many of which still contain abundant suitable habitat. Analyses revealed that many taxa have experienced precipitous declines and local extirpations, with very Herpetological Review 40(3), 2009 few species maintaining or increasing in abundance. These ﬁndings span Mexico’s Isthmus of Tehuantepec, across which there is a complete turnover in the salamander biota. Analyses further revealed that these population declines and extirpations seem to disproportionately affect upper cloud forest and high elevation species. Additionally, within these habitats, species that utilize arboreal microhabitats seem to be relatively unaffected while the terrestrial microhabitat specialists are experiencing most of the population declines and extinctions. Interestingly, the low-elevation species disproportionately utilize arboreal microhabitats. Finally, based on records from southern Mexico, the timing of the declines seems to coincide with dramatic declines in anuran species in the region, suggesting that chytridiomycosis may have played a role in the decline of these salamander species as well. ROVITO, S. M., G. PARRA-OLEA, C. R. VÁSQUEZ-ALMAZÁN, T. J. PAPENFUSS, AND D. B. WAKE. 2009. Dramatic declines in neotropical salamander populations are an important part of the global amphibian crisis. Proceedings of the National Academy of Sciences 106:3231–3236. Correspondence to: David Wake, Museum of Vertebrate Zoology and Department of Integrative Biology, University of California, Berkeley, California 94720-3160, USA; e-mail: wakelab@berkeley.edu. Quantifying the Chameleon’s Ability to Adjust UV Exposure in Response to Low Vitamin D3 Diet Vitamin D3 is a hormone essential to most vertebrates including squamates; however, excessive vitamin D3 is fatal. Therefore, animals should be expected to exhibit behaviors that regulate D3 levels. Previous studies have determined the optimal level of UVB exposure (necessary for the production of D3) for chameleons and have shown that they have the ability to behaviorally compensate for a low D3 diet by increasing UVB exposure time. However, these studies failed to quantify the accuracy and precision of this ability. To address this question, the authors of this study placed six Panther Chameleons (Furcifer pardalis) in outdoor enclosures and divided them into high and low D3 diet treatments. Daily movements were monitored via video camera. Using models soaked in provitamin D3 (a chemical vitamin D3 precursor that converts to vitamin D3 when exposed to UVB), the recorded movements were replicated the following day to allow for quantiﬁcation of the level of D3 production. These experiments revealed the chameleon’s ability to behaviorally compensate for different levels of vitamin D3 to be both highly precise and accurate, with low D3 diet treatment lizards maintaining optimal levels of D3 via increased UVB exposure and those in the high D3 diet treatment similarly maintaining optimal levels of D3 by moderating UVB exposure. KARSTEN, K. B., G. W. FERGUSON, T. C. CHEN, AND M. F. HOLICK. 2009. Panther chameleons, Furcifer pardalis, behaviorally regulate optimal exposure to UV depending on dietary vitamin D3 status. Physiological and Biochemical Zoology 82:218–225. Correspondence to: Kristopher Karsten, Department of Biology, Texas Christian University, Fort Worth, Texas 76129, USA; e-mail: k.karsten@tcu.edu. Malagasy Anuran Diversity Greatly Underestimated Madagascar, together with the oceanic islands of the Indian Ocean, is considered one of 34 biodiversity hotspots by the organization Conservation International and possesses a unique herpetofauna. Malagasy anurans are particularly diverse and unique, constituting ﬁve evolutionary lineages (including two endemic families) and exhibiting 100% species endemicity. One of the primary goals of conservation groups is to document the biodiversity within these hotspots before it is too late. With this goal in mind, the authors conducted surveys and analyses with the goal of assessing Malagasy anuran diversity. A combination of mitochondrial sequence, vocalization, morphological, and geographic data were accumulated for ca. 2850 specimens. Molecular analyses were performed to identify divergent lineages within currently recognized species. If these lineages were found to be morphologically or bioacoustically divergent, they were categorized as Conﬁrmed Candidate Species (CCS); if no morphological or bioacoustic data were available, they were classiﬁed as Unconﬁrmed Candidate Species (UCS). These analyses revealed a veritable cornucopia of undescribed diversity. Assuming the CCS and UCS categories reﬂect legitimate species, this study raises the number of species occurring in Madagascar from 244 to 465 (an increase of 90.6%). These results reiterate how much of the world’s diversity remains to be discovered and emphasizes the importance of quantifying the biodiversity of hotspots. VIEITES, D. R., K. C. WOLLENBERG, F. ANDREONE, J. KÖHLER, F. GLAW, AND M. VENCES. 2009. Vast underestimation of Madagascar’s biodiversity evidenced by an integrative amphibian inventory. Proceedings of the National Academy of Sciences 106:8267–8272. Correspondence to: David Vieites, Museo Nacional de Ciencias Naturales, Consejo Superior de Investigaciones Cientíﬁcas, c/ José Gutierrez Abascal 2, 28006 Madrid, Spain; e-mail: vieites@mncn.csic.es. Box Turtle Abscesses Heal Themselves Cranial abscesses are a frequently observed phenomenon in box turtles (Terrapene). While standard procedures exist for treating this disease in captive animals, the ﬁnal result of such abscesses in nature has remained uncertain. The author reports on ﬁndings resulting from tracking 144 Eastern Box Turtles (T. carolina). Twenty cases of aural abscesses were recorded over the course of a decade. Of these, ﬁfteen cases healed naturally and ﬁve required researcherinduced amputation (after turtle exhibited obvious weight loss). This is the ﬁrst evidence in support of the hypothesis proposing that these abscesses spontaneously disappear in nature. BELZER, W. R. 2008. Spontaneous resolution of aural abscesses in the eastern box turtle, Terrapene carolina carolina. Journal of Herpetological Medicine and Surgery 18:64–66. Correspondence to: William Belzer, Box Turtle Conservation Trust, 304 East Bissell Ave., Oil City, Pennsylvania 16301, USA; e-mail: billbelzer@hotmail.com. Herpetological Review 40(3), 2009 261 Piscivory Correlated with Morphological Evolution in Natricine Snakes The various species of natricine snakes exhibit the full spectrum of percentage of diet composed of ﬁsh. However, much remains to be learned about how morphology and feeding performance are related to obligate piscivory in these snakes. The authors of this study accumulated data on quadrate bone length (as they hypothesize an increase in quadrate length would allow snakes to more easily feed on odd-shaped prey items), ﬁsh-eating performance, and fraction of ﬁsh in diet, and incorporating accepted hypotheses of evolutionary relationships, tested for convergent evolution in natricines. Experiments ascertaining feeding performance involved feeding snakes ﬁsh and timing the event. Results revealed that increased feeding performance was indeed correlated with elongation of the quadrate when the prey is ﬁsh. Furthermore, an ancestral state reconstruction suggested that all major lineages were ancestrally generalists; specialization on ﬁsh or non-ﬁsh evolved several times quickly and recently within those lineages. Finally, an independent contrasts analysis conﬁrmed increased quadrate length is highly correlated with increased percent ﬁsh in diet and scales allometrically with increased head length. This fascinating example of convergent evolution suggests that dietary specialization can have dramatic effects on all aspects of an organism’s biology. VINCENT, S. E., M. C. BRANDLEY, A. HERREL, M. E. ALFARO. 2009. Convergence in trophic morphology and feeding performance among piscivorous natricine snakes. Journal of Evolutionary Biology 22:1203–1211. Correspondence to: Anthony Herrel, UMR 7179 C.N.R.S/M.N.H.N., Département d’Ecologie et de Gestion de la Biodiversité, 57 rue Cuvier, Case postale 55, 75231, Paris Cedex 5, France; e-mail: anthony. herrel@mnhn.fr. Where Does the Ring End? Testing Biogeographic Hypotheses in Ensatina Ensatina salamanders have served as the textbook example of a “ring species” for decades. Historically, it has been suspected the terminus of this ring occurred in the mountains of southern California. However, geologic data suggests that a signiﬁcant barrier to dispersal existed in coastal central California, as the southern Coast Range was ﬁrst an island chain, then (once joined with the mainland) separated from the northern Coast Range by the outlet of the inland sea at what is now Monterey. Under this scenario, dispersal south along the Sierra Nevada, west to the coast, then north to Monterey would be the more likely dispersal scenario and Monterey should be the terminus of the ring. If the terminus is in the south, lineages on either side of Monterey would be closely related and lineages in the mountains of southern California would be deeply divergent. If the terminus is in Monterey, the opposite would be true. To evaluate these two hypotheses, a phylogeographic analysis was conducted using mitochondrial DNA sequences from samples obtained from throughout the distribution of the species. An analysis of the credible set of trees revealed a sisterspecies relationship of the two southern California lineages (and the Monterey terminus hypothesis) could conﬁdently be rejected. 262 Alternatively, greater than 99.9% of the trees were consistent with a southern California terminus and ﬁnd lineages north and south of Monterey to have a sister relationship. Interestingly, a molecular clock analysis revealed that this coastal California clade originated no sooner than nine million years ago, vastly predating the uniﬁcation of the Coast Range. KUCHTA, S. R., D. S. PARKS, R. LOCKRIDGE MUELLER, AND D. B. WAKE. 2009. Closing the ring: historical biogeography of the salamander ring species Ensatina eschscholtzii. Journal of Biogeography 36:982–995. Correspondence to: Shawn Kuchta, Department of Animal Ecology, Lund University, Ecology Building, Sölvegatan 37, SE-223 62 Lund, Sweden; e-mail: shawn.kuchta@zooekol.lu.se. Venom Potency Correlated with Diet in Echis Vipers Given the different physiologies of different prey items, it would follow that major dietary changes would lead to venom evolution in snakes. This hypothesis was tested in snakes of the genus Echis. Previous work suggests that Echis is made up of four species groups. Mitochondrial and nuclear sequence data were generated for representatives of each of the four lineages and a Bayesian phylogenetic analysis was performed to reveal the relationship of these lineages to each other. Additionally, data were accumulated on diet and aspects of the venom (LD50 and time to death) when administered to scorpions. Placing dietary preference on the resulting tree, it was discovered that the ancestral condition for the genus is an arthropod diet, with one Echis lineage secondarily evolving a rodent diet. Additionally, a correlation was revealed between percentage of arthropods in diet and toxicity (LD50) to scorpions. Interestingly, time to death did not appear to be correlated with arthropod diet, suggesting that venom conservation alone is the target of selection. BARLOW, A., C. E. POOK, R. A. HARRISON, AND W. WÜSTER. 2009. Coevolution of diet and prey-speciﬁc venom activity supports the role of selection in snake venom evolution. Proceedings of the Royal Society B 276:2443–2449. Correspondence to: Axel Barlow, School of Biological Sciences, Environment Centre Wales, Bangor University, Bangor LL57 2UW, UK; e-mail: bsu004@bangor.ac.uk. Lampropeltine Relationships Resolved Hypotheses regarding the evolutionary relationships and the resulting taxonomy of the North American lampropeltine snakes have been in ﬂux recently. Using mitochondrial and nuclear DNA sequence data, the authors performed a comprehensive phylogenetic analysis of this group using Old World rat snakes for outgroups. This analysis revealed a monophyletic Lampropeltini and clades largely concordant with morphology (including monophyletic Pantherophis and Pituophis), upon which the original taxonomy was based. Three taxa (Arizona, Rhinocheilus, and Cemophora) were revealed to be closely allied with Lampropeltis (king snakes); a fourth (Stilosoma) was recovered as highly nested within Lampropeltis. Furthermore, several novel relationships were revealed Herpetological Review 40(3), 2009 within Lampropeltis and one species (L. triangulum) was found to be comprised of no fewer than two deeply divergent lineages. Finally, a molecular clock analysis found the origin of Lampropeltini to be concordant with an initial dispersal into North America via Beringia. PYRON, R. A., AND F. T. BURBRINK. 2009. Neogene diversification and taxonomic stability in the snake tribe Lampropeltini (Serpentes: Colubridae). Molecular Phylogenetics and Evolution 52:524–529. Correspondence to: Alexander Pyron, Department of Biology, The Graduate School and University Center, The City University of New York, 365 Fifth Ave., New York, NY 10016, USA; e-mail: rpyron@gc.cuny.edu. Australian Cane Toads More Susceptible to Ant Attack than Native Anurans As invasive species originated in and were adapted to different environmental conditions, they should be expected to exhibit vulnerabilities to certain dangers not shared with native animals, regardless of overall successfulness in the new environment. In this study, the authors investigated the susceptibility of Cane Toads (Bufo [Rhinella] marinus) to ant attack relative to native anurans. Metamorphic Cane Toads, Litoria (ﬁve species), and myobatrachids (two species), all of which co-occur in Australia’s Northern Territory, were measured for various morphometric traits, sprint speed performance, and response to a simulated ant attack. Daily activity patterns were documented both in captivity and in the ﬁeld for meat ants (Iridomyrmex spp.; common predatory ants in ecosystems of interest) and in captivity for the anurans. Furthermore, ants were introduced to anuran terraria and interactions were recorded. Finally, bait was placed in speciﬁc microhabitats within enclosures to look for variation in detectability; bait was also placed in the center of a piece of paper laden with species-speciﬁc anuran secretions and ants were introduced to terraria to determine how the ants responded. The Cane Toads were found to come into contact with ants more frequently than the other anurans because of their diurnal activity and preferred, exposed microhabitats. Additionally, they failed to anticipate an impending attack more frequently than native frogs and the toad’s short hops and frequent reliance on crypsis increased their susceptibility to ant attack. Ants failed to elicit a species-speciﬁc response to the anuran secretions. Now that they have been identiﬁed, these vulnerabilities could perhaps be employed in cane toad eradication projects. WARD-FEAR, G., G. P. BROWN, M. J. GREENLEES, AND R. SHINE. 2009. Maladaptive traits in invasive species: in Australia, cane toads are more vulnerable to predatory ants than are native frogs. Functional Ecololgy 23:559–568. Correspondence to: Richard Shine, School of Biological Sciences A08, University of Sydney, NSW 2006, Australia; e-mail: rics@bio.usyd.edu. au. ZOO VIEW Herpetological Review, 2009, 40(3), 263–273. © 2009 by Society for the Study of Amphibians and Reptiles History of Early French Herpetology. Part I: The Reptile Menagerie of the Museum of Natural History in Paris JAMES B. MURPHY Department of Herpetology, Smithsonian National Zoological Park 3001 Connecticut Ave., N.W., Washington DC 20008, USA e-mail: jbmurphy2@juno.com Today the Jardin des Plantes is interesting to visit purely on an historical perspective as it has two separate buildings devoted to reptiles. The most interesting dates from the Victorian era, while the “new” one looks as if it was constructed in the 1930s. The Victorian building still retains and uses the original ornate legged, glass-fronted, mahogany reptile cases and marble-walled crocodile pits. To see the reptile cages here is to step back in time and view the ﬁrst dawning of modern herpetoculture. —Jon Coote, 2001 The Site.—The Ménagerie, called Les Jardin des Plantes, of the Museum of Natural History in Paris (la ménagerie du Muséum d'histoire naturelle) was created in 1793 and was the ﬁrst of the national menageries. Prior to the inception of the Jardin des Plantes, practically all exotic animal collections were held by kings, queens, rulers, and other powerful political ﬁgures. During the French Revolution, the Jardin des Plantes was appropriated by the revolutionaries from the Jardin du Roi in Versailles. In 1793 this Royal Menagerie was renamed Jardin des Plantes and moved from Versailles to Paris. In Lewis Pyenson and Susan SheetsPyenson’s book, Servants of Nature. A History of Scientiﬁc Institutions, Enterprises, and Sensibilities, they describe it this way: “The splendors of Versailles, including its extensive gardens and menagerie, became one of the most powerful symbols of the ancien régime to the revolutionaries of the 1790s. In the context of this upheaval, a Jacobin mob set out from Paris to liberate the animals, resulting in the senseless slaughter of most of them. The surviving animals went to the Jardin des Plantes in Paris, to serve as the nucleus of a new national collection. Other animals were added that had been seized from private menageries and circuses” (p. 166). It was the ﬁrst truly public collection. If one compares it to the Schönbrunn Menagerie in Vienna, Austria, the latter was a collection that had enclosures surrounded by a central viewing pavilion, designed to provide views of the animals only to those privileged enough to be inside it. The Jardin conversely has broad walkways and avenues, down which ordinary people could, and still can, stroll to admire the animals. This design reﬂects its unique origin—a royal menagerie appropriated by the people for the people. Truly an important change in zoo history! The Jardin is a short walk from the Museum so it is likely that herpetologists from both places regularly interacted with each other. In fact, it is reasonable to assume that herpetologists stationed at the museum would dash over to the menagerie whenever a new shipment of interesting live amphibians and reptiles arrived. In the mid-18th and early 19th century, three famous herpetologists were associated directly or indirectly with the menag- Herpetological Review 40(3), 2009 263 erie: Georges-Louis Leclerc compte de Buffon (Fig. 1), Baron Georges Cuvier (Fig. 2), and Count de Lacepède (Fig. 3). The rich history as a center for experimental zoology is detailed extensively by Gustave Loisel (1912) and other references are provided below. The Terrarium, from 1839 under the charge of André Marie Constant Duméril, was a separate section in the old monkey house, which housed 80 reptiles of 24 species, amphibians, ﬁshes and insects. A common practice in Europe was to encourage travelers to secure specimens for collections; in some cases, written instructions were given to the collectors. Built in the 1870s, the Menagerie’s reptile building today has a large aquatic exhibit which houses an impressive collection of crocodilians. At one end of the building, large tortoises are on display. Madagascan reptiles are well represented. Several extensive graphic displays with lifelike models of reptiles are situated throughout the building. Some of the larger enclosures are being remodeled. In the newer Vivarium building located near the reptile building, smaller amphibians and reptiles are represented in naturalistic settings. Publications.—Between 1834–1854, Constant Duméril (Fig. 4), his son Auguste Henri André Duméril (Fig. 5), and Gabriel Bibron (Fig. 6) published the monumental multi-volume work called Erpétologie générale, ou, Histoire naturelle complète des FIG. 1. Portrait of Georges-Louis Leclerc comte de Buffon (1707–1788) in M. Boitard’s Le Jardin des Plantes. Description et Moeurs des Mammifères in 1845. Pyenson and SheetsPyenson said that “…Buffon added greenhouses and a natural history cabinet” to the garden (p. 154) and that “When the Jardin des Plantes reached its apogee under the old regime, private menageries still ﬂourished in France. Especially important from a scientiﬁc standpoint was the comte de Buffon’s menagerie at Montbard. There Buffon had purchased the picturesque ruins of an old castle. He reconstructed the castle, planted gardens, established laboratories, and founded a zoological park. He stocked the park with a wide range of exotic animals, which served as a basis for writing his multivolume Historie naturelle. At Montbard, he also undertook varied research in experimental zoology, notably in reproduction and hybridization” (p. 168). In Zoological Illustration, David Knight says that Buffon “…summed up the work of pre-Linnaean natural historians” (p. 96). As Director of the Jardin du Roi in Versailles, he became the model of the scientiﬁc collector and lobbied for a formalized relationship between the museum and menagerie. He transformed the king's garden into a scientiﬁcally signiﬁcant museum and research center. Buffon became the curator in 1739 and he expanded the gardens greatly. His massive Histoire naturelle (36 volumes) [other sources say it was published in 44 volumes between 1749 and 1788, with subscribers purchasing volumes ahead of production. Knight (1977) says that it was 44 volumes, but that the last eight were completed by Lacepède] set out to organize all that was then known about the natural world. Credit: Courtesy of Smithsonian Institution Libraries, Washington, DC. 264 reptiles. In one of the volumes—the Atlas—are 120 nice illustrations of amphibians and reptiles. In 1854–55, Auguste Duméril wrote Notice historique sur la Ménagerie des Reptiles du Muséum d'histoire naturelle et observations qui y ont été recueillies, featuring six beautifully hand-colored plus two uncolored plates. This work listed the observations collected at the menagerie, summarized FIG. 2. Portrait of Baron Georges Cuin the Index (Fig. 7). In vier (1769–1832) in M. Boitard’s Le Jarthe Notice historique, din des Plantes. Description et Moeurs Duméril included many des Mammifères in 1845. In 1802, he references to the earlier became titular professor at the Jardin Erpétologie générale so des Plantes. His most famous work is some relevant ones from the Règne animal distribué d'après son organisation (1817; translated into Engthe Erpétologie générale lish as The Animal Kingdom). Credit: Atlas and one from Études Courtesy of Smithsonian Institution Lisur les reptiles are shown braries, Washington, DC. here (Figs. 8–11, 12–20). In three additional papers published by the Museum, Auguste Duméril recorded information about specimens in the Ménagerie. In 1858–61, he provided a list of ﬁshes, reptiles and amphibians, and notes on the menagerie of reptiles. In 1865, reptile and amphibian holdings between 1838–1864 were compiled. New herps in the collection were recorded in the museum archival publication in 1869–70 (Fig. 21). Arrival of the “Victorian” Building.—The London Zoo opened the World’s ﬁrst reptile building in 1849 (see Murphy, 2007 for details). In 1870–1874, an architect Emile Blanchard designed a reptile menagerie with a pavilion (Figs. 22–24). The facility was 30 m long, with two halls which contained smaller squamates and chelonians. Two large center exhibition halls were called “Crocodile Hall” and “Aquarium Hall;” the latter contained freshwater ﬁshes and amphibians. FIG. 3. Portrait of Bernard-GermainOne unusual feature is Etienne de la Ville-sur-Illon comte de that the rear walls of enLacepède (1756–1825). This French closures on the outer wall naturalist, principally known for his are glass. As a result, these herpetological work, published several displays contained living seminal papers on zoo history, philosoplants which thrived in phy, and design. Credit: Courtesy of natural light. The glass- Kraig Adler. Herpetological Review 40(3), 2009 fronted, mahogany cages standing on ornate legs and the marble-walled crocodile pits are still in use (Coote 2001). Bronze sculptures were placed throughout the menagerie: “The Snake Charmer” and “The Crocodile Hunter” by Arthur Bourgeois in front of the reptile pavilion and “Eve” by Guitton near the outdoor crocodile pool. In addition to his regular duties in the museum working with the collecFIG. 4. Portrait of André Marie Contion, Léon-Louis Vaillant stant Duméril (1774–1860). Credit: was in charge of the reptile Courtesy of Kraig Adler. menagerie and aquarium. In 1898, he produced Muséum d'histoire naturelle guide à la ménagerie des Reptiles. On the front cover of this guide are drawings of Count de Lacepède, C. Duméril, A. Duméril, and Vaillant himself (Fig. 25). There is a picture of the ﬂoorplan (Fig. 26) and another one of visitors enjoying a trip to the reptile building (Fig. 26a). Scientiﬁc Discoveries.—Pierre Bernard mentioned that a female python laid 15 eggs, eight of which hatched in Le Jardin (Bernard 1842–1843). In 1841, Achille Valenciennes reported his observations on the incubating female python (called Python bivittatus) at the Menagerie to the French Academy (Valenciennes 1841; Fig. 27). The belief that brooding female pythons produce heat to incubate their eggs was a controversial issue during the nineteenth century (Duméril 1842, 1858). Duméril was convinced that the “cold-blooded nature” of reptiles could not lead to these temperature increases and the issue remained unresolved for many years (see Murphy 2007). Likely the most famous description presented by Auguste Duméril was his discovery in the menagerie that the larval axolotls of Mexico grow, become sexually mature, reproduce, and yet retain larval features such as external gills—the neotony of Ambystoma mexicanum (Figs. 28, 28a). Duméril published papers on reproduction, developmental anomalies, and an amelanotic strain. This FIG. 5. Portrait of Auguste Henri Annow-critically endangered dré Duméril (1812–1870). Credit: Coursalamander is found in tesy of Kraig Adler. the Xochimilco aquatic ecosystem around Mexico City; this taxon is negatively impacted by pollution threats and invasive species, effects currently under study by a number of zoo biologists (Koenig 2008; Johnson et al., Connect [October 2008:6–9]). Collection.—Deleuze (1823) described an enclosure divided into ﬁve parts housing smaller aquatic birds, long-legged birds and gallinaceous fowls, FIG. 6. Portrait of Gabriel Bibron and a large semi-aquatic (1806–1848). Credit: Courtesy of Kraig exhibit for chelonians. Adler. The list of species kept was impressive: Radiated, Geometric, Angulated, and Greek Tortoises; Painted, European Pond, Snapping, and Snake-Necked Turtles; Diamondback Terrapin; a number of undescribed taxa; and, amazingly, a Hawksbill Seaturtle which likely lived in fresh water. There were also FIG. 7. Index from Notice historique sur la Ménagerie des reptiles du Muséum d'histoire naturelle et observations qui y ont été recueillies. Credit: Library of Roy McDiarmid. Herpetological Review 40(3), 2009 265 Timber Rattlesnakes, African Chameleons, and European Olms on exhibit. In 1910, the collection numbered 216 reptiles and 237 amphibians. Longevities included a 60-year-old American Alligator, 35 year old Australian turtle, 22-yearold snapping turtle, three giant tortoises FIG. 8. Illustration of African Spurred Tortoise over 20 years, (Testudo sulcata, now Geochelone sulcata) above M a d a g a s c a n and Common Spider Tortoise (Pyxis arachnoides) boa over 21 from Erpétologie générale. Credit: Collections of years, 14-year- Ernst Mayr Library, Museum of Comparative Zoolold Reticulated ogy, Harvard University. Python and a "molure" python two years older, Japanese salamander for 30 years and a siren for 23 years. Jacques Pellegrin at the museum also published on herp longevities at the menagerie: longevity in ﬁsh and amphibians (Rev. Gen. Sci. Paris 36:510–512, 1925); longevity in reptiles (Rev. Gen. Sci. Paris 37:47–49, 1926); longevity in alligator (Bull. Mus. Nat. Hist. ser. 2, 9:176–177, 1937). One may consult Flower (1925, 1936, 1937) for additional longevities. Concluding Remarks.— Count de Lacepède wrote important papers at the FIG. 9. Illustration of Mata Mata museum detailing the Turtle (Chelus ﬁmbriatus) and Snakemission of zoos, design Necked Turtle from New Holland recommendations, and from Erpétologie générale. The Snakeintegration of science. In Necked Turtle called Chelodina novae1801, he said, “When an hollandiae is the Common Snake Neck animal dies, it is immedi- Turtle (Chelodina longicollis). Credit: Collections of Ernst Mayr Library, Muately brought to the laboseum of Comparative Zoology, Harvard ratory of anatomy. There University. 266 FIG. 10. Illustration of Box Turtle skulls and skeleton from Erpétologie générale. Credit: Library of Roy McDiarmid. the skin is taken and sent to the laboratory of zoology where the professor has it mounted if it is not yet on exhibit.” He clearly understood the proper relationship between a museum and a zoological garden, stressing cooperation between the two institutions FIG. 11. Illustration of crocodilian heads from Erpétologie générale. The scientiﬁc name Alligator lucius is the old name for the American Alligator. The name “Caiman a Tète de Brochet,” is sometimes used in books from the 19th century and is translated as “Caiman with the snout of a pike.” Credit: Library of Roy McDiarmid. Herpetological Review 40(3), 2009 FIG. 11a. In the past, crocodilians were often kept in dense numbers in small enclosures in zoos. In this undated postcard (but possibly from the late 19th or early 20th century), American Alligators are shown in an exhibit at the Jardin surrounded by recurved bars. An American Crocodile was received in 1851 and Émile Wapler from New Orleans donated six American Alligators to the Menagerie the next year where they lived for many years (Duméril, 1854–1855, 1858–1861). Credit: Collection of Smithsonian National Zoological Park. to use animals upon death to the fullest (see Murphy and Iliff 2004). There are several examples which serve to illustrate the value of this living collection. Marie Firmin Bocourt learned how to draw herpetological subjects by watching living models at the Ménagerie when he was young. LéonLouis Vaillant published on reptilian behavior and physiology, based on observations collected at the Ménagerie. Auguste Duméril wrote some of his treatises on the amphibians and reptiles in the Jardin’s collection, often accompanied by exquisite illustrations. The Ménagerie and earlier Terrarium served an important function FIG. 12. Illustration of chameleons for artists and biologists from Erpétologie générale. Auguste from that period—a place Duméril found that European chamewhere living animals leons which began arriving at the Jardin could be observed. As a in August 1851 were delicate captives. result, many illustrations Over the next four years, more than 160 published by French bi- were kept with none living more than 13 months. His lizards fed on a variety ologists began to be more of insects but quickly died. Duméril noprecise: natural poses, ticed that females became egg-bound or ﬁne detail of diagnostic often died after oviposition and no eggs characters, and accurate ever hatched. Credit: Courtesy of Smithcoloration. Improvements sonian Institution Libraries, Washington, in printing technology DC. FIG. 13. Illustration of chameleon skulls and skeleton from Erpétologie générale. Note that the two outer and three inner ﬁngers are not fused on the forefeet and three outer and two inner toes are not fused on the hind feet. Chaméléon ordinaire is the Common Chameleon (Chamaeleo chamaeleon) and the Chaméléon Nez-Fourchu is now Furcifer biﬁdis. Credit: Library of Roy McDiarmid. also occurred which contributed to this precision. Lithography was important in the late 18th century and multicolored lithographic prints replaced hand coloring in the early 19th century. The improvement in animal illustrations in France and elsewhere in Europe took place in less than 100 years. The change to more realistic depictions of animals came about from the availability of better preserved specimens and live animals, advances in printing technology, and the evolution away from the use of patterns that dominated Renaissance art. This last point is signiﬁcant, because plant illustration was already very realistic due to the fact that specimens were more widely available (plants were already being pressed and dried and attached to sheets), and the fact that Renaissance animal illustration was still based heavily on myth, as in the bestiary tradition. Consider the contrast between Albrecht Durer’s beautiful, lifelike and realistic depictions of a rabbit (an animal he had seen alive frequently) and his famous boiler-plate rhino, an animal he was drawing based on a sketch and description, in the context of the bestiary tradition. Lavishly illustrated books were designed to be sold not just to people interested in natural history, but those interested in art as well as neither audience was big enough to support the costs FIG. 14. Illustration of banded phase (sometimes called Bell’s Monitor) of the Lace Monitor (Varanus Bellii, now Varanus varius) and Clouded Monitor (Varanus nebulosus) from Erpétologie générale. Credit: Library of Roy McDiarmid. Herpetological Review 40(3), 2009 267 of publication alone (this point is made in Rifkin and Ackerman 2006. Human Anatomy [From the Renaissance to the Digital Age]. Buffon is a great example of this—his natural history was a very popular book in its day, selling well to the general public as well as naturalists and art collectors. Rifken and Ackerman quote Cuvier as having said that “One must say that without the art of the printer, natural history and anatomy, as they exist today, would not be possible” (p. 271 FIG. 15. Illustration of Mud Snake in their book). More ex(Hydrops abacure, now Farancia abapensive books with handcura) from Erpétologie générale. Credcolored illustrations sold it: Library of Roy McDiarmid. well to art collectors, then as now. Copperplate printing was in use in the mid-16th century, but although copperplates could produce ﬁner lines and hence ﬁner details, the copper plates did not last as long as woodblocks, so woodblocks remained popular with printers for a long time due to their durability. Lithography came out of Germany in the late 18th century, discovered by Alois Seneﬁlder. I have selected an array of interesting plates which reﬂect the essence of living herps and the ﬁne artwork created during this period. These will appear in Part II. All colored originals have been converted to a black-and-white format. Acknowledgments.—This contribution is dedicated to Ken Kawata, retired curator at the Staten Island Zoo in New York. Throughout his professional career, he has stressed the importance of looking at zoo and aquarium history by returning to our earlier roots to improve chances of not duplicating efforts already tried. Kawata has become a mentor to a whole new generation of zoo workers. I am grateful to Emily Becker, Dana Fisher and James Hanken from the Museum of Comparative Zoology, Harvard University for providing some of the illustrations. Polly Lasker, Smithsonian Institution Libraries, searched archival materials to assure that citations were correct. 268 FIG. 16. Illustration of African Rock Python (Python sebae) from Erpétologie générale. Credit: Library of Roy McDiarmid. FIG. 17. Illustration of True Viper heads with horns from Erpétologie générale. Current common and scientiﬁc names are #1 Sand Viper (Vipera ammodytes); #2 Rhinoceros Viper (Bitis nasicornis); #3 possibly Sahara Sand Viper (Cerastes vipera); #4 Many-Horned Adder (Bitis cornuta); #5 Persian Horned Viper (Pseudocerastes persicus). Credit: Library of Roy McDiarmid. Thanks are also due to Jean-Luc Berthier, Leslie Overstreet, and Daria Wingreen. My wife Judith Block, Kraig Adler, Jon Coote, Roy McDiarmid, and John Simmons read early drafts and made many helpful suggestions. FIG. 18. Illustration of snake skulls from Erpétologie générale. Current common and scientiﬁc names are #1 Reticulate Worm Snake (Typhlops reticulatus); #3 Indian Python (Python molurus); #4 Emerald Tree Boa (Corallus caninus). Credit: Library of Roy McDiarmid. Herpetological Review 40(3), 2009 FIG. 19. Drawing of Copperhead (Trigonocephalus contortrix, now Agkistrodon contortrix) by Firmin Bocourt from Études sur les reptiles. Many extralimital species from the United States, South America, and the West Indies were included. REFERENCES Historical Overview ADLER, K. 1989. Herpetologists of the past. In K. Adler (ed.), Contributions to the History of Herpetology, pp. 5–141. SSAR Contributions to Herpetology, Vol. 5. Oxford, Ohio. ________ . 2007. Herpetologists of the past, Part 2. In K. Adler (ed.), Contributions to the History of Herpetology, pp. 3–373. SSAR Contributions to Herpetology, Vol. 21. St. Louis, Missouri. BARATAY, E. 1998. Zoos: histoire des jardins zoologiques en Occident (XVIe–XXe siècle). La Découverte, Paris. FIG. 20. Illustration of frogs from Erpétologie générale. Rhacophore de Reinwardt is Rhacophorus reinwardtii. L'Hylode de la Martinique is Eleutherodactylus martinicensis. Credit: Library of Roy McDiarmid. FIG. 21. List of herpetological specimens from Quatrième notice sur La Ménagerie des Reptiles du Muséum d'histoire naturelle. BELL, C. E. (editor). 2001. Encyclopedia of the World’s Zoos. Chicago, Fitzroy Dearborn Publishers. BERNARD, P. 1842–1843. Le Jardin des Plantes: description complète, historique et pittoresque du Muséum d'histoire naturelle, de la ménagerie, des serres, des galeries de minéralogie et d'anatomie, et de la vallée suisse : moeurs et instincts des animaux, botanique, anatomie comparée : minéralogie, géologie, zoologie (The Jardin des Plantes: a complete, historical and picturesque description of the Museum of Natural History, of the menagerie, the green-houses, the galleries of mineralogy and anatomy and of the Swiss Valley : mores and instincts of the animals, botany, comparative anatomy : mineralogy, geology, zoology). L. Curmer, Paris. [beginning on page 153, Pierre Bernard (1810–1876) discussed the habits and characteristics of reptiles living in the menagerie and included several excellent plates of a scorpion, lizard and FIG. 22. Undated illustration, possibly around mid-1870s, of exterior of reptile building at La Ménagerie Jardin des Plantes. Credit: provided by Jean-Luc Berthier, Jardin des Plantes Archives. Herpetological Review 40(3), 2009 269 FIG. 23. Undated illustration, possibly around mid-1870s, of interior of reptile building at La Ménagerie Jardin des Plantes. Note glass-fronted, mahogany cages standing on ornate legs on right. Credit: provided by Jean-Luc Berthier, Jardin des Plantes Archives. snake. He defended the need for a living animal collection as a valuable vehicle for research and education.]. BODSON, L. 1984. Living reptiles in captivity: A historical survey from the origins to the end of the XVIIITH Century. Acta Zool. Path. Antverpiensia 78:15–32. BOITARD, P. 1845. Le jardin des plantes; description et mœurs des mammifères de la Ménagerie et du Muséum d'histoire naturelle / / par Pierre Boitard ; précédé d'une introd. historique, descriptive et pittoresque par J. Janin. Imprint: Paris, J.-J. Dubochet, 1845. CAP, P.-A. 1854. Le muséum d'histoire naturelle; histoire de la fondation et des développements successifs de l'éstablissement; biographie des hommes célèbres qui y ont contribué par leur enseignement ou par leurs découvertes, histoire des recherches, des voyages, des applications utiles auxquels le muséum a donné lieu, pour les arts, le commerce et l'agriculture, description des galeries, du jardin, des serres et de la ménagerie, / par m. P.-A. Cap et une société de savants et d'aides naturalistesdu muséum. Paris, L. Curmer. COOTE, J. G. 2001. A history of western herpetoculture before the 20th century. In W. E. Becker (ed.), 25th International Herpetological Symposium on Captive Propagation and Husbandry, pp. 19–47. International Herpetological Symposium, Detroit, Michigan. CORTÁZAR, J. 1956. [The axolotl was a big draw of the public, too. In more recent times, a wonderful short story was written by an Argentine (Julio Cortázar) in 1956 about seeing the axolotl and recognizing it as fellow exile—you can read the story at http://www.cis.vt.edu/modernworld/d/axolotl.html ]. DELEUZE, J. P. F. 1823. History and description of the Royal Museum of Natural History: published by order of the administration of that establishment / / translated from the French of M. Deleuze; with three plans and fourteen views of the galleries, gardens, and menagerie. Paris, Printed for A. Royer . . . , by L.T. Cellot . . . KNIGHT, D. M. 1977. Zoological Illustration: An Essay towards a History of Printed Zoological Pictures. Folkstone, Eng., Dawson; Hamden, Conn., Archon Books. FIG. 24. Cover of Le Monde LACEPÈDE, B.-G.-É. 1801. Discours sur Illustré published 24 October les établissements publics destinés 1874 showing reptile display à renfermer des animaux vivants, et and inhabitants at Ménagerie connus sous le nom de ménageries; Jardin des Plantes. Credit: proréimpr. dans Oeuvres de L. (1883) vided by Jean-Luc Berthier, 1:106–111. (Lecture about the pub- Jardin des Plantes Archives. 270 lic installations intended to conﬁne living animals, and known by the name of menageries; reprinted in L.’s works (1833), Part 1, p. 106–111). ________ . 1801. La Ménagerie du Muséum national d'histoire naturelle; ou, Les animaux vivants, peints d'après nature, sur vélin, par le citoyen Maréchal, et gravés au Jardin de Plantes, avec l'agrément de l'Administration, par le citoyen Miger. Avec une note descriptive et historique pour chaque animal, par les citoyens Lacépède et Cuvier (The Menagerie of the National Museum of Natural History or its FIG. 25. Cover of Guidebook for repLiving Animals Painted tile building at Ménagerie Jardin des from Nature on Velin by Citizen Maréchal, a paint- Plantes, by Léon-Louis Vaillant in 1898. er of the Museum and En- On the front cover of this guide are graved in the "Jardin des drawings of Lacepède, C. Duméril, A. Plantes" with Agreement Duméril and Vaillant. Credit: provided of the Administration by by Kraig Adler. Citizen Miger, Engraver and Member of the Royal Academy of Paintings, with a Description and Historical Note by the Citizens Lacepède and Cuvier). Miger, Paris. [Bernard-GermainÉtienne de la Ville-sur-Illon, Compte de Lacepède (1756–1825) also published "Histoire Naturelle des Quadrupèdes Ovipares et des Serpens" in 1788–1789.]. KISLING, V. N. JR. (ed.). 2001. Zoo and Aquarium History: Ancient Animal Collections to Zoological Gardens. CRC Press, Boca Raton, Florida. LOISEL, G. 1912. Histoire des ménageries de l'antiquité à nos jours (History of Menageries from Antiquity to Present Times). O. Doin et ﬁls, Paris. MULLAN, B., AND G. MARVIN. 1999. Zoo Culture. Second Edition. University of Illinois Press, Urbana and Chicago IL. [two photographs of reptile cases and crocodilian enclosure.]. MURPHY, J. B. 2007. Herpetological History of the Zoo and Aquarium World. Krieger Publishing Co., Malabar FL. ________ , AND G. ILIFF. 2004. Count de Lacepède: Renaissance Zoo Man. Herpetol. Rev. 35:220–223. OSBORNE, M. A. 1996. Zoos in the family. The Geoffroy Saint-Hilaire clan and the three zoos of Paris, p. 33–42. In R. J. Hoage, and W. A. Deiss (eds.), New Worlds, New Animals. From Menagerie to Zoological Park in the Nineteenth Century. The Johns Hopkins University Press, Baltimore and London. PEEL, C. V. A. 1903. The Zoological Gardens of Europe. Their History and Chief Features. F. E. Robinson & Co., London. PETZOLD, H.-G. 2008. The Lives of Captive Reptiles. Society for the Study of Amphibians and Reptiles. Contributions to Herpetology, volume 22, Ithaca NY. PUJOULX, J. B. 1803. Promenades au Jardin des plantes, a la ménagerie et dans les galeries du Muséum d'histoire naturelle ... Paris, Librairie Économique. (On pp. 105–127, the biology of amphibians and reptiles is discussed.). PYENSON, L., AND S. SHEETS-PYENSON.1999. Servants of Nature. A History of Scientiﬁc Institutions, Enterprises, and Sensibilities. New York, W. Herpetological Review 40(3), 2009 FIG. 27. Indian Python (Python molurus) incubating eggs at Ménagerie Jardin des Plantes. Illustration reproduced from S. G. Goodrich's Johnson's Natural History, Comprehensive, Scientiﬁc, and Popular, Illustrating And Describing The Animal Kingdom With Its Wonders And Curiosities, From Man, Through All The Divisions, Classes, And Orders, To The Animalculae In A Drop of Water; Showing The Habits, Structure, And Classiﬁcation Of Animals, With Their Relations To Agriculture, Manufactures, Commerce, And The Arts. Volume 2 in 1870. Credit: Courtesy of Smithsonian Institution Libraries, Washington, DC. FIGS. 26, 26a. Title page and floorplan of the Ménagerie Jardin des Plantes, by Léon-Louis Vaillant in 1898. This nice drawing of the building's façade shows the bronze statues at center and right of building. Credit: provided by Kraig Adler. H. Norton and Company. RIFKIN, B. A., AND M. J. ACKERMAN. 2006. Human Anatomy (From the Renaissance to the Digital Age). New York, Harry N. Abrams, Inc. ROUSSEAU, L. 1837. Promenades au Jardin des plantes, comprenant la description 1 de la ménagerie ... 2 du cabinet d'anatomie comparée; 3 des galeries de zoologie, de botanique, de minéralogie et de géologie; 4 de l'École de botanique; 5 des serres et du jardin de naturalisation et des semis; 6 de la bibliothèque, etc. / Par MM. Louis Rousseau . . . et Céran Lemonnier . . . Paris, Chez J.B. Baillière. General BONNATERRE, ABBÉ (PIERRE JOSEPH). 1789. Tableau encyclopédique et méthodique des trois règnes de la nature. Erpétologie. Paris, Chez Panckoucke. ________ . 1790. Tableau encyclopédique et méthodique des trois règnes de la nature. Ophiologie. Paris, Chez Panckoucke. DAUBENTON, L.-J.-M. 1784. Les animaux quadrupèdes ovipares, et les serpens. [Paris: s.n.]. DAUDIN, F. M. 1802–1803. Histoire naturelle, générale et particulière, des reptiles; ouvrage faisant suite à l'Histoire naturelle générale et particulière, composée par Leclerc de Buffon, et rédigée par C. S. Sonnini. Paris, Impr. de F. Dufart, an X–Xi [1802–1803]. D'ORBIGNY, M. C. 1848. Dictionnaire universel d'histoire naturelle. zoologie. atlas. reptiles, poissons et insectes. tome deuxiéme (Universal Dictionary of Natural History. Zoology. Atlas. Reptiles, Fishes and Insects.). Paris. DUMÉRIL, A. H. A. 1842. Sur le développement de la chaleur dans les oeufs des serpents, et sur l'inﬂuence attribuée à l'incubation de la mere; par M. Duméril (Concerning the production of heat in the eggs of snakes, and the inﬂuence attributed to the incubation of the mother; by M. Duméril). Comptes Rendus 14:193–210. ________ . 1852. Inﬂuence exercée sur la temperature des Ophidiens par l'échauffement du milieu qu'ils habitant (Effects on the temperature of ophidians by heating their ambient environment). Annales d. Sci. Nat. Zool. 3rd ser.:17–22. ________ . 1854–55. Notice historique sur la ménagerie des reptiles du Muséum d'histoire naturelle et observations qui y ont été recueillies (Historical notice on the menagerie of reptiles of the Museum of Natural History and observations collected there). Archives du Muséum d'histoire naturelle 7:193–320. ________ . 1858–61. Deuxième notice sur La Ménagerie des Reptiles du Muséum d'histoire naturelle. (Second notice). Archives du Muséum d'histoire naturelle (Archives of the Museum of Natural History) 10:429–460. ________ . 1861. Lettres de M. Auguste Duméril professeur d'herpetologie et d'ichthyologie relatives au Catalogue des Poissons de la collection du Museum d'Histoire Naturelle de Paris et au Catalogue de la Menagerie des Reptiles. . . Arch. Mus. Paris, X:427–460. ________ . 1865. Troisième notice sur La Ménagerie des Reptiles du Muséum d'histoire naturelle. (Third notice). Nouvelles archives du Muséum d'histoire naturelle (New archives of the Museum of Natural History) 1:31–50. ________ . 1865. Observations sur la reproduction dans la ménagerie des reptiles du muséum d'histoire naturelle des axolotls batraciens urodèles a branchies extérieurs du Mexique sur leur développement et sur leurs métamorphoses par M. le professeur Aug. Duméril planche 10. Nouv. Arch. Mus. Hist. Nat. 2:265–291. ________ . 1866. Observations faites a la ménagerie du muséum d'histoire naturelle sur la reproduction des axolotls batraciens urodèles a branchies extérieurs et sur les métamorphoses qu'ils y ont subies. par Herpetological Review 40(3), 2009 271 FIGS. 28, 28a. Two illustrations from the papers by Auguste Duméril describing neoteny in Mexican Axolotls (Ambystoma mexicanum). M. le professeur A. Duméril. Extrait du Bulletin de la Societé Impériale d'Acclimatation. (No. de févier 1866.) Paris Imprimerie de E. Martinet Rue Mignon, 2. ________ . 1867. Métamorphoses des batraciens urodèles à branchies externes du Mexique, dits axolotls, observés à la ménagerie des reptiles du Múséum d'histoire naturelle (Metamorphoses of the urodele batrachians with external gills of Mexico, called axolotls, observed in the menagerie of reptiles of the Museum of Natural History). Ann. Sci. nat. 7: 229–354. ________ . 1867. Expériences démontrant que la vie aquatique des axolotls batraciens urodèles a branchies extérieurs se continue, sans trouble apparent, apres l'ablation des houppes branchiales par M. le professeur Aug. Duméril. Mem. Mus. Hist. Natl, Paris:189–192. ________ . 1869–70. Quatrième notice sur La Ménagerie des Reptiles du Muséum d'histoire naturelle. (Fourth notice). Nouvelles archives du Muséum d'histoire naturelle (New archives of the Museum of Natural History) 5:47–60. ________ . 1870. Création d'une race blanche d'Axolotls à la Ménagerie des Reptiles du Muséum d'histoire naturelle, et remarques sur la transformation de ces Batraciens; par m. Aug. Duméril. Institut Impérial de France. Académie des Sciences. Extrait des Comptes rendus des scéances de l' Academie des Sciences, t. LXX, séance du avril:1–4. ________ , M.-F. BOCOURT, AND F. MOCQUARD. 1870. Études sur les reptiles. Published by Impr. nationale, Paris, which was issued as 3. ptie, 1. section of Recherches zoologiques of the Mission scientiﬁque au Mexique et dans l'Amérique centrale. DUMÉRIL, A. M. C., G. BIBRON, AND A. DUMÉRIL. 1854. Erpétologie générale, ou, Histoire naturelle complète des reptiles. Atlas. Roret, Par- 272 is. DUVERNOY, G.-L. 1842. La Règne Animal par Georges Cuvier, Reptiles volume. Disciple edition. Paris. FLOWER, S. S. 1925. Contributions to our knowledge of the duration of life in vertebrate animals.– II. Batrachians. Proc. Zool. Soc. London 1925:269–289. ________ . 1925. Contributions to our knowledge of the duration of life in vertebrate animals. –III. Reptiles. Proc. Zool. Soc. London 1925:911– 981. ________ . 1936. Further notes on the duration of life in animals. II. Amphibians. Proc. Zool. Soc. London 1936:369–394. ________ . 1937. Further notes on the duration of life in animals. III. Reptiles. Proc. Zool. Soc. London 1937, 107 (ser. A):1–39. GEOFFROY SAINT-HILAIRE, E. 1802. Mémoires d'histoire naturelle / / par E. Geoffroy. Imprint: Paris : Baudouin, Imprimeur de l'Institut national, An XI (1802). ________ . 1802–1810. Résumé sur quelques conditions générales des rochers: et la spécialité de cet organe chez le crocodile / / Par M. Geoffroy-S.-Hilaire. Imprint: [Paris : s.n., between 1802 and 1810]. ________ . 1803?. Observations anatomiques sur le crocodile du Nil. Imprint: [Paris : s.n., 1803?]. ________ . 1809. Mémoire sur les tortues molles, nouveau genre sous le nom de trionyx, et sur la formation des carapaces / / Par M. Geoffroy-SaintHilaire. Imprint: [Paris : s.n., 1809?] ________ . 1835. Études progressives d'un naturaliste, pendant les années 1834 et 1835; / faisant suite a ses publications dans les 42 volumes des mémoires et annales du Museum d'histoire naturelle, par Geoffroy Saint-Hilaire (Étienne). Imprint: Paris, Roret, 1835. ________ , AND I. GEOFFREY SAINT-HILAIRE. 1827. Description des reptiles qui se trouvent en Égypte, pages 115–160, 8 plates. In Description de l'Égypte, ou Recueil des Observations et des Recherches qui ont été faites en Égypte pendant l'Expédition de l'Armée Française. Histoire Naturelle. Tome Premier [Volume 1, part 1] (Description of Reptiles Found in Egypt, pages 115–160, 8 plates. In Description of Egypt, or Collection of Observations and Research Performed in Egypt during the Expedition of the French Army. Natural History. Volume One.). Imprimerie Impériale, Paris. GUICHENOT, A. 1850. Histoire naturelle des reptiles et des poissons / par A. Guichenot. Exploration scientiﬁque de l’Algérie pendant les années 1840, 1841, 1842. Sciences physiques. Zoologie; 5. Paris, Imprimerie nationale. KOENIG, R. 2008. Sanctuaries aim to preserve a model organism’s wild type. Science 322:1456–1457. LACEPÈDE, B.-G.-É. 1788–1789. Histoire naturelle des quadrupèdes ovipares et des serpens. Paris, Hôtel de Thou. LESSON, R. P. 1832–1835?. Illustrations de zoologie, ou Recueil de figures d'animaux peintes d'après nature; / par R.-P. Lesson . . . Ouvrage orné de planches dessinées et gravées par les meileurs artistes . . . (Zoological Illustrations, or Collection of Animal Figures Painted from Nature, by R.-P. Lesson . . . Work Decorated with Plates Designed and Engraved by the Best Artists . . .). A. Bertrand, Paris. SAGRA, R.. DE LA. 1839–1856. Historia fisica, politica y natural de la isla de Cuba. Paris, En libreria de Arthus Bertrand ... SONNINI, C.-N.-S.. 1801. Histoire Naturelle des Reptiles / par C.S. Sonnini et P.A. Latreille. Paris, Imprimerie de Crapelet, An X-[1802]. VAILLANT, L.-L. 1878. Observations anatomo-pathologiques faites sur une Platemys Macquaria, Cuv. Bull. Soc. Philom. Paris 7 ser.2:14– 17. ________ . 1880. Note sur la ponte du Pleurodèle de Waltl observée à la Ménagerie des Reptiles du Muséum d'Histoire naturelle. Bull. Soc. Philom. Paris (ser. 7) 4(3):127–129. ________ . 1880. Sur la ponte et le développement du Pleurodeles Waltlii, Mich. Observés à la Ménagerie des reptiles du Muséum d'histoire naturelle. Extrait du Bull. Societé d'Acclimitation, pp.1–3. ________ . 1881. Note sur un appariel destiné au transport des batrachiens Herpetological Review 40(3), 2009 anoures vivants. Bull. Societé d'Acclimitation 8(3 série):191–193. . 1892. Contribution a l'etude de l'alimentation chez les ophidiens. Recherches Biol. Menag. Reptiles 1:221–223. ________ . 1896. Sur le mode de formation des coprolithes helicoides, d'apres les faites observes a la Menagerie des Reptiles sur les Protopteres. C. R. Acad. Sci. Paris 122:1–2. ________ . 1898. Muséum d'histoire naturelle guide à la ménagerie des Reptiles. Laboratoire d'Herpétologie, Paris. ________ . 1905. Remarques sur le developpement d'une jeune tortue charbonniere (Testudo carbonaria Spix), observee a la menagerie des Reptiles du Museum d'Histoire naturelle. Bull. Mus. Nat. Hist. (3):139– 141. ________ . 1910. La Ménagerie des reptiles au 31 Décembre 1909. Bull. Mus. National d'Hist. Nat. Paris, no. 1:11–13. ________ , AND G. GRANDIDIER. 1910. Histoire physique, naturelle, et politique de Madagascar. Impr. Nationale, Paris. ________ , AND A. PETTIT. 1902. Lésion stomacales observées chez un Python se Séba. Extrait du Bulletin du Muséum d'histoire naturelle, no.8, p.593–595. ________ , AND ________. 1902. Fibrome observé sur un Megalobatrachus maximus, Schlegel, à la ménagerie du Muséum. Extrait du Bulletin du Muséum d'histoire naturelle. No.5:301–305. VALENCIENNES, A. 1841. Observations faites pendant l'incubation d'une femelle du Python à deux raies (Python bivittatus, Kuhl.) pendant les mois de mai et de juin 1841; par M. Valenciennes (Observations made during incubation of a female python on two occasions in (Python bivittatus, Kuhl.) during the months of May and June 1841; by M. Valenciennes). Comptes Rendus 13:126–133. ________ Amplectant pair of Agalychnis moreletii (Morelet’s Treefrog), Belize. Illustration by Peter Stafford. LETTERS TO THE EDITOR Herpetological Review, 2009, 40(3), 273–275. © 2009 by Society for the Study of Amphibians and Reptiles The State of Natural History: A Perspective from the Literature on West Indian Herpetology ROBERT W. HENDERSON Vertebrate Zoology, Milwaukee Public Museum Milwaukee, Wisconsin 53233-1478, USA e-mail: henderson@mpm.edu and ROBERT POWELL Department of Biology, Avila University Kansas City, Missouri 64145, USA e-mail: robert.powell@avila.edu Natural History: The study and description of organisms and natural objects, especially their origins, evolution, and interrelationships.1 Deﬁnitions of natural history vary, with differences attributable to changing historical perspectives (what we now call “biology” was referred to as natural history until the early 20th century) and motive (either laudatory or derogatory). Recent years have seen a spate of papers addressing the “state” of natural history, whether it’s dying or healthy, exactly what encompasses natural history, and what is “good” or “contemporary” natural history. Arnold (2003) and Greene (2005) provided effective, albeit somewhat contradictory, perspectives. Some would suggest that natural history has a long and storied heritage, embodied by great 19th-century naturalists like Alexander von Humboldt and Charles Darwin and their modern-day counterparts like Henry Fitch and John Iverson, whose detailed multi-year studies of the lives of reptiles are primary references for anyone studying those species. In contrast, others propose that natural history today has little relevance beyond a small role in more “conceptual” investigations of our natural world. WordNet (Cognitive Science Laboratory, Princeton University) deﬁnes natural history as “the scientiﬁc study of plants or animals (more observational than experimental) usually published in popular magazines rather than in academic journals.” Detailed observations of natural history may be deemed oldfashioned and out of vogue—but they will be as valid 100 years from now as on the day they were published. Not surprisingly, Fitch (1987) stated this quite clearly: “When an observer is fortunate enough to see and record behavior signiﬁcant in the natural history of a species, his observations should be published.... Even a single observation may constitute a valuable contribution and may be a break-through in understanding the species’ ecology.” Although we are not particularly interested in entering the debate regarding the state or status of natural history, we do have an interest in the subject—in fact, curiosity regarding the lives of animals in their natural (and often altered) habitats led us to our life-long careers—and, based on our recent experience in accu1 http://dictionary.reference.com/browse/Natural+history?jss=1 Herpetological Review 40(3), 2009 273 FIG. 1. The number of publications produced by decade (1740–49 ... 2000–09) that include some natural history information relevant to West Indian amphibians and reptiles. The 1910s and 1920s largely represent the efforts of Thomas Barbour (with E. R. Dunn and K. P. Schmidt contributing in the 1920s); Barbour remained active in the 1930s. The drop-off in the 1940s can be attributed to World War II, although Chapman Grant was active at this time. The 1950s mark the initiation of studies by Albert Schwartz and Ernest Williams in the region. Interest in Antillean herpetology blossomed in the 1960s with Orlando Garrido, James Lazell, A. Stanley Rand, Rodolfo Ruibal, Schwartz, Richard Thomas, and Williams as major contributors. By the 1970s, an ecological focus began to share the spotlight with alpha taxonomy through the contributions of Thomas Schoener, Judy Stamps, and others. Research activity has continued to increase in the region to the present day, with major emphases on conservation, ecology, and phylogenetics, with biologists who live on the islands as well as researchers from North America and Europe making valuable contributions. The number of contributions published subsequent to the previous compendium on West Indian herpetology (Schwartz and Henderson 1991) rivals the number published during the previous 250 years. mulating information for our book on the natural history of West Indian reptiles and amphibians (Henderson and Powell 2009), “natural history” data are being published in a staggering number of journals with an incredibly broad range of foci. Evidently, a great many people remain interested in and are practitioners of natural history. Background.—In 2005, we proposed to the University Press of Florida a book manuscript summarizing the natural history of each of about 700 species of West Indian frogs and reptiles. We soon thereafter signed a contract agreeing to deliver a 500-page manuscript in December 2007. We both have long histories of working in the West Indies and, we thought, a fairly intimate knowledge of the literature pertaining to its herpetofauna. We were mistaken. By the time we submitted the ﬁnal version of the manuscript in February 2009 (after reviews and copyediting), it had expanded to over 1800 pages covering 737 species and we had cited more than 2600 sources of information. Those 2600+ citations were gleaned from 317 different sources, including 74 books, 180 274 book chapters, and more than 300 different scientiﬁc journals and publication series. Details of the natural history of Antillean frogs and reptiles came from many obscure (to us) sources, including, for example, the Journal of Aviation/Aerospace Education and Research, Insectes Sociaux, Letters in Applied Microbiology, and Peptides. Pertinent literature came from publications originating in nations far removed from the West Indies, including Japan, Russia, and Czechoslovakia. We cited publications that appeared between 1740 and 2009 (Fig. 1), but only 18 publications with some germane natural history data appeared in the 1800s, and another 16 during the ﬁrst two decades of the 20th century. The numbers began to increase in the 1920s and 30s with the contributions of Thomas Barbour, Emmett R. Dunn, Gladwyn K. Noble, and Karl P. Schmidt. The 1940s showed a decline in relevant publications, almost certainly attributable to WWII. Albert Schwartz and Ernest E. Williams became active in the islands in the 1950s, and this is reﬂected in the resurgence of activity throughout the Caribbean Region. The 1960s are, we believe, the real turning point for Herpetological Review 40(3), 2009 interest in West Indian herpetology. Schwartz and Richard Thomas, and Williams and his students (most notably A. Stanley Rand) were amazingly proliﬁc, and the number of contributions more than tripled between the 1950s and 60s. The number of contributions containing some natural history information pertinent to the West Indian herpetofauna has steadily increased each decade, with more than 1200 appearing since 1990. Results.—Henderson and Powell (2009) included 737 species accounts, varying in length from a few lines to many pages. In order to make the accounts as complete and detailed as possible, we sought to include data from all of the relevant sources we could locate, regardless of whether the project from which the published information came was systematic in nature, hypothesis-driven and presented in a conceptual context, or was based on an anecdotal observation. We did not consider the impact factor of journals from which we extracted information. If information was available, no matter how obscure the publication outlet, we felt it was our responsibility to ﬁnd it (a process that often entailed considerable help from many colleagues throughout the world). We determined that studies dedicated to some aspect of natural history2 have addressed 43% of West Indian species. “Dedicated” implies an a priori plan to study some facet of natural history, and that the observations were not based solely on fortuitous occurrences. This by no means suggests that we have a good handle on the natural history of 43% of the Antillean herpetofauna. To the contrary, we have a fairly broad knowledge of only about 5% of West Indian species. The most frequently cited publication in our book is the IUCN Red List of Threatened Species (268 citations; with the vast majority on frogs, reﬂecting the 2004 Global Amphibian Assessment); the fourth most was the Catalogue of American Amphibians and Reptiles (147). Because neither of these generally contains original data, we omit them from our list of the top 10 sources of natural history information pertaining to the West Indian herpetofauna: (1) Herpetological Review — 156 citations, (2) Caribbean Journal of Science — 151, (3) Journal of Herpetology — 107, (4) Copeia — 89, (5) Herpetologica — 88, (6) Iguana (now Reptiles & Amphibians) — 69, (7) Breviora (Museum of Comparative Zoology) — 53, (8) Poeyana (Cuba) — 52, (9–tie) Ecology and Iguana Specialist Group Newsletter — 31, and (10) Bulletin of the Museum of Comparative Zoology — 28. Most of the top ten was not a surprise. These publications share a willingness (at least historically) to accept and publish studies involving descriptive natural history. Herpetological Review (HR) devotes a section to brief natural history observations in 2 Our categories of “natural history” included (obviously not all were applicable to all species): abundance, activity, behavior, biomass, burrows (retreats), commensalism, competition, conservation status, defense, desiccation, diet and foraging, disease, distribution, ecomorphology (anoles), ecosystem processes, energetics, growth, habitat, home range, homing, hurricane effects, hybridization, injuries, interspeciﬁc (or interclass) interactions, life expectancy, life history, longevity, miscellany, mortality, movement, nesting, parasites, performance, population size and density, population structure, predation, reproduction, salinity tolerance, sex ratios, site ﬁdelity, size, source of introduced populations, survival, swimming, tail autotomy, territoriality, thermal biology, vision, vocalization. each issue, and the Caribbean Journal of Science (CJS), based in Puerto Rico, is the most important biological journal published in the West Indies. The Journal of Herpetology (JH), although the youngest of the three most inﬂuential herpetological journals in North America, provided the most citations, but the number did not differ all that much from those in Copeia or Herpetologica. Poeyana is published in Cuba, and most of its contributions appeared in the 1960s–1980s; today, Cuban herpetologists publish primarily in “mainstream” journals, including HR, CJS, JH, and others in the top ten. The Museum of Comparative Zoology (MCZ) at Harvard University has a long (>125 years) history of West Indian ﬁeldwork and publications focused on Antillean herpetology. Curators Samuel W. Garman, Thomas Barbour, Ernest E. Williams, and, currently, Jonathon B. Losos have been active in the Caribbean. That the MCZ was the institution that produced the most citations was predictable (87 including six Memoirs of the MCZ). Its closest competitor was the American Museum of Natural History with 20. In the West Indies, Cuban herpetologists were far and away the most proliﬁc publishers, often in one of several institution-based publication series. Based on the Journal Citation Reports, the mean impact factor (a measure of the frequency with which the “average article” in a journal has been cited in a given period of time) for the four publications in our top 10 that were listed in the Reports (HR, CJS, Iguana, Breviora, Poeyana, and the Bulletin of the Museum of Comparative Zoology were not listed) was 1.620 ± 0.807 (range = 0.527–4.822). By comparison, the average of the top 10 publications (based on impact factor) that we cited, but that did not necessarily make the top 10 of the most frequently cited, was 11.219 ± 2.974 (range = 3.934–28.751). The mean number of times the journals in our top 10 were cited was 82.7 ± 14.7 (range = 31–156), and the mean for the top 10 with the highest impact factors was 9.7 ± 2.8 (range = 2–31). Only one publication (Ecology) made both top 10 lists, undoubtedly reﬂecting the reality that high-impact journals are more likely to include review articles, be interdisciplinary (i.e., not focus on a particular geographic region or taxonomic group), and emphasize topics more “conceptual” than descriptive natural history. Despite our efforts to be as complete as possible, we are certain that some citations have been missed. Nevertheless, in addition to expanding our own awareness of what we know (and don’t know) about many West Indian species, we developed a broader appreciation for the scope of “natural history” and how many aspects of the subject are widely integrated into studies that range far beyond descriptions of how animals make a living in their natural (and sometimes unnatural) habitats. LITERATURE CITED ARNOLD, S. J. 2003. Too much natural history, or too little? Anim. Behav. 65:1065–1068. FITCH, H. S. 1987. The sin of anecdotal writing. Herpetol. Rev. 18:68. GREENE, H. W. 2005. Organisms in nature as a central focus for biology. Trends Ecol. Evol. 20:23–27. HENDERSON, R. W., AND R. POWELL. 2009. Natural History of West Indian Reptiles and Amphibians. University Press of Florida, Gainesville. SCHWARTZ, A. AND R. W. HENDERSON. 1991. Amphibians and Reptiles of the West Indies: Descriptions, Distributions, and Natural History. University of Florida Press, Gainesville. Herpetological Review 40(3), 2009 275 ARTICLES Herpetological Review, 2009, 40(3), 276–282. © 2009 by Society for the Study of Amphibians and Reptiles The Taxonomic Status of the Inornate (Unstriped) and Ornate (Striped) Whiptail Lizards (Aspidoscelis inornata [Baird]) from Coahuila and Nuevo León, México JAMES M. WALKER* Department of Biological Sciences, University of Arkansas Fayetteville, Arkansas 72701, USA e-mail: jmwalker@uark.edu JAMES R. DIXON Department of Wildlife and Fisheries Sciences, Texas A & M University College Station, Texas 77843, USA RALPH W. AXTELL Department of Biological Sciences, Southern Illinois University at Edwardsville Edwardsville, Illinois 62026, USA and JAMES E. CORDES Division of Sciences, Louisiana State University at Eunice Eunice, Louisiana 70535, USA *Corresponding author The Little Striped Whiptail (Aspidoscelis [Cnemidophorus] inornata) includes 10 (Wright and Lowe 1993) or 11 (Reeder et al. 2002) subspecies ranging from Arizona, New Mexico, and Texas in the USA, south into Chihuahua, Coahuila, Durango, Nuevo León, San Luis Potosí, and Zacatecas States in México. Although usually faintly to vividly striped dorsally (ornate) over its vast range, individuals and/or populations of inornate (unstriped) A. inornata occur at a minimum of one locality in Chihuahua (Cordes and Walker 1996; Walker et al. 1996) and nine in Nuevo León (Axtell 1961; Wright and Lowe 1993; this study). The genetic basis and taxonomic implication of an inornate morph in Nuevo León has been the subject of considerable debate, dating back more than 150 years, with different whiptail investigators drawing conﬂicting conclusions (Axtell 1961; Baird 1858; Reeder et al. 2002; Wright and Lowe 1993). We discuss taxonomic issues pertaining to A. inornata in the context of a widely distributed polytypic species (Liner 2007; Liner and Casas-Andreu 2008; Wright and Lowe 1993) in which there are numerous unresolved problems pertaining to the signiﬁcance of variation in color pattern (Reeder et al. 2002; Walker et al. 1996; Wright and Lowe 1993). There also exists an alternative classiﬁcation of populations of “Little Striped Whiptail” in the United States based on the evolutionary species concept. Collins (1997) and Crother (2000, 2008) listed three color-pattern variants (i.e., subspecies sensu Wright and Lowe 1993) from New Mexico and Arizona as species (i.e., A. arizonae, A. gypsi, and A. pai) based on apparent allopatric distributions (Wright and Lowe 1993; p. 134) and morphological diagnosibility. Using these criteria, a minimum of three subspecies recognized by Wright and Lowe (1993) from México also could be elevated to species rank (i.e., A. cienegae, A. chihuahuae, and A. paulula). The genetic 276 contributions of A. inornata to the origins of no fewer than two diploid (i.e., A. innotata and A. neomexicana) and six triploid (i.e., A. exsanguis, A. ﬂagellicauda, A. opatae, A. sonorae, A. uniparens, and A. velox) hybrid-derived parthenogenetic species have been discussed by Densmore et al. (1989a, b) and Wright (1993). This study pertains to what Baird (1858) described as two species of whiptail lizards from Pesquería Grande (= Villa de García = García), Nuevo León. He named Cnemidophorus inornatus to reﬂect a “...light greenish olive...” dorsum in one, and C. octolineatus in allusion to “...eight equidistant and approximated light lines” on the dorsum of the other (Baird 1858; p. 255). Opinions differ on whether C. octolineatus should be treated as a synonym of C. inornatus (Axtell 1961; Burger 1950; Walker et al. 1996; Williams 1968) or whether these two represent diagnosable taxa (Reeder et al. 2002; Wright and Lowe 1993). That the different color patterns in these otherwise similar forms represent dimorphism within a subspecies is based on Axtell’s (1961) report of their coexistence and intimate relationship at a site at Villa de García. The case for partitioning C. i. inornatus into two taxa is based on Wright and Lowe’s (1993) report of an association of lizards at a different site near Villa de García consisting of inornate and ornate individuals and their possible intergrades or hybrids. Wright and Lowe (1993; p. 136) concluded that “...the possibility that two distinctive taxa (species or subspecies) exist in the vicinity of the type locality is by no means eliminated [by available data].” Most recently, Reeder et al. (2002) placed all species of Cnemidophorus, except those in the paraphyletic “C. lemniscatus” species group, into the genus Aspidoscelis. However, without comment, they allocated inornate lizards from Nuevo León, México, into Aspidoscelis inornata inornata (= Cnemidophorus inornatus Baird) and revived A. i. octolineata (= C. octolineatus Baird) from the synonymy of C. i. inornatus to accommodate ornate lizards from Coahuila and Nuevo León. Herein, we (1) present data for variation in ornate lizards from Coahuila, (2) data for both ornate and inornate individuals from Nuevo León, (3) list new locality records for inornate lizards from Nuevo León that support recognition of A. i. octolineata as a valid taxon, as listed by Reeder et al. (2002), and (4) redeﬁne A. i. inornata. We studied the inornate and inornate specimens of Little Striped Whiptail (A. inornata), listed by Axtell (1961) and Wright and Lowe (1993) from the vicinity of Villa de García in the course of a reassessment of their distributional, morphological, and taxonomic relationships. We also examined other samples from Coahuila and Nuevo León for the presence of the inornate form. Specimens were from the following collections (institutional acronyms follow Leviton et al. [1985]): LACM, TCWC, TNHC, UAZ, UCM, USNM (specimens referenced from the literature), UTEP (specimen transferred from another collection), and R. W. Axtell’s personal collection (RWA). Each collection site for A. inornata was assigned a code consisting of C (= Coahuila) or NL (= Nuevo León) and a number (i.e., C-1–C-24 and NL-1–NL-17; Appendix 1). Two type specimens (USNM 3032A [lectotype] and 3032B [paralectotype]; Baird 1858; Wright and Lowe 1993) from the type locality, six lizards from NL-1 (Axtell 1961), and nine from NL-2 (Wright and Lowe 1993) have previously served as vouchers for inornate A. inornata from Nuevo León. We studied 42 additional inornate specimens from the state, which were held in LACM, RWA, and TCWC. These specimens were from NL-2 Herpetological Review 40(3), 2009 (10 inornate [IN] and 48 ornate [OR] lizards), northwest of the type locality (four IN and one OR at NL-3 and one IN and no OR at NL-4), and sites accessed by the part of México Highway 53 extending northwest from Mina to the Nuevo León–Coahuila border (27 IN and no OR at NL-5–9). We used Google Earth (www.earth.google.com) to locate collection sites with known coordinates, scan topography, determine elevations, and measure line distances between sites from which A. i. inornata has been collected in Nuevo León. However, the exact spatial relationship of the sites originally visited by RWA (NL-1) and by J. W. Wright and associates (NL-2) could not be determined as no coordinates were included in the catalog entries for Wright’s UAZ and LACM samples. We used the Federal Communications Commission website (www.fcc.gov/mb/audio/bickel/DDDMMSS-decimal.html) to convert data accompanying museum samples for degrees, minutes, and seconds to decimal degrees latitude and longitude. Meristic characters (terminology after Burt 1931; Smith 1946; Walker et al. 1996; Wright and Lowe 1993; Tables 1, 2) included counts of: (1) granules (= scales) at midbody from the right outer row of ventral scales over the body to the left outer row of ventral scales (GAB), (2) dorsal granules longitudinally from the occipitals to the ﬁrst row of caudals (OR), (3) granules between the paravertebral stripes at midbody (PV; only possible for ornate lizards), (4) percent of granules around midbody between paravertebral stripes (PV/GAB x 100; only possible for ornate lizards), (5) femoral pores (FP, summed from both sides), (6) subdigital lamellae of the fourth toe of the left pes (SDL), (7) circumorbitals (COS; summed from both sides), (8) lateral supraocular granules (LSG; summed from both sides), and (9) interlabials (ILS; summed from both sides). Each specimen of A. inornata was sexed (via inspection of femoral pores and cloacal region), and its snout–vent length (SVL) measured to the nearest mm (Table 1). We used ANOVA in JMP v. 7 (SAS Institute, Cary, NC) to generate a mean ± 1 SE for SVL pooled by sex, and for each univariate character of scutellation in each geographical and pooled sample of A. inornata (Tables 1, 2). We selected the treatment “All PairsTukey-Kramer HSD” to determine statistical differences between any two means. This procedure yields post-hoc comparisons, which preserves alpha (P < 0.05) in statistical comparisons of multiple means (Sokal and Rohlf 1981). Chi-square values were computed using SPSS v. 14.1 (SPSS, Chicago, IL) to separately compare ratios of numbers of dorsal color pattern variants designated A–D for the 1962 UAZ and 1975 LACM samples of A. inornata from site NL-2 in Nuevo León. Aspidoscelis i. inornata lacks dorsal stripes and spots (Fig. 1A, B, C). In contrast, A. i. octolineata has 7–11 pale longitudinal dorsal stripes and darker intervening ﬁelds; each side of the body (ventral to dorsal) has a lateral, a dorsolateral, and a paravertebral primary stripe (Fig. 1D, J, L). The middorsal region between the paravertebral stripes in each ornate lizard has 1–4 additional supernumerary stripes (sensu Wright and Lowe [1993], if more than the usual number of two). We used “7+ stripes” to denote the presence of an intermittently joined and split vertebral stripe between the head and tail of a lizard. In ornate A. inornata, the dark ﬁelds between the stripes always lack spots; however, certain ﬁelds may develop an indistinct longitudinal line (supernumerary stripes, sensu Walker et al. [1996]) between the primary stripes. Axtell (1961) described an area ca. 50 × 100 m of wind-formed sand mounds just under 3 m high at NL-1, referenced by him as the type locality of A. inornata, from which he collected six inornate and ﬁve ornate specimens (three ornate lizards were obtained approximately 100 m to the southeast among mounds less that 0.5 m high). He described each of the inornate adults from NL-1 as having a “...completely unicolor brownish olive...” dorsum and each of the ornate adults as being “...distinctly striped with eight light lines on a brownish gray to olive background...” (Axtell 1961; p. 149). None of these specimens was intermediate in color pattern between the inornate and ornate morphotypes. A return to NL-1 by RWA in the 1960s revealed that the area previously inhabited by inornate and ornate A. inornata had been destroyed by construction activities. Wright and Lowe (1993) described a differently constituted association of whiptail lizards at site NL-2 located 1.6 km west and 1.6 km south of Villa de García, where inornate and ornate A. inornata and apparent intergrades (or hybrids) were collected. We agree that adults UAZ 14067–75 are inornate (our variant A; 29.0%) and that adults UAZ 14076–88, 14093–95, 14097–102 are ornate (our variants B–D; 71.0%), the latter having stripes ranging from distinct to indistinct, straight to irregular, and with or without supernumerary stripes. Among the inornate specimens are four females (UAZ 14067, 14069, 14072, 14075; 54–60 mm SVL) and ﬁve males (UAZ 14068, 14070–71, 14073–74; 55–60 mm SVL). Among the 22 ornate specimens in the UAZ sample are four females (UAZ 14078, 14080, 14083, 14095; 57–61 mm SVL) and one male (UAZ 14099; 60 mm SVL) (designated variant B; 22.7%) with gray-tan ﬁelds of only slightly darker hues than the stripes; four of these individuals have 9–10 stripes rather than the usual number of eight, and in some the stripes are atypical. In the ornate group from NL-2 designated variant D (50%), six females (UAZ 14077, 14079, 14082 [Wright and Lowe (1993, ﬁgure 7)], 14093, 14097–98; 58–62 mm SVL) and ﬁve males (UAZ 14081, 14085, 14087 [Wright and Lowe (1993, ﬁgure 7)], 14094, 14101; 56–60 mm SVL) have the pattern of A. i. octolineata comparable to specimens from NL-12, in which dark ﬁelds strongly contrast with seven (one specimen), eight, or nine pale stripes. Last, in the ornate group from NL-2 designated variant C (27.3%) are three females (UAZ 14076, 14086, 14088; 56–58 mm SVL) and three males (UAZ 14084, 14100, 14102; 58–61 mm SVL) with dorsal patterns of intermediate contrast between the ﬁelds and stripes. A large unreported sample of A. inornata collected in LACM from NL-2 in 1975 includes seven (14.0%; LACM 116195–201; Fig. 1A, B, C) inornate and 43 (86.0%; LACM 116152–94) ornate specimens of A. inornata. Among the ornate lizards are six (14.0%, Fig. 1D, E, F) of variant B, nine (20.9%, Fig. 1G, H, I) of variant C, and 28 (65.1%, Fig. 1J, K, L) of variant D. Another unreported LACM sample from NL-2 collected in 1980 includes three (37.5%) inornate and ﬁve (62.5%) ornate specimens of A. inornata. The variation in color pattern described above in samples from NL-1 and NL-2 is not attributable to either sexual dimorphism or ontogenetic development; both males and females over a similar SVL range in both groups have the patterns described. We concur with Wright and Lowe (1993) that some of the variation observed among the ornate specimens from NL-2 likely reﬂects gene exchange between inornate and ornate lizards. The disparate ratios of inornate to ornate lizards in the 1962 (9:31, χ2 = 5.452, P = 0.02) and 1975 (7:44, χ2 = 26.843, P = 0.0005) samples from NL-2 are Herpetological Review 40(3), 2009 277 TABLE 1. Meristic characters (see text for definitions of abbreviations) of Aspidoscelis inornata from Nuevo León, México. Data are (ﬁrst row) mean ± SE, (second row) range and (N) and tabulated by site code (see text for explanation), color pattern variant (IN = inornate A. i. inornata), OR = ornate A. i. octolineata), and museum collection. Character NL-2 (IN) UAZ, LACM NL-2 (OR) UAZ, LACM NL-3 (IN) TCWC NL-5 (IN) TCWC NL-11 (OR) TCWC NL-12 (OR) UCM 71.9 ± 0.74 66–75 (15) 70.9 ± 0.45 64–81 (65) 69.5 ± 1.66 66–74 (4) 69.9 ± 0.77 64–80 (21) 70.8 ± 0.98 66–72 (6) 72.9 ± 1.18 70–77 (7) OR 179.5 ± 2.31 164–191 (15) 183.7 ± 1.20 160–205 (65) 171.5 ± 4.87 160–181 (4) 181.0 ± 1.62 166–191 (21) 173.3 ± 4.59 154–186 (6) 185.1 ± 2.76 176–195 (7) PV — — 16.6 ± 0.23 13–21 (65) — — — — 17.5 ± 0.85 14–20 (6) 14.9 ± 0.61 13–18 (8) PV/GAB — — 23.5 ± 0.33 17.8–30.4 (65) — — — — 24.6 ± 0.93 21.0–27.7 (6) 20.7 ± 0.67 18.3–23.3 (7) FP 34.0 ± 0.51 31–37 (15) 34.6 ± 0.32 28–40 (65) 36.0 ± 0.41 35–37 (4) 36.9 ± 0.56 33–44 (21) 34.8 ± 1.42 29–39 (6) 34.8 ± 0.75 32–38 (8) SDL 29.8 ± 0.46 27–34 (15) 29.6 ± 0.20 26–33 (65) 30.8 ± 0.25 30–31 (4) 31.0 ± 0.29 28–33 (21) 31.7 ± 0.33 30–32 (6) 32.3 ± 0.49 30–35 (8) COS 12.3 ± 0.53 9–15 (15) 12.3 ± 0.26 8–18 (65) 11.0 ± 1.58 7–14 (4) 11.7 ± 0.42 9–15 (21) 10.7 ± 1.02 6–13 (6) 11.5 ± 0.87 9–16 (8) LSG 26.1 ± 1.74 18–41 (15) 26.5 ± 0.66 17–45 (65) 25.8 ± 4.30 19–37 (4) 29.3 ± 1.55 17–49 (21) 26.3 ± 1.63 21–31 (6) 31.3 ± 3.02 22–47 (8) ILS 25.4 ± 1.30 18–38 (15) 25.7 ± 0.67 16–41 (65) 25.8 ± 2.43 21–32 (4) 27.5 ± 1.21 17–41 (21) 25.2 ± 2.61 15–33 (6) 31.4 ± 2.46 23–44 (8) GAB not likely explained by chance or sampling effort. The 1975 sample also includes fewer ornate individuals with unusual stripe characteristics than were present in the 1962 sample. Although the 1962 sample has statistically indistinguishable frequencies of the three ornate variants represented in Fig. 1 (χ2 = 2.818, P = 0.24), this was not the case for the 1975 sample (χ2 = 21.318, P = 0.0005). In Nuevo León, inornate lizards were previously known from two sites at Villa de García, both also inhabited by ornate lizards (Axtell 1961; Wright and Lowe 1993) and one with intergrades or hybrids of the two (Wright and Lowe 1993). We describe a distribution area for the inornate variant in Nuevo León of larger size than those of subspecies of Little Striped Whiptail in México (e.g., A. i. cienegae) and the United States (e.g., A. i. gypsi) (Wright and Lowe 1993, p. 134). This area in Nuevo León extends from Villa de García and La Paz thence north–northwest to southeast of Soledad (spanning 25.8167–26.3333ºN latitude, 100.5833–100.8333ºW longitude, and 693–930 m elevation). The linear distance between the most widely separated sites for inornate lizards (i.e., A. i. inornata) in Nuevo León is ca. 65 km from NL-2 to NL-9. In addition to the two sites noted, both inornate (TCWC 44262–65) and ornate (TCWC 44261) individuals of A. inornata also occur at a third site (NL-3), located 14.1 km NW of Villa de García, Nuevo León. Two juvenile males from NL-3 (TCWC 44263 and 44264) are the smallest (36 and 39 mm SVL, respectively) inornate lizards known to us. In ethanol, both are similar to adults in that they lack stripes and have a gray-brown dorsum. One male (TCWC 44266) from NL-4 at 1.6 km west of La Paz establishes 278 the presence of the inornate variant at a fourth site in the general area of Villa de García. All other samples of inornate lizards were collected well north of Villa de García, the largest sample consisting of 13 adult males (TCWC 43591–92, 43595–98, 43600–01, 43603–04, 43606, 43610–11; 54–59 mm SVL) and eight adult females (TCWC 43593–94, 43599, 43602, 43605, 43607–09; 51–64 mm SVL) from NL-5 at 8.2 km west of Mina. Specimens of A. marmorata with strongly contrasting dorsal patterns also occur at NL-5 and nearby (Hendricks and Dixon 1986). Only inornate specimens of A. inornata are present in samples from NL-6 (N = 1), NL-7 (N = 1), NL-8 (N = 3), and NL-9 (N = 1), which are between 8.2 km northwest of Mina and 17.8 km southeast of Soledad over a road distance of about 42 km. Two samples containing ornate lizards from 0.6 km southeast of Soledad (NL-10, N = 1) and 4.0 km northwest of Soledad (NL-11, N = 6) suggest that A. i. octolineata replaces A. i. inornata to the northwest of NL-9. All 143 specimens of A. i. octolineata that we examined from 24 sites in southwestern Coahuila (Appendix 1) have ornate patterns consisting of 7+ (27.2%), 8 (38.1%), or 9 (34.5%) stripes. No statistical differences were found between the pooled mean SVL for each of inornate females, inornate males, ornate females, and ornate males of our sample of 189 adult A. inornata from Coahuila and Nuevo León (Table 2). Likewise, we found no signiﬁcant differences in scutellation (excluding PV and PV/GAB) between samples of inornate lizards collected at NL-2 in 1962 versus those from 1975 and 1980, between samples of ornate lizards collected at NL-2 in 1962 versus those from 1975 and 1980, or between all Herpetological Review 40(3), 2009 TABLE 2. Summary of meristic characters and SVL (see text for definitions of abbreviasamples of inornate and ornate individuals from NLtions) for Aspidoscelis inornata inornata (inornate) from Nuevo León and A. i. octolineata 2 (Table 2). We suspect that the greater variation in (ornate) from Nuevo León and Coahuila, México. Data are mean ± SE (ﬁrst row) and range GAB, OR, FP, COS, LSG, and ILS in ornate relative and (N) (second row). to inornate lizards from NL-2 is a reﬂection of the larger sample size of ornate lizards (N = 65 vs. 15, Character Nuevo León Nuevo León Coahuila respectively). Sample sizes were sufﬁcient (N ≥ A. i. inornata A. i. octolineata A. i. octolineata 4) to compare scutellation data of inornate lizards from Nuevo León for only three sites: NL-2 (N = GAB 70.5 ± 0.48 71.0 ± 0.38 68.7 ± 0.66 15), NL-3 (N = 4), and NL-5 (N = 21). The only 64–80 (44) 64–81 (80) 57–79 (59) signiﬁcant difference among these samples for the 179.0 ± 1.26 182.6 ± 1.13 172.4 ± 1.63 seven characters we studied was the number of FP OR 160–191 (44) 154–205 (80) 140–178 (59) between NL-2 (34.0 ± 0.51) and NL-5 (36.9 ± 0.56). Among the ornate lizards from Nuevo León, sample PV — 16.5 ± 0.30 14.5 ± 0.33 sizes were sufﬁcient (N ≥ 6) to compare scutellation — 13–21 (81) 7–21 (59) data for three sites as well: NL-2 (N = 65), NL-11 (N = 6), and NL-12 (N = 8). The only signiﬁcant PV/GAB — 23.3 ± 0.30 21.0 ± 0.40 difference for nine characters in these samples — 17.8–30.4 (80) 10.3–26.5 (59) (Table 1) was SDL between NL-2 (29.6 ± 0.20) 35.8 ± 0.37 34.7 ± 0.28 34.8 ± 0.32 and NL-12 (32.3 ± 0.49). We pooled all inornate FP 31–44 (44) 32–40 (81) 30–41 (59) and ornate individuals of A. i. inornata from Nuevo León into separate samples for comparisons of 30.5 ± 0.24 30.0 ± 0.20 30.0 ± 0.32 univarate meristic characters (Table 2). Signiﬁcant SDL 27–34 (44) 26–35 (81) 26–37 (59) differences were found between two characters in the pooled samples: OR (179.0 ± 1.26 for inornate COS 11.8 ± 0.30 12.1 ± 0.24 11.1 ± 0.26 and 182.6 ± 1.13 for ornate lizards) and FP (35.8 ± 7–15 (44) 6–18 (81) 6–15 (59) 0.37 for inornate and 34.8 ± 0.32 for ornate lizards). Examples of signiﬁcant differences between Nuevo LSG 27.6 ± 1.05 26.9 ± 0.63 28.0 ± 0.92 17–49 (44) 17–47 (81) 16–50 (59) León and Coahuila samples of A. i. inornata are apparent in OR and PV (Table 2). 26.3 ± 0.78 26.2 ± 0.34 24.0 ± 0.79 Specimens of A. inornata referenced in this study ILS 17–41 (44) 15–44 (81) 11–48 (59) voucher new distribution records extending the known range of this species in western Nuevo León SVL (females) 58.2 ± 0.71 57.6 ± 0.42 57.2 ± 0.84 well north of Villa de García. Several samples from 52–64 (19) 51–63 (44) 51–64 (16) northwest of Mina, which include only inornate lizards, indicate that true-breeding demes occur in the SVL (males) 56.8 ± 0.46 57.4 ± 0.39 56.1 ± 0.55 absence of ornate lizards in a sizeable area of north51–62 (28) 51–63 (45) 52–65 (37) western Nuevo León. Consequently, we agree with Reeder et al. (2002) that the striped subspecies from Coahuila and several sites in Nuevo León, which was previously taxonomic conclusions. Persistence of this color combination on known as A. i. inornata, should be recognized as A. i. octolineata, matching sand dune formations is an apparent result of selection and that the inornate form in Nuevo León be recognized as A. i. for crypsis (Dixon 1967; Rosenblum 2005, 2006; Rosenblum et inornata. Moreover, our analyses of samples of A. inornata from al. 2004), and may be an apomorphy of the pale stripes and dark UAZ and LACM collected in 1962 and 1975 revealed that the rela- ﬁelds in A. i. llanuras in areas peripheral to WSMR. We infer that tive abundance of the inornate and ornate B–D variants at NL-2 complete loss of stripes in A. i. gypsi by gradualism is unlikely, as might vary over time, possibly as a result of ﬂuctuations in rates that would require loss of the genetic potential for organization of of gene ﬂow among co-occurring populations of the color-pattern dorsal pigment into a lineate (i.e., ornate) pattern. variants. The extent of geographic coexistence between inornate That an unstriped pattern can arise spontaneously in a populaand ornate pattern types in the general vicinity of Villa de García tion of A. inornata is shown by the discovery of such individuals appears to be on the order of a few hectares. at Picacho, west of the Río Conchos, Chihuahua. At Picacho, Elsewhere within the range of Little Striped Whiptail, there well within the range of A. i. heptagramma, there is a population are either populations or individual lizards with dorsal patterns of ornate lizards characterized by unusual striping among which that trend toward the extreme inornate morph. Wright and Lowe inornate individuals are occasionally found (Cordes and Walker (1993) described A. i. gypsi from the White Sands Missile Range 1996; Walker et al. 1996, ﬁgure 2). Repeated visits to this site by (WSMR) in the Tularosa Basin, Otero County, New Mexico, based JEC in different years produced no evidence that the proportion on a dorsal pattern of reduced contrast between stripes and ﬁelds. of inornate individuals is increasing in the population. Although However, Rosenblum (2005, 2006) and Rosenblum et al. (2004) the genetic basis of color expression in A. inornata is inadequately have intensively investigated the evolutionary and ecological im- known despite intensive research on A. i. gypsi by Rosenblum et plications of this blanched dorsum in A. inornata, but did not draw al. (2004), we suspect that the inornate pattern of A. i. inornata Herpetological Review 40(3), 2009 279 FIG. 1. Dorsal color-pattern variation in Aspidoscelis inornata from 1.6 km W, 1.6 km S of Villa de García (NL-2), Nuevo León, México in LACM collection (A, LACM 116201, 60 mm SVL; B, LACM 116195, 62 mm SVL; C, LACM 116197, 56 mm SVL; D, LACM 116152, 60 mm SVL; E, LACM 116174, 57 mm SVL; F, LACM 116167, 55 mm SVL; G, LACM 116166, 60 mm SVL; H, LACM 116169, 58 mm SVL; I, LACM 116168, 61 mm SVL; J, LACM 116177, 57 mm SVL; K, LACM 131727, 61 mm SVL; L, LACM 116172, 60 mm SVL). A–G, J are males; H–I, K–L are females. in Nuevo León appeared suddenly as was reported for specimens with unstriped dorsal pattern at Picacho, Chihuahua (Walker and Cordes 1996). In fact, A. i. inornata is the only taxon in the diverse 280 A. sexlineata species group characterized by absence of stripes at all ontogenetic stages of pattern development. We infer that this attribute in A. i. inornata represents one or more mutations Herpetological Review 40(3), 2009 that have silenced the mechanism involved in the organization of pale dorsal stripes, as was deduced for the few individuals from Picacho, Chihuahua. Although contact zones for A. i. inornata and A. i. octolineata at Villa de García have been known from as early as the 1850s (Baird 1858) until as late as the 1980s (this study), elsewhere in Nuevo León the inornate subspecies has either replaced the ornate subspecies or now inhabits areas where it does not occur. Acknowledgments.—We are grateful to the following curators and collection managers for the opportunity to examine specimens of A. inornata in their care: G. Bradley (UAZ), K. Vaughan (TCWC), D. Cannatella and J. Rosales (TNHC), D. Kizirian and K. Beaman (LACM), and A. de Queiroz and R. Humphrey (UCM). Ernest Liner provided information about his experiences with A. inornata in Nuevo León, J. Wright provided helpful information, and H. Taylor provided advice on statistical procedures. Special appreciation is extended to E. Rosenblum for critically reading a version of the manuscript, without speciﬁc endorsements of any part of it. Constructive criticism by reviewers T. Reeder and C. J. Cole, and associate editor R. Espinoza, were indispensable in sharpening the focus of the paper. LITERATURE CITED AXTELL, R. W. 1961. Cnemidophorus inornatus, the valid name for the Little Striped Whiptail lizard, with the description of an annectant subspecies. Copeia 1961:148–158. BAIRD, S. F. 1858. Descriptions of new genera and species of North American lizards in the museum of the Smithsonian Institution. Proc. Acad. Nat. Sci. Philadelphia 1858:253–256. BURGER, L. W. 1950. New, revived, and reallocated names for North American whiptailed lizards, genus Cnemidophorus. Nat. Hist. Misc. Chicago Acad. Sci. 65:1–9. BURT, C. E. 1931. A study of the teiid lizards of the genus Cnemidophorus with special reference to their phylogenetic relationships. Bull. U. S. Nat. Mus. 154:1–286. COLLINS, J. T. 1997. Standard common and current scientific names for North American amphibians and reptiles. SSAR Herpetol. Circ. 25:1–40. CORDES, J. E., AND J. M. WALKER. 1996. Attempted mating between two color pattern morphs of Cnemidophorus inornatus (Little Striped Whiptail Lizard). Bull. Chicago Herpetol. Soc. 31:43. CROTHER, B. I. (ed.). 2000. Scientific and standard English names of amphibians and reptiles of North America north of Mexico, with comments regarding conﬁdence in our understanding. SSAR Herpetol. Circ. 29:1–82. ________ . 2008. Scientific and standard English names of amphibians and reptiles of North America north of Mexico, with comments regarding conﬁdence in our understanding. SSAR Herpetol. Circ. 37:1–84. DENSMORE, L. D, III, J. W. WRIGHT, AND W. M. BROWN. 1989a. Mitochondrial-DNA analyses and the origin and relative age of parthenogenetic lizards (genus Cnemidophorus). II. C. neomexicanus and the C. tesselatus complex. Evolution 43:943–957. ________ , C. C. MORITZ, J. W. WRIGHT, AND W. M. BROWN. 1989b. Mitochondrial-DNA analyses and the origin and relative age of parthenogenetic lizards (genus Cnemidophorus). IV. Nine sexlineatus-group unisexuals. Evolution 43:969–983. DIXON, J. R. 1967. Aspects of the biology of the lizards of the White Sands, New Mexico. Contrib. Sci., Los Angeles Co. Mus. 129:1–22. HENDRICKS, F. S., AND J. R. DIXON. 1986. Systematics and biogeography of Cnemidophorus marmoratus (Sauria: Teiidae). Texas J. Sci. 38:327–402. LEVITON, A. E., R. H. GIBBS, JR., E. HEAL, AND C. E. DAWSON. 1985. Standards in herpetology and ichthyology: Part I. Standard symbolic codes for institutional resource collections in herpetology and ichthyology. Copeia 1985:802–832. LINER. E. A. 2007. A checklist of the amphibians and reptiles of Mexico. Louisiana State Univ. Occas. Pap. Mus. Nat. Sci. 80:1–60. ________ , AND G. CASAS-ANDREU. 2008. Standard Spanish, English and scientiﬁc names of the amphibians and reptiles of Mexico. SSAR Herpetol. Circ. 38:1–162. REEDER, T. W., T. W., C. J. COLE, AND H. C. DESSAUER. 2002. Phylogenetic relationships of whiptail lizards of the genus Cnemidophorus (Squamata: Teiidae): a test of monophyly, reevaluation of karyotypic evolution, and review of hybrid origins. Amer. Mus. Novitates 3365:1–61. ROSENBLUM, E. B. 2005. The role of phenotypic plasticity in color variation of Tularosa Basin lizards. Copeia 2005:586–596. ________ . 2006. Convergent evolution and divergent selection: lizards at the White Sands ecotone. Amer. Nat. 167:1–15. ________ , H. E. HOEKSTRA, AND M. W. NACHMAN. 2004. Adaptive reptile color variation and the evolution of the Mc1r gene. Evolution 58:1794–1808. SMITH, H. M. 1946. Handbook of Lizards: Lizards of the United States and of Canada. Comstock Publ. Co., Ithaca, New York. 557 pp. SOKAL, R. R., AND F. J. ROHLF. 1981. Biometry. W. H. Freeman and Co., San Francisco, California. 857 pp. WALKER, J. M., J. E. CORDES, F. M. QUIJANO, AND E. H. GARCIA. 1996. Implications of extraordinary variation in the Little Striped Whiptail lizard, Cnemidophorus inornatus Baird in Chihuahua, Mexico. J. Herpetol. 30:271–275. WILLIAMS, K. L. 1968. A new subspecies of the teiid lizard Cnemidophorus inornatus from Mexico. J. Herpetol. 1:21–24. WRIGHT, J. W. 1993. Evolution of the lizards of the genus Cnemidophorus, pp. 27–81. In: Biology of Whiptail Lizards (Genus Cnemidophorus). J. W. Wright and L. J. Vitt (eds.). Oklahoma Museum of Natural History, Norman, OK. ________ , AND C. H. LOWE. 1993. Synopsis of the subspecies of the Little Striped Whiptail lizard, Cnemidophorus inornatus Baird. J. Arizona– Nevada Acad. Sci. 27:129–157. APPENDIX 1 Specimens of Aspidoscelis inornata inornata (IN) and A. i. octolineata (OR) Examined Exclusively ornate form: MÉXICO. Coahuila: UCM 37936: 8.2 km W of Monclova (C-1); TNHC 30301–02: 12.8 km NNE of Monclova (C-2); TCWC 43139–42: 54.1 km S of Monclova (975 m) (C-3); TCWC 43143–44: 5.1 km S of San Lazaro (1097 m) (C-4); TCWC 43145: 28.8 km N of Santa Cruz (C-5); TNHC 30679–82: 4.8 km SSE of Est. Gloria (C-6); TNHC 30688–89: 11.2 km SSE of Est. Gloria (C-7); TNHC 30692: 14.4 km SSE of Est. Gloria (C-8); TCWC 43569–71: 19.5 km SE of Castaños (26.6333ºN, 101.3167ºW, 930 m) (C-9); TCWC 43616–17: 23.0 km S of Castaños (26.6167ºN, 101.3500ºW, 1005 m) (C-10); TCWC 43572–89: 24.6 km SE of Castaños (26.60ºN, 101.2833ºW, 935 m) (C-11); TNHC 30268–69: 25.6 km S of Castaños (C-12); TNHC 30266: 35.5 km S of Castaños (C-13); TCWC 43590: 37.8 km SE of Castaños (26.5000ºN, 101.1333ºW, 1006 m) (C-14); TCWC 43612–15: 1.9 km S of Santa Teresa (26.4333ºN, 101.3500ºW, 1128 m) (C-15); TCWC 43618–21: 30.6 km E of Casas Colorados (26.2167ºN, 101.3833ºW, 1219 m) (C-16); LACM 130634: 24.0 km NW of Santa Cruz (C-17); LACM 130631–33: 25.3 km NW of Santa Cruz (C-18); UCM 37886–92: 7.2 km W of Ojo Caliente (C-19); UCM 37934: 28.3 km N of Saltillo (C-20); UCM 37935: 20.3 km N of Saltillo (C-21); UCM 37893–933: 12.2 km N of Saltillo (C-22); TNHC 30237: 22.4 km NNW of La Rosa (C-23); TNHC 30236: 16 km NNW of La Rosa (C-24). Inornate (IN) and ornate (OR) forms: MÉXICO. Nuevo León: RWA 2779–84 (2783 = UTEP 14506) IN, RWA 2785–93 OR: 30–50 m E of highway and 10–15 m S of bridge over the Río Pesquería at Villa Herpetological Review 40(3), 2009 281 de García (25.8167ºN, 100.5833ºW) (NL-1); UAZ 14067–75, LACM 116195–201, 131729–31 IN; UAZ 14076–88, 14093–95, 14097–102, LACM 116152–94, 131727–28, 131732–34 OR: 1.6 km W, 1.6 km S of Villa de García (NL-2); TCWC 44262–65 IN; TCWC 44261 OR: 14.1 km NW of Villa de García (25.8500ºN, 100.6833ºW, 716 m) (NL-3); TCWC 44266 IN: 1.6 km W of La Paz (25.9333ºN, 100.6500ºW, 716 m) (NL-4); TCWC 43591–611 IN: 8.2 km W of Mina by México Hwy 53 (26.0333ºN, 100.6667ºW, 671 m) (NL-5); TCWC 51814 IN: 8.5 km NNW of Mina (NL-6); TCWC 51815 IN: 14.4 km NNW of Mina (NL-7); RWA 6049–51 IN: 9.2 km N, 10.3 km W of Mina (26.0833ºN, 100.6306ºW, 644 m) (NL8); TCWC 46866 IN: 17.8 km SE of Soledad (26.3333ºN, 100.8333ºW, 930 m) (NL-9); TCWC 46867 OR: 0.6 km SE of Soledad (26.4000ºN, 100.9667ºW, 640 m) (NL-10); TCWC 46868–73 OR: 4.0 km NW of Soledad (26.4333ºN, 101.0167ºW, 701 m) (NL-11); UCM 37937–45 OR: 1.3 km W of Casa Blanca (NL-12); (TCWC 51816–24 OR: 5.6 km W of Aramberri (24.1000ºN, 99.8167ºW) (NL-13); TNHC 30123–24 OR: 19.2 km S of El Canelo (NL-14); TNHC 30169 OR: 1.6 km S of San Roberto (NL-15); TCWC 56790 OR: 22.7 km N of Doctor Arroyo (NL-16); TCWC 56788 OR: 20.3 km N of Doctor Arroyo (NL-17). Herpetological Review, 2009, 40(3), 282–283. © 2009 by Society for the Study of Amphibians and Reptiles Discovery of a Surviving Population of the Montane Streamside Frog Craugastor milesi (Schmidt) JONATHAN E. KOLBY The Conservation Agency, Jamestown, Rhode Island 02835, USA Current Address: 1671 Edmund Terrace Union, New Jersey 07083, USA Operation Wallacea, Hope House, Old Bolingbroke, Lincolnshire PE23 4EX, UK e-mail: j_kolby@hotmail.com and JAMES R. MCCRANIE 10770 SW 164th Street Miami, Florida 33157-2933, USA e-mail: jmccrani@bellsouth.net Amphibian declines are taking place worldwide, but are most prevalent among montane stream-associated species in the neotropics (Stuart et al. 2004). Some of these declines are due to habitat loss, but many are taking place in seemingly pristine habitats (Stuart et al. 2004). Lips et al. (2006) reported the link between the chytrid fungus Batrachochytrium dendrobatidis and a rapid mass mortality and loss of amphibian diversity at a site in Panama. Pounds et al. (2006) provided evidence that the chytrid fungus in combination with global warming may be responsible for the disappearances of various species of the neotropical frog genus Atelopus. The Honduran endemic montane streamside frog Craugastor milesi (Schmidt) was last seen alive in 1983, even though one of its known localities remains in a pristine condition (McCranie and Castañeda 2007; McCranie and Wilson 2002). Subsequently, numerous efforts to ﬁnd this frog at its remaining pristine locality in Parque Nacional Cusuco have failed (McCranie and Castañeda 2007; McCranie and Wilson 2002; Townsend and Wilson 2008), even though it was a common frog at that locality in the late 282 FIG. 1. Adult male Craugastor milesi found 29 June 2008 in Parque Nacional Cusuco, Honduras. 1970s and early 1980s. Consequently, C. milesi was one of only nine New World anuran species classiﬁed as extinct by Young et al. (2004). On 29 June 2008, an adult male Craugastor milesi (Fig. 1) was collected in Parque Nacional Cusuco while conducting a nocturnal herpetological survey. The site was located at an elevation of 1841 m and the weather was cool and breezy with light precipitation. A nearby data logger recorded the air temperature to be 16°C during this period of activity. The specimen was perched on a low branch along the edge of a small montane stream. This stream is a tributary of the larger Rio Cantilles, with which it joins approximately100 m downstream from the collection site. A skin swab was collected from this specimen for the detection of B. dendrobatidis. The sample was processed by PCR analysis and produced a negative result for infection. On 30 June 2008 and 5–6 July 2008, extensive day and nighttime surveys of the same site and surrounding habitat failed to produce additional specimens of Craugastor milesi. During these surveys, four Plectrohyla dasypus larvae were collected from the stream and swabbed for B. dendrobatidis. Two of the four tadpoles exhibited severe oral defects and subsequently tested positive for infection. Surveys along portions of the nearby Rio Cantilles were similarly unsuccessful in locating additional C. milesi, but revealed the presence of B. dendrobatidis in six out of seven additional P. dasypus larvae. Although other Craugastor species could be found widely distributed across Parque Nacional Cusuco, the forest surrounding the C. milesi collection site appeared to be unusually devoid of terrestrial anurans. The rediscovery of Craugastor milesi in Parque Nacional Cusuco is signiﬁcant in light of the current global amphibian extinction crisis. Despite the common global threat of habitat destruction, the pristine condition of Parque Nacional Cusuco during the 1980s negates habitat destruction as a possible explanation for the unexpected disappearance of C. milesi. The most likely agent responsible for C. milesi’s demise, B. dendrobatidis, has recently Herpetological Review 40(3), 2009 been identiﬁed in Parque Nacional Cusuco and may be responsible for the declining populations of several other critically endangered amphibian species in the park (Kolby et al. 2009). If true, the discovery of an uninfected adult C. milesi from within the heart of a B. dendrobatidis-ridden habitat produces hopeful implications for resistance to chytridiomycosis, as it is highly unlikely that this individual successfully evaded infection throughout development while being exposed to contaminated water sources. On 14 July 2007, a specimen with striking resemblances to C. milesi was encountered in direct proximity to the conﬁrmed collection site in 2008, but efforts to capture and positively identify this specimen proved unsuccessful at the time. Although a genetically robust population of C. milesi may exist in Parque Nacional Cusuco, it is too early to estimate the extant population size and determine whether this species is rebounding from the brink of extinction or still in a period of decline. Acknowledgments.—Financial support was provided by the National Geographic Society (Young Explorers Grant C146-08), and Omaha’s Henry Doorly Zoo. Collection and export permits were granted by C. Cárcamo de Martínez of AFE-CODEHFOR, Tegucigalpa. K. Hunter conducted the PCR analysis for B. dendrobatidis. Personnel of Operation Wallacea provided crucial logistical support while in the ﬁeld. LITERATURE CITED KOLBY, J. E., G. E. PADGETT-FLOHR, AND R. FIELD. 2009. Amphibian chytrid fungus (Batrachochytrium dendrobatidis) in Cusuco National Park, Honduras. Dis. Aquat. Org., Spec. Ed. 4. Published online 6 May 2009. doi: 10.3354/dao02055 LIPS, K. R., F. BREM, R. BRENES, J. D. REEVE, R. A. ALFORD, J. VOYLES, C. CAREY, L. LIVO, A. P. PESSIER, AND J. P. COLLINS. 2006. Emerging infectious disease and the loss of biodiversity in a Neotropical amphibian community. Proc. Natl. Acad. Sci. 103:3165–3170. MCCRANIE, J. R., AND F. E. CASTAÑEDA. 2007. Guía de Campo de los Anﬁbios de Honduras. Bibliomania! Salt Lake City, Utah. ________ , AND L. D. WILSON. 2002. The Amphibians of Honduras. Soc. Study Amphib. Reptiles, Contrib. Herpetol. 19:i–x, 1–625. POUNDS, J. A., M. R. BUSTAMANTE, L. A. COLOMA, J. A. CONSUEGRA, M. P. L. FOGDEN, P. N. FOSTER, E. LA MARCA, K. L. MASTERS, A. MERINOVITERI, R. PUSCHENDORF, S. R. RON, G. A. SÁNCHEZ-AZOFEIFA, C. J. STILL, AND B. E. YOUNG. 2006. Widespread amphibian extinctions from epidemic disease driven by global warming. Nature 439:161–167. 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 306:1783– 1786. TOWNSEND, J. H., AND L. D. WILSON. 2008. Guide to The Amphibians & Reptiles of Cusuco National Park, Honduras. Guía de Los Anﬁbios y Reptiles del Parque Nacional Cusuco, Honduras (translated by P. M. KULSTAD). Bibliomania!, Salt Lake City, Utah. YOUNG, B. E., S. N. STUART, J. S. CHANSON, N. A. COX, AND T. M. BOUCHER. 2004. Disappearing jewels: the status of New World amphibians. NatureServe, Arlington, Virginia. Herpetological Review, 2009, 40(3), 283–286. © 2009 by Society for the Study of Amphibians and Reptiles Observations on the Ecology of Trachemys adiutrix and Kinosternon scorpioides on Curupu Island, Brazil LARISSA BARRETO e-mail: lara@ufma.br LENY CLAUDIA LIMA e-mail: lenyclaudiapl@hotmail.com and SUZANE BARBOSA e-mail: zanegb10@yahoo.com.br Universidade Federal do Maranhão Departamento de Oceanograﬁa e Limnologia Av. dos Portugueses s/n, Campus do Bacanga, CEP 65080-040 São Luís- MA, Brazil Brazil has an extensive coastline in the state of Maranhão, including many islands and a continental area called the “Baixada Maranhense,” where several turtle species of the families Chelidae, Pelomedusidae, Kinosternidae, and Emydidae occur. Both Kinosternon scorpioides and Trachemys adiutrix occur on Curupu Island. Kinosternon scorpioides lives in both still and running water, frequently in areas affected by human activity, especially drainage canals (Pritchard and Trebbau 1984). This species is distributed from Mexico to South America, where it thrives along the coast in Colombia, Guiana, Trinidad, and Brazil as well as the interior of the Amazon to Bolivia (Pritchard and Trebbau 1984). The genus Trachemys ranges from North America into Central America (Savage 1966), southern Brazil, Uruguay, and Argentina (Vanzolini 1995) and has the largest geographic distribution of any turtle genus in the New World (Seidel 2002). Trachemys adiutrix is an aquatic species that inhabits permanent and seasonal bodies of water, and may move great distances during the rainy season (Vanzolini 1995). The only published reference on Trachemys in Maranhão is the original description of T. adiutrix from the northeastern region of the state (Vanzolini 1995). Information on the occurrence of K. scorpioides in the coastal region of Brazil consists of one record reported by Pritchard and Trebbau (1984), who observed the species on Marajó Island, Pará state. Because little is known of K. scorpioides and T. adiutrix, we report data on selected aspects of their biology in the coastal zone of northeastern Brazil. We speciﬁcally examined reproductive ecology (nesting behavior and season) and population structure of these two species. MATERIALS AND METHODS Curupu Island (2.4025°S, 44.0219°W) is located ca. 30 km from the center of São Luís, state of Maranhão, Brazil. The island is ringed by extensive sandy beaches, with ﬂooded ﬁelds, marshy vegetation, dunes, freshwater lagoons, and mangroves further inland. The rainy season extends from January to June and the dry season from July to December. Two-day sampling periods were conducted every 15 d throughout the year from August 2000 to March 2002 (Figs. 1, 2). During each sampling period, data were Herpetological Review 40(3), 2009 283 collected between 0500 h and 1200 h. During each ﬁeld visit, we captured turtles in three lagoons located in the central part of the island. These lagoons were located ca. 20 m apart and were ca. 400 m2 in area each. They ranged from oligotrophic to eutrophic and contained an abundance of macrophytes, pteridophytes, macroalgae, and ﬁsh. Turtles were collected using baited funnel traps (one trap per lagoon) constructed of wooden slats with a funnel entrance. These traps were made by the local community using wood stems or wire, and were similar to crayﬁsh traps (Rocha et al. 1997). Traps were baited with fresh ﬁsh and placed in the water 2–3 m from shore the night before the sampling period. We also recorded any sightings of turtles moving across land during each sampling period. Reproductive ecology.—The time period when females were gravid was determined by inguinal palpation (Pritchard and Trebbau 1984), and the nesting period was determined by the presence of nests in the wild. We searched for nests every 15 d in two consecutive days during only the dry season of the sampling period, totaling 20 d of searching. Turtle nests were located each morning (between 0500 h and 0600 h) by following tracks left by nesting females in the sand. We measured the eggs (length and width) of one nest found intact to the nearest mm using dial calipers. The period of hatchling emergence of both species was determined by the presence of hatchlings in traps of local ﬁshermen. All hatchlings caught were measured. They were identiﬁed as hatchlings by their thin carapace, caruncle, and lack of growth rings. Analysis of population structure.—Population structure of the two species was estimated by quantifying the distribution of number of individuals in different size classes and sexes throughout the year. All animals were measured using straightline maximum carapace length (SCL) and plastron lengths to the nearest mm with calipers, sexed, and marked with a unique cut in the center of the marginal scutes (Cagle 1939). All turtles were immediately released at their point of capture after processing. Turtles were individually marked as part of an ongoing research project. The sex of the animals was determined by examination of tail length (for both species) and plastron concavity (only for K. scorpioides) (Pritchard and Trebbau 1984) and, for analysis of size distribution the individuals from each species were grouped FIG. 2. Number of male and female Trachemys adiutrix captured by month during the sampling period from August 2000 to March 2002. into 1 cm size classes. A one-way ANOVA was used to determine whether there was a signiﬁcant difference in mean size (CL) between males and females in each species. The overall sex ratio was compared to an expected sex ratio of unity using a chi-square test. We had no reason to suspect that this population exhibited a 1:1 ratio, and were merely using the 1:1 ratio as a reference point for comparison. All statistical analyses were performed with STATISTIC v. 6.0 (Statsoft 2001) and according to Sokal and Rohlf 1981. We separated mature and immature individuals for the two species based on the minimum size of carapace length of gravid females found in this study, supplemented with information in the literature from closely related species (e.g., Bager 2003; Iverson 1986; Pritchard and Trebbau 1984) and unpublished data of a specialist (R. Vogt). Adult female Kinosternon scorpiodes are sexually mature at 10.4 cm CL and males are mature at 10.1 cm in the Yucatan Peninsula of Mexico (R. Vogt, pers. comm.). Trachemys adiutrix are considered adults in seasonal lakes when males reach 9.9 cm CL and females 10.1, in permanent lakes when males reach 13.7 cm and females reach 16.5 cm (Batistella 2008). For this study, we considered an age estimation of maturity for K. scorpioides and T. adiutrix individuals with minimum carapace length of 10.0 cm for males and females. RESULTS AND DISCUSSION FIG. 1. Number of male and female Kinosternon scorpioides captured by month during the sampling period from August 2000 to March 2002. 284 Turtles were observed crossing dunes, in grassy areas, and in the bottom of lagoons. The bottom of the lagoons are full of organic material and mud. Individuals of both species were observed throughout the sampling period walking across dunes during the early morning (0600 h) and late afternoon (1800 h). Gravid turtles were captured during August (N = 38 for K. scorpioides; N = 4 for T. adiutrix). We found gravid females with maximum and minimum carapace length sizes of 13.7 and 10.0 cm for K. scorpioides, and 15.0 and 12.8 cm for T. adiutrix, respectively. Fifteen nests were observed in the study area, but identiﬁcation to species was not possible because the nests had been ravaged by predators (probably foxes, Cerdocyon thous, T. Oliveira, pers. comm.). The nests were observed in grassy areas 15–20 m from water. All nests were found between August and Herpetological Review 40(3), 2009 December, during the dry season, with the greatest number being recorded in October and December (N = 7 and 5, respectively). In October 2001, one intact T. adiutrix nest was found. This nest was in sandy substrate shaded by grass, was 9.4 cm deep and 11.5 cm in diameter, and contained six elongated eggs with parchmentlike shells. Mean egg size was 3.8 ± 0.08 cm (3.7–3.9) in length and 2.4 ± 0.04 (2.4–2.5) in width. Three K. scorpioides hatchlings had a mean carapace length of 3.5 ± 0.07 cm (3.47–3.61). Hatchling T. adiutrix (N = 23) had a mean SCL of 3.5 ± 0.2 cm (2.79–3.78). All of these hatchlings were captured in January and February. We captured 109 K. scorpioides; the overall sex ratio (45 females: 35 males = 1.28) did not differ signiﬁcantly from unity (χ2 = 4.24, df = 3, P = 0.23). We captured 69 T. adiutrix; 20 females and 15 males. The overall sex ratio was 1.34:1, again not signiﬁcantly different from unity (χ2 = 11.83, df = 6, P = 0.06) (see Figs. 1–2 for details). The most abundant size class of K. scorpioides was the 11–12 cm size class, and for T. adiutrix in the 8, 9, and 15 cm size class (Figs. 3 and 4, respectively). Male K. scorpioides were signiﬁcantly larger than females (F = 12.57, df = 39, P = 0.001). There was no signiﬁcant difference in size between male and female T. adiutrix (F = 0.66, df = 23, P = 0.42). In our study, hatchling K. scorpioides had a mean CL of 3.5 cm and were observed at the onset of the rainy season. We suspect that larger hatchlings had grown prior to ﬁrst capture because smaller hatchling sizes have been reported than those observed in our study (see Pritchard and Trebbau 1984). Rocha and Molina (1990) reported a mean CL of 2.85 cm for hatchlings incubated in captivity. In the coastal area of Maranhão State, Vanzolini (1995) observed that T. adiutrix laid its eggs in sand close to the water, similar to our observations. Molina (1996) observed that T. dorbignyi females in captivity preferred to lay their eggs in a predominantly sandy substrate and in areas without vegetation. The nesting period of T. dorbignyi in captivity extends from August to January with the majority of egg laying being concentrated in October and November (Molina 1995, 1996). Rocha and Molina (1990) observed that the nesting period of K. scorpioides in captivity lasted from March to August during the dry season. In our study area, the majority of nests were discovered during October and December during the dry season. FIG. 3. Size distribution of Kinosternon scorpioides at Curupu Island, Maranhão, Brazil from turtles trapped January 2001 to December 2002. FIG. 4. Size distribution of Trachemys adiutrix at Curupu Island, Maranhão, Brazil from turtles trapped January 2001 to December 2002. The relatively small maximum size of T. adiutrix (16 cm) can probably be linked to the hunting of these animals by local people in the study area on Curupu Island, and not because the entrance of the traps prohibited catching larger turtles. In Curupu, because larger sizes of T. adiutrix are targeted for their meat, hunting may remove the largest animals from the population. In another locality (Caju Island in Maranhão State) where there is no turtle hunting, we found turtles larger than 21 cm (L. Barreto, pers. obs.). Acknowledgments.—Richard C. Vogt and two anonymous revisers provided valuable comments on the manuscript. We thank Conselho Nacional de Pesquisa (CNPq) for its ﬁnancial support during the project in the form of a scholarship. We thank A. C. L. de Castro for statistical analyses and Claudio Gomes, Tiago Ramos, and Anderson Magalhães for ﬁeld support. Thomas Pedersen improved the digital art (graphic) ﬁles. This work conducted under IBAMA Institute license number 02012.0015171/2005-25. We adhered to all appropriate animal care and wildlife collecting laws during this study. LITERATURE CITED CAGLE, F. R.1939. A system of marking turtles for future identification. Copeia 1939:170–173. BAGER, A. 2003. Aspectos da Biologia e Ecologia daTartaruga Tigre D’água, Trachemys dorbigni, (Testudine – Emydidae) no Extremo Sul do Estado do Rio Grande do Sul - Brasil. Universidade Federal do Rio Grande do Sul, Porto Alegre, Rio Grande do Sul. 100 pp. BATISTELA, A. M. 2008. Biologia de Trachemys adiutrix (Vanzolini 1995) no Litoral do Nordeste - Brasil. Instituto Nacional de Pesquisas da Amazônia, Universidade do Amazonas. Manaus, Amazonas. 104 pp. IVERSON, J. B. 1986. Notes on the natural history of the Oaxaca mud turtle, Kinosternon oaxacae. J. Herpetol. 20:119–123. MOLINA, F. B. 1995. Nesting behavior of the Brazilian slider turtle (Trachemys dorbignyi) in captivity, with comments on behavioral enrichment. In G. P. Nabhan, H. E. Lawler, and E. Mellink (eds.), International Herpetological Symposium, 1995, pp. 33–35. Denver Zoological Gardens, Colorado. ________ . 1996. Os quelônios e sua biologia – O tigre d’água Trachemys dorbignyi. Informativo Pró – tartaruga 1996:45– 47. PRITCHARD, P. H. C., AND P. TREBBAU. 1984. Kinosternon scorpioides scorpioides (Linnaeus, 1766). In P. H. C. Pritchard and P. Trebbau (eds.), The Turtles of Venezuela, pp. 239–248. SSAR Contributions to Herpetology No. 2. Oxford, Ohio. ROCHA, M. B. DA, AND F. B. MOLINA. 1990. Reproductive biology of Kinosternon scorpiodes (Testudines: Kinosternidae) in captivity. Tortoises and Turtles 5:8. Herpetological Review 40(3), 2009 285 ROCHA, C. A, W. FRANKLIN JUNIOR, W. P. DANTAS, M. F. FARIAS, AND A. M. E. DE OLIVEIRA. 1997. Fauna e flora acompanhantes da pesca da lagosta no Nordeste do Brasil. In C. A. Rocha (ed.), Boletim TécnicoCientíﬁco Cepene 5:15–28. SAVAGE, J. M. 1966. The origins and history of the Central American herpetofauna. Copeia 1966:719–766. SEIDEL, M. E. 2002. Taxonomic observations on extant species and subspecies of slider turtles, genus Trachemys. J. Herpetol. 36:285– 292. SOKAL, R. R., AND F. J. ROHLF. 1981. Biometry. 2nd ed. W. H. Freeman, New York, New York. 588 pp. STATSOFT, INC. 2001. Statistica (data analyses software system) version 6.0. www.statsoft.com. VANZOLINI, P. E. 1995. A new species of turtle, genus Trachemys, from the state of Maranhão, Brazil (Testudines, Emydidae). Rev. Brasil. Biol. 55:111–125. Herpetological Review, 2009, 40(3), 286–288. © 2009 by Society for the Study of Amphibians and Reptiles Foraging Ecology of Spotted Turtles (Clemmys guttata) in Ontario, Canada MEGAN L. RASMUSSEN Department of Biology, Laurentian University 935 Ramsey Lake Road, Sudbury, Ontario, Canada, P3E 2C6 e-mail: ml_rasmussen@laurentian.ca JAMES E. PATERSON Department of Integrative Biology, University of Guelph 50 Stone Road East, Guelph, Ontario, Canada, N1G 2W1 e-mail: patersoj@uoguelph.ca and JACQUELINE D. LITZGUS* Department of Biology, Laurentian University 935 Ramsey Lake Road, Sudbury, Ontario, Canada, P3E 2C6 * Corresponding author; e-mail: jlitzgus@laurentian.ca The conservation of animal species relies on the identiﬁcation and protection of suitable habitat. While general habitat features are important to the success of vertebrate ectotherms because they provide structures for thermoregulation (Blouin-Demers and Weatherhead 2001; Downes and Shine 1998) and predator avoidance (Spencer 2002), biotic elements such as food resource use and distribution are often overlooked in studies of habitat selection. Qualitative descriptions of diet do little to further the understanding of resource use and requirements, which ultimately affect the ﬁtness of individuals. The recent use of stable isotope analysis has shed light on the relationship between foraging ecology and nutrient assimilation in some turtle species (Aresco and James 2005; Bulte and Blouin-Demers 2008; Hatase et al. 2006; Seminoff et al. 2007). Detailed observations of species-speciﬁc foraging behavior assist in the implementation of stable isotope analysis by focusing sampling efforts on likely prey, reducing potential costs and time spent in the ﬁeld. The Spotted Turtle (Clemmys guttata) is a species for which diet and foraging behavior have been qualitatively described (Ernst 1976; Milam and Melvin 2001; Surface 1908). Stomach content analyses of 27 Spotted Turtles revealed three individuals with 286 plant material, while all individuals contained aquatic and terrestrial invertebrates in their guts (Surface 1908). These ﬁndings were supported by Ernst’s (1976) observations of Spotted Turtles ingesting both plant and animal materials. Spotted Turtles are noted to be opportunistic omnivores, with a highly variable list of diet items including cranberries, earthworms, aquatic insect larvae, crustaceans, snails, salamanders, ﬁsh, birds, algae, and tadpoles (Ernst 1976). The relative importance of these food items has not been reported. Spotted Turtles are cold-tolerant and most active in the early spring (Ernst 1976, 1982; Haxton and Berrill 1999; Litzgus and Mousseau 2004; Milam and Melvin 2001; Ward et al. 1976), but they have not been recorded foraging in water cooler than 14°C (Ernst 1976). The broad distribution of this species, as well as differing habitat use between northern and southern populations (Haxton and Berrill 1999; Litzgus and Mousseau 2004; Milam and Melvin 2001; Ward et al. 1976), indicate that detailed dietary and foraging choices should be documented to further our understanding of resource use and habitat selection. In Canada, Spotted Turtles are listed as an endangered species (COSEWIC 2004), and declines and local extirpations have been documented within Ontario despite habitat protection (Browne and Hecnar 2007). Recovery plans should take into account resource distribution, as well as general habitat qualities, in order to create effective management guidelines for this species. The purpose of our study was to quantify dietary choices in a population of Spotted Turtles and describe foraging behaviours observed in natural settings. Materials and Methods.—The foraging ecology of Spotted Turtles in a population on the shores of Lake Huron in Ontario, Canada was documented during a radio telemetry study in 2007 and 2008. The study site consists of a 95-ha mosaic of Great Lakes coastal wetlands bounded by forested swamps, upland forests, and minimal human development. One hundred and seventeen individuals (54 females, 49 males, 14 juveniles) were captured and marked (Cagle 1939), with an equal distribution of adult males and females (χ2 = 0.24, df = 1, p > 0.05). Fifteen individuals (5 males, 10 females) were located 2–4 times a week from April to August 2007 and 2008 using radio telemetry, with up to 16 additional females located daily during June of both years. Transmitters (SI-2F and SI-2T, 12g, Holohil Systems Ltd, Carp, Ontario) were attached to the rear marginal scutes using copper wire and putty epoxy. Turtles without transmitters were also observed opportunistically. Behavior was characterized for each turtle sighting over the entire study period. Foraging behavior was deﬁned as any aquatic or semi-aquatic activity in which the turtle was actively searching, grabbing, or biting at items. Target items were identiﬁed by observing the item during the foraging period, and any unidentiﬁable items were recorded as “unknown.” For each individual turtle observed foraging, ambient air (10 cm from the ground in shade), substrate surface (direct sunlight on substrate), and water temperature (mid-depth of water column) were measured to 0.1°C using a digital thermometer. Temperature-sensitive transmitters allowed the estimation of turtle body temperature from the pulse rate. Results.—Two hundred and twenty-seven foraging observations were made during the study (Table 1), all within 2–40 cm of water. The greatest proportion of foraging observations were of turtles taking aquatic invertebrates (74%), followed by ﬁsh (~16%). Foraging was observed throughout the active season (April to Au- Herpetological Review 40(3), 2009 TABLE 1. Observations of foraging by Spotted Turtles (Clemmys guttata) in Ontario, Canada over the course of gust) in both years; however, a two-year (2007, 2008) radio telemetry study. Targets indicated by * were consumed as carrion. observations of foraging were more numerous in the earlier 2007 2008 Total % of Foraging Observations months (Fig. 1). While the Target Item (excluding unidentiﬁed targets) number of observers (MLR and JEP) remained constant Aquatic Invertebrates Total 13 19 32 74.0 over both years of the study, Freshwater snails 8 8 16 37.2 it is likely that experience Trichoptera 1 4 5 11.6 gained in 2007 contributed to Leech 1 0 1 2.3 an increase in overall sightings in 2008. To account for Crayﬁsh 1 0 1 2.3 this, monthly observations Unidentiﬁed 2 7 9 20.9 are presented as percentages Vegetation 0 1 1 2.3 of the total observations per Rainbow Trout (Onchorhynchus mykiss)* 0 1 1 2.3 year (Fig. 1). Carp (Cyprinus carpio)* 0 2 2 4.6 In 2007, water temperatures 2 2 4 9.3 recorded during foraging Small Ciprinids and other minnows* Leopard Frog (Rana [Lithobates] pipiens)* 1 0 1 2.3 ranged from 13°C to 33.8°C. 0 1 1 2.3 In 2008, water temperatures Tadpoles (Bufo [Anaxyrus] americanus) ranged from 7.7°C to 31.7°C. For individuals outﬁtted with radio transmitters in 2008, estimated body temperature during that inactive individuals were not choosing cooler microclimates, foraging ranged from 9.9°C to 31.9°C. Prey items chosen were and therefore may not be aestivating (Litzgus and Brooks 2000). not related to differences in water temperatures (F = 0.44, df = Alternatively, higher August temperatures may allow turtles to 3,164, p = 0.72) or estimated turtle body temperatures (F = 0.09, forage in more heavily-vegetated waters reducing our ability to df = 3,113, p = 0.96). observe foraging behavior later in the season. Capital energy acDiscussion.—The large proportion of invertebrates in Spotted quired during foraging late in the summer may help supplement Turtle diets observed in our study agrees with ﬁndings of the stom- income energy gains during the limited time available for foraging ach content analysis by Surface (1908), and foraging observations between emergence and reproductive activities the following year by Ernst (1976). When consuming hard-shelled snails, individuals (Gregory 2006). would repeatedly bite and release the snail, eventually consuming Prior to our study, Spotted Turtles were not recorded foraging in the soft body and discarding the hard shell. While the majority waters cooler than 14°C (Ernst 1976). In both years, we observed of animal materials were assumed to be consumed post-mortem, individuals foraging in water cooler than this minimum. In 2007, tadpoles were consumed live. Stomach content analysis by Sur- we observed foraging at 13°C and made multiple observations face (1908) indicated the presence of land-dwelling invertebrates. of foraging in water temperatures as low as 7.7°C in 2008. We Whether these items were captured on land or accidentally fell into the water was unknown. Foraging for terrestrial prey was conﬁrmed once during our study. An individual male was observed in shallow water (3 cm) positioning himself vertically using his front legs, and biting at the remains of a spider’s web hung between short rushes. Capture and consumption of the spider or items in the spider web was not directly observed. This observation would suggest that while Spotted Turtles are conﬁned to consuming prey items aquatically, these prey items could be captured in a terrestrial environment. Observations of foraging behavior were most numerous in May, June, and July, with few observations in August (Fig. 1). Foraging is probably most important in early spring after emergence from hibernation, and during the mating and nesting seasons. The energetic demands of sperm production, egg production, copulation, and nesting (Hughes and Brooks 2006; Morreale et al. 1984; Pearse and Avise 2001) require a large effort to be expended in resource gathering early in the season for turtles at northern latitudes. The lack of foraging observations in August could be due to periods FIG. 1. The percentage of foraging behavior observations for Spotted of inactivity. Aestivation has been observed in Spotted Turtles and Turtles (Clemmys guttata) in Ontario, Canada for the active seasons of is thought to conserve energy and water during high temperatures 2007 (black, N = 58 observations) and 2008 (white, N = 170 observain the summer months (Ernst 1982; Perillo 1997). However, tions). In April 2007, a lack of researcher presence resulted in no data temperature data gathered in one Ontario population documented being collected, denoted as “n.d.” on the graph. Herpetological Review 40(3), 2009 287 did not detect prey choice differences at different environmental temperatures, and this can be explained by the apparently opportunistic nature of foraging. The ability of Spotted Turtles to forage at low temperatures may give them a ﬁtness advantage in the unpredictable and short active seasons typical of northern climates. The acquisition of resources during the active season is vital to an individual’s survival from year to year, and also to their ability to produce offspring (Litzgus and Brooks 1998; Litzgus et al. 2008). If Spotted Turtles are restricted by variable water levels in their shallow wetland habitats, then the ability to forage successfully in the cooler months of April and May would certainly be advantageous. Our study quantitatively highlights the relative importance of food items in the diet of a population of Spotted Turtles. The importance of vegetative matter in Spotted Turtle diets, at least in our population, appears to be minimal. Close observation revealed that although they may appear to be targeting aquatic vegetation, it is in fact most likely that individuals are seeking small freshwater snails and benthic invertebrates within the matrix of aquatic vegetation. This vegetative material may not contribute to energy stores. Stable isotope analysis combined with stomach content analysis in Snapping Turtles (Chelydra serpentina) showed that although large amounts of vegetation may be found within the stomach, little of this is translated into tissues (Aresco and James 2005). It appears as though Spotted Turtles are most dependent on macroinvertebrates, with a strong tendency to detect and consume carrion. Future studies of Spotted Turtle foraging ecology should examine the importance of various diet items through stomach content analysis coupled with radio-isotope analysis to explore the relative importance of various food items in tissues, as well as the turtle’s trophic position in aquatic ecosystems. This information should be used to supplement habitat selection studies in dictating protection and enhancement guidelines for the continued presence of this species in our ecosystems. Acknowledgments.—All work was conducted in compliance with CCAC guidelines and under approved Laurentian University Animal Care Protocol #2004-11-01. We thank the local landowners for allowing us access to their land. The research was supported by funding from Ontario Power Generation and NSERC. LITERATURE CITED ARESCO, M. J., AND F. C. JAMES. 2005. Ecological relationships of turtles in northern Florida lakes: a study of omnivory and the structure of a lake food web. Final Report. Florida Fish and Wildlife Conservation Commission, Tallahassee, Florida, USA. BLOUIN-DEMERS, G., AND P. J. WEATHERHEAD. 2001. An experimental test of the link between foraging, habitat selection and thermoregulation in black rat snakes Elaphe obsoleta obsoleta. J. Anim. Ecol. 70:1006–1013. BROWNE, C. L., AND S. J. HECNAR. 2007. Species loss and shifting population structure of freshwater turtles despite habitat protection. Biol. Conserv. 138:421–429. BULTE, G., AND G. BLOUIN-DEMERS. 2008. Northern map turtles (Graptemys geographica) derive energy from the pelagic pathway through predation on zebra mussels (Dreissena polymorpha). Freshwat. Biol. 53:497–508. CAGLE, F. R. 1939. A system of marking turtles for future identification. Copeia 1939:170–172. 288 COSEWIC. 2004. COSEWIC assessment and update status report on the spotted turtle Clemmys guttata in Canada. Committee on the Status of Endangered Wildlife in Canada. Ottawa, Canada. DOWNES, S., AND R. SHINE. 1998. Heat, safety or solitude? Using habitat selection experiments to identify a lizard’s priorities. Anim. Behav. 55:1387–1396. ERNST, C. H. 1976. Ecology of the spotted turtle, Clemmys guttata (Reptilia, Testudines, Testudinidae), in southeastern Pennsylvania. J. Herpetol. 10:25–33. ________ . 1982. Environmental temperatures and activities in wild spotted turtles, Clemmys guttata. J. Herpetol. 16:112–120. GREGORY, P. T. 2006. Influence of income and capital on reproduction in a viviparous snake: direct and indirect effects. J. Zool. 270:414–419. HATASE, H., K. SATO, M. YAMAGUCHI, K. TAKAHASHI, AND K. TSUKAMOTO. 2006. Individual variation in feeding habitat use by adult female green sea turtles (Chelonia mydas): Are they obligate neritic herbivores? Oecologia 149:52–64. HAXTON, T., AND M. BERRILL. 1999. Habitat selectivity of Clemmys guttata in central Ontario. Can. J. Zool. 77:593–599. HUGHES, E. J., AND R. J. BROOKS. 2006. The good mother: does nestsite selection constitute parental investment in turtles? Can. J. Zool. 84:1545–1554. LITZGUS, J. D., F. BOLTON, AND A. I. SCHULTE-HOSTEDDE. 2008. Reproductive output depends on body condition in spotted turtles (Clemmys guttata). Copeia 2008:86–92. ________ , AND R. J. BROOKS. 1998. Reproduction in a northern population of Clemmys guttata. J. Herpetol. 32:252–259. ________ , AND ________. 2000. Habitat and temperature selection of Clemmys guttata in a northern population. J. Herpetol. 34:178–185. ________ , AND T. A. MOUSSEAU. 2004. Home range and seasonal activity of southern spotted turtles (Clemmys guttata): Implications for management. Copeia 2004:804–817. MILAM, J. C., AND S. M. MELVIN. 2001. Density, habitat use, movements, and conservation of spotted turtles (Clemmys guttata) in Massachusetts. J. Herpetol. 35:418–427. MORREALE, S. J., J. W. GIBBONS, AND J. D. CONGDON. 1984. Significance of activity and movement in the yellow-bellied slider turtle (Pseudemys scripta). Can. J. Zool. 62:1038–1042. PEARSE, D. E., AND J. C. AVISE. 2001. Turtle mating systems: behaviour, sperm storage, and genetic paternity. J. Hered. 92:206–211. PERILLO, K. M. 1997. Seasonal movements and habitat preferences of spotted turtles (Clemmys guttata) in north central Connecticut. Chelonian Conserv. Biol. 2:445–447. SEMINOFF, J. A., K. A. BJORNDAJ, AND A. B. BOLTEN. 2007. Stable carbon and nitrogen isotope discrimination and turnover in pond sliders Trachemys scripta: Insights for trophic study of freshwater turtles. Copeia 2007:534–542. SPENCER, R. J. 2002. Testing nest site selection: fitness trade-offs and predation risk in turtles. Ecology 83:2136–2144. SURFACE, H. A. 1908. First report on the economic features of the turtles of Pennsylvania. Zool. Bull. Div. Zool. Pennsylvania Dept. Agric. 6:105–196. WARD, F. P., C. J. HOHMANN, J. F. ULRICH, AND S. E. HILL. 1976. Seasonal micro habitat selections of spotted turtles Clemmys guttata in Maryland USA elucidated by radio isotope tracking. Herpetologica 32:60–64. Herpetological Review 40(3), 2009 Herpetological Review, 2009, 40(3), 289–293. © 2009 by Society for the Study of Amphibians and Reptiles Herpetofaunal Conservation in the Rainforest: Perceptions of Ecotourists TIFFANY M. DOAN Department of Biology, Central Connecticut State University 1615 Stanley Street, New Britain, Connecticut 06050, USA e-mail: tiffperu@yahoo.com The Tambopata region of Peru has the highest overall species diversity of reptiles and amphibians recorded in the world (Doan and Arizábal 2002). Notable amphibians include ﬁve species of poison dart frogs and 45 tree frog species (Doan and Arizábal 2002). Three large reptile species are under threat of extinction due to harvesting for eggs or skins: the black caiman (Melanosuchus niger), the spectacled caiman (Caiman crocodilus), and the yellow-spotted river turtle (Podocnemis uniﬁlis) (Kirkby et al. 2000). Ecotourists visiting the Tambopata region have the opportunity to see many herpetofaunal species during their visit. The rise of ecotourism in Latin America has brought a mix of beneﬁts and problems for local people, national economies, and natural habitats (Groom et al. 1991). In the Tambopata region of Peru, the number of rainforest tourist lodges has increased greatly in recent years, with a corresponding inﬂux of foreign visitors (Groom et al. 1991; Kirkby 2002). As tourist lodges seek greater proﬁts by attracting more visitors, the local habitats may be stressed or altered (Doan 2000; Fennell 1999; Grossberg et al. 2003; Mieczkowski 1995; Orams 2002; Wheeler 1997). Moreover, tourist operations may affect the behavior (Arianoutsou, 1988; Grossberg et al. 2003) or populations (Kirkby and Cornejo Farfán 2000) of fauna in an attempt to create a memorable experience for visitors. Offering guided hikes into the rainforest and boat journeys on rivers allows tourists to encounter many animal species. In some cases guides or other lodge personnel may capture certain animals so that tourists can observe the animals up close. In most cases the animals are then released back into their natural habitat. In addition, many tourist operators capture and restrain animals so that they remain near the lodge, forming a zoo-like atmosphere for personal encounters with animal species (Orams 2002; pers. observ.). Capturing, restraining, and provisioning wild animals have been hypothesized to be detrimental to wild animal populations (Orams 2002). The practice of capturing or caging animals at ecotourist lodges may have two impacts on the rainforest experience of the ecotourists that visit such lodges. The ﬁrst effect would be a positive one, where the tourist is able to view animals that are not easily viewed or view animals at close proximity that are normally wary of the presence of humans. In this case, the tourism experience is enhanced by close contact with animals. However, as many international ecotourists are well educated, environmentally conscious individuals (Sekercioglu 2002), it is possible that the practices of capturing and provisioning animals would be perceived negatively, because tourists may be aware of the potential detrimental effects on the captive or tame animals and their populations. The perceptions of ecotourists toward the treatment of wild animals may differ depending on the animal group concerned. Tourists are thought to have deep affection for animals such as monkeys and birds, but lower affection for less desirable animals such as turtles or frogs (Pough et al. 2004; Tremblay 2002). Many bird watchers are content to devote much time and money to viewing interesting bird species. Moreover, the thoughts of ecotourists about the conservation status of less desirable animals such as reptiles and amphibians may be far removed from their feelings about well-liked mammals and birds. Tourists who are highly concerned about the well being of mammals and birds may have little or no interest or concern for reptiles and amphibians. Tambopata, Peru currently has twelve large tourist lodges lining the Madre de Dios and Tambopata Rivers, in addition to several smaller lodges (Kirkby 2002). Among these lodges, the policies and practices of capturing, restraining, and provisioning wild animals differ from an absolute ban on these practices, to common use of such practices, including the viewing of captive and tame animals as one of the primary foci of the rainforest lodge experience. As pointed out by Budowski (1976), ecotourists may not always be lucky enough to encounter the large charismatic mammals or birds that they wish to see, but they have a good chance to see the smaller creatures, such as reptiles and amphibians. To determine if the conservation practices of rainforest tourist lodges in southeastern Peru matched the wishes of their ecotourists, tourists were requested to ﬁll out a questionnaire that queried their attitudes towards a rather unpopular group of animals: reptiles and amphibians. METHODS Study Area.—The study took place in the Tambopata Province of the Madre de Dios Department of southeastern Peru. The four tourist lodges included in the study lie in the buffer zone surrounding the Reserva Nacional Tambopata. This region has seen a marked increase in tourism in the past 25 years and tourism has become one of the most important industries for the region (Groom et al. 1991; Kirkby 2002). The habitat in the Tambopata Province is classiﬁed as a Tropical/Subtropical Moist Forest according to the Holdridge system (Gentry and León 1997; Holdridge et al. 1971) and receives an average of 2400 mm of rain annually (Duellman 2005). The majority of tour visits to each of the four lodges last two nights, although longer tours are available. A typical tour package includes retrieval of the visitors at the airport in Puerto Maldonado on the ﬁrst morning, followed by bus and boat transport to the lodge. An orientation talk and afternoon hike complete the ﬁrst day. The second morning guides take groups out for a longer hike that may include a canoe ride on an oxbow lake. The second afternoon is typically free and the second night may offer a lecture by one of the guides or visiting scientists and/or a boat excursion on the river to view caimans, other nocturnal wildlife, and the stars. During the nocturnal boat trip guides may capture small caimans and bring them aboard the boat to show the tourists, after which the caimans are released back into the river. The next morning the tourists are taken back to the airport for their return ﬂight to Cusco or Lima. Ecotourist Questionnaire.—During 1997 and 1998 tourists at four ecotourism lodges in the Tambopata Province, Peru were asked to ﬁll out a questionnaire during the last night of their rainforest visit. Both English and Spanish versions of the Herpetological Review 40(3), 2009 289 questionnaire were offered. All guests were invited to ﬁll out the questionnaire but a small percentage (<5%) declined to partake in the survey. The four lodges were: EcoAmazonía (EA) and Reserva Amazónica (RA; formerly named Cuzco Amazónico; Duellman and Koechlin 1991) on the Madre de Dios River and Explorer’s Inn (EI; Erwin 1984) and Libertador Tambopata Lodge (LTL; formerly named Tambopata Jungle Lodge; Yu et al. 1997) on the Tambopata River. The questionnaire asked tourists to rate familiarity, viewing desirability, and feelings about the conservation status of the avifauna and herpetofauna (bird questionnaire results are not presented in this paper). They were additionally asked if they would like the lodges to capture animals during guided trips or to keep animals as pets of the lodge. The herpetofaunal portion of the English questionnaire is depicted in Figure 1. Not all respondents answered every question; therefore, the results of each question are based on different total numbers. Chisquare goodness-of-ﬁt statistical tests were conducted for question numbers 16–23. For each of these, the number of agree and strongly agree responses were tested versus disagree and strongly disagree responses. For these statistical tests, neutral responses were not used (they were a miniscule fraction of responses on most questions). Chi-square tests were also used for question 24, with one test used to determine if there were signiﬁcant differences in “1” responses (largest threat) and another test used to determine if there were differences in “6” responses (least threat). RESULTS In total 129 questionnaires were ﬁlled out by ecotourists at the four sites included in the study (EA 26; RA 14; EI 13; LTL 26; the remainder did not indicate which lodge they had visited). Nineteen of the questionnaires completed were the Spanish version and 110 were the English version. Ecotourists that ﬁlled out questionnaires ranged in age from 14–82 years and were from 17 different countries (Argentina, Australia, Belgium, Canada, France, Germany, Ireland, Israel, Italy, Luxemburg, The Netherlands, New Zealand, Peru, Spain, Switzerland, United Kingdom, and United States). The mean age of the respondents was 41.2 years and the most common nationalities were United States (29.09%), Australia (16.36%), Canada (12.73%), and the United Kingdom (10.00%). All other nationalities were represented by less than 10 respondents. Of the tourists that indicated their gender, 56 were male and 56 female. Thirty-two respondents (28.6%) stated that they have a scientiﬁc background or qualiﬁcations. A summary of the questionnaire results is given in Table 1. Familiarity and Local Experience.—Out of the reptile and amphibian groups about which tourists were asked to rate familiarity, tree frogs were the most familiar animals. Tourists were least familiar with poison dart frogs and coral snakes. Tourists claimed to have seen a wide variety of reptiles and amphibians during their stay. Although crocodiles do not occur in the area, four tourists claimed to have seen both caimans and crocodiles during their lodge visit. Desire to View.—Tourists wished to see a large variety of reptiles and amphibians. The most popular animals were caimans, followed closely by boa constrictors, but no tourist wished to see venomous snakes and two respondents stated they did not wish to 290 see any reptiles or amphibians at all. When asked if they would like to see more reptiles and amphibians than they had seen during their stay, the large majority of tourists stated that they would. The difference between respondents who answered agree or strongly agree versus those who answered disagree or strongly disagree was highly signiﬁcant, both for amphibians and reptiles. Although not regularly offered by any of the lodges included in this study (but sometimes offered on an ad hoc basis), the large majority of tourists (91.1%) would like to participate in a night hike to view nocturnal animals, including most reptiles and amphibians. Caiman Viewing.—Just over half (59.8%) of tourists who responded participated in a nocturnal boat journey to view caimans (primarily spectacled caimans, Caiman crocodilus). All of the respondents to this question (n = 67) but one saw caimans during their trip. The number of caimans seen by individuals ranged from one to ten. Pets and Capture of Reptiles and Amphibians.—Contrary to the beliefs of most of the lodge operators and guides (pers. comm.), the large majority of tourists (91.8%) did not wish to see reptiles and amphibians kept as pets and did not approve (94.2%) of guides capturing caimans during the nocturnal boat ride. Conservation.—The large majority of tourists (95.4%) agreed that reptile and amphibian conservation is as important as bird and mammal conservation and perceived that deforestation was seen as the most important threat (57.1% ranked it number 1), whereas tourism and global warming were seen as least threatening to reptile and amphibian species (41.9% and 32.4%, respectively, ranked them number 6). More Information Needed.—Seventy-three percent of responding tourists felt that they needed additional information regarding venomous snakes in the area. Suggestions by the lodge tourists were a nighttime presentation in the dining room and that in general more information about all snakes, both venomous and nonvenomous, would be welcome. DISCUSSION The highly signiﬁcant results of this study were remarkable. Contrary to expectations and the beliefs of tourist operators at the four lodges included in this study, visitors had strong interest in reptiles and amphibians. Despite their low level of familiarity prior to the visit, nearly all visitors expressed interest in observing more reptiles and amphibians, and most tourists wished to participate in a night hike. None of the four tourist lodges included in this study regularly offer night hikes to their visitors. Tourist operators may think that tourists would rather rest at night or would be fearful of walking in the forest at night, but the results of this study clearly show the contrary. Tourist operators could increase the unique experience of the rainforest visit for their tourists by offering night hikes, allowing their guests to see not only many more amphibians and reptiles, but also give them the chance to see other nocturnal animals, such as tarantulas, opossums, bats, and owls. Also, tourists would have the opportunity to understand the differences in the ecology of nocturnal animals versus diurnal ones. A signiﬁcant majority of visitors would like to view more reptiles and amphibians in their natural setting and did not wish to have Herpetological Review 40(3), 2009 FIG. 1. The English language version of the herpetofaunal questionnaire. Questions 1–12 surveyed the tourists about birds and are not included in this study. Herpetological Review 40(3), 2009 291 TABLE 1. Results of select questions of the questionnaire. n is total responding for each question. χ2 p 109 90.000 < 0.001 4 2 110 72.516 < 0.001 Disagree Strongly disagree 4 4 112 60.844 < 0.001 4 4 12 Disagree Strongly disagree 33 57 110 68.612 < 0.001 Strongly agree Agree Neutral 2 4 8 Disagree Strongly disagree 28 69 111 80.398 < 0.001 Reptiles and amphibians should be as protected as birds and mammals: Strongly agree Agree Neutral 84 19 1 Disagree Strongly disagree 1 4 109 88.926 < 0.001 Largest threat to herpetofaunal conservation: Deforestation Water pollution Collection for pet trade 72 23 13 Hunting Global warming Tourism 13 5 0 126 163.333 < 0.001 Least threat to herpetofaunal conservation: Tourism Global warming Collection for pet trade 62 48 14 Hunting Water pollution Deforestation 12 10 2 107 119.242 < 0.001 Tourists request more information about venomous reptiles: Yes 73 No 27 100 Question Answers n Tourists are familiar with: Tree Frogs Pit Vipers River Turtles Caimans 81 40 68 72 Crocodiles Poison Dart Frogs Boa Constrictors Coral Snakes 76 27 71 35 105 During their stay tourists saw: Caimans Turtles Lizards Tree frogs 85 68 67 44 Nonvenomous snakes Poison dart frogs Venomous snakes Crocodiles 25 10 5 4 109 Tourists would most like to see: Caimans Boa constrictors Poison dart frogs Tree frogs Turtles 39 25 9 6 6 Crocodiles Lizards Venomous snakes No reptiles or amphibians 4 1 0 2 104 Tourists would like to see more amphibians: Strongly agree Agree Neutral 49 41 19 Disagree Strongly disagree 0 0 Tourists would like to see more reptiles: Strongly agree Agree Neutral 48 41 15 Disagree Strongly disagree Tourists would like to participate in a night walk: Strongly agree Agree Neutral 42 40 22 The lodge should keep captive reptiles and amphibians: Strongly agree Agree Neutral The lodge should capture caimans to show the tourists: them captured or kept as pets. Two of the tourist lodges included in this study (EA and RA) have recently devoted much of their effort into raising animals as pets or attracting them to the lodge with food. Although the animals they currently restrain and attract have been mammals and birds, it is conceivable that they would also keep animals such as tortoises or frogs as pets. The results of this study clearly show that keeping reptiles or amphibians as pets would be contrary to the wishes of the ecotourists. 292 In addition, respondents were concerned about the conservation of reptiles and amphibians. They cared about the conservation status of these strange, unfamiliar animals, just as they would for more charismatic mammals and birds. Ecotourists considered deforestation to be the most important threat to reptile amphibian conservation by far. Few tourists considered water pollution, collection for the pet trade, hunting, global warming, or tourism to be important threats. While deforestation is obviously an important threat for Herpetological Review 40(3), 2009 all rainforest species (Duellman and Trueb 1986; Laurance and Bierregaard 1997), collection for the pet trade of animals such as boa constrictors, and hunting of caimans and river turtle eggs also are deleterious to local populations (Duellman 2005; Kirkby 2002; Pough et al. 2004). In addition, water pollution has been shown to impact amphibian populations in some areas (Duellman and Trueb 1986; Stebbins and Cohen 1995), and global warming has been hypothesized to be contributing to the global amphibian population decline (Pough et al. 2004; Stebbins and Cohen 1995). Tourists in Tambopata need to receive more education about the potential threats to the herpetofauna to be fully informed about conservation issues affecting local species. The results of this study seemed to be consistent over all groups of people who responded to the questionnaire—nationalities, genders, and ages, and did not depend upon which lodges the tourists were visiting. This suggests that these results may be broadly applicable to other rainforest lodges and, perhaps, to other areas of the world. Ecotourism has the potential to be a sustainable form of utilizing rainforest resources (Groom et al. 1991; Scace 1993), and the encouraging results of this study suggest that the visitors to Tambopata may contribute to sustainability by promoting responsible ecotourism. Acknowledgments.—My greatest thanks go to the lodge visitors that agreed to take the time to complete the questionnaire. I would like to thank Huw Lloyd and Wilfredo Arizábal Arriaga for the ideas to begin this study and for help implementing it. I would also like to thank the lodge owners, managers, and guides and EcoAmazonía, Reserva Amazónica, Explorer’s Inn, and Libertador Tambopata Lodge for permitting me to conduct the study with their guests. I would like to thank Chris Blair and Carol L. Spencer for their comments on the manuscript. This study was carried out during Project Tambopata, which was funded by the Percy Sladen Memorial Fund, the Tambopata Reserve Society (UK), the Grand Circle Foundation, the Lindeth Charitable Trust, the Albert Reckitt Trust, and the Anglo-Peruvian Society. LITERATURE CITED ARIANOUTSOU, M. 1988. Assessing the impacts of human activities on nesting of loggerhead sea-turtles (Caretta caretta L.) on Zakynthos Island, Western Greece. Environ. Cons. 15:327–334. BUDOWSKI, G. 1976. Why save tropical rain forests? Some arguments from campaigning conservationists. Amazoniana 4:529–538. DOAN, T. M. 2000. The effects of ecotourism in developing nations: an analysis of case studies. J. Sustainable Tourism 8:288–304. ________ , AND W. ARIZÁBAL ARRIAGA. 2002. Microgeographic variation in species composition of the herpetofaunal communities of Tambopata Region, Peru. Biotropica 34:101–117. DUELLMAN, W. E. 2005. Cusco Amazónico: The Lives of Amphibians and Reptiles in an Amazonian Rainforest. Comstock Publ. Assoc., Ithaca, New York. 472 pp. ________ , AND J. E. KOECHLIN. 1991. The Reserva Cuzco Amazonico, Peru: biological investigations, conservation, and ecotourism. Occas. Pap. Mus. Nat. Hist., Univ. Kansas 142:1–38. ________ , AND L. TRUEB. 1986. Biology of Amphibians. Johns Hopkins University Press, Baltimore, Maryland. 670 pp. ERWIN, T. L. 1984. Tambopata Reserved Zone, Madre de Dios, Perú: history and description of the Reserve. Rev. Peruana Entomol. 27:1–8. FENNELL, D. A. 1999. Ecotourism: An Introduction. Routledge, London. 315 pp. GENTRY, A. H., AND B. LEÓN. 1997. Tambopata region, Peru. In S. D. Davis, V. H. Heywood, O. Herrera-MacBryde, J. Villa-Lobos, and A. C. Hamilton (eds.), Centres of Plant Diversity: a Guide and Strategy for their Conservation, Volume 3—The Americas, pp. 355–359. WWF and IUCN, Oxford. GROOM, M. J., D. PODOLSKY, AND C. MUNN. 1991. Tourism as a sustainable use of wildlife: a case study of Madre de Dios, south-eastern Peru. In J. Robinson and K. Redford (eds.), Neotropical Wildlife Use and Conservation, pp. 393–412. University of Chicago Press, Chicago, Illinois. GROSSBERG, R., A. TREVES, AND L. NAUGHTON-TREVES. 2003. The incidental ecotourist: measuring visitor impacts on endangered howler monkeys at a Belizean archaeological site. Environ. Cons. 30:40–51. HOLDRIDGE, L. R., W. C. GRENKE, W. H. HATHEWAY, T. LIANG, AND J. A. TOSI. 1971. Forest Environments in Tropical Life Zones: A Pilot Study. Pergamon Press, New York, New York. 780 pp. KIRKBY, C. A. 2000. The impact of trail-use by tourists on the mammal fauna of Tambopata, south-eastern Peru. In C. A. Kirkby, T. M. Doan, H. Lloyd, A. Cornejo Farfán, W. Arizábal Arriaga, and A. Palomino Marín (eds.), Tourism Development and the Status of Neotropical Lowland Wildlife in Tambopata, South-eastern Peru: Recommendations for Tourism and Conservation/El Desarrollo Turístico, su Impacto sobre la Fauna Neotropical de Tambopata, Sureste del Perú: Recomendaciones para el Turismo y la Conservación, pp. 14–62. Tambopata Reserve Society, London. ________ . 2002. Estándares Ecoturísticos para la Reserva Nacional Tambopata, el Parque Nacional Bahuaja Sonene y sus Zonas de Amortiguamiento, Madre de Dios, Perú. WWW-Oﬁcina del Programa del Perú, Puerto Maldonado, Peru. 32 pp. ________ , T. M. DOAN, H. LLOYD, A. CORNEJO FARFÁN, W. ARIZÁBAL ARRIAGA, AND A. PALOMINO MARÍN (EDS.). 2000. Tourism Development and the Status of Neotropical Lowland Wildlife in Tambopata, South-eastern Peru: Recommendations for Tourism and Conservation/El Desarrollo Turístico, su Impacto sobre la Fauna Neotropical de Tambopata, Sureste del Perú: Recomendaciones para el Turismo y la Conservación. Tambopata Reserve Society, London. 154 pp. LAURANCE, W. F., AND R. O. BIERREGAARD. 1997. Tropical Forest Remnants. University of Chicago Press, Chicago, Illinois. 632 pp. MIECZKOWSKI, Z. 1995. Environmental Issues of Tourism and Recreation. University Press of America, Lanham, Maryland. 552 pp. ORAMS, M. B. 2002. Feeding wildlife as a tourism attraction: a review of issues and impacts. Tourism Manag. 23:281–293. POUGH, F. H., R. M. ANDREWS, J. E. CADLE, M. L. CRUMP, A. H. SAVITZKY, AND K. D. WELLS. 2004. Herpetology. Pearson Education, Upper Saddle River, New Jersey. 736 pp. SCACE, R. C. 1993. An ecotourism perspective. In G. Nelson, R. Butler, and G. Wall (eds.), Tourism and Sustainable Development: Monitoring, Planning, Managing, pp. 59–82. University of Waterloo, Waterloo, Canada. SEKERCIOGLU, C. H. 2002. Impacts of birdwatching on human and avian communities. Environ. Cons. 29:282–289. STEBBINS, R. C., AND N. W. COHEN. 1995. A Natural History of Amphibians. Princeton University Press, Princeton, New Jersey. 315 pp. TREMBLAY, P. 2002. Tourism wildlife icons: attractions or marketing symbols? J. Hospitality and Tourism Management 9:164–180. WHEELER, B. 1997. Tourism’s troubled times: responsible tourism is not the answer. In L. France (ed.), The Earthscan Reader in Sustainable Tourism, pp. 61–67. Earthscan Publications, London. YU, D. W., T. HENDRICKSON, AND A. CASTILLO. 1997. Ecotourism and conservation in Amazonian Perú: short-term and long-term challenges. Environ. Cons. 24:130–138. Herpetological Review 40(3), 2009 293 Herpetological Review, 2009, 40(3), 294–301. © 2009 by Society for the Study of Amphibians and Reptiles Distribution and Natural History Notes on Some Poorly Known Frogs and Snakes from Peninsular Malaysia CHAN KIN ONN Institute for Environment and Development (LESTARI) Universiti Kebangsaan Malaysia, 43600 Bangi Selangor Darul Ehsan, Malaysia e-mail: kin_onn@yahoo.com and NORHAYATI AHMAD School of Environment and Natural Resource Sciences Faculty of Science and Technology, Universiti Kebangsaan Malaysia 43600 Bangi, Selangor Darul Ehsan, Malaysia e-mail: yati_68@yahoo.co.uk Despite having been studied for many years, the status and knowledge of the herpetofauna of Peninsular Malaysia is far from being complete or fully understood. Although relatively recent new species discoveries have added signiﬁcantly to the overall biodiversity of the region (Chan and Grismer 2008; Chan et al. 2009; Das et al. 2007; Das and Lim 2001; Grismer and Chan 2008; Grismer et al. 2006 and references therein; Grismer et al. 2008a, b, c; Grismer et al. 2009; Grismer and Norhayati 2009; Leong and Lim 2003; Matsui and Jaafar 2006; McLeod and Norhayati 2007; Van Rooijen and Vogel 2008; Vogel and Van Rooijen 2007; Wood et al. 2008, 2009), information on existing species are still inadequate or lacking altogether. Alongside these new ﬁndings, intensive and systematic ﬁeld work has also uncovered a number of rare species (e.g., Cyrtodactylus sworderi, Grismer et al. 2007; Enhydris pahangensis, Chan, in press) which have not been recorded since their discoveries in the early 1900s. Many of these species lack complete descriptions and/or proper color illustrations and next to nothing is known about their life history as they have rarely been observed in the wild. This paper addresses some of these issues by providing expanded descriptions, color photographs, life history observations, and new locality records of the more poorly known frogs and snakes of Peninsular Malaysia. MATERIALS AND METHODS Color photographs were taken for vouchers and color pattern comparisons. Measurements were taken with Mitutoyo digimatic caliper to the nearest 0.1 mm. Measurements taken for frogs are as follows: snout–vent length (SVL); tibia length (TiL), measured from the center of the knee to the centre of the ankle. For snakes: snout–vent length (SVL); tail length (TaL), from cloaca to tail tip; number of dorsal scale rows (DoS), counted at one head length posterior to head, at midbody, and one head length anterior to vent; number of ventral scale rows (Ven); number of supralabials (SupL), with the number of SupL in contact with the orbit in brackets; number of infralabials (InfL). Voucher specimens or photographs are provided to support new locality records. All specimens have been deposited at the museum of the Department of Wildlife and National Parks Peninsular Malaysa (DWNP) and the herpetological collection at Universiti Kebangsaan Malaysia (UKM). HC refers to the Herpetological Collection of Universiti Kebangsaan Malaysia 294 (UKM), Bangi, Selangor, Peninsular Malaysia; UKMDPC refers to Universiti Kebangsaan Malaysia Digital Photograph Collection; LSUDPC refers to the La Sierra University Digital Photograph Collection. Institutional abbreviations follow Leviton et al. (1985), except we retain ZRC for USDZ, following conventional usage. Measurements taken on voucher specimens are presented in Table 1. RESULTS Species accounts Rhacophorus robinsoni Boulenger 1903 Robinson’s Treefrog Fig. 1 Rhacophorus robinsonii Boulenger 1903 in Annandale and Robinson (eds.), Fasciculi Malayenses, 2(1):136. Type locality: “Bukit Besar. 2500 feet”, Pattani, Thailand. Rhacophorus (Rhacophorus) robinsonii Ahl 1931:157 Rhacophorus pardalis robinsoni Wolf 1936:208 Rhacophorus robinsoni Inger 1954:372; Berry 1975:109; Manthey & Grossman 1997:138; Das & Norsham 2007:70 Description. SVL male 59 mm; female 82 mm; Vomerine teeth in two long oblique series touching the inner edge of the choanae; snout obtusely pointed; canthus rostralis sharp; tympanum distinct, as high as wide; supratympanic fold distinct, outlined with black, extends horizontally posterior to the orbit, slants, and ends abruptly posterior to the tympanum. Finger tips expanded into large disks bearing circum-marginal and transverse ventral grooves; width of three outer ﬁngers equal to tympanum diameter; broad webbing reaches base of disks of outer three ﬁngers but only halfway to disk of ﬁrst ﬁnger; nuptial pad present in males. Toe disks smaller than those of ﬁngers, bearing circum-marginal and transverse ventral grooves; completely webbed to disks on all toes; a small inner metatarsal tubercle; no outer metatarsal tubercle. Skin above smooth; slightly granular on ﬂanks and throat; belly covered with large, smooth, rounded granules; arms and tarsus without ﬂaps of skin; no anal ﬂap. Color in life. Pale brown or dark chocolate brown to light green above; two thin, black bars on the head, one crossing the snout, and another longer bar between the orbits; dorsum scattered with dark blotches; with or without a white spot below the tympanum; sides of head a darker shade of brown; ﬂanks, inner and outer sides of limbs yellowish with black and whitish-blue reticulations; limbs bearing dark cross-bars; interdigital webbing black with ﬁne brown veins; throat whitish, mottled with brown; belly white with dark reticulations. Juveniles: yellow to orange interdigital webbing; underside of the body yellowish. Life history. Rhacophorus robinsoni inhabits primary or old, undisturbed secondary forests and has been found on tree tops as high as 10 m above the forest ﬂoor (Grismer et al. 2004) and on leaves as low as 1 m above the ground. Like many of its other congeners, frogs descend lower to the ground following heavy rainfall to breed along stagnant puddles or ponds. However, one specimen from Ulu Gombak was found sitting on a leaf during an extremely dry night when no other frogs were abroad and Grismer et al. (2004) recorded a gravid female during the day, crouched in Herpetological Review 40(3), 2009 the interstices of truck tracks on a logging road far from any body of water. Distribution in Peninsular Malaysia. Kuala Teku, Pahang (Berry 1975); Gombak, Selangor (Yong 1977); Temenggor, Perak (Grismer et al. 2004); Cameron Highlands, Pahang; Kampung Chennah, Negeri Sembilan (this report). Notes. In Peninsular Malaysia, Rhacophorus robinsoni has only been recorded three times, the ﬁrst being from Kuala Teku, Pahang in the early 1900’s (Boulenger 1912) and the second from Gombak, Selangor (Yong 1977). A third specimen was reported by Grismer et al. (2004) from Temenggor Forest Reserve, Perak, but was left unidentiﬁed (Rhacophorus sp.). This species has since been found at Cameron Highlands, Pahang (UKMDPC 1.0002) and Kampung Chennah, Negeri Sembilan (HC 191,192; Fig. 1). Given the current distribution pattern of R. robinsoni which ranges from southern Thailand to its southernmost record in Negeri Sembilan, it is likely that this species has a wider distribution range and probably occurs throughout Peninsular Malaysia. Theloderma leprosa Tschudi 1838 Fig. 2 Theloderma leprosa Tschudi 1838, Classiﬁcation der Batrachier, 32: 73. Type locality: “Sumatra”, Indonesia. Hyla leprosa Schlegel 1858:55 Theloderma leprosum Boulenger 1884:21 Polypedates leprosus Günther 1887:315 Rhacophorus leprosus Boulenger 1890:324; Berry 1975:100 Theloderma leprosa Das & Norsham 2007: 71 Description. SVL 65 mm; Vomerine teeth in two short series touching the inner front edge of the choanae; head broader than long; canthus rostralis visible, angular; lores distinctly concave; nares protruding from snout with an internarial sulcus; tympanum distinct, ¾ eye diameter, ellipsoidal, covered with small, rounded warts not bearing asperities; supratympanic fold absent. Finger tips expanded into disks bearing circum marginal grooves; ﬁngers free of web; outer metacarpal tubercle present; nuptial pad present in males. Disks of toes smaller than those of ﬁngers, bearing circum marginal grooves; toes nearly entirely webbed, reaching disks of third and ﬁfth, third phalanx of fourth; subarticular tubercles well developed; inner metatarsal tubercle present. Entire upper surface covered with prominent warts of varying sizes, all bearing granular asperities; a series of large, ridge-like warts on back and hind limbs; throat with raised granules bearing apical asperities; chest and belly covered with large, smooth, rounded granules. Chocolate to grayish brown above; asperities on warts whitish which gives it a speckled appearance; underside of body, lower side of ﬂanks and inner side of limbs black with whitish and pale blue reticulations; interdigital webbing and disks orange. Life history. Theloderma leprosa has been observed in tree-holes as high as 3 m above ground (pers. obs.) The size of these large thelodermids (Max. SVL 65 mm) may play a role in habitat stratiﬁcation within this group, with smaller congeners (T. asperum, T. horridum, and T. licin) and other tree-hole dwellers (Metaphrynella pollicaris and Phrynella pulchra) occupying smaller tree holes and cavities closer to the ground. Thus far, frogs have only been found in primary and old secondary forests where bigger trees are presumably able to provide the speciﬁc microhabitat require- ments for these large tree-hole breeders. Berry (1975) noted that a specimen was found in a hole made by cicadas in the ground while Dring (1979) found a specimen on a palm leaf 1.5 m above ground at 790 m in elevation. Theloderma leprosa appears to be a high altitude species, occurring at elevations of approximately 800 m and above sea level. Distribution in Peninsular Malaysia. Bukit Larut, Perak; Gunung Angsi, Negeri Sembilan; Gunung Tapis, Pahang (Berry 1975); Gunung Lawit, Terengganu (Dring 1979); Cameron Highlands, Pahang; Gunung Besar Hantu, Negeri Sembilan (this report). Notes. Theloderma leprosa was ﬁrst discovered in the Malay Peninsula by Mr. L. Wray from Larut Hills [Bukit Larut], Perak at 4000 feet elevation (Boulenger 1912). Subsequent locality records were from Gunung Angsi, Negeri Sembilan (at 2600 feet) and Gunung Tapis, Pahang (Berry 1975). Dring (1979) reports this species from Gunung Lawit, Terengganu. Since then, this species has also been discovered in Cameron Highlands, Pahang and Gunung Besar Hantu, Negeri Sembilan (HC 281; Fig. 2). Thus far, T. leprosa has been shown only to occur at high elevations in all three of Peninsular Malaysia’s main mountain ranges, Bukit Larut on Banjaran Bintang; Cameron Highlands, Gunung Besar Hantu and Gunung Angsi on Banjaran Titiwangsa; Gunung Lawit and Gunung Tapis on Banjaran Timur. Following such trends, this species is likely to occur on other mountain tops throughout those mountain ranges. Hylarana siberu (Dring, McCarthy & Whitten 1990) Fig. 3 Rana siberu Dring, McCarthy & Whitten 1990. Indo-Malayan Zoology, 6(1989): 119-132. Type locality: “Teitei Bulak, Sabeuleleu, Siberut [Island], 1° 21´S, 98° 59´E”, Mentawai Islands, Indonesia. Rana (Pulchrana) siberu Dubois 1992:326 Pulchrana siberu Frost et al. 2006:369 Hylarana siberu Che et al. 2007:1 Description. SVL male 39.5 mm; female 44.9 mm; Skin above smooth, slightly rugose on back, ﬂanks and underside of body. Tympanum visible, slightly higher than wide, about ½ eye diameter; supratympanic fold absent. Nuptial pads absent in males; humeral glands enlarged. Fingers with rudimentary web; outer metatarsal tubercle round, inner one oval. Web reaching one half phalanx of ﬁrst toe and ﬁrst phalange of second toe, one half phalanx between second and third toes, base of disk of third toe and second phalanx of fourth toe, two half phalanges between fourth and ﬁfth toes; inner metatarsal tubercle oval. Dorsum completely black without markings; complete or nearly complete reddish/orange dorsolateral stripe from rostrum, along the canthal region, lateral margin of upper eyelid, along body and terminating at sacral region; series of small, white spots on upper labial; ﬂanks and upper side of limbs with round, creamy yellow spots, sometimes connected together forming short, roundish streaks; throat and underside of limbs whitish, heavily stippled with dark brown; belly whitish with dark brown reticulations Life history. Hylarana siberu inhabits lowland primary forest and has been found along the edge of a temporary pool in a swampy area away from streams. Another specimen was caught in a pit-fall trap away from streams as well which may indicate that this species is not a riparian as is its putatively close relative Herpetological Review 40(3), 2009 295 Hylarana picturata which it most closely resembles. Distribution in Peninsular Malaysia. Lakum Forest Reserve, Pahang (Leong and Lim 2004); Kuala Gandah, Pahang (this report). Notes. Hylarana siberu was previously known from the Indonesian Island of Siberut (west Sumatra) and Sumatra only (Dring et al. 1990). The ﬁrst record of this species in Peninsular Malaysia was of a single adult male from the Lakum Forest Reserve, Pahang (Leong and Lim 2004). This paper reports the second known specimen of an adult female (DWNP 1189; Fig. 3) from Kuala Gandah, Pahang, approximately 25 km from Lakum Forest Reserve. In Peninsular Malaysa, H. siberu most closely resembles H. picturata but differs by having an immaculate dorsum vs. a spotted dorsum; continuous, unbroken dorsolateral stripe vs. discontinuous dorsolateral stripe formed by closely linked spots; and reduction in webbing on toes vs. nearly full webbed. Reptilia Calliophis gracilis Gray 1835 Spotted Coral Snake Fig. 4 Calliophis gracilis Gray 1835. Illustrations of Indian Zoology, chieﬂy selected from the collection of Major - General Hardwicke. Vol. 2. London (1833–1834): 263 pp. Callophis gracilis Günther 1864 Elaps nigromaculatus Cantor 1847:1029 Callophis gracilis Boulenger 1912:203 Calliophis gracilis Tweedie 1983:110 Calliophis gracilis Welch 1994:39; Slowinski, Boundy & Lawson 2001 Description. Body slender, relatively long, up to 740 mm total length; tail very short, terminating in a blunt point; dorsal scales arranged in 13/13/13 rows; ventrals 303–324; subcaudals 21–28; six supralabials, 3rd and 4th touching the orbit; ﬁve infralabials. Body gray above with a thin, black, vertebral stripe connected to a series of small black spots. Similar lateral stripe connects a series of larger, black spots which alternate with smaller, vertebral spots. Top of head with dark, symmetrical markings. Underside of body with alternating black and yellow bands. Underside of tail banded black and orange. Life history. A specimen from Cameron Highlands was found by Orang Asli collectors above 700 m elevation. At Bukit Lagong, Selangor, a subadult specimen was observed in the afternoon, swimming across a dammed-up portion of stream at 350 m elevation. Distribution in Peninsular Malaysia. Penang, southern Perak, Selangor (Boulenger 1912); Fraser’s Hill, Pahang (Leong and Lim 2003); Cameron Highlands, Pahang (this report) Notes. According to Cantor (1847), Calliophis gracilis is common in the hills of Penang but no reports of this species have been made from that island since his early observations. In Selangor, this snake is known from Gombak and Bukit Lagong, Kepong. Xenelaphis ellipsifer Boulenger 1900 Ocellated Brown Snake Fig. 5 Xenelaphis ellipsifer Boulenger 1900. Description of new reptiles and batrachians from Bomeo. Proc. Zool. Soc. London: 296 182–187. Type locality: Sarawak river. Xenelaphis ellipsifer Tweedie 1983:48; Manthey & Grossmann 1997: 396; Malkmus et al. 2002:369 Description. Maximum recorded length 2320 mm; 1-2 preoculars, one subocular, two postoculars; one loreal; 8-9 supralabials; 2+2 or 1+2 temporals; dorsal scales arranged in 17/17/15 rows; 186–203 ventrals; anal plate divided; 129–134 subcaudals. Brown above, paler on the sides with a series 18-20 black, circular or ellipsoidal markings, each crossing the back and touching the ventrals on either side; the circles are bordered externally with paler brown or white and in between each circle, lower on the sides is an irregularly shaped black spot; top of head and neck rusty brown with longitudinal black markings on side of neck; sides of head and neck yellow; belly creamy white. Life history. Xenelaphis ellipsifer inhabits primary and old secondary forests from lowlands to 1100 m in elevation. Adults probably feed on ﬁsh and rodents while juveniles prefer ﬁsh, frogs and lizards (Malkmus et al. 2002). According to the Orang Asli collectors who found this species in Kampung Chennah, Negeri Sembilan, the snake was abroad at night, foraging in and along a moderately ﬂowing stream and was quick to escape when approached. The same snake was observed along the same stream a number of instances which might suggest a narrow home range. Distribution in Peninsular Malaysia. Bukit Larut, Perak; Cameron Highlands, Pahang (Tweedie 1983); Gombak, Selangor (pers. comm.); Kampung Chennah, Negeri Sembilan (this report). Notes. Xenelaphis ellipsifer was not known from Peninsular Malaysia until 1954 when a specimen was found in a collection from Bukit Larut, Perak. It was then the ﬁfth specimen of X. ellipsifer known to science, with previous records being from Borneo and Sumatra (Tweedie 1983). Lim (1958) reported a skin obtained from Cameron Highlands in 1957. Since then, X. ellipsifer has not been seen until Sheperd and Auliya (pers. comm.) reported a sighting of this species at Gombak, Selangor. Following that, another specimen was found by Orang Asli collectors in the lowland forests of Kampung Chennah, Negeri Sembilan which constitutes a new locality record (HC 194; Fig. 5). Oligodon signatus (Günther 1864) Barred Kukri Snake Fig. 6 Simotes signatus Günther 1864. The Reptiles of British India. London (Taylor & Francis), xxvii + 452 pp. FIGURES 1–8, OPPOSITE PAGE. FIG. 1. Rhacophorus robinsoni from Kampung Chennah, Negeri Sembilan (HC 191). FIG. 2. Theloderma leprosa from Gunung Besar Hantu, Negeri Sembilan (HC 281). FIG. 3. Hylarana siberu from Kuala Gandah, Pahang (DWNP 1189). FIG. 4. Calliophis gracilis from Cameron Highlands (HC 193). FIG. 5. Xenelaphis ellipsifer from Kampung Chennah, Negeri Sembilan (HC 194). FIG. 6. Oligodon signatus from Kampung Chennah, Negeri Sembilan (HC 196). FIG. 7. Asthenodipsas malaccanus from Gunung Besar Hantu, Negeri Sembilan (HC 195). FIG. 8. Enhydris pahangensis from Nenasi Forest Reserve, Pahang (HC 198). Herpetological Review 40(3), 2009 Herpetological Review 40(3), 2009 297 Oligodon signatus Wall 1923; Tweedie 1985:53; Hendrickson 1966:67; Manthey & Grossman 1997:373 Description. Maximum length 600 mm; seven supralabials, third and fourth contact orbit; loreal usually present, rarely absent on one side of the head; dorsal scales arranged in 17/15/15 rows; 141–157 ventrals; 47–60 subcaudals. Chocolate brown above with a series of reddish, chevron-like markings mid-dorsally that begin just posterior to the nape until the tip of tail. Markings are spaced approximately 8–10 scales apart at midbody. These markings are larger and more elongate at the anterior portion of the body and decrease in size towards the tail. Underside of body pinkish red. Life history. This semi-fossorial snake inhabits lowland to upper hill forests. Distribution in Peninsular Malaysia. Selangor; Malacca (Batchelor 1958); Gunung Panti Forest Reserve, Johor (Yong 2006); Cameron Highlands, Pahang; Kampung Chennah, Negeri Sembilan (This report). Notes. In Peninsular Malaysia, Batchelor (1958) reports Oligodon signatus from Selangor and Malacca. Yong (2006) reports a specimen from Gunung Panti Forest Reserve, Johor, which was positively identiﬁed from a roadkill. Two additional localities reported here are Kampung Chennah, Negeri Sembilan (HC 196; Fig. 6) and Cameron Highlands, Pahang (UKMDPC 1.0077). The latter represents the northern most record for this species. Asthenodipsas malaccanus Malayan Slug Snake Fig. 7 Asthenodipsas malaccana Peters 1864. Über neue Amphibien (Typhloscincus, Typhlops, Asthenodipsas, Ogmodon). Mber. k. preuss. Akad. Wiss., Berlin: 271–276. Amblycephalus malaccanus Boulenger 1892; De Rooij 1917 Pareas malaccanus Robinson & Kloss 1920; Grandison 1972, 1978 Internatus malaccanus Rao & Yang 1992 Asthenodipsas malaccanus Iskandar & Colijn 2002; Grossman & Tillack 2004 Description. Body cylindrical with a very weak vertebral ridge; 6–8 supralabials, 3rd and 4th, rarely only 3rd touching orbit; 5–7 infralabials; dorsal scales arranged in 15/15/15 rows; 154–180 ventrals; 26–56 subcaudals. Ground color of dorsal surface of body and tail greyish brown; top and sides of head and supralabials dusky white smudged with irregularly shaped, gray to brown markings; iris red; dull, white, vertebral line extending posteriorly approximately 14 scale rows connects white head markings with ﬁrst white, dorsal body band; dorsum overlain with approximately 28 irregularly shaped to squarish, white vertebral blotches; some blotches occur only on one or the other side of the vertebral region, covering 3–5 scale rows; others straddle the vertebral region covering 5–7 scale rows; series of broken to continuous white markings on vertebral scales connect dorsal blotches; brown stippling on centralmost scales of blotches give blotches a smudged appearance; sides of body and tail immaculate; infralabials, gular region and throat white with dark grey-brown mottling; ventral scales grayish brown, slightly lighter than color of dorsum; white midventral line extends from throat to tip of tail. 298 Life history. A gravid female carrying three eggs was found above 800 m elevation on Gunung Besar Hantu, Negeri Sembilan during the month of September (HC 195; Fig. 7). Distribution in Peninsular Malaysia. Asahan Estate, Malacca; Gunung Benom, Pahang; Perak; Selangor (Tweedie 1983); EndauRompin National Park, Johor (LSUDPC 4860); Gunung Besar Hantu, Negeri Sembilan (this report). Notes. Asthenodipsas malaccanus ranges from Thailand, through Peninsular Malaysia to Indonesia (Sumatra and the Mentawai Islands). Stuebing and Inger (1999) reported this species from Borneo, however, we have examined a series of eight specimens of A. malaccanus from Thailand, Indonesia, and Peninsular Malaysia, including the holotype (see appendix) and found them to differ notably from the Bornean population by having a weak vertebral ridge vs. a strong ridge; and a blackish dorsum with white, squarish, paravertebral blotches vs. a tan dorsum with dark, vertical bars. Therefore, we consider the A. malaccanus of Borneo to be a potentially new species. Enhydris pahangensis Pahang Water Snake Fig. 8 Enhydris pahangensis Tweedie 1946. Tweedie, M.W. F. 1946. A new snake from the Malay Peninsula. Ann. Mag. nat. Hist. (11) 13 (98):142–144. Description. Males up to 300 mm SVL; females up to 392 mm SVL; dorsal scales arranged in 25/25/21 rows; 129–132 ventrals; 47–55 subcaudals; eight supralabials, 4th touching the orbit; 11 infralabials. Grayish or yellowish brown above with a series of paired, dark spots beginning from the nape to the base of tail. Spots are usually half a scale wide and may not always be parallel to each other. Anterior portion of the head darkly stippled, slightly darker than ground body color. Labials white, heavily stippled with black. A creamcolored lateral stripe on each side of the body extends from the side of the head to the tail tip. On the body, this stripe covers four of the lowest dorsal scale rows, reduced to three towards the vent. The stripe is bordered above and below by a black zig-zag line, the line bordering the ventral scales is thicker and more pronounced. Ventrals white, edged with black, with an antero-medial dark blotch on each ventral scale. Chin scales white, mottled with black. Life history. This species inhabits primary or undisturbed secondary forests and has been found among decaying leaves in a stagnant portion of a small and slow moving stream. Distribution in Peninsular Malaysia. Kuala Tahan, Pahang (Tweedie 1983); Sungai Kura, Hulu Terengganu (Chan, in press); Nenasi Forest Reserve, Pahang (this report). Notes. In Peninsular Malaysia, Enhydris pahangensis most resembles E. plumbea but differs by having 25 dorsal scale rows vs. 19; a creamy lateral stripe distinctly bordered above and below by a black zig-zag line vs. a creamy lateral stripe lacking distinct zig-zag borders; labials, chin scales, and anterior portion of snout mottled with dark gray vs. no mottling; ventral and subcaudal scales white, edged with black and an antero-medial dark blotch on each ventral scale vs. immaculate white. Previously, E. pahangensis was only known from the type specimen from Kuala Tahan, Pahang. Subsequently, a second specimen was collected from Hulu Terengganu (Chan, in press), thereby expanding its Herpetological Review 40(3), 2009 Rhacophorus robinsoni Rhacophorus robinsoni Theloderma leprosa Hylarana siberu Calliophis gracilis Species “Kampung Chennah, Negeri Sembilan” “Kampung Chennah, Negeri Sembilan” “Kampung Chennah, Negeri Sembilan” “Kampung Chennah, Negeri Sembilan” “Kampung Chennah, Negeri Sembilan” “Kuala Gandah, Pahang” “Cameron Highlands, Pahang” Locality Adult Male Juvenile Adult Male Sex 315 850 37.7 59.2 30.4 61.1 SVL 16.9 25.7 13.9 33.1 TiL 88 30 351 TaL 17/15/15 17/17/15 13/13/13 DoS 311 191 Ven Mus. No. Adult Female Adult Male Adult Female 384 HC 191 HC 192 HC 281 DWNP 1189 HC 193 HC 194 Adult Female (gravid) Adult Female (gravid) 39 25/25/21 25/25/21 15/15/15 131 132 156 170 47 55 54 28 25 130 SubC 6 (3+4) SupL InfL 8 5 11 11 5 10 8 (4) 7 (3+4) 7 (3+4) 8 (4) 8 (4) Herpetological Review 40(3), 2009 550 69 68.5 BATCHELOR, D. M. 1958. Some notes on the snakes of Asahan, Malacca. Malayan Nat. J. 12:103–111. BERRY, P. Y. 1975. The Amphibian Fauna of Peninsular Malaysia. Tropical Press, Kuala Lumpur, Malaysia. HC 196 HC 195 300 392 LITERATURE CITED TABLE 1. Measurements taken for examined specimens. See methods and materials for abbreviations. Blank = not applicable. Xenelaphis ellipsifer Oligodon signatus Adult Male Adult Female Acknowledgments.—We thank L. Lee Grismer and Johan van Rooijen for their critical insights, comments, and contributions. HC 197 HC 198 Herpetofaunal surveys generally result in skewed observation frequencies with some species observed much more frequently than others (e.g., Das 1996; Murphy et al. 1994; Ziegler 2002) and several factors can contribute to the perceived rarity of a particular species. First, some species are rarely seen simply because of their association with speciﬁc, isolated, habitats like forest canopies, karst formations, mountain tops or islands (e.g., Chan and Grismer 2008; Chan et al. 2009; Grismer and Chan 2008; Grismer 2007, 2008; Grismer et al. 2008a, b, c; Stuebing and Tan 2002). Surveys to these locations are usually more infrequent resulting in fewer reports. Other factors include elusive lifestyles or morphology (nocturnal or fossorial habits and camouﬂage) and the fact that some species do actually occur in low densities. These factors are difﬁcult to disentangle in practice unless very strict survey methodologies are applied. However, species abundance patterns in many taxa are known to correspond roughly to a log-normal distribution with some species being abundant and many occurring in low densities (e.g., Limpert et al. 2001; Longino et al. 2002; Loreau 1992). Not surprisingly, such uneven species abundance patterns also appear to hold true for reptiles and amphibians. For instance, Lloyd et al. (1968) carried out a very thorough herpetological survey in a lowland rainforest in Sarawak and concluded that the herpetofaunal species abundance is fairly uneven, with other workers coming to similar conclusions (e.g., Dash and Mahanta 1993; Inger and Colwell 1977; Karns et al. 2005). Knowledge on the herpetofauna of Peninsular Malaysia has seen great progress in recent years with new locality records and new species growing at an unprecedented rate (Chan and Grismer 2008; Chan et al. 2009; Grismer 2008; Grismer and Chan 2008; Grismer and Pan, 2008; Grismer et al. 2004, 2006; Wood et al. 2008, 2009). Such reports are crucial for biogeographic studies and focus on the importance and utilization of regional checklists which are not to be underestimated. Grismer and Pan (2008) highlighted regions of Peninsular Malaysia most understudied and suggested areas where focused research efforts should continue and Grismer et al. (2009) stressed the importance of looking more closely at species that have already been collected. This two-pronged approach to studying the diversity of this region’s herpetofauna has already proved invaluable (see the review by Grismer et al. 2009) and will only generate more discoveries as time passes. “Gunung Besar Hantu, Negeri Sembilan” “Nenasi Forest Reserve, Pahang” “Nenasi Forest Reserve, Pahang” DISCUSSION Asthenodipsas malaccanus Enhydris pahangensis Enhydris pahangensis distribution to the neighboring state. We report of a third locality from Nenasi Forest Reserve, Pahang (HC197, HC198; Fig. 8). Due to its close resemblance to E. plumbea whose identity it has probably been masquerading under most of the time, this species is likely to encompass a wider distribution which might be revealed upon closer examination of existing museum specimens. 299 BOULENGER, G. A. 1912. A vertebrate fauna of the Malay Peninsula from the Isthmus of Kra to Singapore including the adjacent islands. Reptilia and Batrachia. Taylor & Francis, London. 294 pp. CANTOR, T. 1847. Catalogue of reptiles inhabiting the Malayan Peninsula and islands, collected or observed by Theodore Cantor, Esq., M.D. Bengal Medical Service. J. Thomas, Calcutta. 157 pp. CHAN, K. O. In press. The rediscovery of Enhydris pahangensis Tweedie, 1946. Hamadryad. ________ , AND L. L. GRISMER. 2008. A new species of Cnemaspis Strauch 1887 (Squamata: Gekkonidae) from Selangor, Peninsular Malaysia. Zootaxa 1877:49–57. ________ ________ , , A. NORHAYATI, AND B. DAICUS. 2009. A new species of Gastrophrynoides (Anura: Microhylidae): an addition to a previously monotypic genus and a new genus for Peninsular Malaysia. Zootaxa 2124:63–68. DAS, I. 1996. Spatio-temporal resource utilization by a Bornean rainforest herpetofauna: Preliminary results. Tropical Rainforest Research: 315–323. ________ , AND K. K. P. LIM. 2001. A new Bufo (Anura: Bufonidae) from the peat swamps of Selangor, Peninsular Malaysia. Rafﬂes Bull. Zool. 49(1):1–6. ________ , Y. NORSHAM, AND J. SUKUMARAN. 2007. A new species of Microhyla (Anura: Microhylidae) from the Malay Peninsula. Hamadryad 31(2):304–314. DASH, M. C., AND J. K. MAHANTA. 1993. Quantitative analysis of the community structure of tropical amphibian assemblages and its signiﬁcance for conservation. J. Biosc. 18(1):121–139. DRING, J. C. M. 1979. Amphibians and reptiles from northern Trengganu, Malaysia, with descriptions of two new geckos: Cnemaspis and Cyrtodactylus. Bull. Brit. Mus. (Nat. Hist.) 34:181–241. ________ , C. J. MCCARTHY, AND A. J. WHITTEN. 1990. The terrestrial herpetofauna of the Mentawai Islands, Indonesia. Indo-Malayan Zool. 6:119–132. GRISMER, L. L. 2007. A new species of small montane forest floor skink (genus Sphenomorphus Fitzinger 1843) from southern Peninsular Malaysia. Herpetologica 63:544–551. ________ . 2008. A new species of insular skink (genus Sphenomorphus Fitzinger 1843) from the Langkawi Archipelago, Kedah, West Malaysia with the ﬁrst report of the herpetofauna of Pulau Singa Besar and an updated checklist of the herpetofauna of Pulau Langkawi. Zootaxa 1691:53–66. ________ , AND K. O. CHAN. 2008. A new species of Cnemaspis Strauch 1887 (Squamata: Gekkonidae) from Pulau Perhentian Besar, Terengganu, Peninsular Malaysia. Zootaxa 1771:1–15. ________ , AND A. NORHAYATI. 2009. A new insular species of Cyrtodactylus (Squamata: Gekkonidae) from the Langkawi Archipelago, Kedah, Peninsular Malaysia. Zootaxa 1924:53–68. ________ ________ , , AND K.O. CHAN. 2009. It’s raining lizards! More than 30 geckos found and still counting. Malay. Nat. 62(3):12–19. ________ , AND K. A. PAN. 2008. Diversity, endemism, and conservation of the amphibians and reptiles of southern Peninsular Malaysia and its offshore islands. Herpetol. Rev. 39:270–281. ________ , K.O. CHAN, N. NUROLHUDA, AND M. SUMONTHA. 2008a. A new species of karst dwelling gecko (genus Cnemaspis Strauch 1887) from the border region of Thailand and Peninsular Malaysia. Zootaxa 1875:51–68. ________ ________ ., , J. L. GRISMER, P. L. JR. WOOD, AND B. DAICUS. 2008b. Three new species of Cyrtodactylus (Squamata: Gekkonidae) from Peninsular Malaysia. Zootaxa 1921:1–23. ________ ., J. L. GRISMER, P. L. JR. WOOD, AND K. O. CHAN. 2008c. The distribution, taxonomy, and redescription of the geckos Cnemaspis afﬁnis (Stoliczka 1887) and C. ﬂavolineata (Nicholls 1949) with descriptions of a new montane species and two new lowland, karst-dwelling species from Peninsular Malaysia. Zootaxa 1931:1–24. ________ , J. SUKUMARAN, J. L. GRISMER, T. M. YOUMANS, P. L. WOOD, JR., AND 300 R. JOHNSON. 2004. Report on the herpetofauna from the Temenggor Forest Reserve, Perak, West Malaysia. Hamadryad 29(1):15–32. ________ , P. L. WOOD, JR., AND T. M. YOUMANS. 2007. Redescription of the gekkonid lizard Cyrtodactylus sworderi (Smith, 1925) from southern Peninsular Malaysia. Hamadryad 31(2):250–257. ________ , T. M. YOUMANS, P. L. WOOD, JR., AND J. L. GRISMER. 2006. Checklist of the herpetofauna of the Seribuat Archipelago, West Malaysia, with comments on biogeography, natural history and adaptive types. Rafﬂes Bull. Zool. 54(1):157–180. INGER, R. F., AND R. K. COLWELL. 1977. Organization of contiguous communities of amphibians and reptiles in Thailand. Ecol. Monogr. 47:229–253. KARNS, D. K., J. C. MURPHY, H. K. VORIS, AND J. S. SUDDETH. 2005. Comparison of semi-aquatic snake communities associated with the Khorat Basin, Thailand. Nat. Hist. J. Chulalongkorn Univ. 52:73–90. LEONG, T. M., AND K. K. P. LIM. 2003. Herpetofaunal records from Fraser’s Hill, Peninsular Malaysia, with larval descriptions of Limnonectes nitidus and Theloderma asperum (Amphibia: Ranidae and Rhacophoridae). Rafﬂes Bull. Zool. 51:123–136. ________ , AND B. L. LIM. 2004. Rana siberu Dring, Mccarthy & Whitten, 1990 — a ﬁrst record for Peninsular Malaysia (Amphibia: Anura: Ranidae). Rafﬂes Bull. Zool. 52:261–263. LEVITON, A. E., S. C. ANDERSON, R. H. GIBBS, E. HEAL, AND C. E. DAWSON. 1985. Standards in herpetology and ichthyology: Part I. Standard symbolic codes for institutional resource collections in herpetology and ichthyology. Copeia 1985:802–832. LIM, B. L. 1958. Colour patterns of some Malayan snakes. Malayan Nat. J. 12:116–118. LIMPERT, E., W. H. STAHEL, AND M. ABBT. 2001. Log-normal distributions across the sciences: keys and clues. Bioscience 51:341–352. LLOYD, M., R. F. INGER, AND F. W. KING. 1968. On the diversity of reptile and amphibian species in a Bornean rain forest. Am. Nat. 102:497–515. LONGINO, J. T., J. CODDINGTON, AND R. K. COLWELL. 2002. The ant fauna of a tropical rainforest: estimating species richness three different ways. Ecology 83:689–702. LOREAU, M. 1992. Species abundance patterns and the structure of groundbeetle communities. Ann. Zool. Fennici 28:49–56. MALKMUS, R., U. MANTHEY, G. VOGEL, P. HOFFMANN, AND J. KOSUCH. 2002. Amphibians and Reptiles of Mount Kinabalu (North Borneo). A.R.G. Ganther Verlag, Rugell. 404 pp. MATSUI, M., AND I. JAAFAR. 2006. A new cascade frog of the subgenus Odorrana from Peninsular Malaysia. Zool. Sci. 23:647–651. MCLEOD, D., AND A. NORHAYATI. 2007. A new species of Theloderma (Anura: Rhacophoridae) from southern Thailand and Peninsular Malaysia. Russ. J. Herpetol. 141:65–72. MURPHY, J. C., H. K. VORIS, AND D. L. KARNS. 1994. A field guide to the snakes of the Danum Valley, a Bornean tropical rainforest ecosystem. Bull. Chicago Herpetol. Soc. 29:133–151. STUEBING, R. B., AND R. F. INGER. 1999. A Field Guide to the Snakes of Borneo. Natural History Publications, Kota Kinabalu, Sabah. 254 pp. ________ , AND F. L. TAN. 2002. Notes on the fire-lipped keelback Rhabdophis murudensis (Smith, 1925) (Ophidia: Colubridae: Natricinae) from northern Borneo. Rafﬂes Bull. Zool. 50:227–230. TWEEDIE, M. W. F. 1983. The Snakes of Malaya. Third ed. Singapore National Printers (Pte) Ltd., Singapore. 167 pp. VAN ROOIJEN, J., AND G. VOGEL. 2008. Contributions to a review of the Dendrelaphis pictus complex (Serpentes: Colubridae) – 1. Description of a sympatric species. Amphibia-Reptilia 29:101–115. VOGEL, G., AND J. VAN ROOIJEN. 2007. A new species of Dendrelaphis (Serpentes: Colubridae) from Southeast Asia. Zootaxa 1394:25–45. WOOD, P. L. JR., J. L. GRISMER, L. L. GRISMER, A. NORHAYATI, K. O. CHAN, A. BAUER. 2009. Two new montane species of Acanthosaura Gray, 1831 (Squamata: Agamidae) from Peninsular Malaysia. Zootaxa 2012:28–46. ________ , L. L. GRISMER, A. NORHAYATI, AND S. JULIANA. 2008. Two new species Herpetological Review 40(3), 2009 of torrent-dwelling toads Ansonia Stoliczka, 1870 (Anura: Bufonidae) from Peninsular Malaysia. Herpetologica 63:321–340. YONG, D. L. 2006. Preliminary list of larger vertebrates of Panti Forest Reserve, South Johor (2002–2006). Singapore Avifauna 20:26–35. YONG, H. S. 1977. Rediscovery of Rhacophorous robinsoni, an Asian frog in Peninsular Malaysia. Malayan Nat. J. 30:59–61. ZIEGLER, T. 2002. Die Amphibien und Reptilien eines TieflandfeuchtwaldSchutzgebietes in Vietnam. Münster (Natur und Tier), 342 S. APPENDIX Specimens examined in museums; most of these are listed by Duellman (2001:704). A review of available specimens of P. hazelae listed by Duellman (2001:1125), however, revealed several misidentiﬁed specimens and other problems. The status of some specimens of P. hazelae listed by Duellman (2001:1125) is given in Table 1. The specimens from Puerto de Gallo, Sierra Madre del Sur, Guerrero (CAS 143106–07 ) are referable to Plectrohyla mykter. The two records from the vicinity of Putla, Oaxaca (Sierra Yacuyacua) are more problematic. Duellman (2001:1125, ﬁg. 386) listed and mapped two specimens from “16 kilometers southwest of Cuquila” in the Asthenodipsas malaccanus: ZMB 5041, Malacca; BMNH 1924.10.23.7, Labong Tendai, Sumatra, Indonesia; BMNH 1967.2276, Gunung Benom, Malaysia; SMF 32580, Perak, Malaysia; ZMH R06836, Lebong-Tandai, Benkoelen, Sumatra, Indonesia; ZFMK 45131, Satun, Thailand; ZMB 50673, Thung Song, Thailand; ZRC 2.2746, Asahan Estate, Melaka. Calliophis gracilis: HC 193, Cameron Highlands, Pahang, Peninsular Malaysia. Enhydris pahangensis: HC 197,198, Nenasi Forest Reserve, Pahang, Peninsular Malaysia. Hylarana siberu: DWNP 1189, Kuala Gandah, Pahang, Peninsular Malaysia. Oligodon signatus: HC 196, Kampung Chennah, Negeri Sembilan, Peninsular Malaysia. Rhacophorus robinsoni: HC 191, 192, Kampung Chennah, Negeri Sembilan, Peninsular Malaysia. Theloderma leprosa: HC 281, Gunung Besar Hantu, Negeri Sembilan, Peninsular Malaysia. Xenelaphis ellipsifer: HC 194, Kampung Chennah, Negeri Sembilan, Peninsular Malaysia. Herpetological Review, 2009, 40(3), 301–302. © 2009 by Society for the Study of Amphibians and Reptiles On the Distribution and Status of Plectrohyla hazelae (Taylor, 1940) (Amphibia: Hylidae) from Oaxaca, Mexico JOSEPH R. MENDELSON III* and EDWARD H. KABAY Department of Herpetology, Zoo Atlanta 800 Cherokee Ave SE, Atlanta, Georgia 30315-1440, USA *Corresponding author; e-mail: jmendelson@zooatlanta.org Duellman (2001) reviewed and updated the distributional status of all hylid species in Mesoamerica that were known at the time. We examined the updated distributional map for P. hazelae (Duellman 2001:ﬁg. 386, and presented here as Fig. 1a). This same geographic information is also reﬂected on the map accompanying the account of P. hazelae appearing on the online IUCN Red List (http://www. iucnredlist.org/), which is the same information compiled by the Global Amphibian Assessment (Stuart et al. 2004). Based on these sources, the prevailing concept of the distribution of this species (Fig. 1a) is that it occurs disjunctly in three montane areas in Guerrero and Oaxaca, Mexico, namely the Sierra Madre del Sur, Sierra Yucuyacua, and Sierra Aloapaneca. The type locality is on Cerro San Felipe, in the Sierra Aloapaneca, and a small number of specimens from various localities on those slopes are deposited FIG. 1a (top). The distribution of Plectrohyla hazelae, as presented by Duellman (2001:ﬁg. 386), showing putative localities for the species in the Sierra Madre del Sur, Sierra Yucuyacua, and the Sierra Aloapaneca or Guerrero and Oaxaca, Mexico. This ﬁgure does not indicate the record (KU 13707) from near the town of Tamazulapan, Oaxaca, in the Sierra Mixes; this record was listed by Duellman (2001) but does not appear on this map (see text). This ﬁgure has been modiﬁed from the original (Duellman, 2001:ﬁg. 386) to exclude locality records of species not relevant to the current paper. FIG. 1b (bottom). Our current concept of the distribution of Plectrohyla hazelae, indicating that it is once known to have once existed only on the slopes of Cerro San Felipe, in the Sierra Aloapaneca of Oaxaca, Mexico. The species was last seen alive in 1975 (see text). The question mark indicates the locality for the specimen (KU 137037) from near Tamazulapan, in the Sierra Mixes of Oaxaca, Mexico, that is here referred to Plectrohyla sp. indet. Herpetological Review 40(3), 2009 301 TABLE 1. Identifications and notes on specimens referred to Plectrohyla hazelae by Duellman (2001:1125). Specimens Locality New Identiﬁcation Comments CAS 143106–07 Guerrero: Puerto de Gallo Plectrohyla mykter Juveniles MSU (uncatalogued) Oaxaca: 16 km SE Cuquila n/a Specimens lost (see text) USNM 224499 Oaxaca: 4.7 km W La Cumbre Plectrohyla hazelae Last known specimen of P. hazelae; collected in 1975 KU 13707 Oaxaca: between Tlahuitoltepec and Tamazulapan Plectrohyla sp. indet. Metamorphic individual; identity uncertain. Unclear if this record was incorrectly mapped by Duellman (2001: ﬁg.386), or whether it was not mapped (see text). collections of Michigan State University; however, these specimens cannot be located under any taxonomic name (L. Abraczinskas, in litt.). Subsequent research indicated that two hylid frogs were collected at that locality by Robert G. Webb on 21 July 1968 (Field Numbers: RGW 4784, 4785; original locality written as “10 mi southwest Cuquila”); squamate specimens collected there at the same time are present in the MSU collection. Webb (in litt.) veriﬁed the collection of the specimens from his ﬁeld notes and recalls physically showing them to W. E. Duellman in some informal setting, perhaps at a meeting. Duellman, however, (in litt.) has no written record, nor recollection, of the specimens. All other material collected by Webb during that era was deposited at MSU, but our search (www.herpnet.org) and queries to other collection managers, including KU, found no amphibians bearing locality information referencing the tiny village of Cuquila. Evidently, the specimens were never catalogued in a museum and are missing. Based on the lack of voucher specimens, and our own opinion that this is an unlikely locality for a species otherwise known only from the slopes of Cerro San Felipe, we believe this locality record to be in error. An additional specimen (KU 137037) reported by Duellman (2005:1125) from Oaxaca “btw Tlahuitoltepec & Tamazulapan” is also problematic. The specimen is a barely post-metamorphic individual, and thus difﬁcult to identify. However, the presence of nearly complete webbing on the feet (versus about half-webbed in P. hazelae) indicates that it is not referable to P. hazelae; we here refer it to Plectrohyla sp. indet., pending review of other specimens from the region. An additional inconsistency exists in the map presented by Duellman (2001:ﬁg. 386) in that it does not show a locality in the Sierra Mixes, where KU 137037 was collected. It is possible that this locality was confused with another village named Tamazulapan existing in the Sierra Yacuyuna, near the towns of Tlaxiaco and Putla; however, the map shows only one symbol for that region, so it is not clear if that symbol refers to KU 137037 and/or the missing MSU specimens, or was intended to indicate both localities (because, as it turns out, the village of Tamazulapan in this region is near Cuquila). In either case, the symbol should be removed from the distributional map of P. hazelae because there is no evidence that the species occurs there. In Fig. 1b, we illustrate our concept of the distribution of this species, which can now be summarized simply as the immediate vicinity of Cerro San Felipe, in the Sierra Aloapaneca of Oaxaca, Mexico. We also note that specimens (TNHC 28598–99; Oaxaca: 13 km NW Ixtlán 302 de Juárez) referred to P. hazelae by Duellman (2001:704) do not represent that species and are perhaps referable to Ecnomiohyla miotympanum. Finally, we sadly report that recent intensive amphibian surveys on Cerro San Felipe have uncovered no specimens of Plectrohyla (G. Parra-Olea, T. Papenfuss, and D. Wake, in litt.). Whereas adults of Plectrohyla spp. sometimes can be difﬁcult to locate, the stream dwelling tadpoles are obvious, and present year-round (JRM, personal observation; J. A. Campbell, personal communication; Lips et al., 2004). We estimate this species became extinct sometime probably not very long after 1975—just as proposed by Lips et al. (2004)—when the last known specimen (USNM 224499) was collected on Cerro San Felipe by our colleague Roy W. McDiarmid. Acknowledgments.— Carl Franklin, Jonathan Campbell, Darrel Frost, Linda Abraczinskas, Robert Webb, William Duellman, Karen Lips, Amanda McDaniel, Jason Brock, David Wake, Ted Papenfuss, and Gabriela Parra-Olea all kindly provided considerable assistance and information for this project. We are grateful to the curators of FMNH, USNM, KU, TNHC, and AMNH for permission to review specimens from those collections. LITERATURE CITED DUELLMAN, W. E. 2001. Hylid Frogs of Middle America. Society for the Study of Amphibians and Reptiles, Ithaca, New York, 1170 pp, 92 plates. LIPS, K. R., J. R. MENDELSON III, A. MUÑOZ-ALONSO, L. CANSECO-MÁRQUEZ, AND D. G. MULCAHY. 2004. Amphibian population declines in montane southern Mexico: resurveys of historical localities. Biol. Cons. 119:555–564. 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 306:1783–1786. Specimens examined.—Ecnomiohyla cf. miotympanum: MEXICO: OAXACA: Ixtlán de Juárez, 8 mi [by road] NW, 7600 ft (TNHC 28598, 28599). Plectrohyla mykter: MEXICO: GUERRERO: Puerto de Gallo, 8200 ft (CAS 143106, 143107). Plectrohyla hazelae: MEXICO: OAXACA: 2 km S El Punto, 2390 m (KU 100969); Cerro San Felipe, about 10 miles north of Oaxaca, Oaxaca (FMNH 126323, 108509–13, 126419); Cerro San Felipe (USNM 114576, 114577); La Cumbre, 4.6 mi W of, on road to Cerro San Felipe (USNM 224499). Plectrohyla sp. indet.: MEXICO: OAXACA: btw Tlahuitoltepec and Tamazulapan, 2200 m (KU 13707). Herpetological Review 40(3), 2009 TECHNIQUES Herpetological Review, 2009, 40(3), 303–304. © 2009 by Society for the Study of Amphibians and Reptiles A Method for Constructing an Adjustable Platform to Obtain Lateral Photographs of Larval Anurans MATTHEW C. SCHACHT and LANCE D. MCBRAYER* Department of Biology, 202 Georgia Avenue Georgia Southern University, Statesboro, Georgia 30460, USA * Corresponding author; e-mail: lmcbrayer@georgiasouthern.edu Digitally photographing specimens and using computer software to obtain morphological measurements is a commonly used technique in larval anuran research. Lateral photographs of specimens can be used to obtain measures such as snout-vent length, body height, tail length, tail height, tail musculature length, and tail musculature height (Dayton et al. 2005; Lardner 2000; Relyea 2001). However, obtaining quality lateral photographs of larval anurans can be difﬁcult due to their general body plan (a rounded body and laterally compressed tail). When a specimen is laid on its side in a lateral view, the tail remains ﬂat and in the horizontal plane, but the curvature of the body often causes the specimen to bend so that the body and tail are no longer aligned in the horizontal plane (Figs. 1a and 1b). If the specimen is bent when photographed, measurements of total length or snout-vent length are likely to be underestimates (Fig. 1b). Thus, it is necessary to compensate for the lateral curvature of the body in order to maximize the accuracy of measurements obtained from lateral photographs. McDiarmid and Altig (1999) describe a modiﬁed aquarium apparatus for photographing larval anurans. While this apparatus may be ideal for photographing specimens for voucher or identiﬁcation purposes, its application in morphological studies may be suboptimal due to its large size, need for clear water to submerse specimens in, and requirement of an electronic ﬂash that is independent of the camera. Because morphological studies often require large sample sizes, decreasing the amount of time it takes to prepare a specimen for photographing is important for efﬁciency purposes. We describe a simple method for constructing an adjustable platform that enables researchers to consistently and rapidly obtain quality lateral photographs of formalin ﬁxed, or anesthetized, tadpoles. This platform supports specimens in a manner that prevents bending caused by the lateral curvature of the body (Figs. 1c, 1d), and can accommodate variously sized species or broad ontogenetic series of specimens. Platform construction can be completed in less than 30 minutes with the following materials: two standard microscope slides (3 in. x 1 in.); two hollow tubes, one with a slightly larger diameter than the other; and epoxy glue. The microscope slides act as a platform on which the tail and tip of snout of rest. The hollow tubes are glued beneath the microscope slides and allow the platform to be adjusted. For tubes, we used a drinking straw and the shaft of a FIG. 1. (a) Side view illustration showing the bending of a larval anuran specimen caused by the lateral curvature of the body. The dashed line is provided to enhance visualization of the degree of bending. (b) Specimen a, viewed from above. The dashed line shows where the outline of the head would be if the specimen’s body and tail were aligned in a ﬂat plane. (c) Side view of a specimen supported in a manner that prevents the lateral curvature of the body. (d) Specimen c, when viewed from above. disposable ink pen. The smaller diameter tube (i.e., the straw) is cut into two 3-inch pieces and glued parallel to the short edge of the slide approximately one-half inch from each end. The pieces of straw should protrude at least one-half inch from the long edge of the microscope slide. The larger tubes (pen shafts) are then glued to the second microscope slide in the same manner as previously described, with the exception that the circular openings of the shafts should not protrude from underneath the microscope slide (Fig. 2). The microscope slides are aligned and the two pieces of drinking straw (or tube with smaller diameter) are inserted into the pen shafts (tube with larger diameter). The larger diameter of the disposable ink pen allows the stage to be quickly and easily manipulated to accommodate specimens of different sizes. A scale bar can be glued to the surface of either microscope slide so that it will be in the ﬁeld of view with the specimen and in the same plane as the specimen when it is photographed from directly above (Fig. 2). Specimens are positioned on the platform so that only the extreme anterior of the body (snout) is touching the slide on the left, and the tail is supported by the slide FIG. 2. Larval anuran specimen properly positioned on the completed platform. Selected dimensions of the platform are provided. The ﬁnished platform rests on a ﬂat surface and photographs may be obtained by positioning a camera directly above. Herpetological Review 40(3), 2009 303 on the right (Figs. 1c, 1d, and 2). From the author’s experience, the apparatus is easily manipulated and facilitates rapid handling and photographing of specimens. We have found this apparatus to be both durable and efﬁcient, as it has been used to obtain over 500 photographs of specimens, with an approximate handling time of under 20 seconds for each specimen. Further, this platform is especially useful when photographing smaller species of larval anurans (e.g., Bufo terrestris, < 40 mm total length), or very small specimens at early developmental stages. Simple modiﬁcations such as adding an additional microscope slide to one side of the platform may be necessary in order to photograph very large specimens (e.g., Rana catesbeiana) or late developmental stages of large species. Acknowledgments.—We thank Steve Williams, Tim Gowan, Dana Nayduch, and three reviewers for their valuable critiques of earlier versions of this manuscript. The research associated with this manuscript was approved by the Georgia Southern University IACUC, and funded in part by grants from Georgia Southern University. LITERATURE CITED DAYTON, G. H., D. SAENZ, K. A. BAUM, R. B. LANGERHANS, AND T. J. DEWITT. 2005. Body shape, burst speed and escape behavior of larval anurans. Oikos 111:582–591. LARDNER, B. 2000. Morphological and life history responses to predators in larvae of seven anurans. Oikos 88:169–180. MCDIARMID, R. W., AND R. ALTIG. 1999. Research: materials and techniques. In R. W. McDiarmid and R. Altig (eds.), Tadpoles: The Biology of Anuran Larvae, pp. 7–23. University of Chicago Press, Chicago, Illinois. RELYEA, R. A. 2001. Morphological and behavioral plasticity of larval anurans in response to different predators. Ecology 82:523–540. Herpetological Review, 2009, 40(3), 304–306. © 2009 by Society for the Study of Amphibians and Reptiles Baiting Differentially Inﬂuences Capture Rates of Large Aquatic Salamanders, Siren and Amphiuma CLINTON P. SMITH1 DENISE R. GREGOIRE and MARGARET S. GUNZBURGER2 United States Geological Survey, Florida Integrated Science Center 7920 NW 71st Street, Gainesville, Florida 32653-3701, USA 1 Corresponding author, present address: Center for Forest Research, Mississippi State University, Mississippi State, Mississippi 39762, USA e-mail: sscpsmith@hotmail.com 2 Present address: Nokuse Plantation, 13292 County Highway 3280 Bruce, Florida 32455, USA Understanding the potential biases associated with a sampling method are necessary before undertaking a study of the ecology of any organism (Dodd 2003; Gunzburger 2007). Trapping is an effective way to determine whether a species is present in a certain habitat, and many forms of passive traps for aquatic salamanders 304 have been developed, including minnow traps and other funnel traps, leaf litterbags, and box traps (Dodd 2003; Heyer et al. 1994). Baiting passive traps to attract organisms may increase the number of animals captured, but it may also introduce sampling bias to a study (Sorensen 2003). Baiting might attract more animals to an area than normally would be found there, which could lead to an overestimate of abundance. Another bias could be a potential change in behavior or survival of the target species due to predators that are also attracted to the bait. There also could be differences among target species in their food preferences and response to different types of bait. The Greater Siren (Siren lacertina) and Two-toed Amphiuma (Amphiuma means) are large aquatic salamanders that are relatively poorly studied, perhaps because of their cryptic behavior and the perceived difﬁculty in sampling their habitats. These species co-occur throughout most of their geographic ranges in the Southeastern Coastal Plain of the United States (Petranka 1998), though inter-speciﬁc interactions may affect microhabitat occupancy (Snodgrass et al. 1999). Both salamanders are primarily nocturnal, with daylight activities restricted to either dense aquatic vegetation or hiding in burrows in the substrate (Duellman and Schwartz 1958; Freeman 1958; Knepton 1954). Limited data on their movement patterns suggest these salamanders have relatively small home ranges (Sorensen 2004) with occasional short dispersal across land during heavy rain or ﬂooding (Aresco 2002; Carr 1940; Gibbons and Semlitsch 1991). The diet of these salamanders may have signiﬁcant inﬂuence on the effectiveness of baiting to attract these species to traps. Although there is considerable overlap in the diet of these species, diet studies suggest Amphiumas are more carnivorous than Sirens. This is supported by stable isotope analysis that placed Amphiumas at a higher trophic position than Sirens in a North Florida lake (Aresco and James 2005). Dietary analyses of Amphiumas, based on foraging habits or stomach contents, have shown they are entirely carnivorous, feeding on insects (both aquatic and terrestrial), many types of amphibians (including conspeciﬁcs), reptiles, ﬁsh, crayﬁsh, mollusks, and spiders (Chaney 1951; Hamilton 1950; Hargitt 1892; Lee 1969; Machovina 1994). In contrast, plant material has often been found in dietary analyses of Sirens (Davis and Knapp 1953; Dunn 1924), although other studies have shown Sirens ingest invertebrates, especially mollusks, and ﬁsh along with vegetation (Burch and Wood 1955; Hanlin 1978; Moler 1994). Siren lacertina are able to effectively digest plant material, as the gastrointestinal structure of S. lacertina is similar to other vertebrates that use fermentative digestion (Pryor et al. 2006). Differential movement patterns also may affect the capture of these salamanders, with more mobile species having a greater likelihood of encountering a passive trap than sedentary species. Unfortunately, there have been few studies examining movements of these species within habitats. A study conducted in Missouri of Siren intermedia noted most movements were > 10 m (75% of the population), with a mean home range of 94.8 m2 (Frese et al. 2003). Another study found > 77% of the movements of S. intermedia in Texas were < 6 m (Gehlbach and Kennedy 1978). The objective of this study was to evaluate the effect of baiting traps on capturing Siren spp. and Amphiuma means. A previous study (Sorensen 2004) found that baiting did not inﬂuence the capture rates of Siren spp. and A. means in crayﬁsh traps, but we Herpetological Review 40(3), 2009 believe that study may have been biased by the close proximity (one meter apart) of baited and unbaited traps. Chemical cues from the bait (canned sardines in oil) may have pervaded the entire trapping area (Sorensen 2004). The difference in trap spacing used for this study should eliminate the biases that may have been involved in the previous study. Materials and Methods.—This study was conducted during two trapping intervals in June–July 2006 at Lake Suggs, a 34 ha darkwater lake at the Ordway Swisher Biological Station in Putnam County, Florida (USA). Our study used a blocked design, with four blocks of 10 traps for a total of 40 traps. Traps were deployed along a single transect with one trap every ﬁve meters (195 m long transect). The transect was set ca. 3–4 m from the shoreline in dense submergent aquatic vegetation (Limnobium spongia, Pontedaria cordata, and Hydrocotyle sp.). Two blocks (one half of the transect line) were baited in the ﬁrst trapping interval (5–9 June 2006) and the other two blocks were baited in the second trapping interval (3–7 July 2006). The remaining two blocks in each trapping interval did not have bait added to them. Baiting an entire block of traps and separating each block by 5 m allowed us to prevent contamination of unbaited traps by chemical cues from nearby baited traps. Because it is unlikely salamanders are distributed equally along the length of the transect, our design allowed us to evaluate separately the effect of baiting and block location on the number of animals captured. We used modiﬁed crayﬁsh traps (Lee Fisher International, Tampa, Florida) lined with 0.5 mm mesh which were attached to a PVC pipe anchored into the substrate (Johnson and Barichivich 2004). Traps were baited with canned sardines in olive oil; holes were punched in the can and the can was suspended inside the trap with a length of cord. Bait was not replaced daily, but rather one can of bait was left in each baited trap for the entire trap interval. Traps were set for four nights per interval (320 total trap-nights) and checked every morning. We analyzed capture rates of each salamander species separately because we do not know the actual population size of either species nor the proportion that were captured in the traps, so we cannot evaluate the relative efﬁcacy of the traps across species. Capture data could not be made normal with transformation, so we conducted Kruskal-Wallis one-way ANOVAs for each species with number captured as the dependent variable and presence of bait, sampling interval (June or July), and block as factors. Results.—A total of 147 individuals (new captures and recaptures) were caught during the two sampling intervals, including 87 Amphiuma means, 56 Siren lacertina, 3 S. intermedia, and 1 small Siren sp. that escaped prior to identiﬁcation. The number of new captures of unmarked individuals was 106, comprised of 52 A. means, 50 S. lacertina, 3 S. intermedia, and 1 unidentiﬁed Siren sp. Numbers of all Siren species were pooled for data analysis. The mean catch per unit effort (CPUE; captures/trap-night) for the two-month duration of this study was 0.27 A. means/trap-night and 0.18 Siren spp./trap-night. This CPUE was higher than the previous study (0.11 A. means /trap-night and 0.10 Siren spp./trap-night; Sorensen 2004). Baited traps captured 72% (N = 106) of the animals whereas unbaited traps captured 28% (N = 41). Several individuals from the two-month sampling period were recaptured more than once, and multiple salamanders were captured in the same trap on several occasions. Multiple salamanders were captured in the same trap on 23 occasions during the two-month sampling period: six traps with multiple A. means, three traps with multiple Siren spp., and 14 traps with a combination of A. means and Siren spp. Sorenson (2003) reported multiple captures in the same trap on seven occasions over a 12-month period. The results of the Kruskal-Wallis analyses showed that the effects of block and sampling interval were not signiﬁcant for either species. Signiﬁcantly more Amphiuma were caught in baited traps relative to unbaited traps (Mann-Whitney U = 1492, p < 0.001); there was no correlation of capture rates of Siren with baiting (Fig. 1). Discussion.—Baiting of traps did not have the same effect on capture rates of these two species of salamanders. The dietary differences of these species accurately predict their response to bait: The carnivorous Amphiuma means is strongly attracted to bait, while the omnivorous Siren spp. were apparently not attracted to bait. Although not signiﬁcant, both this study and Sorensen (2003) demonstrated a trend towards higher capture rates of S. lacertina in unbaited traps (Fig. 1). The high capture rate of A. means could be due to the more predaceous behavior of A. means or other factors such as the ability of A. means to detect and locate bait. Amphiuma means can be caught on a trotline baited with dead crayﬁsh (Chaney 1951), suggesting that chemoreception may play a role in locating prey by these salamanders. In contrast, a laboratory experiment suggested that chemical cues do not seem to play a role in prey location by S. intermedia (Sullivan et al. 2000). Although we did not conduct comparisons across species, Willson et al. (2005) noted higher numbers of Siren spp. captures (N = 228) in commercial funnel traps compared with A. means (N = 14). Baiting was not part of their experiment, but the lack of bait within traps could explain the higher capture rates of Siren spp. over A. means. If baiting does attract A. means, then Siren spp. may be avoiding these traps because S. lacertina has been documented as a prey item in A. means stomach analyses (Machovina 1994). Our study demonstrates that the decision of whether or not to use baited traps is likely to inﬂuence the outcome of trapping efforts for FIG. 1. Mean and standard error of catch per unit effort (number of captures/trap/night) of Amphiuma means and Siren sp. in 20 baited and 20 unbaited crayﬁsh traps in two four-night sampling intervals. Herpetological Review 40(3), 2009 305 these species of aquatic salamanders. We suggest that researchers conducting rapid assessment surveys for large aquatic salamanders, in which the response variable is detection or non-detection of a species, should use both baited and unbaited traps to increase the likelihood of trapping both Siren spp. and Amphiuma spp. if they are present. In contrast, if researchers are more interested in conducting long-term demographic studies, using unbaited traps may result in a more accurate and unbiased reﬂection of the relative abundance of the two species. We suggest researchers may minimize the likelihood of making critical errors in interpreting population data by conducting even a short-term series of trials designed to understand sampling biases prior to initiating longerterm studies. Acknowledgments.—We thank S. Coates and M. Sunquist for access to the Ordway Swisher Biological Station under permit OR-05-07. We thank K. Sorensen for advice and access to the previous data collected at this site. C. K. Dodd, Jr., W. Barichivich, and J. Staiger provided comments on earlier drafts of this manuscript. This research was supported by the USGS Amphibian Research and Monitoring Initiative and a USGS Bureau Venture Capital Fund Grant to MSG. Research was conducted under USGS Florida Integrated Science Center IACUC Protocol 2006-04. The use of trade, product, or ﬁrm names does not imply endorsement by the U.S. Government. LITERATURE CITED ARESCO, M. J. 2002. Amphiuma means (Two-toed amphiuma). Overland migration. Herpetol. Rev. 33:296–297. ________ , and F. C. James. 2005. Ecological relationships of turtles in northern Florida lakes: a study of omnivory and the structure of a lake food web. Final Report. Florida Fish and Wildlife Conservation Commission, Tallahassee, Florida. BURCH, P. R., AND J. T. WOOD. 1955. The salamander Siren lacertina feeding on clams and snails. Copeia 1955:255–256. CARR, A. F. JR. 1940. A contribution to the herpetology of Florida. Univ. Florida Publ. Biol. Sci. 3:1–118. CHANEY, A. H. 1951. The food habits of the salamander Amphiuma tridactylum. Copeia 1951:45–49. DAVIS, W. B., AND F. T. KNAPP. 1953. Notes on the salamander Siren intermedia. Copeia 1953:119–121. DODD, C. K., JR. 2003. Monitoring Amphibians in Great Smoky Mountains National Park. U.S. Geological Survey Circular No. 1258. 117 pp. DUELLMAN, W. E., AND A. SCHWARTZ. 1958. Amphibians and reptiles of southern Florida. Bull. Florida State Mus. 3:182–325. DUNN, E. R. 1924. Siren, a herbivorous salamander? Science 59:145. FREEMAN, J. R. 1958. Burrowing in the salamanders Pseudobranchus striatus and Siren lacertina. Herpetologica 14:130. FRESE, P. W., A. MATHIS, AND R. WILKINSON. 2003. Population characteristics, growth, and spatial activity of Siren intermedia in an intensively managed wetland. Southwest. Nat. 48:534–542. GEHLBACH, F. R., AND S. E. KENNEDY. 1978. Population ecology of a highly productive aquatic salamander (Siren intermedia). Southwest. Nat. 23:423–430. GIBBONS, J. W., AND R. D. SEMLITSCH. 1991. Guide to the Reptiles and Amphibians of the Savannah River Site. University of Georgia Press, Athens, Georgia. 192 pp. GUNZBURGER, M. S. 2007. Evaluation of seven aquatic sampling methods for amphibians and other aquatic fauna. Appl. Herpetol. 4:47-63. HAMILTON, W. J. 1950. Notes on the food of the congo eel, Amphiuma. Nat. Hist. Misc. 62:1–3. HANLIN, H. G. 1978. Food habits of the greater siren Siren lacertina, in an Alabama coastal plain pond. Copeia 1978:358–360. 306 HARGITT, C. W. 1892. On some habits of Amphiuma means. Science 20:159. HEYER, W. R., M. A. DONNELLY, R. W. MCDIARMID, L. C. HAYEK, AND M. S. FOSTER. 1994. Measuring and Monitoring Biological Diversity: Standard Methods for Amphibians. Smithsonian Institution Press, Washington, D.C. 388 pp. JOHNSON, S. A., AND W. J. BARICHIVICH. 2004. A simple technique for trapping Siren lacertina, Amphiuma means, and other aquatic vertebrates. J. Freshw. Ecol. 19:263–269. KNEPTON, J. C., JR. 1954. A note on the burrowing habits of the salamander Amphiuma means means. Copeia 1954:68. LEE, D. S. 1969. Observations on the feeding habits of the congo eel. Florida Nat. 42:95. MACHOVINA, B. L. 1994. Ecology and life history of the salamander Amphiuma means in Everglades National Park. Unpubl. Masters thesis, Florida International University, Miami, Florida. MOLER, P. E. 1994. Siren lacertina (Greater Siren). Diet. Herpetol. Rev. 25:62. PETRANKA, J. W. 1998. Salamanders of the United States and Canada. Smithsonian Institution Press, Washington, D.C. 587 pp. PRYOR, G. S., D. P. GERMAN, AND K. A. BJORNDAL. 2006. Gastrointestinal fermentation in greater sirens (Siren lacertina). J. Herpetol. 40:112–117. SNODGRASS, J. W., J. W. ACKERMAN, A. L. BRYAN, Jr., AND J. BURGER. 1999. Inﬂuence of hydroperiod, isolation, and heterospeciﬁcs on the distribution of aquatic salamanders (Siren and Amphiuma) among depression wetlands. Copeia 1:107–113. SORENSEN, K. 2003. Trapping success and population analysis of Siren lacertina and Amphiuma means. Unpubl. Masters thesis, University of Florida, Gainesville, Florida. SORENSEN, K. 2004. Population characteristics of Siren lacertina and Amphiuma means in North Florida. Southeast. Nat. 3:249–258. SULLIVAN, A. M., P. W. FRESE, AND A. MATHIS. 2000. Does the aquatic salamander, Siren intermedia, respond to chemical cues from prey? J. Herpetol. 34:607–611. WILLSON, J. D., C. T. WINNE, AND L. A. FEDEWA. 2005. Unveiling escape and capture rates of aquatic snakes and salamanders (Siren spp. and Amphiuma means) in commercial funnel traps. J. Freshw. Ecol. 20:397–403. Bolitoglossa mexicana (Mexican Mushroom-tongues Salamander), Guatemala. Illustration by Peter Stafford. Herpetological Review 40(3), 2009 AMPHIBIAN DISEASES Herpetological Review, 2009, 40(3), 307–308. © 2009 by Society for the Study of Amphibians and Reptiles Reassessment of the Historical Timeline for Batrachochytrium dendrobatidis Presence in Honduras and Conservation Implications for Plectrohyla dasypus JONATHAN E. KOLBY* The Conservation Agency, Jamestown, Rhode Island 02835, USA Operation Wallacea, Hope House, Old Bolingbroke Lincolnshire PE23 4EX, UK and GRETCHEN E. PADGETT-FLOHR Department of Zoology, Southern Illinois University Carbondale, Illinois 62918, USA e-mail: gpadgettﬂohr@aol.com *Present address: 1671 Edmund Terrace, Union, New Jersey 07083, USA e-mail: j_kolby@hotmail.com The amphibian chytrid fungus Batrachochytrium dendrobatidis (Bd) has been identiﬁed as an exceptionally devastating amphibian pathogen, capable of acting as both the proximate and ultimate causes of amphibian extinction (Wake and Vredenberg 2008). As described by Skerrat et al. (2007), Bd is a highly transmissible pathogen, which has recently been recognized as an invasive species and listed as notiﬁable by the World Organization for Animal Health (OIE: http://www.oie.int/eng/normes/fcode/en_chapitre_ 2.4.1.htm#rubrique_batrachochytrium_dendrobatidis). In recent FIG. 1. Ventral anterior surface of the larval Plectrohyla dasypus collected in 1996 (USNM 523472) which was examined histologically for Batrachochytrium dendrobatidis infection. Specimen displayed gaps in tooth rows and erosion of jaw sheaths. years, signiﬁcant advances in ﬁeld and laboratory techniques have enabled scientists to detect Bd within amphibian species using non-lethal methods (Annis et al. 2004, Boyle et al. 2004, Hyatt et al. 2007) which has increased our ability to test amphibians that would otherwise not be testable due to regulatory restrictions on take of endangered species. Detection of Bd in extant populations is an important component in assessing the status of a population or species because it reveals an elevated risk for subsequent population declines. Therefore, identifying the distribution of Bd-infected amphibians can help guide the investment of resources towards areas in most need of frequent monitoring. However, contemporary Bd occurrence does not provide data as to when the pathogen was introduced into a geographic area. Contemporary surveys are often unable to show the length of time between the arrival of Bd and the onset of population declines. Developing a timeline of Bd introduction in a speciﬁc area requires multiple surveys that encompass a timeframe ranging from pre-Bd introduction through post-Bd arrival. Bd was identiﬁed only 11 years ago, and by the time it was described and testing techniques for infection were made available, Bd had already become widespread (Stuart et al. 2004); thus for many localities, determining the timeline for Bd introduction requires retrospective analyses of archived specimens. As we demonstrate in this article, such retrospective studies need not be expansive to produce valuable data regarding the earliest known occurrence of Bd in a species or geographic location. In 2007, we conﬁrmed the presence of Bd within Cusuco National Park, Honduras (Kolby et al. 2009). A species of particular concern was Plectrohyla dasypus, an amphibian endemic to Cusuco National Park which displayed an alarmingly high prevalence (78%) of Bd. This species is listed as Critically Endangered by the IUCN Red List of Threatened Species, in part “… because of a drastic population decline, estimated to be more than 80% over the last ten years, inferred from the apparent disappearance of most of the population (probably due to chytridiomycosis)” (Cruz et al. 2004). Although current and historic observations suggest the involvement of Bd in these declines, a cause and effect relationship has not been deﬁnitively demonstrated. This is especially relevant for P. dasypus, now a candidate for ex situ management, as the future of the species is dependent upon an accurate identiﬁcation of the causative factors driving the impending extinction. Therefore, the answer to the following question carries profound importance: Has Bd been present in Cusuco National Park, Honduras for at least 10 years? Field surveys conducted in 2007 revealed a high incidence of oral defects in Bd-infected P. dasypus larvae (Kolby et al. 2009), leading us to suspect that oral defects in this species may often be indicative of Bd infection. The earliest published record of larval oral defects in this species originated from a series of ﬁve larvae (USNM 523472) collected from Cusuco National Park in 1996 (McCranie and Wilson 2002). Morphological examination of this larval series was conducted at the Smithsonian Museum of Natural History on 27 February 2008. Using a 20x hand lens, oral defects were observed in two of the ﬁve specimens, which included gaps in tooth rows and erosion of jaw sheaths. We selected one of these two specimens for histological examination to diagnose Bd (Fig. 1). We excised the complete oral disc intact including the underlying soft tissue and stored the samples in 70% ethanol. Tissues Herpetological Review 40(3), 2009 307 habitat alteration, and xenobiotics) could similarly precipitate an amphibian population decline, the testing of potential cause-effect relationships related to these variables is crucial in determining whether an ex situ conservation management plan will be necessary and feasible to prevent extinction of P. dasypus. Acknowledgments.—We thank R. McDiarmid and the National Museum of Natural History, Smithsonian Institution, for permission to examine the referenced specimens. LITERATURE CITED FIG. 2. Top: Batrachochytrium dendrobatidis infection in the toothrow of larval Plectrohyla dasypus viewed via light microscopy at 200×. Bottom: B. dendrobatidis infection viewed at 400×. Dark staining in some cells indicates bacterial colonization of empty zoosporangia (arrow). were parafﬁn-embedded, serial-sectioned in 4-μm increments and haematoxylin-eosin (H&E) stained for examination under light microscopy. The specimen was found to be infected with Bd when serial sections were viewed by light microscopy. Bd thalli and empty zoosporangia were clearly distinguishable in the toothrows and were characterized by thick cell walls and slight ovoid shape (Fig. 2). Our examination conﬁrmed the presence of Bd in Cusuco National Park, Honduras in 1996, seven years earlier than is currently documented for the country (Puschendorf et al. 2006). Since the population decline of P. dasypus is believed to have commenced in approximately 1994 (Cruz et al. 2004), our ﬁndings chronologically align the presence of Bd with the onset of enigmatic decline. Analysis of samples collected in 2007 (Kolby et al. 2009) revealed a high prevalence of Bd and conﬁrmed infection in a recently metamorphosed P. dasypus which displayed behavioral symptoms of amphibian chytridiomycosis (lethargy, loss of righting reﬂex, etc.) immediately preceding death. Integrating our contemporary ﬁeld survey data with retrospective histological analysis, it now seems reasonable to consider Bd as a potential proximate and ultimate factor in the decline of P. dasypus over the past decade. Since a multitude of other factors (e.g., global climate change, 308 ANNIS, S. L., F. P. DASTOOR, H. ZIEL, P. DASZAK, AND J. E. LONGCORE. 2004. A DNA-based assay identiﬁes Batrachochytrium dendrobatidis in amphibians. J. Wildl. Dis. 40(3):420–428. BOYLE, D. G., D. B. BOYLE, V. OLSEN, J. A. T. MORGAN, AND A. D. HYATT. 2004. Rapid quantitative detection of chytridiomycosis (Batrachochytrium dendrobatidis) in amphibian samples using real-time Taqman PCR assay. Dis. Aquat. Org. 60:133–139. CRUZ, G., L. D. WILSON, AND F. CASTEÑEDA. 2004. Plectrohyla dasypus. In IUCN 2007. 2007 IUCN Red List of Threatened Species. <www. iucnredlist.org>. Accessed 11 March 2007. HYATT, A. D., D. G. BOYLE, V. OLSEN, D. B. BOYLE, L. BERGER, D. OBENDORF, A. DALTON, K. KRIGER, M. HERO, H. HINES, R. PHILLOTT, R. CAMPBELL, G. MARANTELLI, F. GLEASON, AND A. COLLING. 2007. Diagnostic assays and sampling protocols for the detection of Batrachochytrium dendrobatidis. Dis. Aquat. Org. 73:175–192 KOLBY, J. E., G. E. PADGETT-FLOHR, AND R. FIELD. 2009. Amphibian chytrid fungus (Batrachochytrium dendrobatidis) in Cusuco National Park, Honduras. Dis. Aquat. Org., Spec. Ed. 4. Published online 6 May 2009. doi: 10.3354/dao02055 MCCRANIE, J. R., AND L. D. WILSON. 2002. The Amphibians of Honduras. Soc. Study Amphib. Reptiles, Contrib. Herpetol. 19:i–x, 1–625. PUSCHENDORF, R., F. CASTAÑEDA, AND J. R. MCCRANIE. 2006. Chytridiomycosis in wild frogs from Pico Bonito National Park, Honduras. EcoHealth 3:178–181. SKERRATT, L. F., L. BERGER, R. SPEARE, S. CASHINS, K. R. MCDONALD, A. D. PHILLOTT, H. B. HINES, AND N. KENYON. 2007. Spread of chytridiomycosis has caused the rapid global decline and extinction of frogs. EcoHealth 4:125–134. 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 306:1783– 1786. WAKE, D. B. AND V. T. VREDENBURG. 2008. Are we in the midst of the sixth mass extinction? A view from the world of amphibians. Proc. Natl. Acad. Sci. USA 105 (Suppl):11466–11473. Herpetological Review 40(3), 2009 Herpetological Review, 2009, 40(3), 309–311. © 2009 by Society for the Study of Amphibians and Reptiles Local and Regional Patterns of Amphibian Chytrid Prevalence on the Osa Peninsula, Costa Rica CAREN S. GOLDBERG* Fish and Wildlife Resources, University of Idaho Moscow, Idaho, 83844-1136, USA TANYA J. HAWLEY Department of Biology, University of Miami Coral Gables, Florida, 33124-0421, USA and LISETTE P. WAITS Fish and Wildlife Resources, University of Idaho Moscow, Idaho, 83844-1136, USA *Corresponding author: cgoldberg@vandals.uidaho.edu In Costa Rica, 34% of known amphibian species are classiﬁed as threatened; chytridiomycosis (infection caused by Batrachochytrium dendrobatidis (Bd)) is listed as a known or possible cause of decline for more than half of these species (Stuart et al. 2004). While Bd is known to be associated with extirpations of amphibian populations at high elevations in Latin America (Lips et al. 2006), this pathogen also exists at low elevations (Puschendorf et al. 2006), where its effects on amphibian populations are unclear. The Osa Peninsula, on the southern Paciﬁc coast of Costa Rica, contains a mixture of lowland tropical forest and cattle pastures (Sanchez-Azofeifa et al. 2002). Annual precipitation in this area reaches 5500 mm per year, with a dry season from December to April and a wet season from May to November (Sanchez-Azofeifa et al. 2002). Mapping of Bd outbreaks from northwest to southeast in Latin America indicates that this area may have been exposed between 1988 and 1993 (Lips et al. 2006); however, ecological niche modeling does not predict that the Osa Peninsula includes the fundamental niche of this fungus (Ron 2005). While no rapid declines of frog populations have been documented in the Osa Peninsula, population declines of some species of the genera Craugastor and Allobates are suspected to have occurred in recent years (M. Sasa, pers. comm.). We tested frogs for Bd along three streams in this region to quantify the prevalence of this pathogen and investigate local patterns of infection. Methods.—We surveyed three streams on the Osa Peninsula between 27 May and 8 June 2006 (Fig. 1). The ﬁrst stream was Quebrada Aguabuena in the Rincón area on the north side of the peninsula (70 m elev., 1.4 km survey length; 8.7001°N, 83.5234°W), the second was Quebrada Bilingual on a private farm near Piro on the southeastern side of the peninsula (30 m elev., 0.8 km survey length; 8.4292°N, 83.3583°W), and the third was Quebrada Cameronal near the Sirena station in Corcovado National Park, on the south-central side of the peninsula (25 m elev., 0.9 km survey length; 8.4292°N, 83.3583°W). Over 1000 researchers and students visited the Rincón area between 1962 and 1973 because the Osa (Rincon) Field Station was located there (Christen 1995), and the Sirena station currently receives a large number of international visitors per year. Piro has had very few visitors outside of local families. Due to heavy rain and low activity of frogs on the stream at Sirena, we supplemented our sample by surveying higher ground and collected samples from six frogs along the stream at a later date (21–22 September 2006). We sampled each site on several consecutive nights, from just after sundown until frog encounters became less frequent (usually around midnight). We hand captured each frog using fresh, powder-free latex gloves. We swabbed the underside of each frog, including toes, with a wooden-handled swab (Puritan Medical Products, Guilford, Maine, USA) for approximately 20 passes and preserved the swab in salt solution (20% DMSO, 0.25 M EDTA, 100 mM Tris, pH 7.5 and NaCl to saturation; Seutin et al. 1991). These tubes were stored at room temperature until DNA extraction. Locations of captured frogs were recorded with a Garmin GPS 12XT (Garmin Ltd., Olathe, Kansas, USA) or determined using compass and measuring tape from GPS locations. We extracted DNA from the swabs using a DNeasy Blood & Tissue Kit (Qiagen, Valencia, California, USA), adapting the tissue protocol by heating the AE solution to 55ºC before elution and incubating the sample in the AE solution at 70ºC for ﬁve minutes before the ﬁnal centrifugation. We used an Applied Biosystems 7500 Fast Real-Time PCR System (Applied Biosystems, Foster City, California, USA) to test for the presence of Bd using the protocol of Boyle et al. (2004) with the following changes: total reaction volume was 20μl, probe concentration was 125 nM, extracted DNA was not diluted, and one out of the three wells for each sample contained the internal control recommended by Hyatt et al. (2007) to ensure there was no inhibition of the reaction. International quantiﬁcation standards were obtained from the Australian Animal Health Laboratory (Geelong, Victoria, Australia). Any sample testing positive at fewer than all three wells was rerun. A second test with any positive wells conﬁrmed a positive result, while a second test with no positive wells was regarded as a negative result. A negative control was included in each extraction to conﬁrm that there was no contamination. We calculated overall prevalence of infected individuals at each site and used binomial probabilities to calculate the upper bound of a 95% conﬁdence interval by ﬁnding the highest level of prevalence where we had up to a 5% chance of not detecting the pathogen when no Bd was detected at a site. For sites where we detected Bd, we determined conﬁdence intervals using the relationship between the F distribution and the binomial distribution (Zar 1984). For these calculations, we assumed that our procedure was 100% accurate in detecting infection on a sampled individual. Ecological group was determined using breeding and habitat descriptions from Savage (2002). Results.—Across the three sites, we sampled 178 individuals and found 9 samples that tested positive for Bd (Table 1). Overall prevalence of frogs testing positive for Bd was highest at Sirena and lowest at Piro, where we found no infected individuals; however, 95% conﬁdence intervals overlapped for all sites. Frogs testing positive were spread among ecological groups (Table 1). Six of seven frogs testing positive for Bd at Rincón were found within 120 m of a dirt road crossing the stream; the remaining individual (Smilisca sordida) was found 760 m downstream from the road crossing (Fig. 1). Discussion.—We found a low level of Bd infection (approximately 8% prevalence) at two sites on the Osa Peninsula of Costa Herpetological Review 40(3), 2009 309 individuals at a third site. This estimate of prevalence is similar to that found in live adults by Puschendorf et al. (2006) in an apparently pre-decline amphibian community in Braulio Carrillo National Park, Costa Rica (5.9%) and by Lips et al. (2003) in a declining amphibian community in Las Tablas, Costa Rica (14.3%). Both survey sites on the Osa Peninsula with Bd-positive individuals have a history of large numbers of international visitors; the third has received few visitors from outside of the local area. However, our sample size was not large enough to determine if prevalence differed among sites and more data would be required to determine whether frequency of human visitation and Bd infection prevalence are correlated. Within-stream patterns of Bd infection may be structured by differences in habitat use and behavior among species (Rowley and Alford 2007; Woodhams et al. 2003). However, the few Bdinfected frogs we found were distributed across ecological groups that differed in microhabitat and mode of reproduction. FIG. 1. Location of the three sampling areas on the Osa peninsula, Costa Rica, and Bd status of This pattern of infection across species individuals sampled May–September 2006. Large map displays Bd status of individual anurans in groups is consistent with a survey of the Rincón sampling area. Dashed lines indicate dirt roads. Braulio Carrillo National Park, Costa Rica (Puschendorf et al. 2006), and does Rica, a lowland region not predicted to contain this pathogen by not follow the pattern found in southeast Queensland, Australia, the niche model of Ron (2005), and we detected no Bd-positive where frogs breeding in permanent water were more likely to be TABLE 1. Anurans tested for Bd on the Osa Peninsula, Costa Rica, in May, June, and September 2006. For each site (Rincón, Piro, Sirena), number of each species sampled (N) and number testing positive for Bd (+) are presented, followed by the quantiﬁcation of the amount of Bd collected in genome equivalents in parentheses. Conﬁdence intervals for prevalence were calculated using the binomial distribution. * Indicates samples collected in September 2006. Species Allobates talamancae Cochranella granulosa Cochranella pulverata Craugaster ﬁtzingeri Ecological group Terrestrial stream-breeder Stream-associated arboreal breeder Stream-associated arboreal breeder Terrestrial direct-developer N Rincón + N Piro + N Sirena + 6 0 1 9 0 0 0 2 6 4 20 2 0 0 0 0 0 0 0 5* 0 0 0 2 (177, 638) Hyalinobatrachium valerioi Stream-associated arboreal breeder 7 2 (15, 510) 13 0 0 0 4 0 (22, 125) Leptodactylus savagei Terrestrial pond-breeder 8 0 4 0 Rhinella marina Smilisca sordida Terrestrial pond-breeder Terrestrial stream-breeder 5 55 0 3 10 3 0 0 (1 sample*) 16 0 0 0 (289, 863, 1012) Prevalence (95% Conﬁdence interval) 310 0.077 (0.031–0.184) Herpetological Review 40(3), 2009 0 (0–0.047) 0.080 (0.010–0.395) infected than frogs breeding in ephemeral or terrestrial habitats (Kriger and Hero 2007). Along the Rincón stream, we found most infected individuals near a dirt road crossing, with the exception of one S. sordida individual found further downstream. While this concentration of infected frogs may be located near the road for unrelated reasons, the pattern could indicate that a reservoir for this pathogen is located outside of the stream and stream frogs are potentially exposed through the movement of vehicles or animals traveling along the road. Acknowledgments. –We thank D. Bellanero, J. Manning, and D. Matlaga for help in the ﬁeld and J. Kerby, C. Richards, C. Anderson, J. Adams, and T. Edwards for their help with developing lab protocols. J. Evans, M. Murphy, and two reviewers provided helpful comments on an earlier draft of this manuscript. Funding was provided by NSF-IGERT grant no. 0114304 for C.S.G. and by an EPA fellowship and a Kushlan/Frohring grant from the University of Miami to T.J.H. Field protocols were approved by the University of Idaho (ACUC Protocol 2004-42) and MINAE in Costa Rica (INV-ACOSA—005-06). LITERATURE CITED BOYLE, D. G., D. B. BOYLE, V. OLSEN, J. A. T. MORGAN, AND A. D. HYATT. 2004. Rapid quantitative detection of chytridiomycosis (Batrachochytrium dendrobatidis) in amphibian samples using real-time Taqman PCR assay. Dis. Aquat. Org. 60:141–148. CHRISTEN, C. A. 1995. Tropical ﬁeld ecology and conservation initiatives on the Osa Peninsula, Costa Rica, 1962–1973. In Y. Chantelin and C. Bonneuil (eds.), 20th Century Sciences: Beyond the Metropolis, Volume 3, Nature and Environment, pp. 366–427. Orsom Éditions, L’Institut Français de Recherche Scientiﬁque pour le Développement en Coopération, Paris, France. HYATT, A. D., D. G. BOYLE, V. OLSEN, D. B. BOYLE, L. BERGER, D. OBENDORF, A. DALTON, K. KRIGER, M. HERO, H. HINES, R. PHILLOTT, R. CAMPBELL, G. MARANTELLI, F. GLEASON, AND A. COLLING. 2007. Diagnostic assays and sampling protocols for the detection of Batrachochytrium dendrobatidis. Dis. Aquat. Org. 73:175–192. KRIGER, K. M., AND J. M. HERO. 2007. The chytrid fungus Batrachochytrium dendrobatidis is non-randomly distributed across amphibian breeding habitats. Divers. Distrib. 13:781–788. LIPS, K. R., F. BREM, R. BRENES, J. D. REEVE, R. A. ALFORD, J. VOYLE, C. CAREY, L. LIVO, A. P. PESSIER, AND J. P. COLLINS. 2006. Emerging infectious disease and the loss of biodiversity in a Neotropical amphibian community. Proc. Natl. Acad. Sci. USA 103:3165–3170. ________ , D. E. GREEN, AND R. PAPENDICK. 2003. Chytridiomycosis in wild frogs from southern Costa Rica. J. Herpetol. 37:215–218. PUSCHENDORF, R., F. BOLAÑOS, AND G. CHAVES. 2006. The amphibian chytrid fungus along an altitudinal transect before the ﬁrst reported declines in Costa Rica. Biol. Cons. 132:136–142. RON, S. 2005. Predicting the distribution of the amphibian pathogen Batrachochytrium dendrobatidis in the New World. Biotropica 37:209–221. ROWLEY, J. J. L., AND R. A. ALFORD. 2007. Behaviour of Australian rainforest stream frogs may affect the transmission of chytridiomycosis. Dis. Aquat. Org 77:1–9. SANCHEZ-AZOFEIFA, G. A., B. RIVARD, J. CALVO, AND I. MOORTHY. 2002. Dynamics of tropical deforestation around national parks: remote sensing of forest change on the Osa Peninsula of Costa Rica. Mt. Res. Dev. 22:352–358. SAVAGE, J. M. 2002. Amphibians and Reptiles of Costa Rica: A Herpetofauna between Two Continents, between Two Seas. University of Chicago Press, Chicago, Illinois. 934 pp. SEUTIN, G., B. N. WHITE, AND P. T. BOAG. 1991. Preservation of avian blood and tissue samples for DNA analysis. Can. J. Zool. 69:82–90. 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 306:1783–1786. WOODHAMS, D. C., R. A. ALFORD, AND G. MARANTELLI. 2003. Emerging disease of amphibian cured by elevated body temperature. Dis. Aquat. Org 55:65–67. ZAR, J. H. 1984. Biostatistical Analysis. 2nd ed. Prentice Hall, Englewood Cliffs, New Jersey. 718 pp. Herpetological Review, 2009, 40(3), 311–313. © 2009 by Society for the Study of Amphibians and Reptiles Occurrence of Batrachochytrium dendrobatidis in an Anuran Community in the Southeastern Talamanca Region of Costa Rica DANIEL SAENZ* CORY K. ADAMS JOSH B. PIERCE Southern Research Station, Forest Service, U. S. Department of Agriculture 506 Hayter Street, Nacogdoches, Texas 75965-3556, USA and DAVID LAURENCIO Department of Biological Sciences, 331 Funchess Hall, Auburn University Auburn, Alabama 36849, USA * Corresponding author; e-mail: dsaenz@fs.fed.us Soon after the discovery of the amphibian disease chytridiomycosis, caused by the pathogenic fungus Batrachochytrium dendrobatidis (Bd, Longcore et al. 1999), it became apparent that Bd was a major threat to amphibians resulting in mass die-offs and population declines throughout the world (Berger et al. 1998; Blaustein and Keisecker 2002; Daszak et al. 2003; McCallum 2005; Rachowicz et al. 2006). Evidence suggests that Bd infected amphibian populations in Central America as early as the 1980s (Pounds et al. 1997). Work done in Central America has implicated that a wave of Bd has moved through the montane regions of Central America and was associated with major declines in amphibian populations and species richness (Lips et al. 2006; Puschendorf et al. 2006). However, in some cases, Bd also may occur in amphibian communities with little or no effect on populations (Berger et al. 1998; Brem and Lips 2008; Garner et al. 2006). We sampled amphibians for the presence of Bd in the Kèköldi Indigenous Reserve in the lower elevations of the Talamanca region, Costa Rica, to determine Bd infection rates in an area not previously surveyed. Also, we attempted to determine which species might be at greatest risk from Bd. Although our study site had not been sampled prior to this study, Bd likely reached this region in the early 1990s (see Lips et al. 2006). We sampled anurans for Bd from 6 to 17 January 2008 in the 3538-ha Kèköldi Indigenous Reserve, located in southeastern Costa Rica near Hone Creek in Limon Province (Fig. 1). The habitat in the reserve is mostly secondary forest, underplanted with cacao trees; however some primary lowland tropical rainforest occurs at higher elevations in the reserve. The climate at Kèköldi is hot and Herpetological Review 40(3), 2009 311 TABLE 1. List of anuran species tested for the presence of Batrachochytrium dendrobatidis (Bd) within Kèköldi Indigenous Reserve, Costa Rica. Family Species Aromobatidae Allobates talamancae 0/2 Bufonidae Incilius coniferus Rhinella marinus 0/1 0/1 Centrolenidae Hyalinobatrachyum valerioi 0/5 Craugastoridae Craugastor bransfordii Craugastor crassidigitus Craugastor gollmeri Craugastor megacephalus Craugastor noblei Craugastor sp. 1/13 3/12 0/1 0/12 1/2 0/3 Dendrobatidae Dendrobates auratus Oophaga pumilio Phyllobates lugubris Silverstoneia ﬂotator 1/9 1/16 1/4 0/15 Eleutherodactylidae Diasporus diastema 1/16 Hylidae Agalychnis callidryas Smilisca phaeota Smilisca sordida 0/1 0/1 1/3 Leptodactylidae Leptodactylus savagei 0/6 Ranidae Lithobates warszewitschii 0/2 Strabomantidae Pristimantis cerasinus 0/1 FIG. 1. Study site location, Kèköldi Indigenous Reserve, Costa Rica, where 20 frog species were examined for the amphibian chytrid fungus, Batrachochytrium dendrobatidis. humid year-round, with an annual mean temperature of 26ºC and average rainfall of approximately 2500 mm per year. We searched for adult frogs along the trails and streams in the reserve and captured them by hand. Each individual was handled with a new pair of sterile nitrile gloves. We sampled for Bd by rubbing a sterile cotton swab on the dorsum, ventral surfaces and feet of each frog for approximately 30 sec, then the animal was released. The swab was then immediately placed in a sterile microcentrifuge tube containing 1 ml of 70% ethanol and later sent to Pisces Molecular Lab (Boulder, Colorado, USA) for PCR analyses. Global positioning coordinates and elevation were taken at each capture site using a Garmin® GPS unit. During 12 days of sampling at Kèköldi, no sick or dead frogs were observed. We sampled 126 adult frogs of 20 different species, from 10 different Families. Of these 20 species, only 8 tested positive for Bd. Ten of the 126 individuals tested positive for an overall detection rate of 7.9% for the anuran community. Too few individuals were sampled to determine Bd prevalence per species. Only one species, Craugastor crassidigitus, had more than one individual (3 of 12) test positive for Bd (Table 1). Anurans were sampled at elevations ranging from 29 to 150 m. We had too few positive samples to test for effects of elevation on Bd detection rates. Numerous studies have documented the presence of Bd in Central America (Brem and Lips 2008; Lips 1998; Lips et al. 2003; Picco and Collins 2007; Puschendorf et al. 2006). Puschendorf et al. (2006) demonstrated through histological examination of museum specimens that the fungus was present in Braulio Carrillo National Park, Costa Rica, by 1986 at almost all elevations sampled. Declining species richness in amphibian communities along a north-tosouth transect in Central America has been linked to Bd (Lips et al. 2006). According to the timeline in Lips et al. (2006) tracking the epidemic wave of Bd in Costa Rica, the fungus probably reached the Kèköldi Reserve in the early to mid 1990s. Since Bd likely moved through Kèköldi 15 to 20 years prior to this study, it is possible that an initial epizootic event took place resulting in declines in amphibian species richness and abundance followed by a rebound to an enzootic state (Brem and Lips 2008). It is also possible that the fungus never reached an 312 No. animals infected / examined epizootic state in the low elevation Kèköldi Reserve because the warm air temperatures are less than optimal for Bd growth and infection of amphibians (Kriger and Hero 2006; Longcore et al. 1999; Retallick et al. 2004). Evidence for enzootic Bd includes the relative low incidence of detection in PCR samples and the fact that no sick or dead frogs were encountered. Frogs appeared to be very abundant in the reserve; however follow-up population and Bd sampling is needed to conﬁrm whether the Bd is currently epizootic or enzootic. Acknowledgments.—We thank R. Schaefer and M. Kwiatkowski for useful comments on an earlier draft of this manuscript. We thank T. Hibbitts, D. Henderson, T. Krause, and L. Larencio for assisting with ﬁeld sampling. LITERATURE CITED BERGER, L., R. SPEARE, P. DASZAK, D. E. GREEN, A. A. CUNNINGHAM, C. L. GOGGIN, R. SLOCOMBE, M. A. RAGAN, A. D. HYATT, K. R. MCDONALD, H. B. HINES, K. R. LIPS, G. MARTANTELLI, AND H. PARKES. 1998. Chytridiomycosis causes amphibian mortality associated with population Herpetological Review 40(3), 2009 declines in the rain forests of Australia and Central America. Proc. Nat. Acad. Sci. 95:9031–9036. BLAUSTEIN, A. R., AND J. M. KEISECKER. 2002. Complexity in conservation: lessons from the global decline of amphibian populations. Ecol. Lett. 5:597–608. BREM, F. M. R., AND K. R. LIPS. 2008. Batrachochytrium dendrobatidis infection patterns among Panama amphibian species, habitats and elevations during epizootic and enzootic stages. Dis. Aquat. Org. 81:189–202. DASZAK, P., A. A. CUNNINGHAM, AND A. D. HYATT. 2003. Infectious disease and amphibian declines. Diversity and Distributions 9:141–150. GARNER, T. W. J., M. W. PERKINS, P. GOVINDARAJULU, D. SEGLIE, S. WALKER, A. A. CUNNINGHAM, AND M. C. FISHER. 2006. The emerging amphibian pathogen Batrachochytrium dendrobatidis globally infects introduced populations of the North American bullfrog, Rana catesbeiana. Biol. Lett. 2:455–459. KRIGER, K. M., AND J. M. HERO. 2006. Large-scale seasonal variation in the prevalence and severity of chytridiomycosis. J. Zool. (Lond) 271:352–359. LIPS, K. R. 1998. Decline of a tropical montane amphibian fauna. Conserv. Biol. 21(1):106–117. ________ , F. BREM, R. BRENES, J. D. REEVE, R. A. ALFORD, J. VOYLES, C. CAREY, L. LIVO, A. P. PESSIER, AND J.P. COLLINS. 2006. Emerging infectious disease and the loss of biodiversity in a neotropical amphibian community. PNAS 103(6):3165–3170. ________ , D. E. GREEN, AND R. PAPENDICK. 2003. Chytridiomycosis in wild frogs from Southern Costa Rica. J. Herpetol. 37(1):215–218. LONGCORE, J. E., A. P. PESSIER, AND D. K. NICHOLS. 1999. Batrachochytrium dendrobatidis gen. et sp. nov., a chytrid pathogenic to amphibians. Mycologia 91(2):219–227. MCCALLUM, H. 2005. Inclusion of chytridiomycosis as the agent in widespread frog declines. Conserv. Biol. 19(5):1421–1430. PICCO, A. M., AND J. P. COLLINS. 2007. Fungal and viral pathogen occurrence in Costa Rican amphibians. J. Herpetol. 41(4):746–749. POUNDS, J. A., M. P. L. FOGDEN, J. M . SAVAGE, AND G. C. GORMAN. 1997. Test of null models for amphibian declines on a tropical mountain. Conserv. Biol. 11(6):1307–1322. PUSCHENDORF, R., F. BOLANOS, AND G. CHAVES. 2006. The amphibian chytrid fungus along an altitudinal transect before the ﬁrst reported declines in Costa Rica. Biol. Conserv. 132:136–142. RACHOWICZ, L. R., R. A. KNAPP, J. A. T. MORGAN, M. J. STICE, V. T. VRENDENBURG, J. M. PARKER, AND C. J. BRIGGS. 2006. Emerging infectious disease as a proximate cause of amphibian mass mortality. Ecology 87(7):1671–1683. RETALLICK, R. W. R., H. MCCALLUM, AND R. SPEARE. 2004. Endemic infection of the amphibian chytrid fungus in a frog community post-decline. PLoS 2(11):1965–1971. Herpetological Review, 2009, 40(3), 313–316. © 2009 by Society for the Study of Amphibians and Reptiles Detecting Batrachochytrium dendrobatidis in the Wild When Amphibians Are Absent JOEL G. WIXSON and KEVIN B. ROGERS* Aquatic Research Group, Colorado Division of Wildlife P.O. Box 775777, Steamboat Springs, Colorado 80477, USA *Corresponding author; e-mail: kevin.rogers@state.co.us Once common in the southern Rocky Mountains of North America, sharp declines in Boreal Toad (Anaxyrus boreas boreas) populations precipitated their listing as a state endangered species in Colorado, USA (Loefﬂer 2001) and consideration for listing under the Endangered Species Act (U.S. Fish and Wildlife Service 2005). The amphibian chytrid fungus (Batrachochytrium dendrobatidis, hereafter Bd) has been implicated in these declines (Livo 2000; Muths et al. 2003; Scherer et al. 2005). Interest in reintroducing A. b. boreas into historical habitats (Loefﬂer 2001) has spurred the need to develop a test for the presence of Bd. Reintroduction efforts are time consuming and costly, and their success may hinge on the occurrence of Bd at a potential site. As such, it is imperative that disease status be considered when evaluating potential reintroduction efforts. Currently our ability to detect Bd at a site relies on resident amphibians being present, yet they are not at many promising potential reintroduction locations. Since Bd can persist at a location even in the absence of amphibian species (Longcore et al. 1999; Rowley et al. 2007; Speare et al. 2001), we suspect that amphibians may not be the only host, and that infection can be maintained through other alternate hosts or environmental reservoirs. We hope that by testing these non-amphibian sources, the Bd status at potential reintroduction sites can be evaluated. Rowley et al. (2007) did not detect Bd in retreat sites of rain forest stream frogs, while Lips et al. (2006) did ﬁnd Bd DNA on stream boulders but not in ﬁltered water samples. Others have detected Bd in ﬁltered water samples (Kirshtein et al. 2007; Walker et al. 2007), but their approaches do not always perform well in waters carrying high organic loads that rapidly clog ﬁlters (Cossel and Lindquist 2009) or cause PCR inhibition (Kirshtein et al. 2007). Our initial efforts toward ﬁnding alternative Bd hosts focused on insects, because they are readily available and chytrid fungi can degrade chitin, a component of aquatic insect exoskeletons (Johnson and Speare 2003; Powell 1993). These early surveys were unable to conﬁrm the presence of Bd in samples of Dytiscidae, Coenagrionidae, Hydrophilidae, or Notonectidae from two ponds known to harbor the fungus (Rogers et al. 2004). Samples of Corixidae, algae, snails, and clams taken from a third pond with infected Boreal Chorus Frogs (Pseudacris maculata) were also negative for Bd DNA (Rogers and Wood 2005). In an effort to establish a more rigorous examination of potential alternate hosts, we initiated a study to explore the feasibility of using sentinel cages and ﬁsh following reports that Bd could be found on the scales of Fathead Minnows (Pimephales promelas) that were exposed to Bd in the laboratory (R. Retallick, pers. comm., GHD, Australia). Feathers and keratin were included Herpetological Review 40(3), 2009 313 in this ﬁeld study as well when others demonstrated the ability to culture Bd on sterile duck feathers (Johnson and Speare 2005) or 1% keratin agar (Piotrowski 2004) in vitro. Methods.—Sentinel experiments were conducted in both a midelevation site in the town of Steamboat Springs, Colorado, and a high elevation site on the Grand Mesa, Colorado after the spring thaw in 2005 when prevalence of Bd infection was greatest (K. Rogers, unpubl. data). This occurred in May for the low elevation site in Steamboat Springs, and in July for the high elevation sites on the Grand Mesa. Bd presence at both sites was conﬁrmed by swabbing resident P. maculata following Livo (2004). DNA was extracted from the samples using a standard spin column protocol. All sample DNA preparations were assayed for the presence of the Bd ribosomal RNA Intervening Transcribed Sequence (ITS) region by 45 cycle single-round PCR ampliﬁcation (Annis et al. 2004) that was modiﬁed for greater speciﬁcity and sensitivity at Pisces Molecular, Boulder, Colorado. Cages (0.125 m3) were constructed of 4 × 4 cm pine boards and 3 mm mesh to house sentinel animals. Cages were deployed in water less than 60 cm deep, and secured to the bottom with metal stakes. A protective hardware cloth (25 mm mesh) was attached to the outside of each cage to protect them from predators. Sentinel ﬁsh were sampled for Bd at 1, 3, 7, and 14 days following introduction to the cages, and mortalities were noted. Fish were swabbed on their right ﬂanks one day after exposure. After 3, 7 and 14 days of exposure, 10 ﬁsh of each species from each pen were euthanized with MS-222, then swabbed, scraped, and ﬁn clipped. A cotton swab (Puritan cotton-tipped applicators, VWR International, West Chester, Pennsylvania), was stroked 20 times unidirectionally across the left ﬂank of each sentinel ﬁsh, then preserved in 70% ethanol (Livo et al. 2004) for subsequent PCR screening. The skin scrapes followed a similar protocol but used a sharpened wooden dowel (Livo 2004). Paired and caudal ﬁns were removed and preserved in 70% ethanol. In addition to sampling sentinel ﬁsh, six mallard (Anas platyrhynchos) ﬂank feathers were taped together at the stalk and suspended in the surface ﬁlm with a string attached to the outside of the cage. A feather was collected on each sampling day by clipping the exposed end of the feather and placing the complete piece in a 2-mL microcentrifuge tube containing 70% ethanol, then processed with the same PCR procedure. The ﬁrst study was conducted in a small temporary springﬂooded pond (Trafalger Pond) next to the Yampa River within the city limits of Steamboats Springs, Colorado (2051 m elev.; 40.47445°N, 106.83017°W). Skin swabs from 20 resident adult P. maculata collected during the breeding season suggest this pond has harbored Bd since at least 2004 (30% prevalence, K. Rogers, unpubl. data). Thirty Rainbow Trout (Oncorhynchus mykiss), 30 P. promelas, and 30 Goldﬁsh (Carassius auratus) were used as sentinel ﬁsh in each of four cages spread throughout the pond, in addition to six Mallard ﬂank feathers suspended outside of each cage. The second study was conducted in the Kannah Creek drainage on the Grand Mesa near Grand Junction, Colorado (3268 m elev.; 39.04420°N, 108.02992°W). Dozens of small ponds in this drainage are home to robust populations of P. maculata. Ponds with perennial water also support Tiger Salamanders (Ambystoma tigrinum). Bd was ﬁrst detected in this drainage in 2003 in P. maculata 314 collected from a 1.0 ha pond (Pond 4) used in this study (Rogers and Banulis 2004). One cage with 30 P. promelas was deployed in each of two additional 0.5 ha ponds, hereafter referred to as Lands End and Cow Camp. Pond 4 received two cages, each with 30 P. promelas. Six Mallard ﬂank feathers were installed outside of each of the four cages. In addition, we explored baiting Bd with pure keratin (VWR International, West Chester, Pennsylvania). Keratin tea bags were constructed from paper coffee ﬁlters, cut in half and sewed together. Five bags were fastened to each cage with 3 g of keratin per bag. A bag was removed from each cage on every sampling occasion, and a portion of the contents preserved in 70% ethanol. Skin swabs from 20 adult P. maculata were collected from each of these three ponds the day after this 14-day experiment to evaluate the prevalence of Bd in 2005. Results.—In an effort to reduce assay costs in the ﬁrst study, only the feathers collected at 1, 3, 7, and 14 days following exposure along with ﬁsh skin swabs collected one day following exposure on O. mykiss and three days following exposure on P. promelas and C. auratus were submitted for analysis. None of these 16 feather samples or 120 ﬁsh samples suggested that Bd was present. Because the ability to use sentinel organisms at this site did not appear promising and processing samples was costly, the remaining samples were archived. In the second study, despite a substantial number of Bd-positive samples from the P. maculata collected at the end of our experiment (prevalence of Bd ranging from 25–30% in all three ponds), only six of 350 ﬁsh swab, scrape, and ﬁn samples were Bd positive. These included ﬁve swabs collected one day after exposure on P. promelas in Cow Camp and a single swab from Lands End, also collected one day after exposure that returned a very weak positive signal. None of the ﬁsh swabs, scrapes, or ﬁn clips collected 3, 7, or 14 days after exposure yielded positive results. Feathers and raw keratin were equally ineffective, as all 32 samples failed to register any evidence of Bd over the course of the experiment. Discussion.—Caged ﬁsh, feathers, and keratin were ineffective at sampling Bd in ponds known to have amphibians with the disease. Although swabbing the ﬂanks of P. promelas exposed for one day in a Bd-positive environment yielded Bd-positive results, ﬁsh sampling was clearly much less sensitive than sampling P. maculata in the same environment. The fact that the majority of the positive results came from the single cage at Cow Camp, and that positives were found after only one day of exposure but not after a week or two weeks makes the use of P. promelas as sentinel organisms problematic. Rather than Bd actually infecting the host, this suggests that the cage was fortuitously deployed in an area with Bd zoospores, and that they simply adhered to the ﬁsh when sampled on that ﬁrst day. Chytrid spores have a short-lived free-swimming stage that only lasts about 24 hrs before encysting (Piotrowski et al. 2004). Even with this 24 hr active period the spore can only swim 2 cm (Piotrowski et al. 2004). Thus we may have simply been lucky in our placement of the Cow Camp cage in particular. There does not appear to be any particular afﬁnity by the zoospores toward P. promelas, as subsequent samples revealed no indication of Bd presence. Given the inconsistent nature of the results, it is doubtful that using sentinel P. promelas would be a viable approach for screening potential amphibian reintroduction sites for the presence of the fungus. Although sentinel ﬁsh have been used to test for pathogens like whirling disease (Koel et al. Herpetological Review 40(3), 2009 2006; Thompson et al. 1999), the construction and deployment of cages remains labor intensive, particularly in sites that are difﬁcult to access. This is an additional consideration, given the apparently low sensitivity of P. promelas as a sentinel to detect Bd. Because the majority of positive samples came from the same cage on the same day, contamination of the samples is a concern. The frog samples used to conﬁrm the presence of the fungus were collected following the experiment, and stored in a different location making them an unlikely source of contamination. Using latex gloves between samples, and sterilizing equipment with an open ﬂame further minimized contamination risk. Contamination occurring in the original source stock of P. promelas was also ruled out, as most positive signals came from a single cage, and only early in the study. Samples collected after 3, 7, and 14 days were all Bd-negative. Given that others have been successful growing chytrid fungi on a 1% keratin agar (Piotrowski et al. 2004) or on feathers (Johnson and Speare 2005) in vitro, we were surprised that none of the keratin or feather samples yielded a positive PCR result. It was suggested that perhaps the Bd fungus did not colonize the interior of the keratin bags, but rather just the outside, which was not sampled. Subsequent tests using agar-impregnated swabs rolled in raw keratin however also failed to bait in Bd following three days of exposure (K. Rogers, unpubl. data). If Bd has a patchy distribution in a pond environment, it would be difﬁcult to sample with ﬁxed organisms or objects. A more effective approach would be to release amphibians targeted for a reintroduction effort, then subsequently collecting survivors the following year to sample for Bd. Using a non-tethered organism allows the target to move through the environment as it would following a repatriation effort, encountering pathogens along the way. Repatriation efforts require the ability to produce large numbers of offspring for subsequent release; a portion of this captive production could be used to assay potential ﬁeld sites for Bd. If a captive broodstock is not available, fertilized eggs could be secured from the wild, washed to minimize risk of Bd transfer, then raised to the larval stage for release (Rogers and Banulis 2004). This approach requires that target amphibians be produced during pilot studies prior to implementing full repatriation efforts to determine the suitability of potential translocation sites. Acknowledgments.—Our gratitude is extended to C. Slubowski who shepherded the sentinel cages on the Grand Mesa and helped collect samples. F. B. Wright III is thanked for assisting with sample collection and providing many helpful ideas. We also appreciate the time of G. and J. Wixson who supplied the necessary tools and advice in building the cages. Substantial improvements to the manuscript were provided by D. Olson, J. Rowley, and an anonymous reviewer. All applicable institutional animal care guidelines were followed during the implementation of this project. LITERATURE CITED ANNIS, S. L., F. P. DASTOOR, H. ZIEL, P. DASZAK, AND J. E. LONGCORE. 2004. A DNA-based assay identiﬁes Batrachochytrium dendrobatidis in amphibians. J. Wildl. Dis. 40:420-428. COSSEL, J. O. JR., AND E. D. LINDQUIST. 2009. Batrachochytrium dendrobatidis in arboreal and lotic water sources in Panama. Herpetol. Rev. 40:45-47. JOHNSON, M. L., AND R. SPEARE. 2003. Survival of Batrachochytrium dendrobatidis in water: quarantine and disease control implications. Emerg. Infect. Dis. 9:922–925. ________ , AND R. SPEARE. 2005. Possible modes of dissemination of the amphibian chytrid Batrachochytrium dendrobatidis in the environment. Dis. Aq. Org. 65:181–186. KIRSHTEIN, J. D., C. W. ANDERSON, J. S. WOOD, J. E. LONGCORE, AND M. A. VOYTEK. 2007. Quantitative PCR detection of Batrachochytrium dendrobatidis DNA from sediments and water. Dis. Aq. Org. 77:11–15. KOEL, T. M., D. L. MAHONY, K. L. KINNAN, C. RASMUSSEN, C. J. HUDSON, S. MURCIA, AND B. L. KERANS. 2006. Myxobolus cerebralis in native cutthroat trout of the Yellowstone Lake ecosystem. J. Aq. Animal Health 18:157–175. LIPS, K. R., F. BREM, R BRENES, J. D. REEVE, R. A. ALFORD, J. VOYLES, C. CAREY, L. LIVO, A. P. PESSIER, AND J. P. COLLINS. 2006. Emerging infectious disease and the loss of biodiversity in a neotropical amphibian community. Proc. Natl. Acad. Sci. 103:3165–3170. LIVO, L. J. 2000. Amphibious assault. Colorado Outdoors 49(6):26–29. ________ . 2004. Methods for obtaining Batrachochytrium dendrobatidis (Bd) samples for PCR testing. In K. B. Rogers (ed.), Boreal Toad Research Report 2003, pp. 64–68. Colorado Division of Wildlife, Fort Collins, Colorado. http://wildlife.state.co.us/Research/Aquatic/BorealToad/. ________ , J. WOOD, S. ANNIS, C. CAREY, J. EPP, AND M. S. JONES. 2004. Evaluation of techniques for detecting Batrachochytrium dendrobatidis from amphibians for PCR testing. In K. B. Rogers (ed.), Boreal Toad Research Report 2003, pp. 15–22. Colorado Division of Wildlife, Fort Collins, Colorado. http://wildlife.state.co.us/Research/Aquatic/BorealToad/. LOEFFLER, C. 2001. Conservation plan and agreement for the management and recovery of the southern Rocky Mountain population of the boreal toad (Bufo boreas boreas). Boreal Toad Recovery Team. 76 pp. Colorado Division of Wildlife, Fort Collins, Colorado. http://wildlife. state.co.us/Research/Aquatic/BorealToad/. LONGCORE, J. E., A. P. PESSIER, and D. K. NICHOLS. 1999. Batrachochytrium dendrobatidis gen. et sp. nov., a chytrid pathogenic to amphibians. Mycologia. 91:219–227. MUTHS, E., P. S. CORN, A. P. PESSIER, AND D. E. GREEN. 2003. Evidence for disease-related amphibian decline in Colorado. Biol. Conserv. 110:357–365. PIOTROWSKI, J. S., S. L. ANNIS, AND J. E. LONGCORE. 2004. Physiology of Batrachochytrium dendrobatidis, a chytrid pathogen of amphibians. Mycologia 96(1):9–15. POWELL, M. J. 1993. Looking at mycology with a Janus face: a glimpse of chytridiomycetes active in the environment. Mycologia 85:1–20. ROGERS, K. B., AND T. BANULIS. 2004. Repatriation of boreal toads Bufo boreas on the Grand Mesa, Colorado. In K. B. Rogers (ed.), Boreal Toad Research Report: 2003, pp. 2–12. Colorado Division of Wildlife, Fort Collins, Colorado. http://wildlife.state.co.us/Research/Aquatic/ BorealToad/. ________ , S. RITTMANN, AND J. WOOD. 2004. A look at aquatic macroinvertebrates as reservoirs of Batrachochytrium dendrobatidis infection. In K. B. Rogers (ed.), Boreal Toad Research Report: 2003, pp. 52–54. Colorado Division of Wildlife, Fort Collins, Colorado. http://wildlife. state.co.us/Research/Aquatic/BorealToad/. ________ , AND J. WOOD. 2005. Looking for reservoirs of Batrachochytrium dendrobatidis infection. In T. Jackson (ed.), Report on the Status and Conservation of the Boreal Toad Bufo boreas boreas in the Southern Rocky Mountains: 2004, pp. 59–60. Colorado Division of Wildlife, Fort Collins, Colorado. http://wildlife.state.co.us/Research/Aquatic/ BorealToad/. ROWLEY, J. J. L., L. F. SKERRATT, R. A. ALFORD, AND R. CAMPBELL. 2007. Retreat sites of rain forest stream frogs are not a reservoir for Batrachochytrium dendrobatidis in northern Queensland, Australia. Dis. Aq. Org. 74:7–12. SCHERER, R. D., E. MUTHS, B. R. NOON, AND P. S. CORN. 2005. An evaluation of weather and disease as causes of decline in two populations of boreal toads. Ecol. Appl. 15:2150–2160. Herpetological Review 40(3), 2009 315 SPEARE R., R. ALFORD, K. APLIN, L. BERGER, AND 10 OTHERS. 2001. Nomination for listing of amphibian chytridiomycosis as a key threatening process under the Environment Protection and Biodiversity Conservation Act 1999. In R. Speare (ed.), Developing Management Strategies to Control Amphibian Diseases: Decreasing the Risks Due to Communicable Diseases, pp. 185–196. School of Public Health and Tropical Medicine, James Cook University, Townsville. THOMPSON , K. G, R. B. NEHRING, D. C. BOWDEN, AND T. WYGANT. 1999. Field exposure of seven species or subspecies of salmonids to Myxobolus cerebralis in the Colorado River, Middle Park, Colorado. J. Aq. Anim. Health 11:312–329. U.S. FISH AND WILDLIFE SERVICE. 2005. Revised 12-month ﬁnding for the Southern Rocky Mountain distinct population segment of the boreal toad (Bufo boreas boreas). Federal Register 70:56880–56884. WALKER, S. F., M. B. SALAS, D. JENKINS, T. W. J. GARNER, A. A. CUNNINGHAM, A. D. HYATT, J. BOSCH, AND M. C. FISHER. 2007. Environmental detection of Batrachochytrium dendrobatidis in a temperate climate. Dis. Aq. Org. 77:105–112. Herpetological Review, 2009, 40(3), 316–319. © 2009 by Society for the Study of Amphibians and Reptiles First Report of Ranavirus Infecting Lungless Salamanders MATTHEW J. GRAY* DEBRA L. MILLER** and JASON T. HOVERMAN Center for Wildlife Health, Department of Forestry, Wildlife and Fisheries University of Tennessee, 274 Ellington Plant Sciences Building Knoxville, Tennessee 37996, USA **Veterinary Diagnostic and Investigational Laboratory University of Georgia, 43 Brighton Road Tifton, Georgia 31793-1389, USA *Corresponding author; e-mail: mgray11@utk.edu Ranaviruses are a group of pathogens belonging to the genus Ranavirus (Family Iridoviridae) that have been linked to catastrophic die-offs of larval amphibians in North America and elsewhere (Daszak et al. 2003). They also have been identiﬁed as the etiologic agent in the mass mortality of adult Common Frogs (Rana temporaria) and Common Toads (Bufo bufo) in the United Kingdom (Cunningham et al. 1996; 2007a,b). In the United States, ranaviruses are responsible for the majority of disease-related mortality events in amphibians (Green et al. 2002; Muths et al. 2006). There is evidence that ranaviruses may be an emerging infectious disease (Storfer et al. 2007), possibly due to novel strain introduction (Picco and Collins 2008) or increased occurrence of anthropogenic stressors on the landscape (Forson and Storfer 2006; Gray et al. 2007). Recognizing the potential threat of ranaviruses to global amphibian biodiversity, the World Organization for Animal Health (OIE) recently listed this pathogen as a notiﬁable disease, requiring proof of Ranavirus-negative results before commercial shipment of amphibians (OIE 2008). The OIE identiﬁes ﬁeld surveillance as a critical component of risk assessment for ranaviruses (OIE 2008). Although surveillance for the amphibian pathogen Batrachochytrium dendrobatidis has become widespread (e.g., Chatﬁeld et al. 2009; Rothermel et al. 2008), testing amphibians for Ranavirus occurs less frequently. 316 The Southern Appalachian Mountain Range of North America represents a global hotspot for salamander biodiversity (Dodd 2004; Petranka 1998). In particular, lungless salamanders (Family Plethodontidae) occur in high abundance and biomass (Peterman et al. 2008; Petranka and Murray 2001) and are important components of the ecosystem (Davic and Welsh 2004). Known die-offs of Eastern Newts (Notophthalmus viridescens), Spotted Salamanders (Ambystoma maculatum), Pickerel Frogs (Lithobates palustris), and Wood Frogs (L. sylvaticus) occurred in the Southern Appalachian Mountains due to ranaviruses in 1999 and 2001 (Converse and Green 2005; Green et al. 2002). Despite these mortality events, surveillance for Ranavirus in Southern Appalachian amphibians has been nonexistent. Our goal was to test for the presence of Ranavirus in lungless salamander communities located in the Southern Appalachian Mountains. We tested for Ranavirus in salamander communities at three sites that differed in elevation and report prevalence by species and for each site. Methods.—We captured adult lungless salamanders for Ranavirus testing on 21 April 2007 at three locations in the Great Smoky Mountains National Park, Tennessee: 1) Ash Hopper Branch (456 m elev.; 35.6836°N, 83.5375°W); 2) Chimney Tops Seeps (831 m elev.; 35.6367°N, 83.4928°W); and 3) Indian Gap Seeps (1537 m elev.; 35.61°N, 83.45°W). At all sites, we searched 1 h for salamanders in streams, seeps, and under terrestrial ground cover items (e.g., logs, rocks). We placed captured salamanders in individual 1-L plastic containers and processed up to 10 individuals per species per site. We swabbed the oral cavity and the cloaca twice each. We wore disposable gloves and changed them between animals to minimize the likelihood of cross contamination among samples. We put each swab in separate microcentrifuge tubes, placed the tubes on dry ice, and froze the swabs at -70°C within 10 h of collection. We swabbed 21, 21, and 27 salamanders at Ash Hopper Branch, Chimney Tops, and Indian Gap sites, respectively. All salamanders were released at their approximate capture location within 1 h of capture, and containers, footwear, and equipment were disinfected with 2% Nolvasan® prior to moving between sites (Bryan et al. 2009). We shipped frozen swabs overnight on dry ice to the University of Georgia Veterinary Diagnostic and Investigational Laboratory for Ranavirus testing. We used conventional polymerase chain reaction (PCR) to test for the occurrence of Ranavirus. We extracted genomic DNA from swabs using a QIAamp DNA Mini Kit (Qiagen Inc., Valencia, California). A heminested PCR targeting a 500-bp region of the major capsid protein (MCP) gene was used following the protocol by Kattenbelt et al. (2000). The PCR products were resolved by gel electrophoresis for determination of Ranavirus occurrence. We randomly chose one sample per species with a distinct PCR-positive band, cut the band from the gel, and submitted the isolated product to SeqWright DNA Technology Services, Houston, Texas, for automated sequencing in the forward and reverse directions. We then performed a GenBank BLAST search (http://www.ncbi. nlm.nih.gov/Genbank.html) on the sequences to verify that positive PCR results were Ranavirus. Additionally, real-time quantitative PCR (qPCR) was performed following the procedure by Picco et al. (2007) on samples used for sequencing to further support our ﬁndings. We summarized the positive test results for each species, and tallied Ranavirus prevalence among species for each sampling Herpetological Review 40(3), 2009 TABLE 1. Prevalence of Ranavirus detected from oral and cloacal swabs using PCR for 10 species of adult lungless salamander, Plethodontidae, in the Great Smoky Mountains National Park, Tennessee, USA, April 2007. Species Desmognathus conanti D. imitator D. monticola D. ocoee D. quadramaculatus D. santeetlah D. wrighti Eurycea wilderae Gyrinophilus porphyriticus Plethodon jordani Total No. Infected/ Tested Prevalence Accession Number1 13/14 11/13 1/1 2/2 7/8 11/13 1/2 8/12 1/2 1/2 56/69 0.93 0.85 1.0 1.0 0.88 0.85 0.5 0.67 0.5 0.5 0.81 GQ326559 GQ326560 GQ326561 GQ326562 GQ326563 GQ326564 NS GQ326565 GQ326566 NS 1 GenBank accession number of the DNA sequence used for Ranavirus determination; NS = not sequenced because insufﬁcient genomic DNA was ampliﬁed for sequencing. site. We tested for the differences in Ranavirus prevalence among sites using logistic analysis and calculated odds-ratio statistics to estimate the likelihood of Ranavirus infection for each site (Stokes et al. 2000). All analyses were performed using the SAS® system (SAS Institute, Cary, North Carolina) with α = 0.05. Results.—We captured 10 species of lungless salamanders and all species included individuals that tested positive for Ranavirus. The qPCR results matched the conventional PCR results. We were able to sequence the PCR products from eight of the infected species. GenBank BLAST searches on the sequences revealed identities of 94–96% and 97–100% with ATV and FV3, respectively. Overall prevalence was 81% among species and sites (Table 1). Ranavirus prevalence differed among sites (χ2 = 7.14, P = 0.028; Fig. 1). Salamanders were 10 and 3 times more likely to be infected at Ash Hopper Branch (odds ratio = 0.1) and Chimney Tops Seeps (odds ratio = 0.3), respectively, compared to the Indian Gap Seeps. Discussion.—We report the ﬁrst documentation of Ranavirus infecting salamander species in the family Plethodontidae. We documented Ranavirus infecting 10 species of lungless salamander at an overall prevalence of 81% among species. The high prevalence of Ranavirus in adult lungless salamanders may indicate that they are tolerant of infection. Brunner et al. (2004) reported that postmetamorphic Tiger Salamanders (Ambystoma tigrinum) could be asymptomatic carriers of ranaviruses. However, no studies have quantiﬁed the pathogenicity of ranaviruses for lungless salamanders or whether virulence differs among developmental stages. Studies with anurans and ambystomatid salamanders suggest that adult stages are less susceptible to Ranavirus than larval stages (Collins et al. 2004; Gantress et al. 2003), thus adult amphibians have the greatest likelihood of serving as a Ranavirus reservoir. It also is possible that the high Ranavirus prevalence was evidence that a die-off was occurring or imminent. Laboratory studies with ranaviruses have shown that infection and mortality rates typically track each other in highly susceptible species (Brunner et al. 2007; Cunningham et al. 2007b). None of the individuals that we swabbed had gross signs of ranaviral disease (e.g., edema, erythema) and dead salamanders were not observed. However, it is important to note that diseased individuals often do not exhibit external signs and die-offs in the wild are rarely observed (Green et al. 2009). Inasmuch as this is the ﬁrst documentation of ranaviruses infecting lungless salamanders, more research is needed to determine the threat that ranaviruses pose to this diverse group of amphibians. Although positive results from swab specimens can be an indication of virus infection (Driskell et al. 2009), they also may reﬂect virions attached to skin surfaces (Green et al. 2009). In a separate controlled study with tadpoles, we determined that PCR test results for Ranavirus from swabs have about a 10% falsepositive rate for systemic infection when compared with test results from liver tissue (Miller, Gray, and Hoverman, unpubl. FIG. 1. Ranavirus prevalence in lungless salamanders (Plethodontidae) among three sampling locations that differed in elevation, Great Smoky Mountains National Park, Tennessee, USA, April 2007. Bars with unlike letters are different (P = 0.03) by Wald’s chi-square test; standard error bars calculated as: Herpetological Review 40(3), 2009 317 data). We did not euthanize salamanders in our study so it is unknown if positive swabbed individuals had systemic infections. Nonetheless, given the high prevalence of Ranavirus across species, systemic infection of some individuals was possible. Pathogen prevalence differed among sites, with prevalence increasing with decreasing site elevation. Salamanders at 456 m above sea level were 10 times more likely to be infected with Ranavirus than those at 1537 m elev. Greater pathogen prevalence at lower elevation is opposite of the trends reported by previous studies with Ranavirus and Batrachochytrium dendrobatidis (Brem and Lips 2008; Gahl and Calhoun 2008), which have related differences in infection among elevations to changes in temperature. Inasmuch as stream temperature is relatively constant among elevations in the Southern Appalachian Mountains (Swift and Clinton 1997), more optimal temperature for virus replication or temperature-induced stress probably were not factors responsible for higher prevalence at lower elevation. Higher prevalence at lowelevation sites may reﬂect greater human access. All sites could be accessed by trails but the sampling location for Ash Hopper Branch was <200 m from the U.S. National Park Service Sugarlands Visitor Center and a campground was <100 m from the Chimney Tops Seeps. In contrast, the Indian Gap seeps were approximately 400 m from a minor parking lot. Greater human access could increase the likelihood of infection by increasing the level of stressors in the environment or by introducing Ranavirus virions that are attached to footwear or recreational gear (Green et al. 2009). It also is possible that greater infection at low-elevation sites was a result of watershed position. Most of the salamanders that we collected were in or adjacent to ﬂowing streams. Ranavirus virions transmit effectively in water (Harp and Petranka 2006), and they probably remain active outside the host for several months (Langdon 1989). Thus, greater prevalence of Ranavirus at lower elevation may be a consequence of cumulative downstream transport of virions and higher contact rates with the pathogen. The possible impacts of factors associated with elevation and human access on exposure frequency and susceptibility of lungless salamanders to ranaviruses need to be explored. Our results indicate that plethodontid salamanders are suitable hosts for ranaviruses. The pathogenicity of ranaviruses to plethodontid salamanders remains unknown and is a part of ongoing experimental challenges (Gray, Miller, and Hoverman, unpubl. data). Given the high prevalence of Ranavirus in Southern Appalachian salamanders, we recommend that surveillance efforts be expanded in this region and perhaps elsewhere. Future efforts should include isolating and molecularly characterizing ranaviruses infecting plethodontid salamanders to determine if they are phylogenetically similar to previous isolates. Given the possible threat of ranaviruses to lungless salamanders, we recommend that researchers take biosecurity precautions to reduce the likelihood of transporting Ranavirus virions among watersheds. Bryan et al. (2009) reported that 1% Nolvasan®, 1% Virkon S®, and 3% bleach were effective at inactivating Ranavirus after 1 min contact duration. Mud and debris should be removed from footwear and gear prior to applying disinfectant (Green et al. 2009). Acknowledgments.—Our research was supported by the Tennessee Wildlife Resources Agency, the University of Tennessee Institute of Agriculture, and the Tifton Veterinary Diagnostic and Investigational Laboratory of the University of Georgia. All research procedures followed 318 approved University of Tennessee Institutional Animal Care and Use Committee protocol (1712), and ﬁeld activities were authorized by the United States National Park Service (GRSM-2007-SCI-0014) and the Tennessee Wildlife Resources Agency (1990). We thank L. Whittington for assistance with molecular testing, E. C. Burton and A.C. Schmutzer for help with collecting samples, P. E. Super for coordinating sampling logistics in the Great Smoky Mountains National Park, and two anonymous referees for providing comments on initial drafts of the manuscript. LITERATURE CITED BREM, F. M. R., AND K. R. LIPS. 2008. Batrachochytrium dendrobatidis infection patterns among Panamanian amphibian species, habitats and elevations during epizootic and enzootic stages. Dis. Aquat. Org. 81:189–202. BRYAN, L. K., C. A. BALDWIN, M. J. GRAY, AND D. L. MILLER. 2009. Efﬁcacy of select disinfectants at inactivating Ranavirus. Dis. Aquat. Org. 84:89–94. BRUNNER, J. L., D. M. SCHOCK, AND J. P. COLLINS. 2007. Transmission dynamics of the amphibian ranavirus Ambystoma tigrinum virus. Dis. Aquat. Org. 77:87–95. ________ ________ , , E. W. DAVIDSON, AND J. P. COLLINS. 2004. Intraspeciﬁc reservoirs: complex life history and the persistence of a lethal ranavirus. Ecology 85:560–566. CHATFIELD, M. W. H., B. B. ROTHERMEL, C. S. BROOKS, AND J. B. KAY. 2009. Detection of Batrachochytrium dendrobatidis in amphibians from the Great Smoky Mountains of North Carolina and Tennessee, USA. Herpetol. Rev. 40:176–179. COLLINS, J. P., J. L. BRUNNER, J. K. JANCOVICH, AND D. M. SCHOCK. 2004. A model host-pathogen system for studying infectious disease dynamics in amphibians: tiger salamanders (Ambystoma tigrinum) and Ambystoma tigrinum virus. Herpetol. J. 14:195–200. CONVERSE, K. A., AND D. E. GREEN. 2005. Diseases of tadpoles. In S. K. Majumdar, J. E. Huffman, F. J. Brenner, and A. I. Panah (eds.), Wildlife diseases: Landscape epidemiology, spatial distribution and utilization of remote sensing technology, pp. 72–88. Pennsylvania Academy of Science, Easton, Pennsylvania. CUNNINGHAM, A. A., A. D. HYATT, P. RUSSELL, AND P. M. BENNETT. 2007a. Emerging epidemic diseases of frogs in Britain are dependent on the source of ranavirus agent and the route of exposure. Epidemiol. Infect. 135:1200–1212. ________ ________ ________ , , , AND ________. 2007b. Experimental transmission of a ranavirus disease of common toads (Bufo bufo) to common frogs (Rana temporaria). Epidemiol. Infect. 135:1213–1216. ________ , T. E. S. LANGTON, P. M. BENNETT, J. F. LEWIN, S. E. N. DRURY, R. E. GOUGH, AND S. K. MACGREGOR. 1996. Pathological and microbiological ﬁndings from incidents of unusual mortality of the common frog (Rana temporaria). Philos. Trans. R. Soc. Lond. B 351:1539–1557. DASZAK, P., A. A. CUNNINGHAM, AND A. D. HYATT. 2003. Infectious disease and amphibian population declines. Divers. Distrib. 9:141–150. DAVIC, R. D., AND H. H. WELSH. 2004. On the ecological roles of salamanders. Annu. Rev. Ecol. Evol. Syst. 35:405–434. DODD, C. K., JR. 2004. The amphibians of Great Smoky Mountains National Park. University of Tennessee Press, Knoxville, Tennessee. DRISKELL, E. A., D. L. MILLER, S. L. SWIST, AND Z. S. GYIMESI. In press. PCR detection of Ranavirus in adult anurans from the Louisville Zoological Garden. J Zoo Wildlife Med. FORSON, D. D., AND A. STORFER. 2006. Atrazine increases ranavirus susceptibility in the tiger salamander, Ambystoma tigrinum. Ecol. Appl. 16:2325–2332. GAHL, M. K., AND A. J. K. CALHOUN. 2008. Landscape setting and risk of Ranavirus mortality events. Biol. Conserv. 141:2679–2689. GANTRESS, J., G. D. MANIERO, N. COHEN, AND J. ROBERT. 2003. Development and characterization of a model system to study amphibian immune responses to iridoviruses. Virology 311:254–262. Herpetological Review 40(3), 2009 GRAY, M. J., D. L. MILLER, A. C. SCHMUTZER, AND C. A. BALDWIN. 2007. Frog virus 3 prevalence in tadpole populations inhabiting cattle-access and non-access wetlands in Tennessee, USA. Dis. Aquat. Org. 77:97–103. GREEN, D. E., M. J. GRAY, AND D. L. MILLER. 2009. Disease monitoring and biosecurity. In C. K. Dodd (ed.), Amphibian Ecology and Conservation: A Handbook of Techniques. Oxford University Press, Oxford, United Kingdom. ________ , K. A. CONVERSE, AND A. K. SCHRADER. 2002. Epizootiology of sixtyfour amphibian morbidity and mortality events in the USA, 1996–2001. Ann. New York Acad. Sci. 969:323–339. HARP, E. M., AND J. W. PETRANKA. 2006. Ranavirus in wood frogs (Rana sylvatica): potential sources of transmission within and between ponds. J. Wildl. Dis. 42:307–318. KATTENBELT, J. A., A. D. HYATT, AND A. R. GOULD. 2000. Recovery of ranavirus dsDNA from formalin-ﬁxed archival material. Dis. Aquat. Org. 39:151–154. LANGDON, J. S. 1989. Experimental transmission and pathogenicity of epizootic hematopoietic necrosis virus (EHNV) in redﬁn perch, Perca ﬂuviatilis L. and 11 other teleosts. J. Fish. Dis. 12:295–310. MUTHS, E., A. L. GALLANT, E. H. C. CAMPBELL, W. A. BATTAGLIN, D. E. GREEN, J. S. STAIGER, S. C. WALLS, M. S. GUNZBURGER, AND R. F. KEARNEY. 2006. The Amphibian Research and Monitoring Initiative (ARMI): 5-year report. U.S. Geol. Surv. Sci. Investig. Rep. 2006–5224. OIE (WORLD ORGANIZATION FOR ANIMAL HEALTH). 2008. Chapter 2.4.2: Infection with Ranavirus. Aquatic Animal Health Code. http://www. oie.int/eng/en_index.htm (accessed 12 May 2009). PETERMAN, W. E., J. A. CRAWFORD, AND R. D. SEMLITSCH. 2008. Productivity and signiﬁcance of headwater streams: population structure and biomass of the black-bellied salamander (Desmognathus quadramaculatus). Freshwat. Biol. 53:347–357. PETRANKA, J. W. 1998. Salamanders of the United States and Canada. Smithsonian Institution, Washington, D.C. ________ , AND S. S. MURRAY. 2001. Effectiveness of removal sampling for determining salamander density and biomass: a case study in an Appalachian streamside community. J. Herpetol. 35:36–44. PICCO, A. M., J. L. BRUNNER, AND J. P. COLLINS. 2007. Susceptibility of the endangered California tiger salamander, Ambystoma californiense, to Ranavirus infection. J. Wildl. Dis. 43:286–290. ________ , AND J. P. COLLINS. 2008. Amphibian commerce as a likely source of pathogen pollution. Cons. Biol. 22:1582–1589. ROTHERMEL, B. B., S. C. WALLS, J. C. MITCHELL, C. K. DODD, JR., L. K. IRWIN, D. E. GREEN, V. M. VAZQUEZ, J. W. PETRANKA, AND D. J. STEVENSON. 2008. Widespread occurrence of the amphibian chytrid fungus Batrachochytrium dendrobatidis in the southeastern USA. Dis. Aquat. Org. 82:3–18. STOKES, M. E., C. S. DAVIS, AND G. G. KOCH. 2000. Categorical data analysis using the SAS® system. SAS Institute, Cary, North Carolina. STORFER, A., M. E. ALFARO, B. J. RIDENHOUR, J. K. JANCOVICH, S. G. MECH, M. J. PARRIS, AND J. P. COLLINS. 2007. Phylogenetic concordance analysis shows an emerging pathogen is novel and endemic. Ecol. Lett. 10:1075–1083. SWIFT , L. W., JR., AND P. P. CLINTON. 1997. Stream temperature climate in a set of Southern Appalachian streams. In J. E. Cook and B. P. Oswald (eds.), Proceedings of the First Biennial North American Forest Ecology Workshop, pp. 316–335. North Carolina State University, Raleigh, North Carolina. Herpetological Review, 2009, 40(3), 319–321. © 2009 by Society for the Study of Amphibians and Reptiles Amphibian Chytrid Fungus in Western Toads (Anaxyrus boreas) in British Columbia and Yukon, Canada BRIAN G. SLOUGH 35 Cronkhite Road, Whitehorse, Yukon Territory, Y1A 5S9, Canada e-mail: slough@northwestel.net The amphibian chytrid fungus, Batrachochytrium dendrobatidis (Bd), appears to have a patchy pattern of occurrence, particularly at the northernmost extent of its distribution (Aanensen et al. 2009). In northwestern North America, Bd has been detected in Western Toads, Anaxyrus boreas, in southeast Alaska, USA (Adams et al. 2007), the Peace River district in northeast British Columbia, Canada (Raverty and Reynolds 2001), and in southwest British Columbia (Deguise and Richardson 2009). Bd also was found in Wood Frogs, Lithobates sylvaticus, in Kenai National Wildlife Refuge, Alaska (Reeves and Green 2006, Reeves 2008), and in Northern Red-legged Frogs, Rana aurora, from Vancouver Island, British Columbia (Adams et al. 2007). Conversely, Bd was not detected in Columbia Spotted Frogs (Rana luteiventris), Wood frogs, or Western Toads collected at several locations including the Chilkoot Trail National Historic Site, northwest British Columbia (Adams et al. 2007). Likewise, Bd was not found in Wood Frogs in Denali National Park (Chestnut et al. 2008), Innoko National Wildlife Refuge, or Tetlin National Wildlife Refuge, Alaska (Reeves 2008). However, small sample sizes and low prevalence may have affected detection at some sites (see Skerratt et al. 2008). Amphibians within a large portion of the range of Western Toads at its northern limit in northern British Columbia (Matsuda et al. 2006) and the southeast Yukon Territory, Canada (Slough and Mennell 2006) have not been sampled for Bd. To address this gap, I sampled amphibians for Bd from 4 distinct populations in this region in 2007 and 2008, including Tutshi Lake, Atlin Lake and the Nakina River in northwest British Columbia, and the Coal River in the southeast Yukon Territory (Table 1). Amphibians were located using visual encounter surveys and were captured by hand or dip net, which had been cleaned and dried between captures and disinfected with bleach between sites. Each animal was held separately prior to swabbing and was handled with a new pair of vinyl medical examination gloves and swabbed with a sterile 15 cm cotton-tipped swab (AMG Medical Inc., Montréal, QC) 5-times each below the throat, between the toes of the front feet, along the belly, thighs, around the drink patch and cloaca and between the toes of the hind feet. Each swab was air-dried for 5 min and placed tip down in a 1.5 ml microcentrifuge tube. Excess handle-length of the swab was broken off and the tube was sealed and stored frozen. Swabs were analyzed for Bd with a real-time PCR assay at either the British Columbia Ministry of Agriculture, Fisheries and Food, Animal Health Centre (2007 samples) or the University of Victoria, British Columbia (2008 samples) using the technique described by Boyle et al. (2004). In 2007, seven of 20 toads from Atlin Lake and the Nakina River tested positive for Bd, and the Tutshi Lake samples were Bd-negative (Fig. 1). A Wood Frog from Atlin Lake and two Long-toed Herpetological Review 40(3), 2009 319 FIG. 1. Locations in British Columbia and Yukon, Canada where amphibians were sampled for the presence (+) of Batrachochytrium dendrobatidis (Bd) in 2007 and 2008. “–“ indicates locations where Bd was not detected. Salamanders (Ambystoma macrodactylum) from the Nakina River also tested negative. One of the Bd-positive Western Toads from Atlin Lake had sloughing skin on a thigh; otherwise no other individuals had visual signs of the disease such as lethargy or redness on the legs. In 2008, all 10 toad metamorphs from Atlin Lake, an adult toad, and six Wood Frogs from the Coal River, Yukon Territory, tested Bd-positive. Western Toad tadpoles from the Coal River were Bd-negative. The pattern of patchy Bd occurrence at high northern latitudes is supported by these additional data from northern British Columbia and Yukon Territory. Bd was present in three Western Toad populations sampled in northwestern British Columbia and Yukon Territory but was not detected at Tutshi Lake (Fig. 1), the northwestern-most site surveyed, which coincidentally is only 10 km east of the Chilkoot Trail National Historic Site where amphibians previously sampled also were Bd-negative (Adams et al. 2007). There is concurrent Western Toad breeding site monitoring at one of the study sites, the Atlin Warm Springs, at Atlin Lake, where there were eggs, larvae or metamorphs present demonstrating breeding activity in survey years 1988, 1996, 1998, 1999, 2001, 2005 and 2008, but no breeding activity or adults were observed in 2006, 2007 and 2009 (B. Slough, unpubl. data). This is a small population, with no more than 25 adults and 8 egg clutches observed at one time, however it has demonstrated long-term persistence. Other observations of toads at this site were made in 1924 (Slevin 1928; MVZ 9478–9483, Museum of Vertebrate Biology, University of California, Berkeley), 1952 (Cook 1977), 1962 (RBCM 1273, Royal British Columbia Museum, Victoria), 1979 (CMNAR 21852, Canadian Museum of Nature, Ottawa), 1980 (RBCM 1389) and 1993 (Mennell 1997). These data indicate that the breeding population may have suffered recent losses, and although Bd may be contributing to this, several alternative or interacting factors warrant consideration (e.g., climate change; Slough 2009), especially due to the location of this area at the margin of the species’ range. The general status rank of the Western Toad is Sensitive in British Columbia and the Yukon (Canadian Endangered Species Conservation Council 2006), and it is protected under the federal Species at Risk Act after being designated Special Concern (COSEWIC 2002). The effect of Bd on northern amphibian populations, especially TABLE 1. Incidence of Batrachochytrium dendrobatidis (Bd) in amphibians sampled in northwest British Columbia and Yukon, Canada. The species examined were Anaxyrus boreas (ANBO), Lithobates sylvaticus (LISY), and Ambystoma macrodactylum (AMMA). Location Date Latitude °N, Longitude °W Positive Bd Samples Negative Bd Samples Tutshi River uplands 20 June 2007 59.772, -134.931 0 1 juvenile ANBO Tutshi Lake 3 June 2007 59.817, -134.799 0 8 adult ANBO Atlin Lake, Torres Channel 23 June 2007 59.398, -133.951 1 adult ANBO 0 Atlin Lake, near Grifﬁth Island 23 & 28 June 2007 59.286, -133.824 59.284, -133.823 2 adult ANBO 1 adult ANBO Atlin Lake, 12-Mile Point, Teresa Island 28 June 2007 59.436, -133.695 2 juvenile 1 adult ANBO 1 adult ANBO Atlin Lake, Warm Springs Homestead 19 April 2007 59.406, -133.579 0 1 adult ANBO Atlin Lake, Warm Springs Homestead 26 June 2007 59.406, -133.579 0 1 adult LISY Atlin Lake, Warm Springs 10 May 2008 59.404, -133.573 10 metamorph ANBO 0 Nakina River, near conﬂuence with Sloko River 1 July 2007 59.011, -133.141 59.008, -133.151 1 adult ANBO 1 adult ANBO 2 adult AMMA Coal River Springs and Coal River 23–24 July 2008 60.156, -127.435 1 adult ANBO 6 adult LISY 30 ANBO larvae 320 Herpetological Review 40(3), 2009 Western Toads, warrants further investigation. Acknowledgments.—I thank L. Friis, S. Leaver, and P. Govindarajulu ofEcosystems Branch, British Columbia Ministry of Environment, Victoria, and T. Jung of the Fish and Wildlife Branch, Yukon Department of Environment, Whitehorse, for their support. Samples were obtained with the assistance of S. Badhwar, K. Slough, P. Kukka and C. Lausen. M. Connor, Habitat Steward, Taku River Tlingit First Nation, Atlin, BC coordinated sampling on the Nakina River by D. Milek, D. Adler, T. McCrea, C. Peacock, W. Tyson, S. Cota and A. Kaplan. D. Olson and an anonymous reviewer kindly provided comments on the manuscript. LITERATURE CITED AANENSEN, D. M., B. G. SPRATT, AND M.C. FISHER. 2009. Bd-Maps. Imperial College, London, UK. Available at: www.spatialepidemiology. net/Bd-maps/. (accessed 18 May 2009). ADAMS, M. J., S. K. GALVAN, D. REINITZ, R. A. COLE, S. PAYRE, M. HAHR, AND P. GOVINDARAJULU. 2007. Incidence of the fungus, Batrachochytrium dendrobatidis, in amphibian populations along the northwest coast of North America. Herpetol. Rev. 38:430–431. BOYLE, D. G., D. B. BOYLE, V. OLSEN, J. A. T. MORGAN, AND A. D. HYATT. 2004. Rapid quantitative detection of chytridiomycosis (Batrachochytrium dendrobatidis) in amphibian samples using real-time Taqman PCR assay. Dis. Aquat. Org. 60:141–148. CANADIAN ENDANGERED SPECIES CONSERVATION COUNCIL . 2006. Wild Species 2005: The general status of species in Canada. Available at: http://www. wildspecies.ca/wildspecies2005/. (accessed 18 May 2009). CHESTNUT, T., J. E. JOHNSON, AND R. S. WAGNER. 2008. Results of amphibian chytrid sampling in Denali National Park, Alaska, USA. Herpetol. Rev. 39:202–204. COOK, F. R. 1977. Records of the boreal toad from the Yukon and northern British Columbia. Can. Field-Nat. 91: 185–186. COSEWIC. 2002. COSEWIC assessment and status report on the western toad Bufo boreas in Canada. Committee on the Status of Endangered Wildlife in Canada. Ottawa. vi + 31 pp. DEGUISE, I., AND J. S. RICHARDSON. 2009. Prevalence of the chytrid fungus (Batrachochytrium dendrobatidis) in western toads in southwestern British Columbia, Canada. Northwest. Nat. 90:35–38. MATSUDA, B. M., D. M. GREEN, AND P. T. GREGORY. 2006. Amphibians and reptiles of British Columbia. Royal BC Museum Handbook, Victoria, British Columbia. 266 pp. MENNELL, L. 1997. Amphibians in southwestern Yukon and northwestern British Columbia. Herpetol. Conserv. 1:107–109. RAVERTY, S., AND T. REYNOLDS. 2001. Cutaneous chytridiomycosis in dwarf aquatic frogs (Hymenochirus boettgeri) originating from Southeast Asia and in a western toad (Bufo boreas) from northeastern British Columbia. Can. Vet. J. 42:385–386 REEVES, M. K. 2008. Batrachochytrium dendrobatidis in wood frogs (Rana sylvatica) from three national wildlife refuges in Alaska, USA. Herpetol. Rev. 39:68–70. ________ , AND D. E. GREEN. 2006. Rana sylvatica (wood frog): chytridiomycosis. Herpetol. Rev. 37:450. SKERRATT, L. F., L. BERGER, H. B HINES, K. R. MCDONALD, D. MENDEZ, AND R. SPEARE. 2008. Survey protocol for detecting chytridiomycosis in all Australian frog populations. Dis. Aquat. Org. 80:85–94. SLEVIN, J. R. 1928. The amphibians of western North America. an account of the species known to inhabit California, Alaska, British Columbia, Washington, Oregon, Idaho, Utah, Nevada, Arizona, Sonora, and Lower California. Occas. Pap., California Acad. Sci. 16:1–152. SLOUGH, B. G. 2009. Yukon Territory. In D. H. Olson (coord. ed.), Herpetological Conservation in Northwestern North America. Northwest. Nat. 90: in press. ________ , AND R. L. MENNELL. 2006. Diversity and range of amphibians of the Yukon Territory. Can. Field-Nat. 120:87–92. Herpetological Review, 2009, 40(3), 321–323. © 2009 by Society for the Study of Amphibians and Reptiles Chytridiomycosis-Associated Mortality in a Rana palustris Collected in Great Smoky Mountains National Park, Tennessee, USA MEGAN TODD-THOMPSON University of Tennessee, Department of Ecology and Evolutionary Biology 569 Dabney Hall, Knoxville, Tennessee 37996, USA DEBRA L. MILLER* University of Georgia, Veterinary Diagnostic and Investigational Laboratory 43 Brighton Road, Tifton, Georgia 31793, USA e-mail: millerdl@uga.edu PAUL E. SUPER Appalachian Highlands Science Learning Center Great Smoky Mountains National Park, P.O. Box 357 Lake Junaluska, North Carolina 28745, USA and MATTHEW J. GRAY University of Tennessee, Center for Wildlife Health 274 Ellington Plant Sciences Building, Knoxville, Tennessee 37996, USA *Corresponding author Batrachochytrium dendrobatidis (Bd) has been associated with global amphibian population declines and occasionally with species extinctions (Daszak et al. 1999). Further, recent evidence suggests that surviving amphibian communities may persist with sublethal infections (Rettalick et al. 2004). Although Bd has been detected in amphibians in Great Smoky Mountains National Park (GSMNP; Chatﬁeld et al. 2009), USA, mortality associated with chytridiomycosis has not been reported. The Southern Appalachian Mountain Range, where GSMNP occurs, is known for its amphibian species richness (Dodd 2004). Within the GSMNP, Cades Cove, Tennessee, is one of the most highly visited sites by tourists, and thus an area where anthropogenic stressors (e.g., introduction of chemical agents or novel pathogens) may impact native species. Although amphibian mortality was observed in Cades Cove from 1999 to 2001, these mortality events were attributed to pathogens other than Bd (Dodd 2004; Rothermel et al. 2008). On 5 April 2009, a recently deceased adult male Rana palustris was found in a pond in Cades Cove (35.606667ºN, 83.816944ºW) while conducting ﬁeld surveillance for Ranavirus. The animal was collected and shipped overnight on ice to the University of Georgia, Veterinary Diagnostic and Investigational Laboratory for diagnostic evaluation. Gross examination revealed only mild (ca. 48 h under refrigeration post ﬁeld collection) postmortem autolysis. Grossly there was diffuse erythema (reddening) on the ventromedial legs and random petechia (pin-point hemorrhages), ecchymosis (red blotches), and irregular patches of discoloration elsewhere on the ventral body (Fig. 1A). Vascular congestion was noted in the kidneys but other organs appeared normal. Sections of all organs were collected and ﬁxed in 10% phosphate-buffered formalin, embedded in parafﬁn, cut at 4 μm sections, stained with hematoxylin and eosin, and examined microscopically. Additionally, sections of skin lesions and toes were collected for qPCR and conventional PCR for Bd, and sections of skin lesions, liver and kidney were collected for Herpetological Review 40(3), 2009 321 FIG. 1. A) Gross changes observed in Rana palustris found dead in Great Smoky Mountains National Park, Tennessee, USA. There is diffuse erythema on the ventromedial legs (black arrowheads) and random petechia, ecchymosis (white arrow), and irregular patches of discoloration (black arrows) elsewhere on the ventral body. The white arrowhead indicates an area of postmortem tissue sampling. B) Photomicrograph of a hematoxylin and eosin (H&E) stained section of skin from the frog showing epidermal proliferation. The area to the left of the arrow is more proliferative and the area to the right is more normal. C) Closer view of the proliferative skin area in B (H&E stain), showing the organisms consistent with Batrachochytrium dendrobatidis zoospores (arrows). D) Photomicrograph of an H&E stained section of skeletal muscle subjacent to the skin lesions, and showing swollen ﬁbers that had lost cross-striations and were often fragmented (arrows), and occasional contraction bands (arrowheads). E) Gel electrophoresis of the PCR products isolated from skin and toe samples collected from the frog pictured in A. The 300-bp products were consistent with Bd. Lanes 1 and 2 are Frog Sample 1, Lanes 3 and 4 are Frog Sample 2, Lane 5 is the Positive Bd Control, Lane 6 is the Negative Tissue Control, and 7 is Negative Control, Lane 8 is the molecular weight marker. qPCR and conventional PCR for Ranavirus. Chytrid qPCR was performed using the method described by Boyle et al. (2004), and conventional PCR was performed following the protocol by Annis et al. (2004). Ranavirus qPCR was performed using the protocol by Picco et al. (2007), and conventional PCR was performed following the protocol by Mao et al. (1997). Histologically, the epidermis was multifocally thickened (2–4 cell layers; Fig. 1B) with intracytoplasmic 3–5 μm round organisms (Fig. 1C) consistent with Bd organisms (thalli). This type of lesion was consistently found in the epidermis overlying the digits and in the skin lesions observed grossly. The skeletal muscles subjacent to the skin lesions had swollen ﬁbers and patches of 322 myoﬁber disarray, fragmentation, and loss of cross-striations (Fig. 1D). Conventional PCR was positive for Bd (Fig. 1E), but negative for Ranavirus. Likewise, qPCR was positive for Bd, but negative for Ranavirus. The histological and molecular ﬁndings were consistent with chytridiomycosis caused by the amphibian pathogen Bd. Sensitivity to Bd varies by amphibian species (Blaustein et al. 2005), and may be partially attributed to differences in skin antimicrobial properties (Woodhams et al. 2004). Ultimately, chytridiomycosis is believed to interfere with osmoregulation (Voyles et al. 2007), which leads to muscular degeneration and eventually death (Miller et al. 2008). Other factors (e.g., environmental contaminants, Herpetological Review 40(3), 2009 climate change) are hypothesized to exacerbate the effects of or make amphibians more susceptible to this pathogen (Bosch et al. 2007; Davidson et al. 2007). We are uncertain what factors contributed to this R. palustris developing chytridiomycosis. Rothermel et al. (2008) found that 23 of 35 R. palustris collected from the southeastern United States were positive for Bd; however, chytridiomycosis was not found. In a Canadian survey, Bd prevalence in R. palustris was found to be low (3%; Ouellet et al. 2005). Given the ﬁndings from these studies and the lack of detection of other pathological changes in the R. palustris in our study, an exogenous stressor or combination of stressors (e.g., chemical exposure, habitat change, breeding) may have contributed to the development of chytridiomycosis in this case. The ﬁnding of chytridiomycosis in this R. palustris veriﬁes the need for continued surveillance and testing of amphibian populations in the GSMNP. Given the wealth of amphibian biodiversity in the GSMNP, this location serves as an ideal place to study the variation in pathogen prevalence and species susceptibility of amphibian populations. The co-occurrence of two emerging amphibian pathogens (Ranavirus [Dodd 2004] and Bd) in Cades Cove warrants continued monitoring of amphibian breeding ponds in this highly visited area of GSMNP. Acknowledgments.—We thank Ben Dyer for assistance in ﬁeld collections, and the staff of the University of Georgia Veterinary Diagnostic and Investigational Laboratory for assistance in tissue processing, especially Lisa Whittington, Diane Rousey, and Kim Bridges. This study was conducted with permission from the National Park Service (#GRSM-2008SCI-0056) and an approved University of Tennessee animal care and use protocol (#1763). Funding for ﬁeld collection was provided by the Ecology and Evolutionary Biology Department at the University of Tennessee. DODD, C. K. 2004. The Amphibians of Great Smoky Mountains National Park. University of Tennessee Press, Knoxville. 284 pp. MAO, J. D., R. P. HEDRICK, AND V. G. CHINCHAR. 1997. Molecular characterization, sequence analysis, and taxonomic position of newly isolated ﬁsh iridoviruses. Virology 229:212–220. MILLER, D. L., S. RAJEEV, M. BROOKINS, J. COOK, L. WHITTINGTON, AND C. A. BALDWIN. 2008. Concurrent infection with Ranavirus, Batrachochytrium dendrobatidis, and Aeromonas in a captive anuran colony. J. Zoo Wildl. Med. 39:445–449. OUELLET, M., I. MIKAELIAN, B. D. PAULI, J. RODRIGUE, AND D. M. GREEN. 2005. Historical evidence of widespread chytrid infection in North American amphibian populations. Conserv. Biol. 19:1431–1440. PICCO, A. M., J. L. BRUNNER, AND J. P. COLLINS. 2007. Susceptibility of the endangered California tiger salamander, Ambystoma californiense, to Ranavirus infection. J. Wildl. Dis. 43:286–290. RETTALICK, R.W., H. MCCALLUM, AND R. SPEARE. 2004. Endemic infection of the amphibian chytrid fungus in a frog community post-decline. PLoS. Biol. 2(11):e351. ROTHERMEL, B. B., S. C. WALLS, J. C. MITCHELL, C. K. DODD, L. K. IRWIN, D. E. GREEN, V. M. VAZQUEZ, J. W. PETRANKA, D. J. STEVENSON. 2008. Widespread occurrence of the amphibian chytrid fungus Batrachochytrium dendrobatidis in the southeastern USA. Dis. Aq. Org. 82:3–18 WOODHAMS, D. C., L. A. ROLLINS-SMITH, C. CAREY, L. REINERT, M. J. TYLER, R. A. ALFORD. 2006. Population trends associated with skin peptide defenses against chytridiomycosis in Australian frogs. Oecologia 146:531–540. VOYLES, J., L. BERGER, S. YOUNG, R. SPEARE, R. WEBB, J. WARNER, D. RUDD, R. CAMPBELL, L. F. SKERRATT. 2007. Electrolyte depletion and osmotic imbalance in amphibians with chytridiomycosis. Dis. Aq. Org. 77(2):113–118. LITERATURE CITED ANNIS, S. L., F. P. DASTOOR, H. ZIEL, P. DASZAK, AND J. E. LONGCORE. 2004. A DNA-based assay to identify Batrachochytrium dendrobatidis in amphibians. J. Wildl. Dis. 40:420–428. BLAUSTEIN, A. R., J. M. ROMANSIC, E. A. SCHEESSELE, B. A. HAN, A. P. PESSIER, AND J. E. LONGCORE 2004. Interspecific variation in susceptibility of frog tadpoles to the pathogenic fungus Batrachochytrium dendrobatidis. Conserv. Biol. 19:1460–1468. BOSCH, J., L. M. CARRASCAL, L. DURAN, S. WALKER, AND M. C. FISHER. 2007. Climate change and outbreaks of amphibian chytridiomycosis in a montane area of Central Spain; is there a link? Proc. R. Soc. London Biol. Sci. 274:253–260. BOYLE, D. G., D. B. BOYLE, V. OLSEN, J. A. MORGAN, AND A. D. HYATT. 2004. Rapid quantitative detection of chytridiomycosis (Batrachochytrium dendrobatidis) in amphibian samples using real-time Taqman PCR assay. Dis.Aq. Org. 60:141–148. CHATFIELD, M. W. H., B. B. ROTHERMEL, C. S. BROOKS, AND J. B. KAY. 2009. Detection of Batrachochytrium dendrobatidis in amphibians from the Great Smoky Mountain of North Carolina and Tennessee, USA. Herpetol. Rev. 40:176–179. DASZAK, P., L. BERGER, A. A. CUNNINGHAM, A. D. HYATT, D. E. GREEN, AND R. SPEARE. 1999. Emerging infectious diseases and amphibian population declines. Emerg. Infect. Dis. 5:735–748. DAVIDSON, C., M. F. BENARD, H. B. SHAFFER, J. M. PARKER, C. O’LEARY, J. M. CONLON, AND L. A. ROLLINS-SMITH. 2007. Effects of chytrid and carbaryl exposure on survival, growth, and skin peptide defenses in foothill yellow-legged frogs. Environ. Sci. Technol. 41:1771–1776. Anolis fraseri (Polychrotidae). Reserva Forestal Bosque de Yotoco, Department of Valle del Cauca, Colombia. Illustration by Fernando Vargas Salinas, based on a photograph by FVS. Herpetological Review 40(3), 2009 323 HERPETOLOGICAL HUSBANDRY Herpetological Review, 2009, 40(3), 324–330. © 2009 by Society for the Study of Amphibians and Reptiles Nutrient Composition of Whole Crayﬁsh (Orconectes and Procambarus Species) Consumed by Hellbender (Cryptobranchus alleganiensis) ELLEN S. DIERENFELD* Department of Animal Health and Nutrition Saint Louis Zoo, St. Louis, Missouri 63110, USA KEVIN J. MCGRAW School of Life Sciences, Arizona State University Tempe, Arizona 85287, USA KEVIN FRITSCHE Division of Animal Sciences, University of Missouri Columbia, Missouri 65211, USA JEFFREY T. BRIGGLER Missouri Department of Conservation Jefferson City, Missouri 65102, USA and JEFF ETTLING WildCare Institute, Saint Louis Zoo St. Louis, Missouri 63110, USA *Current Address and corresponding author: Novus International, Inc., St. Charles, Missouri 63304, USA e-mail: edierenfeld@aol.com Hellbenders (Cryptobranchus alleganiensis) are one of the largest salamanders in North America, averaging 29–51 cm in total length, but reaching up to 70 cm (Nickerson and Mays 1973; Conant and Collins 1998; Johnson 2000). The Eastern Hellbender (C. a. alleganiensis) ranges from New York State south to Georgia, and west to Missouri, whereas the Ozark subspecies (C. a. bishopi) is only found in a few locations in south central Missouri and northern Arkansas (Firschein 1951; Pﬁngsten 1990; Petranka 1998). Serious population declines have targeted the Ozark Hellbender as a conservation priority species by numerous institutions (e.g. government agencies, universities, zoos, etc.). These institutions have spent considerable effort to recover the species in Missouri (Wheeler et al. 2003; Briggler et al. 2007). Due to the drastic decline in number of wild Hellbenders in Missouri, captive propagation was one necessary component to the recovery of the species in which the Ron Goellner Center for Hellbender Conservation at the Saint Louis Zoo was created. Numerous Hellbenders were removed from the wild and moved to their new artiﬁcial stream raceway to facilitate long-term propagation efforts. As with many propagation efforts, various aspects of Hellbender biology are being investigated in detail at the St. Louis Zoo, including ecological nutrition and digestive physiology, to better address long-term health needs of this captive breeding stock. Hellbenders are fully aquatic, spending most of their life under ﬂat rocks in stream bottoms (Nickerson and Mays 1973; Johnson 2000). Diet consists of a variety of aquatic prey including small ﬁsh and insects, but a majority of the diet comprises 324 whole crayﬁsh, consumed in their entirety (Nickerson and Mays 1973; Peterson et al. 1989; Petranka 1998). The primary purpose of this study was two-fold: 1) to obtain details on a preferred food item of Hellbenders, for which no nutritional composition data exist (crayﬁsh plus shell) as a basis for assessing nutritional status in the wild; and 2) to compare native food composition with substitute food items used in captivity. A secondary goal was to investigate possible factors that may underlie any differences noted in chemical composition. These initial data might provide useful information for optimizing nutritional health and dietary management of Hellbender and other crayﬁsh-eating species (e.g., storks; Negro and Garrido-Fernandez 2000). Samples.—Three species of native crayﬁsh were used in this study. Long-pincered Crayﬁsh (Orconectes longidigitus), Spothanded Crayﬁsh (O. punctimanus), and Ringed Crayﬁsh (O. neglectus) were opportunistically collected from two locations in close proximity on the North Fork of the White River, Missouri, USA in August 2006 and September 2007 (N = 21 total samples). Crayﬁsh were transported in buckets of stream water and maintained overnight before being identiﬁed to species, weighed, measured, and processed for nutrient analysis. Three individual crayﬁsh were not positively identiﬁed and were included in samples as unidentiﬁed. Feeder crayﬁsh obtained locally in 2005 and 2006 comprised a single species, Northern Crayﬁsh (Orconectes virilis), from Paul’s Bait Shop (PBS, N = 9; St. Louis, Missouri, USA) or directly from Ozark Fisheries (OF, N = 7; Stoutland, Missouri USA). In 2007, Procambarus paeninsulanus (Peninsula Crayﬁsh) from Florida (FL, N = 9) were added as a substitute feeder species, and O. virilis were also collected for analysis from ponds on the grounds of the Saint Louis Zoo as part of a larger population study for potential sustainable harvest (SLZ, N = 21). SLZ crayﬁsh of three body sizes (see below) were sampled from two sites (North and South Lakes) during either summer (Jun–Aug) or fall (Sep–Oct) 2007. Purchased crayﬁsh were shipped or transported in water overnight, and processed within 24 h of receipt. Crayﬁsh acquired on zoo grounds were collected in the morning from traps set the previous day using canned cat food as bait, and sampled within 24 h. Because Hellbenders consume crayﬁsh in toto, an additional subset of trapped crayﬁsh from SLZ (N = 24) were dissected to determine percent contribution of nutrients from shell vs. remainder of the body by weighing and analyzing portions separately, and water and substrate samples were taken for chemical analysis at the two sites. Laboratory Processing/Analyses.—Animals were weighed to the nearest 0.1 g, and carapace and tail-to-rostrum length (TR) measured to the nearest 0.01 cm with calipers. Crayﬁsh were categorized as small (<2.5 cm total length), medium-sized (2.5 to 5 cm total length), or large (>5 cm). All crayﬁsh were euthanized by decapitation, then head and body were homogenized together in a food processor (Toastmaster Chopster). Subsamples of homogenate (0.5 g in duplicate) were taken immediately and extracted via previously described methods for meat samples (Barker et al. 1998) to determine vitamin A (as retinol), E (as αtocopherol), and total carotenoids. Extracts were sealed in cryovials, stored in an ultracold freezer (-20°C), and sent to Arizona State University for analysis via HPLC following the methods Herpetological Review 40(3), 2009 of McGraw et al. (2006). Pigment extracts were resuspended in 200 μl mobile phase (42:42:16% (v/v/v) methanol : acetonitrile : dichloromethane) and injected into a Waters Alliance 2695 HPLC system (Waters Corporation, Milford, MA) ﬁtted with a Waters YMC Carotenoid 5.0 μm column (4.6 mm × 250 mm) and a builtin column heater set at 30°C. A three-step gradient solvent system was used to analyze both xanthophylls and carotenes in a single run, at a constant ﬂow rate of 1.2 ml min-1: ﬁrst, isocratic elution with the aforementioned mobile phase for 11 min, followed by a linear gradient up to 42:23:35% (v/v/v) methanol : acetonitrile : dichloromethane through 21 min, held isocratically at this condition until 26 min, and ﬁnishing with a return to the initial isocratic condition from 26 to 29.5 min. Data were collected from 250 to 600 nm using a Waters 2996 photodiode array detector. Pigments were identiﬁed by comparing their respective retention times and absorbance maxima (λmax) to those of reference carotenoids run as external standards. Subsamples (1.0 g) from 10 randomly-selected farmed crayﬁsh (Orconectes virilis) were frozen for determination of fatty acid content at the University of Missouri-Columbia; a frozen Paciﬁc Krill (Euphasia paciﬁca) sample was also sent for comparison since krill are often used in feeding very small salamanders (J. Ettling, pers. comm.). Total lipids were extracted with chloroform: methanol (2:1 v/v) after samples were homogenized in 10 mM EDTA. Lipids were methylated with freshly made 5% methanolic HCl. Fatty acid methyl esters were extracted with hexane. Pigments and residual water were removed from each sample with the addition of 0.1 g each of anhydrous sodium sulfate and activated charcoal. Methyl esters were analyzed by gas chromatography using an HP 5890A instrument with a 30-meter capillary column (Omegawax 250; Supelco, Bellefonte, PA) according to the methods of Sukhija and Palmquist 1988. Individual fatty acids were identiﬁed using retention times of standards (i.e., Omegawax, PUFA-II, PUFA-I; Supelco). Remaining sample homogenates were freeze-dried (Labconco Freezone 4.5) to determine water content, and dried, ground samples were sent for proximate analysis (crude protein, crude fat, ﬁber fractions, ash) and minerals to Dairy One Forage Lab (Ithaca, NY). Substrate mineral content was determined at the Amino Acid Laboratory, University of California (Davis, California; data not presented). Differences in nutrient composition by species (stream-caught) or source/location within species (pondreared) were evaluated with Mann-Whitney tests at a signiﬁcance level of 0.05 (Snedecor and Cochran 1967). For the STZ crayﬁsh samples only, percentage data were arcsine transformed to approximate normal distribution, and seasonal, body size, and sampling location differences in nutrient composition were compared by analysis of variance. Wild-captured crayﬁsh sampled from Hellbender habitats varied widely in size compared to those utilized as food for captive salamanders at the Saint Louis Zoo (Table 1). Regardless of species, size ranges encompassed the small and medium body size criteria preferred by zoo managers. Speciﬁc preferences of crayﬁsh prey size have not been determined for Hellbenders; presumably crayﬁsh must be suitably sized to trigger a feeding response, yet not too large to defend themselves against ingestion. Although similar in carapace length to those obtained from commercial suppliers, Northern Crayﬁsh from the zoo ponds in this study contained ash-free dry weight (AFDW) similar to that reported from wild-caught Virginia O. virilis (~3.7 g vs. 3.8 g, respectively; Mitchell and Smock 1991) and almost twice the AFDW calculated for the ﬁsheries-reared crayﬁsh (~2.0 g from PBS, ~1.5 g from OF). Body sizes did not differ between O. virilis samples collected for dissection at the two STZ sites, hence data were combined (mean weight = 25.2 ± 5.4 g). Of total body composition, shell comprised 61.5 ± 5.7% of whole body mass, whereas meat and body ﬂuids reﬂected the remaining 38.5 ± 5.7% (wet basis). Shells averaged 31.2 ± 4.3% dry matter (DM) (or 68.8% water), and meat averaged 17.8 ± 2.1% DM, approximately 82.2% water. Proximate Composition.—Overall, crayﬁsh provided crude protein concentrations in excess of the requirements determined for domestic carnivores, ﬁsh and shellﬁsh (ca. 15 to ~40% of DM; Zootrition 2006). Protein and fat, as fractions of whole body mass, are typically inversely related when evaluating composition of whole vertebrate prey. This pattern was not apparent in crayﬁsh, possibly due to the much greater contribution of both ash and carbohydrate fractions (from shells) in crayﬁsh compared to vertebrate prey species. Total ash reported in whole vertebrate prey DM ranges from 7.5 (juvenile hamsters) to 17.5% (snowshoe hare) as opposed to 30–45% in crayﬁsh, whereas carbohydrate content is generally negligible (Dierenfeld et al. 2002). Additionally, the crude fat content of most whole prey (range 5–45% of DM, but typically 15–35%, depending on species and age of prey; Dierenfeld et al. 2002) is considerably higher than recommended minimum dietary levels (~5 to 10% of dry matter, DM) for domestic carnivores (NRC 2006). Crayﬁsh contained some of the lowest fat concentrations measured across a wide variety of whole prey consumed by carnivores, with both wild-caught and farmed crayﬁsh averaging <5% fat (DM basis, Tables 2 and 3). This may be important for overall energetics and nutritional balance, as amphibians and other poikilotherms have a lower metabolic rate than homeotherms, hence require fewer calories per unit body mass. Thus low-fat, lower nutrient density dietary ingredients may be entirely appropriate for Hellbenders, but actual nutritional requirements remain uncharacterized for amphibians and most reptiles. As with all species—but perhaps particularly so for low-energy amphibian species—the feeding of higher-fat dietary ingredients in captive management programs should be carefully controlled to minimize risk of obesity or health conditions associated with excess body condition. Nonetheless, some fat, and particularly essential fatty acids (EFAs) are required in the diet, hence our analysis provides baseline information on fatty acid content of whole crayﬁsh (Table 4). Fatty acids are the primary constituent of most lipids, and can be deposited in animal tissues with minimal modiﬁcation from diets; fatty acid signatures are often used to differentiate and understand trophic interactions in aquatic ecosystems (Iverson et al. 2002). Although the essential fatty acid (EFA) requirements of salamanders have not been determined to our knowledge, minimum dietary recommendations of domestic carnivores for two of the omega-6 EFAs, linoleic acid (18:2n6) and arachidonic (20:4n6), have been set at 0.5 and 0.2% of dietary DM, respectively (NRC 2006). Crayﬁsh in this study contained 9% of total FA as linoleic, and 6.5% as arachidonic acid (see Table 4). Converting FA data reported here to a comparable dietary DM basis yields 0.3% Herpetological Review 40(3), 2009 325 TABLE 1. Size (mean ± SD; range in parentheses) parameters of crayfish sampled to determine nutrient composition of whole crayfish available as food to Ozark Hellbender (Cryptobranchus alleganiensis bishopi). nd = not determined. Species N Mass (g) Carapace Length (cm) Tail–Rostrum Length (cm) Longpincered Crayﬁsh (Orconectes longidigitus) 7 53.20 ± 20.33 (14.54–75.14) 8.00 ± 3.79 (4.6–13.6) 11.22 ± 2.04 (8.00–13.40) Ringed Crayﬁsh (Orconectes neglectus) 7 10.80 ± 3.45 (3.18–14.56) 3.00 ± 0.48 (2.23–3.37) 6.19 ± 1.01 (4.53–7.17) Spothanded Crayﬁsh (Orconectes punctimanus) 4 23.12 ± 6.31 (13.62–31.04) 5.96 ± 2.91 (3.57–9.70) 7.97 ± 0.47 (7.60–8.50) Unidentiﬁed (Orconectes species) 3 8.35 ± 5.33 (3.95–14.27) 2.69 ± 0.55 (2.11–3.20) 5.46 ± 1.11 (4.26–6.43) Northern Crayﬁsh - Local bait shop (Orconectes virilis) 9 12.11 ± 3.56 (6.19–16.69) 7.68 ± 0.82 (6.0–8.9) nd Northern Crayﬁsh - Ozark Fisheries (Orconectes virilis) 7 9.64 ± 2.13 (5.42–16.88) 7.14 ± 0.50 (6.0–8.4) nd Northern Crayﬁsh - SLZ ponds (Orconectes virilis) 21 21.04 ± 8.77 (7.74–38.64) 7.39 ± 2.27 (3.93–18.19) 10.41 ± 2.71 (6.88–15.00) Peninsula Crayﬁsh - Florida (Procambarus paenenisulanus) 9 6.13 ± 2.85 (2.92–9.66) 2.94 ± 0.83 (2.10–4.83) 5.55 ± 0.79 (4.26–6.43) Wild-Caught Farmed linoleic acid (3% crude fat (Table 3) * 0.09), and 0.2% arachidonic (3% fat *0.065). Assuming that amphibian carnivores have similar fatty acid requirements as domestic animals, and since fat has ~2 times the caloric value of carbohydrates or protein, crayﬁsh would provide about 0.5% of dietary energy as linoleic acid, a value that would be considered borderline for meeting EFA requirements in the absence of any preformed arachidonic acid (AA). However, the presence of 0.2% AA determined in the crayﬁsh should be sufﬁcient to meet the requirements for omega6 polyunsaturated FA for most animals that might be consuming crayﬁsh as a primary food source. Marine krill, however, appear to provide insufﬁcient levels of both EFAs (Table 4). Both whole crayﬁsh and whole krill contained relatively high concentrations of polyunsaturated fatty acids (PUFA) (22–32% of total fatty acids as PUFA). PUFA can be more prone to oxidative degradation compared with more saturated fats, thus may require higher levels of dietary antioxidants. By comparison, saturated fats were much higher in the marine krill (45.4%) compared with fresh water crayﬁsh (32.4%), comprising almost half of total fatty acids quantiﬁed. Additionally, the omega-3:omega-6 fatty acid ratios measured in crayﬁsh compared with krill vary considerably (Table 4), with omega-6 fatty acids about 10-fold higher in crayﬁsh compared with krill; these two feed types are not nutritional equivalents. In particular, omega-3:omega-6 ratios/composition may have health implications for captive feeding programs of Hellbenders, particularly regarding immune function (Fritsche 2006), which should be investigated further. Interestingly, meat from farmed crayﬁsh contained higher concentrations of all fatty acids compared with meat from wild crayﬁsh reported in the USDA Nutrient Database (USDA 2009), further suggesting that 326 captive dietary management may certainly inﬂuence composition, and should be considered when raising species as food for other animals. Regarding carbohydrate (CHO), fractions, while most vertebrate whole prey are not considered to contain substantial concentrations of simple sugars, dietary ﬁber (measured as crude ﬁber, acid detergent ﬁber, and/or neutral detergent ﬁber), may be measurable in gastrointestinal contents of herbivorous prey species. In the case of crayﬁsh, the chitinous exoskeleton is a chemical matrix comprising polysaccharide units (similar to plant cellulose) and protein. It is unclear whether the exoskeleton provides a source of dietary nutrients to Hellbenders; chitinase enzyme activity has not been reported, and digestibility trials have not been performed. Hellbenders have the ability to regurgitate exoskeleton fragments—identiﬁcation of such samples is a technique routinely used for feeding ecology studies (Peterson et al. 1989). The dietary ﬁber (CHO) measured in crayﬁsh samples (~14 to 24–30% of DM, depending on fraction considered from Tables 2 and 3) derives from the exoskeleton, and likely provides physical ﬁll and/or a source of mineral nutrition to Hellbenders consuming whole crayﬁsh, rather than readily available calories. Mineral Content.—In general, the wild-caught crayﬁsh (Table 2) contained more inorganic constituents (ash) compared with commercially-reared crayﬁsh (Table 3; 47.16 versus 36.81%, respectively) and thus may provide better dietary mineral levels. The shell contributed 79.6% of measured ash, ~90% of Ca, and 31–64% of other minerals (with the exception of S, where 80% was present in meat). Northern Crayﬁsh sampled from SLZ were more similar to wild-captured crayﬁsh than were those sampled from either PBS or OF, and differed signiﬁcantly (P < 0.05) in Herpetological Review 40(3), 2009 TABLE 2. Nutrient composition (mean ± SD) of wild captured crayfish collected from two locations in close proximity on the North Fork of the White River, Missouri, Aug 2006 and Sept 2007. Data represent whole crayﬁsh; all nutrients except water presented on a dry matter basis. Species N Orconectes longidigitus 7 Orconectes neglectus 7 Orconectes punctimanus 4 Orconectes species 3 67.40 ± 10.07 41.53 ± 3.95 4.47 ± 1.51 14.25 ± 0.49 23.36 ± 4.00 14.70 ± 2.28 42.77 ± 5.09 65.84 ± 10.31 38.33 ± 4.17 4.44 ± 1.32 15.20 ± 0.14 23.02 ± 5.33 14.41 ± 2.47 46.11 ± 5.15 64.25 ± 3.59 38.08 ± 1.83 3.38 ± 1.33 14.20 (N = 1) 24.42 ± 2.78 16.01 ± 3.08 47.43 ± 4.02 71.59 ± 1.57 34.45 ± 1.77 2.75 ± 1.20 na 20.95 ± 2.62 15.42 ± 0.73 52.33 ± 1.59 15.65 ± 2.23 0.73 ± 0.01 0.33 ± 0.05 0.96 ± 0.15 0.52 ± 0.05 0.50 ± 0.04 0.35 ± 0.05 0.49 ± 0.03 38.86 ± 8.40 59.29 ± 26.31 82.29 ± 23.94 0.47 ± 0.28 75.86 ± 8.30 20.27 ± 5.07 0.57 ± 0.10 0.26 ± 0.02 0.93 ± 0.14 0.51 ± 0.09 0.55 ± 0.15 0.31 ± 0.07 0.58 ± 0.05 27.57 ± 5.09 71.57 ± 34.98 85.57 ± 15.13 0.40 ± 0.35 78.57 ± 6.32 15.96 ± 4.51 0.64 (N = 1) 0.30 ± 0.03 0.77 ± 0.08 0.53 ± 0.07 0.47 ± 0.08 0.30 ± 0.05 0.41 (N = 1) 27.00 ± 7.70 73.00 ± 37.72 93.50 ± 38.52 0.20 ± 0.14 65.25 ± 11.09 19.49 ± 0.59 na 0.28 ± 0.02 0.78 ± 0.04 0.54 ± 0.07 0.53 ± 0.11 0.24 ± 0.01 na 39.00 ± 11.31 38.50 ± 6.36 86.50 ± 23.33 0.20 ± 0.00 84.50 ± 3.54 95.57 ± 143.25 318,567± 477,500 267.72 ± 129.08 30.90 ± 12.96 ab 123.43 ± 124.61 411,433 ± 415,367 261.98 ± 141.30 26.43 ± 19.35 ab 160.14 ± 164.89 533,800 ± 549,633 248.26 ± 83.33 43.52 ± 11.74a 10.36 ± 5.34 34,533 ± 17,800 162.21 ± 66.62 21.14 ± 3.08b Nutrient Water, % Crude Protein, % Crude Fat, % Crude Fiber, % Neutral Detergent Fiber, % Acid Detergent Fiber, % Ash, % Minerals Calcium, % Chloride, % Magnesium, % Phosphorus, % Potassium, % Sodium, % Sulfur, % Cobalt, mg/kg Copper, mg/kg Iron, mg/kg Manganese, mg/kg Molybdenum, mg/kg Zinc, mg/kg Fat-Soluble Vitamins Vitamin A, mg/kg, as retinol Vitamin A, IU/kg Vitamin E, mg/kg, α-tocopherol Total Carotenoids, mg/kg ab Means in the same row without a common superscript letter differ (Mann-Whitney test; P < 0.05). total ash, Ca, Cu, and Fe from those groups (Table 3). Other signiﬁcant differences between the SLZ-pond and ﬁshery-pond reared crayﬁsh (Mg, K, S levels) may reﬂect differences in water or substrate quality, and/or result from unspeciﬁed holding and feeding schedules at the bait shop. Macromineral requirements (as a percentage of dietary DM) for growing mammal and bird species (Ca, 0.4 to 1.2%; Mg, 0.03 to 0.1%; P, 0.3 to 0.6%; K, 0.2 to 1.4%; and Na, 0.05 to 0.4%; Zootrition 2006) appear to be met by any of the crayﬁsh analyzed in this study. Wide variability among and within samples in trace element composition is evident and likely due to multiple factors including limited sample size, differing original dietary trace mineral levels, and possibly habitat/substrate variability. Mineral concentrations in water and substrate did not differ between sampled sites at SLZ (data not shown), but neither water, substrate, nor food samples were obtained from any other crayﬁsh group, so source of variation is unknown at this time. Dietary requirements for Cu range from about 3 to 5 mg/kg DM for domestic carnivores (NRC 2006); and high levels can be toxic. In particular, the high Cu level documented in some crayﬁsh (>100 mg/kg) is of potential health concern. Although LC50 is not directly comparable to LD50, toxicity (based on survival) has been determined between 40 and ~800 μg/L for Ambystoma spp. salamanders (Sparling et al. 2000), a magnitude lower than dietary levels measured in this study. While Cu LD50 has not been reported for salamanders, dietary tolerance of this nutrient is 15 and 40 mg/kg for sheep and cattle, respectively, and 100 mg/kg for ﬁsh (NRC 2005). Northern Crayﬁsh could have been exposed to copper from commercial ﬁsh foods and ﬁsh wastes in conﬁned ponds, which may have affected their overall composition. Copper sulfate is often added as an algaecide and may contribute to crayﬁsh body composition (38% of Cu was found in the shell); hence, aquatic environmen- Herpetological Review 40(3), 2009 327 TABLE 3. Nutrient composition (mean ± SD) of pond-reared crayfish consumed by Ozark Hellbender (Cryptobranchus alleganiensis bishopi) at the Saint Louis Zoo (2005–2007). Data represent whole crayﬁsh; all nutrients except water presented on a dry matter basis. Species Orconectes virilis 9 Paul’s Bait Shop Orconectes virilis 7 Ozark Fisheries Orconectes virilis 21 St. Louis Zoo Procambarus paeninsulanus 9 Florida Water, % Crude Protein, % Crude Fat, % Crude Fiber, % 74.31 ± 4.16 54.08 ± 4.14 2.96 ± 1.47 20.04 ± 7.38a 77.58 ± 3.23 54.80 ± 5.68 3.70 ± 0.80 68.02 ± 5.08 42.00 ± 4.98 3.03 ± 1.41 70.71 ± 4.57 47.84 ± 5.29 4.90 ± 1.62 Neutral Detergent Fiber, % Acid Detergent Fiber, % Ash, % 17.27 ± 0.95 13.70 ± 0.66a 35.69 ± 4.42a 13.78 ± 2.79b 21.70 ± 4.70 11.55 ± 0.64b 31.90 ± 3.69a na 28.98 ± 3.31 13.79 ± 1.53 ab 45.36 ± 5.71b na 31.94 ± 14.19 15.44 ± 3.84 34.27 ± 3.20 12.00 ± 1.79a 1.22 ± 0.20 0.40 ± 0.04a 1.16 ± 0.04 0.68 ± 0.11a 0.73 ± 0.06 0.44 ± 0.06ab 0.99 ± 0.29 107.44 ± 12.19a 133.78 ± 53.60a 108.44 ± 101.15 0.17 ± 0.08a 87.78 ± 6.04 11.85 ± 1.87a 1.29 ± 0.01 0.28 ± 0.07b 1.11 ± 0.06 0.96 ± 0.15b 0.78 ± 0.10 0.47 ± 0.03a 1.22 ± 0.12 110.57 ± 18.38a 104.00 ± 34.79a 55.29 ± 13.34 0.53 ± 0.23b 84.86 ± 11.32 16.31 ± 2.31b na 0.37 ± 0.04a 1.07 ± 0.16 0.64 ± 0.14a 0.62 ± 0.09 0.33 ± 0.07b na 34.24 ± 13.51b 315.71 ± 134.60b 43.19 ± 23.56 0.67 ± 0.46b 71.81 ± 9.45 12.25 ± 0.88 na 0.35 ± 0.02 0.92 ± 0.06 0.81 ± 0.08 0.60 ± 0.09 0.40 ± 0.08 na 46.75 ± 9.74 89.25 ± 84.13 9.88 ± 2.17 0.20 ± 0.08 66.38 ± 4.27 482.23 ± 70.95a 1,607,433 ± 236,500a 320.16 ± 118.80a 55.37 ± 15.29 878.22 ± 206.11b 2,927,400 ± 687,033b 485.65 ± 190.76b 56.18 ± 38.79 7.97 ± 2.51c 26,567 ± 8,367c 353.84 ± 136.49ab 32.35 ± 15.32 8.30 ± 2.69 27,667 ± 8,967 252.07 ± 116.61 48.75 ± 17.41 N Nutrient Minerals Calcium, % Chloride, % Magnesium, % Phosphorus, % Potassium, % Sodium, % Sulfur, % Cobalt, mg/kg Copper, mg/kg Iron, mg/kg Manganese, mg/kg Molybdenum, mg/kg Zinc, mg/kg Fat-Soluble Vitamins Vitamin A, mg/kg, as retinol Vitamin A, IU/kg Vitamin E, mg/kg, α-tocopherol Total Carotenoids, mg/kg ab Means in the same row without a common superscript letter differ (Mann-Whitney test; P < 0.05). tal quality is critical both directly and indirectly to the health of the Hellbender. Similarly, iron requirements for carnivores range from ~30 to ~100 mg/kg DM (NRC 2006), and excessive levels may interfere with bioavailability of other trace minerals. Recommended levels of bioavailable Mn and Zn (5 mg/kg, and 10 to 50 mg/kg, for cats and dogs, respectively; NRC 2006) would likely be met by crayﬁsh as a dietary staple, but nutrient requirements and dietary interactions in salamander nutrition remain to be investigated. Fat-Soluble Nutrients.—Vitamin A levels tend to increase with age/maturity in vertebrate prey through accumulation in body stores, particularly liver. Body size and vitamin A content were uncorrelated in our study; however, vitamin A levels were exceptionally high in farmed (OF) and baitshop (PBS) sourced crayﬁsh (Table 3; 1.6 to 2.9 million IU/kg DM [0.3 μg retinol = 1 IU]) 328 as well as in 3 of 4 wild crayﬁsh species (~300,000 to 500,000 IU vitamin A/kg DM). The unidentiﬁed species and the SLZ pond-caught crayﬁsh contained vitamin A concentrations ranging between ~26,000 and 40,000 IU/kg DM. Dietary requirements for vitamin A in domestic carnivores are approximately 4000 to 11,000 IU/kg DM (NRC 2006); presumed upper safe limits of this nutrient range from 33,000 IU/kg (dogs) to 100,000 IU/kg DM (cats) (NRC 1987). Clearly more information is needed on safe levels of this nutrient for salamanders (and crayﬁsh), but it appears likely that diets fed to the farmed animals may have contributed to the high body levels found in this study. Vitamin E concentrations were quite high in all crayﬁsh sampled; crayﬁsh meat has been previously shown to be a good source of this nutrient (USDA 2009) and may provide an important dietary antioxidant. Vitamin E levels in crayﬁsh (160–486 Herpetological Review 40(3), 2009 TABLE 4. Fatty acid composition (mean ± SD) of whole Missouri native crayﬁsh (Northern Crayﬁsh, Orconectes virilis), compared with whole Paciﬁc Krill (Euphasia paciﬁca). Fatty Acid Name Fatty Acid* Crayﬁsh (N = 10) Krill (N = 1) % of total fatty acids Lauric acid Myristic acid Palmitic acid Palmitoleic acid Stearic acid Oleic acid Linoleic acid γ-linolenic acid α-linolenic acid Dihomo- γ -linolenic acid Arachidonic acid Eicosapentaenoic acid (EPA) Docosahexaenoic acid (DHA) 12:0 14:0 16:0 16:1 18:0 18:1n9 18:2n6 18:3n6 18:3n3 20:3n6 20:4n6 20:5n3 22:6n3 %PUFA n3:n6 4.8 ± 1.0 3.6 ± 0.9 19.0 ± 3.6 6.6 ± 3.2 5.0 ± 1.3 24.8 ± 3.8 9.0 ± 2.9 0.7 ± 0.3 4.0 ± 2.6 1.0 ± 1.0 6.5 ± 1.9 8.1 ± 1.9 2.6 ± 1.3 31.9 0.85:1 2.5 15.6 25.6 10.6 1.7 20.3 1.9 0.3 0 0 0.3 13.5 6 22.0 7.8:1 * First number denotes the # of carbons in the fatty acid; 2nd # denotes the total # of double bonds; the number following “n” refers to the position of the ﬁrst double bond relative to the methyl- or “omega”–end of the fatty acid. mg/kg DM, calculated 240–729 IU/kg) consistently exceeded dietary requirements established for domestic carnivores (30–50 IU/kg; 1 mg natural source vitamin E = ~ 1.5 IU) fed diets containing moderate levels of polyunsaturated fats (PUFAs). High dietary PUFA (as found in crayﬁsh), however, may increase the vitamin E requirement up to 5-fold due to the high oxidative potential of these fats; vitamin E can be depleted as it is utilized as a biological antioxidant. Vitamin E deﬁciency has been shown to adversely impact reproduction and immune function in other species (Dierenfeld and Traber 1992). Hence these high concentrations of vitamin E may be beneﬁcial to health of Hellbenders, but require further investigation. The only nutrient category that differed signiﬁcantly among wild-caught crayﬁsh species was total carotenoid concentration (Table 2), and unfortunately, this was compared with the species that remains unidentiﬁed. Carotenoid content measured is likely an artifact of small sample size, but may reﬂect real differences in feeding habits among crayﬁsh species. O. punctimanus appears much broader in its habitat and possibly feeding requirements compared to other species, (MOFWIS 2008) and thus may be able to consume a diet containing higher levels of carotenoid pigments, but must be quantiﬁed further in more controlled studies. Diets fed in the ﬁsheries pond(s) quite likely also contributed to the high levels of vitamins and carotenoids measured in the crayﬁsh. It remains unclear, however, whether these higher nutrient levels are beneﬁcial or detrimental to the salamanders consuming crayﬁsh, as both vitamin A toxicity and deﬁciency can negatively impact health and reproduction. Vitamin A concentrations are lower in native crayﬁsh or pond-captured crayﬁsh compared with those sampled from farm-reared populations, but ranges varied by levels of magnitude, and, in general were higher than documented dietary requirements for carnivorous species in all samples – regardless of origin. Some of these concentrations are in ranges considered toxic for other species; to date, salamander dietary vitamin A nutrient requirements are unknown. Controlled feeding trials should be conducted and/or tissue levels of fat-soluble nutrients should be monitored to determine nutrient interactions and evaluate status. One potentially negative factor associated with low-fat diets (as found in crayﬁsh) may be lower absorption of fat-soluble vitamins; this possibility should be investigated in future studies with Hellbenders. Sources of variability in nutrient parameters measured in the SLZ crayﬁsh found season (summer vs. fall) to be signiﬁcant only in crude fat content (P < 0.01); location signiﬁcantly impacted Cu (P < 0.01), Zn ( P < 0.01), and P (P < 0.001) levels, even though trace mineral and water phosphate levels did not vary between the two sites (data not shown). Body size (hence, age/particular molt) had the greatest impact on both proximate and mineral nutrient composition of Northern crayﬁsh at the Saint Louis Zoo, affecting crude protein (P < 0.001), ash (P < 0.001), Ca (P < 0.05), K (P < 0.001), Na (P < 0.05), P (P < 0.05), Cu (P < 0.001), Mn (P < 0.05), and Zn (P < 0.01) content. Larger crayﬁsh contained higher levels of proximate nutrients (protein, fat, and ash) as well as macrominerals Ca, K, and Na compared to smaller animals. Phosphorus content, however, was higher in smaller crayﬁsh, as were levels of trace minerals (Cu, Mn, and Zn). Locally harvested crayﬁsh represented a nutritionally superior food for captive Hellbender compared to ﬁsheries-reared individuals, and were closer in chemical composition to species found in native habitats. Sustainable local harvest should be explored as a viable alternative feeding option and as a way of reducing fossil fuel (transportation) costs. These data provide preliminary detail on the nutrient composition of whole crayﬁsh eaten by Hellbender, but clearly species, size, and habitat of origin impact the nutritional content of crayﬁsh (and likely other aquatic prey species) eaten by Hellbenders. The chemical composition of native, as well as substitute, food items must be considered integral to the development of optimal diets for this species, and monitoring prey composition can provide strong bioindicators of habitat quality. In addition to crayﬁsh, Hellbenders also eat small ﬁsh, lamprey, worms, insects, snails, mollusks, tadpoles, and ﬁsh entrails in nature (Nickerson and Mays 1973; Peterson et al. 1989; Petranka 1998); in captivity, they are fed black worms, krill, and ﬁsh along with crayﬁsh (J. Ettling, pers. comm.). Intake, utilization, growth response trials, and health assessments to these varied whole prey items, and mixed diets, should provide valuable management guidelines that can assist in captive propagation, habitat evaluation, conservation and ultimate recovery of these unique and threatened salamanders. Characterization of both ex situ and in situ diets remains a critical component underlying health, reproduction, and recovery program management. Acknowledgments.—We thank R. Goellner for his support in early stages of this investigation, the ﬁeld conservation team including K. Goellner, K. Larson, J. Kiger, Y. Huang, J. Utrup, R. Junge and M. Wanner for assisting with the capture of wild-caught crayﬁsh for this study, and A. Braddy, C. Chavez, C. Resimius, T. Rivers, V. Spradling, and S. Herpetological Review 40(3), 2009 329 Tomasic for assisting with sample collection, preparation, analysis, and data organization. S. Blake provided helpful statistical guidance. LITERATURE CITED BARKER, D., M. P. FITZPATRICK, AND E. S. DIERENFELD. 1998. Nutrient composition of selected whole invertebrates. Zoo Biol. 17:223–134. BRIGGLER, J. T., J. UTRUP, C. DAVIDSON, J. HUMPHRIES, J. GROVES, T. JOHNSON, J. ETTLING, M. WANNER, K. TRAYLOR-HOLZER, D. REED, V. LINDGREN, AND O. BYERS (EDS.). 2007. Hellbender population and habitat viability assessment: ﬁnal report, IUCN/SSC Conservation Breeding Specialist Group, Apple Valley, Minnesota. 118 pp. CONANT, R., AND J. T. COLLINS. 1998. A Field Guide to Reptiles and Amphibians of Eastern and Central North American. Houghton-Mifﬂin Co., Boston, Massachusetts. 616 pp. DIERENFELD, E. S., H. L. ALCORN, AND K. L. JACOBSEN. 2002. Nutrient composition of whole vertebrate prey (excluding ﬁsh) fed in zoos. National Agricultural Library Z7994.Z65, 20 pp. http://www.nal.usda.gov/awic/ zoo/WholePreyFinal02May29.pdf. Accessed: 03 January 2009. ________ , AND M. G. TRABER. 1992. Vitamin E status of exotic animals compared with livestock and domestics. In L.Packer and J. Fuchs (eds.), Vitamin E in Health and Disease, pp. 345–360. Marcel Dekker, Inc., New York. FIRSCHEIN, I. L. 1951. The range of Cryptobranchus bishopi and remarks on the distribution of the genus Cryptobranchus. Amer. Midl. Nat. 45:455–459. FRITSCHE, K. 2006. Fatty acids as modulators of the immune response. Ann. Rev. Nutr. 26:45–73. IVERSON, S. J., K. J. FROST, AND S. L. C. LANG. 2002. Fat content and fatty acid composition of forage ﬁshes and invertebrates in Prince William Sound, Alaska: factors contributing to among and within species variability. Mar. Ecol. Prog. Ser. 241:161–181. JOHNSON, T. R. 2000. The Amphibians and Reptiles of Missouri. Missouri Department of Conservation, Jefferson City, Missouri. MCGRAW, K. J., P. M. NOLAN, AND O. L. CRINO. 2006. Carotenoid accumulation strategies for becoming a colorful house ﬁnch: analyses of plasma and liver pigments in wild molting birds. Funct. Ecol. 20:678–688. [MOFWIS]. MISSOURI FISH AND WILDLIFE INFORMATION SYSTEM. 2008. Accessed: 03 January 2009. URL: http://mdc4.mdc.mo.gov/applications/mofwis. MITCHELL, D. J., AND L. A. SMOCK. 1991. Distribution, life history, and production of crayﬁsh in the James River, Virginia. Amer. Midl. Nat. 126:353–363. [NRC]. NATIONAL RESEARCH COUNCIL. 1987. Vitamin Tolerance of Animals. National Academies Press, Washington, DC. 96 pp. ________ . 2005. Mineral Tolerance of Animals. 2nd revised edition. National Academies Press, Washington, DC. 496 pp. ________ . 2006. Nutrient Requirements of Dogs and Cats. National Academies Press, Washington, DC. 424 pp. NEGRO, J. J., AND J. GARRIDO-FERNANDEZ. 2000. Astaxanthin is the major carotenoid in tissues of white storks (Ciconia ciconia) feeding on introduced crayﬁsh (Procambarus clarkii). J. Comp. Biochem. Physiol. B 126:347–352. NICKERSON, M. A., AND C. E. MAYS. 1973. The Hellbender: North American “Giant Salamander”. Milwaukee Public Museum 1:1–106. PETERSON, C. L., J. W. REED, AND R. F. WILKINSON. 1989. 1989. Seasonal food habits of Cryptobranchus alleganiensis (Caudata: Cryptobranchidae). Southwest. Nat. 34:438–441. PETRANKA, J. W. 1998. Salamanders of the United States and Canada. Smithsonian Institution Press, Washington DC. 584 pp. PFINGSTEN, R. A. 1990. The status and distribution of the hellbender, Cryptobranchus alleganiensis, in Ohio. Herpetol. Rev. 21:48–51. SNEDECOR, G. W., AND W. G COCHRAN. 1967. Statistical Methods. 6th ed. Iowa State University Press, Ames, Iowa. 593 pp. 330 SPARLING, D. W., G. LINDER, AND C. A. BISHOP. 2000. Ecotoxicology of Amphibians and Reptiles. SETAC Press, Pensacola, Florida. 904 pp. SUKHIJA, P. S., AND D. L. PALMQUIST. 1988. Rapid method for determination of total fatty acid content and composition of feedstuffs and feces. J. Agr. Food Chem. 36:1202–1206. USDA National Nutrient Database. 2009. http://www.nal.usda.gov/fnic/ foodcomp/search/. Accessed: 03 January 2009. WHEELER, B. A., E. PROSEN, A. MATHIS, AND R. F. WILKINSON. 2003. Population declines of a long-lived salamander: a 20+ year study of hellbenders, Cryptobranchus alleganiensis. Biol. Conserv. 109:151– 156. ZOOTRITION. 2006. Diet analysis software. Version 2.6 Saint Louis Zoo, St. Louis, Missouri. NATURAL HISTORY NOTES The Natural History Notes section is analogous to Geographic Distribution. Preferred notes should 1) focus on observations in the ﬁeld, with little human intrusion; 2) represent more than the isolated documentation of developmental aberrations; and 3) possess a natural history perspective. Individual notes should, with few exceptions, concern only one species, and authors are requested to choose a keyword or short phrase which best describes the nature of their note (e.g., Reproduction, Morphology, Habitat, etc.). Use of ﬁgures to illustrate any data is encouraged, but should replace words rather than embellish them. The section’s intent is to convey information rather than demonstrate prose. Articles submitted to this section will be reviewed and edited prior to acceptance. Electronic submission of manuscripts is requested (as Microsoft Word or Rich Text format [rtf] ﬁles, as e-mail attachments). Figures can be submitted electronically as JPG ﬁles, although higher resolution TIFF or PDF ﬁles will be requested for publication. Please DO NOT send graphic ﬁles as imbedded ﬁgures within a text ﬁle. Additional information concerning preparation and submission of graphics ﬁles is available on the SSAR web site at: http://www.ssarherps.org/HRinfo.html. Manuscripts should be sent to the appropriate section editor: Charles W. Painter (amphibians; charles.painter@state.nm.us); James Harding (turtles; hardingj@msu. edu); Jackson D. Shedd (crocodilians, lizards, and Sphenodon; Jackson. Shedd@gmail.com); and John D. Willson (snakes; willson@uga.edu); and. Standard format for this section is as follows: SCIENTIFIC NAME, STANDARD ENGLISH NAME (if available, for the United States and Canada as it appears in Crother [ed.] 2008. Scientiﬁc and Standard English Names of Amphibians and Reptiles of North America North of Mexico. SSAR Herpetol. Circ. 37:1–84, available from SSAR Publications Secretary, ssar@herplit.com; for Mexico as it appears in Liner and Casas-Andreu 2008, Standard Spanish, English and Scientiﬁc Names of the Amphibians and Reptiles of Mexico. Herpetol. Circ. 38:1–162), KEYWORD. DATA on the animal. Place of deposition or intended deposition of specimen(s), and catalog number(s). Then skip a line and close with SUBMITTED BY (give name and address in full—spell out state names—no abbreviations). (NCN) should be used for common name where none is recognized. References may be brieﬂy cited in text (refer to this issue for citation format). One additional note about the names list (Crother 2008) developed and adopted by ASIH-HL-SSAR: The role of the list is to standardize English names and comment on the current scientiﬁc names. Scientiﬁc names are Herpetological Review 40(3), 2009 hypotheses (or at least represent them) and as such their usage should not be dictated by a list, society, or journal. Recommended citation for notes appearing in this section is: Medina, P., and R. L. Joglar. 2008. Eleutherodactylus richmondi: reproduction. Herpetol. Rev. 39:460. CAUDATA — SALAMANDERS AMBYSTOMA CALIFORNIENSE (California Tiger Salamander). COLORATION AND PATTERN. During the winter of 2003–2004, we conducted a survey for California Tiger Salamanders on farm lands owned by the City of Santa Rosa and two private properties in Sonoma County, California, USA. We used 17 pit-fall drift fence arrays (1360 pitfalls) to capture salamanders. During the course of our survey we captured 114 adult (57 male, 57 female) and 65 subadult California Tiger Salamanders on the city-owned Kelly Farm, an adjacent property on Duer Road, and another property at the north end of the former Santa Rosa Air Center. Animals were not marked and some may have been recaptured during the course of the study. Here we report on observations of the color and color pattern of the salamanders we captured. California Tiger Salamanders typically have a black ground color with distinctive rounded or irregular yellowish to whitish spots and bars (Petranka 1998. Salamanders of the United States and Canada. Smithsonian Institution Press, Washington, D.C. Fig. 1. (A) Adult male Ambystoma californiense (SVL 103 mm; TL 225 mm) exhibiting spotted color pattern typical of the Sonoma County population. (B) Subadult (SVL 70 mm; TL 105 mm) lacking typical spotted pattern. 587 pp.; Stebbins 2003. Western Reptiles and Amphibians 3rd ed. Houghton Mifﬂin Co., Boston, Massachusetts, 533 pp.). All the individuals captured in our survey, however, had a chocolate brown ground color with no gray or black tones (Fig. 1A). One subadult individual captured in our survey completely lacked pale spots or bars and was a uniform dark brown above and below (Fig. 1B); all other subadults captured during our study exhibited a typical spotting pattern. To the best of our knowledge California Tiger Salamanders with a chocolate brown ground color have not been described in the literature. Individuals of this species completely lacking a pale spotting pattern similarly have not been reported. Submitted by ERIC LICHTWARDT and TIMOTHY LACY, LSA Associates, Inc., 157 Park Place, Point Richmond, California 94801, USA (e-mail eric.lichtwardt@lsa-assoc.com). TARICHA RIVULARIS (Red-bellied Newt). ENDOPARASITES. Taricha rivularis occurs in northern coastal California from Sonoma and Lake counties north to Humboldt County (Stebbins 2003. A Field Guide to Western Reptiles and Amphibians, 3rd ed. Houghton Mifﬂin, Boston, Massachusetts. 533 pp.). To our knowledge there are no reports of helminths in T. rivularis. The purpose of this note is to establish the initial helminth list for T. rivularis. One adult male T. rivularis hand collected by MDD in Mendocino Co., California (123.8058°N, 39.4416°W, WGS 84, elev. 26 m) on 23 March 2009 was sacriﬁced and the digestive tract removed, opened, and examined under a dissecting microscope. Fourteen nematodes were found in the intestine, which were ﬁxed in alcohol, cleared in glycerol on a microscope slide under a coverslip, and studied using a compound microscope. Of these, six were female nematodes from the small intestine identiﬁed as Oswaldocruzia pipiens and eight (ﬁve male, three female) from the large intestine were identiﬁed as Cosmocercoides variabilis. Nematodes were deposited in the United States National Parasite Collection (USNPC), Beltsville, MD: Oswaldocruzia pipiens (USNPC 101993); Cosmocercoides variabilis (USNPC 101992). The T. rivularis was not accessioned. Cosmocercoides variabiis is a cosmocercid nematode found in North American salamanders, frogs, lizards, snakes, and turtles (Baker 1987. Occas Paps. Biol. Memor. Univ. Newfoundland 11:1–325). Some uncertainty exists for its hosts because of confusion between C. variabilis and C dukae, a molluscan parasite. Vanderburgh and Anderson (1987. Can. J. Zool. 65:1662–1665) demonstrated that the two species are distinct. The major difference in morphology for the two species is the number of rosette papillae in the male: C. dukae with 12 pairs, C. variabilis with 14–20 pairs. We have assigned these specimens to C. variabilis because males have 16 pairs of papillae. Infection occurs by skin penetration (Baker 1978. J. Parasitol. 64:765–766). All North American specimens of Oswaldocruzia have been referred to O. pipiens by Baker (1977. Can. J. Zool. 55:104–109). Oswaldocruzia pipiens is a strongylid nematode that is widely distributed and has been reported from frogs, toad, salamanders, lizards, and tortoises (Baker 1987, op. cit.). In the case of amphibians, transmission occurs by skin penetration (Baker 1978. Can. J. Zool. 56:1026–1031). Taricha rivularis represents a new host record for O. pipiens and C. variabilis. Herpetological Review 40(3), 2009 331 Specimens were collected under authority of California state ﬁshing license 029486-09 issued to Murray D. Dailey. Submitted by STEPHEN R. GOLDBERG, Department of Biology, Whittier College, Whittier, California 90608, USA (e-mail: sgoldberg@whittier.edu); CHARLES R. BURSEY, Department of Biology, Pennsylvania State University, Shenango Campus, Sharon, Pennsylvania 16146, USA (e-mail:cxb13@psu.edu); and MURRAY D. DAILEY, PO Box 182, The Sea Ranch, California 95497, USA (e-mail: daileym@mcn.org). ANURA — FROGS LIMNONECTES IBANORUM (Rough-backed River Frog). DIET. Limnonectes ibanorum is endemic to the island of Borneo, where it occurs in rainforest at elevations between 50 and 900 m (Inger 1966. Field. Zool. 52:1–402). Although this large frog (SVL to 13 cm) is common along clear, rocky streams of intermediate width (10–30 m) (Inger and Stuebing 2005. A Field Guide to the Frogs of Borneo. Natural History Publications [Borneo], Kota Kinabalu. 201 pp.), almost nothing is known about its ecology. According to Inger and Stuebing (2005, op. cit.), L. ibanorum feeds on large insects, crabs, and small lizards. The diet of its large (SVL > 9 cm) Bornean congeners, i.e. Limnonectes ingeri, L. leporinus, and L. malesianus, also includes frogs (Inger and Stuebing 2005, op. cit.; Malkmus et al. 2002. Amphibians and Reptiles of Mount Kinabalu [North Borneo]. Koeltz Scientiﬁc Books, Königstein. 424 pp.), but there are no reports of predation of frogs by L. ibanorum. On 20 Sept 2007 at 2045 h, I observed an adult L. ibanorum (SVL 11.5 cm) catching and devouring an adult Hylarana raniceps (Fig. 1). The prey was ingested head ﬁrst. The specimen of L. ibanorum was caught afterwards, measured, and released. The observation took place on the bank of the Nyipa River, Camp 1, Gunung Mulu National Park, Sarawak, Malaysia (ca. 150 m elev.). I thank the Sarawak Forest Department for permission to conduct research in Gunung Mulu National Park. Fieldwork was supported by a grant from the German Academic Exchange Service (DAAD). Submitted by JONAS MAXIMILIAN DEHLING, Department of Animal Ecology and Tropical Biology, Biocentre, University of Würzburg, Am Hubland, D-97074 Würzburg, Germany; e-mail: Jonas.M.Dehling@stud-mail.uni-wuerzburg.de. TESTUDINES — TURTLES CHRYSEMYS PICTA BELLII (Western Painted Turtle). AS INTRODUCED PREDATOR. The Western Painted Turtle occurs naturally in the central and northwestern United States and adjacent southern Canada but has been widely introduced to water bodies outside its natural range. Established populations and individual records in California presumably are introduced (Lever 2003. Naturalized Reptiles and Amphibians of the World. Oxford University Press; Stebbins 2003. A Field Guide to Western Reptiles and Amphibians. Houghton Mifﬂin Co., Boston, Massachusetts). At 1600 h on 15 February 2007 an adult (782.8 g) Chrysemys picta bellii was found in Tracy Lake, San Mateo Co., California. When ﬁrst observed, the turtle was in shallow water with a dead Paciﬁc Chorus Frog (Pseudacris regilla) in its mouth. The condition of the frog suggests it was captured alive, rather than scavenged. It is perhaps signiﬁcant that this turtle was eating a native frog in Tracy Lake, which supports a breeding population of the threatened California Red-legged Frog (Rana draytonii). The Western Pond Turtle (Actinemys marmorata), which is listed by California as a Species of Concern, has also been observed in this lake. The introduced Red-eared Slider (Trachemys scripta elegans) has been established in this area for years, but to our knowledge this is the ﬁrst report of C. p. bellii for this area. The impact of introduced turtles on native amphibians through direct predation remains unknown. The impact of Painted Turtles as a competitor of Western Pond Turtles is also unknown, but the latter are also known to feed on native anurans (Bury 1986. J. Herpetol. 20:515–521). Frogs have been previously noted in the diet of C. picta (Ernst et al. 1994. Turtles of the United States and Canada. Smithsonian Inst. Press, Washington, DC) but may not be a signiﬁcant food source. No frogs were detected in the stomachs of 46 C. picta collected in Michigan (Knight and Gibbons 1968. Am. Midl. Nat. 80[2]:558–562). The turtle and frog were collected under California Department of Fish and Game Scientiﬁc Collecting Permit 008139 and deposited in the collections at the California Academy of Sciences. Submitted by JOHN KUNNA and KAREN SWAIM, Swaim Biological, Inc., 4435 First St. PMB 312, Livermore, California 94551, USA (e-mail: john.kunna@gmail.com). FIG. 1. Limnonectes ibanorum ingesting Hylarana raniceps, Gunung Mulu National Park, Sarawak, Malaysia. 332 EMYDOIDEA BLANDINGII (Blanding’s Turtle). PREDATION. Neonate turtles, including Emydoidea blandingii, are known to have many predators. The dissection of Bullfrog (Lithobates [= Rana] catesbeianus) stomachs occasionally reveals the remains of hatchling turtles of various species, and L. catesbeianus has previously been observed in unsuccessful pursuit of hatchling E. blandingii (Bleakney 1963. Can. Field Nat. 77:67–76). Herein we report evidence of the successful predation of a hatchling E. blandingii by an adult L. catesbeianus. On 28 August 2006, 12 hatchling E. blandingii emerged from Herpetological Review 40(3), 2009 a protected nest at Winous Point Marsh, Ottawa County, Ohio, USA. The hatchlings were outﬁtted with 0.5 g radio transmitters and released at the nest site the following day. On 11 September we tracked one of these hatchlings to an agricultural ditch ca. 440 m W of the nest. Over several minutes we followed this individual’s signal as it evaded our capture by rapidly moving back and forth across the ﬂooded ditch. The submerged animal then emerged onto land revealing itself as an adult L. catesbeianus. We proceeded to track the frog over land, by sight and telemetry, through thick multi-ﬂora roses until it disappeared into a large animal burrow. We tracked the radio signal back to the frog the next day, but our attempts to capture the animal proved unsuccessful. We were unable to pick up the radio signal the following day due to presumed exhaustion of the transmitter’s battery. The radio transmitter included a thin vinyl coated antenna measuring ca. 9 cm long. The antenna was not observed protruding from the mouth of the frog and it is uncertain what effect it may have ultimately had on the predator. Although L. catesbeianus is known to consume nearly any animal it can ﬁt into its mouth, this is believed to be the ﬁrst report of E. blandingii having been consumed by this voracious and often abundant species. Submitted by JAMES C. SPETZ (e-mail: jimspetz@hotmail. com), and RICHARD SPENCE, Cleveland Metroparks, 4550 Valley Parkway, Fairview Park, Ohio 44126, USA. EMYDOIDEA BLANDINGII (Blanding’s Turtle). HATCHLING DIET. The behavior of hatchling Emydoidea blandingii in the wild is poorly understood. Previous studies have focused on the movements and habitat use of hatchlings after nest emergence (Butler and Graham 1995. Chelon. Conserv. Biol. 1:187–196; McNeil et al. 2000. Chelon. Conserv. Biol. 3:661–664). Herein, we report an observation of feeding behavior by a hatchling E. blandingii during its ﬁrst weeks post emergence from the nest. On 18 September 2007, six hatchling E. blandingii emerged from a protected nest at Winous Point Marsh, Ottawa Co., Ohio, USA. We attached radio transmitters to the hatchlings and released them at the nest site the same day. We tracked the hatchlings daily and GPS points were taken at each location. One hatchling tracked over the course of 29 days moved an average of 20 m/day. During this time it spent three days in a shallow, muddy agricultural ditch. The animal then exited this ditch and moved a short distance to a small vernal pool where it spent another three days along the shallow margins. The ditch and vernal pool were both located along a narrow strip of shrubby, wooded land between two agricultural ﬁelds. On 15 October we tracked this individual to the top of a dike adjacent to the vernal pool. The animal appeared lethargic and did not move again. We found it dead and intact in the same location two days later. No obvious indication of the cause of death was observed, but a tear in the skin was noted anterior to the left front leg where ﬂy eggs had recently been deposited. Dissection of the digestive tract of this animal revealed the chitinous remains of a small adult dytiscid beetle located within the colon along with some other unidentiﬁed material. The remains of the beetle measured ca. 5 mm long when articulated. The stomach contained an unidentiﬁed pink gelatinous material. Small perforations were observed in the wall of the stomach which were not believed to be due to researcher handling. The cause of these perforations and whether they resulted in the death of this animal could not be determined with certainty. However, it seems possible that they were associated with the death of this animal and may have occurred during consumption of the observed food item. Two additional hatchlings salvaged after predation and partial consumption (in 2006 and 2007) were also dissected and material was observed within the colon of each individual. The stomach was missing from both individuals and was presumably removed when the head of each had been consumed during predation. The colonic material included small bits of vegetation, grains of sand, and some unidentiﬁable material. Specimens will be deposited at The Cleveland Museum of Natural History. Observations of narrow growth annuli on juvenile E. blandingii have previously hinted at hatchling feeding activity (Pappas et al. 2000. Chelon. Conserv. Biol. 3:557–568), but this is believed to be the ﬁrst conﬁrmation of post-emergence feeding activity by hatchling E. blandingii prior to their ﬁrst winter dormancy. Submitted by JAMES C. SPETZ (e-mail: jimspetz@hotmail. com), and RICHARD SPENCE, Cleveland Metroparks, 4550 Valley Parkway, Fairview Park, Ohio 44126, USA. GLYPTEMYS INSCULPTA (Wood Turtle). ECTOPARASITES. Although freshwater turtles are known to be parasitized with leeches of the genus Placobdella (e.g., Watermolen 1996. J. Fresh. Ecol. 11:211–217), several families of biting ﬂies (Diptera) are ectoparasites of turtles as well. For example, biting midges in the genus Culicoides (Diptera: Ceratopogonidae) are well-known pests of vertebrates, including humans. Females of most Culicoides species require blood meals in order to produce one or more clutches of eggs [Blanton and Wirth 1979. Sand Flies (Culicoides) of Florida (Diptera: Ceratopogonidae), (in) Arthropods of Florida & Neighboring Land Areas. 10. 204 pp]. Currently, there are some 1300 known species of Culicoides [Borkent and Wirth 1997. World Species of Biting Midges (Diptera: Ceratopogonidae). Bull. Amer. Mus. Nat. Hist. 233:1–237.]. However, only one of the 152 Nearctic species (Culicoides testudinalis) is known to obtain blood meals from turtles. Herein, we report eight instances of C. testudinalis parasitizing the semi-aquatic turtle, Glyptemys insculpta in June of 2003 and 2004. On 5 June 2003 at 0835, one of us (JM) observed two adult male G. insculpta (# 138 and # 1009) feeding on an unidentiﬁed species of slug, within 4 m of each other, at an undisclosed study site in Nova Scotia, Canada. Both of these turtles were noted to have many midges on their carapaces. Later, at 1110, an adult female G. insculpta (# 1011) that was covered with hundreds of small midges on its carapace, head, and limbs was observed and photographed (Figs. 1–2). At 1151, another adult female (# 1012) was found to be parasitized as well. On 17 June 2003 at 1230, an adult female (# 1113) was noted to have 13 midges; all but three were swollen with blood meals. On 19 June 2003 at 1125, another adult female (# NT39) was reported to be parasitized by midges, but no additional details were recorded. Two additional observations were made in 2004. On 16 June 2004 at 0905, an adult female (#1194) was noted to have hundreds Herpetological Review 40(3), 2009 333 of midges, many of which were on the carapace engorged with blood . Several midges were feeding upon the tissues surrounding the eyes as well. A ﬁnal adult female (# 1177) was observed to be parasitized on 17 June 2004 at 0945. Although approximately 10 midges were associated with this G. insculpta, only one was engorged with blood. The mean carapace length (range) of the aforementioned eight G. insculpta was 195 mm (183–213 mm). The estimated minimum age of these turtles, based on carapacial growth annuli, ranged from 20–26 years. Several dozen of the midges from one G. insculpta (# 1011) were collected and subsequently sent to WLG for identiﬁcation. Nine unengorged midges were cleared in phenol-alcohol and mounted on microscope slides in phenol-Canada balsam by the methods of Wirth and Marston (1968. Ann. Entomol. Soc. Amer. 61: 783–784). Voucher specimens of slide-mounted C. testudinalis will be deposited in the Canadian National Collection of Insects (Ottawa, Ontario) and Florida State Collection of Insects (Gainesville, Florida). All slide-mounted midges are identical to three other females that were collected, mounted, and identiﬁed as C. testudinalis by Wirth; a paratype from Falls Church, Fairfax Co., VA, and two others from Prince Georges and Montgomery counties, Maryland in the synoptic collection of ceratopogonids maintained by WLG. The female holotype of C. testudinalis was collected on 6 June 1953 by Neill Richmond while it fed upon a Wood Turtle (G. insculpta) in Coburn, Centre County, Pennsylvania (Wirth and Hubert. 1962. Ann. Entomol. Soc. Amer. 55:182–195). Of the 78 female paratypes of this ectoparasitic midge, two were found biting Terrapene carolina at Patuxent Wildlife Refuge, Prince Georges County, Maryland by John Scanlon on 12 July 1958; whereas, three others were found “feeding on turtle” by R.C. Shannon on 23 May 1939 at Dead Run, Fairfax County, Virginia. Notably, of the several hundred C. testudinalis observed on the turtles, most individuals appeared to have pierced the sulci between carapacial scutes or the margins of growth annuli with their ﬁnetoothed mandibles in order to reach capillaries with their tubular mouthparts. This is consistent with observations of the biting midge Leptoconops bezzii feeding on the sulci between the scutes of FIG. 2. Adult Wood Turtle (Glyptemys insculpta) from Fig. 1 infested with hundreds of adult female biting midges (Culicoides testudinalis) on its carapace, head, neck, and front leg. The swollen, reddish abdomens of some midges are engorged with blood from the turtle. Testudo graeca in western Syria and Lebanon (Široký et al. 2007. Parasitol. Res. 101:485–489). Moreover, the leech, Placobdella ornata, is also known to obtain blood from the sulci between the scutes that overly the carapace bones (Siddall and Gaffney 2004. J. Parasitol. 90:1186–1188). Approximately 100 midges on one G. insculpta (# 1011) appeared to be fully engorged with blood, and a few that had completed feeding were resting on vegetation (Fig. 1). The highest densities of midges occurred between marginal and costal scute interfaces (Fig. 1); however, a much greater proportion of the midges on the anterior portion of the carapace were engorged (Fig. 2). Midges might initially be attracted to the head of a turtle, the source of exhaled CO2. It is unknown if turtles suffer signiﬁcant ill effects from the bites of Culicoides, but the G. insculpta in question were not observed attempting to dislodge any midges with forelimbs or engaged in any other erratic movements. Submitted by WILLIAM L. GROGAN, JR., Department of Biological Sciences, Salisbury University, Salisbury, Maryland, 21801, USA (e-mail: wlgrogan@salisbury.edu); RAYMOND A. SAUMURE, Research Division, The Springs Preserve, 1001 South Valley View Boulevard, Las Vegas, Nevada, 89107, USA (e-mail: insculpta@gmail.com); JODY MACEACHERN, Lakehead University, 9417A 83 St. NW, Edmonton, Alberta, T6C 2Z8, Canada (e-mail: jmaceach@lakeheadu.ca); LAUREN ALLEN, 841 Bridges Street, Halifax, Nova Scotia, B3H 2Z6, Canada (email: Lauren@nsnt.ca); and MARK D. PULSIFER, 190 Beech Hill Road, R.R. #7 Antigonish, Antigonish County, Nova Scotia, B2G 2L4, Canada (e-mail: pulsifmd@gov.ns.ca). FIG. 1. Posterior portion of adult Wood Turtle (Glyptemys insculpta) infested with female biting midges (Culicoides testudinalis). Note that most of the midges are feeding in the sulci of successive scute layers and several engorged midges are resting on vegetation. 334 GLYPTEMYS MUHLENBERGII (Bog Turtle). LONGEVITY. Between May 1969 and June 1982, William Kimmich conducted mark recapture surveys of Bog and Spotted Turtles (Clemmys guttata) at several sites in southeastern Pennsylvania to gain information about population dynamics of these turtles. (The northern populations of the Bog Turtle were listed as a Threatened species by the U.S. Fish and Wildlife Service in November 1997.) During Herpetological Review 40(3), 2009 this time period Kimmich marked approximately 100 Bog Turtles with small drill holes in the marginal scutes at one of these sites. Kimmich worked independently on this study, but provided his data and volunteer services to The Nature Conservancy after it acquired this property in 1989. Carl Ernst conducted research on Bog Turtles at this site between May 1982 and May 1989 and marked several additional Bog Turtles with square and triangular notches. Ernst also contributed data to the Conservancy for this analysis. In 1992 the Nature Conservancy, using staff and contractors, reinitiated mark/recapture surveys and conducted a short radio telemetry study at the site in 1992. Turtles marked during and after 1992 were marked exclusively with triangular notches in the marginal scutes. Turtles were sexed, and measurements of the carapace length and width, plastron length and width, and shell height were recorded. Shell wear and injuries were also noted at subsequent captures. These characteristics, as well as the conﬁguration of markings on the marginal scutes, were used for the data comparison. At least 29 turtles (9 males and 20 females) that were marked as adults prior to 1982 were recaptured after 1992. Positive identiﬁcation of some of these turtles has not been made because of erosion of holes to the edges of the scutes, the possible addition of marks to some turtles, predator damage to the marginal scutes, and drill holes ﬁlling with mud, making their detection difﬁcult. Fourteen turtles were conﬁrmed to be positive matches between the two data sets. Three of these turtles were recaptured in 1993, 1994 and 1997 respectively, and eleven of these turtles were recaptured after 2000. The most recent of these captures was made during the 2008 ﬁeld season. The average annual growth rate of Bog Turtles is rapid during the ﬁrst years after hatching, and gradually decreases as the turtle ages. Ernst (1977. Herpetologica 33: 241-246) documented annual growth rates of 34.6 percent at hatchling with a gradual decline to 5.2 percent at age twelve, indicating that Bog Turtles reach their maximum size at some time after age 12. Comparison of Kimmich’s data and that collected by The Nature Conservancy reveals an average size variation of 0.56 mm between initial and subsequent captures, indicating that these turtles were fully grown adults when ﬁrst captured, and were presumed to be at least thirteen years old at ﬁrst capture. Counting the number of scute annuli appears to be a reasonable method of determining approximate ages of Bog Turtles up to about 10 to 15 years, after which time growth may nearly cease and their burrowing habit will begin to wear away the annuli. While young Bog Turtles have distinct annuli, the annuli on the shells of turtles that were calculated to be more than 40 years old had been worn completely smooth. The earlier data did not note annuli count or wear and it is possible that several of these turtles are considerably older than the calculated ages. Many factors inﬂuence the speed and amount of wear on the shell, and shell wear may vary greatly between sites. The ages of turtles in these studies were estimated using the aforementioned criteria. One turtle is estimated to be at least 25 years old, two turtles to be at least 35 years old, seven turtles to be at least 38 years old, three turtles to be at least 45 years old, and one turtle is estimated to be at least 49 years old. Notably, one female whose age is calculated to be more than 48 years old had all toes missing on both left feet at both captures, and thus had survived for at least 35 years in the wild in this condition. Submitted by GEORGE C. GRESS, The Nature Conservancy, 2411 South Fifth Avenue, Lebanon, Pennsylvania 17042, USA; e-mail: ggress@tnc.org. GOPHERUS AGASSIZII (Desert Tortoise). MORTALITY. In 1990, populations of Gopherus agassizii in the Mojave Desert were listed as Threatened under the US Endangered Species Act (Fish and Wildlife Service 1990. Federal Register 55:12178–12191). This action was deemed necessary because of the increase in habitat loss and degradation in the Mojave Desert due to the rapid growth of many cities and suburban communities (Boarman and Beaman 2002. The Sensitive Plant and Animal Species of the Western Mojave Desert. U.S. Geological Survey, Western Ecological Research Center, Sacramento, California). Although anthropogenic activities undoubtedly contribute to G. agassizii mortality, many natural causes of mortality have also been reported. Many of the natural dangers to G. agassizii appear to affect young tortoises more than older individuals. Among the many threats are predation, parasites, disease, dehydration, and being crushed when dens collapse (Luckenbach 1982. In R. B. Bury [ed.], North American Tortoises: Conservation and Ecology, pp. 1–37. Wildl. Res. Rep. 12, U.S. Fish and Wildlife Service, Washington, DC). Here we report the ﬁrst observations of G. agassizii mortality caused by individuals falling into and being trapped in rock fractures on a basalt ﬂow. The habitats where G. agassizii can be found in the Mojave Desert vary from sandy valleys ﬁlled with Creosote Bush (Larrea tridentata), to rocky bajadas and hillsides (Riedle et al. 2008. Copeia 2008:414–420). Studies of G. agassizii in the Sonoran Desert suggest that preferred den sites may occur in rugged terrain with high densities of boulders (Barrett 1990. Herpetologica 46:202–206). Several reserves have been established with the aim of protecting critical tortoise habitat. The Red Cliffs Desert Reserve, Washington Co., Utah, was established in 1996 to protect a large tract of desert habitat suitable for G. agassizii and other desert wildlife. This reserve contains a diverse suite of habitats, from Blackbrush (Coleogyne ramosissima) covered mesas, to sandstone and basalt rock outcrops, to creosote bush ﬂats. On two occasions on separate basalt ﬂows in the Red Cliffs Desert Reserve, carcasses of adult G. agassizii were found. These individuals had apparently fallen headﬁrst into fractures in the rocks, become stuck, and died from exposure, likely from overheating. Both individuals had their hind legs in the air and their forelegs and head dangling down into the crack. No noticeable marks were found on the tortoise carcasses that would suggest they were killed and placed in the cracks by predators. It is unknown what caused the tortoises to wander onto the basalt ﬂows. This ﬁnding is interesting because tortoises in the Red Cliffs Desert Reserve often traverse steep sandstone outcrops with little difﬁculty (pers. obs.). It is likely a rare event for a tortoise to be trapped by falling into a crack in the rocks, yet given that two individuals were trapped and died in a similar manner, this may be a more common event than previously thought. Submitted by JOSEPH S. WILSON, Utah State University, Biology Department, 5305 Old Main Hill, Logan, Utah 84322, USA (e-mail: jwilson@biology.usu.edu); and SETH TOPHAM Herpetological Review 40(3), 2009 335 32 East 300 North, St. George, Utah 84770, USA (e-mail: BSTopham@hotmail.com). HYDROMEDUSA MAXIMILIANI (Brazilian Snake-necked Turtle). ALGAL COLONIZATION. Algae of the genus Basicladia (Chlorophyta, Cladophoraceae) are often noted growing on the shells of freshwater turtles, which offer the algae an attractive substrate for colonization (Edgren et al. 1953. Ecology 34:733–740; Ducker 1958. Hydrobiology 10:157–174; Semir et al. 1988. Cienc. cult. 40:885–888). The relationship between algae and turtles has been described as commensal (assuming the turtles receive little or no beneﬁt from the algae) or mutualistic (the turtle using the algae as camouﬂage, perhaps while foraging, and the algae using the turtle as a safe and mobile substrate) (Edgren et al., op. cit.; Niel and Allen 1954. Ecology 35:581–584; Proctor 1958. Ecology, 39:634–645; Dixon 1960. Texas J. Sci. 12:36–38). In November 2004 we captured seven Hydromedusa maximiliani in Reserva Biológica Municipal Santa Cândida (21.6888889°S, 43.3444444°W, 770 m elev.), Juiz de Fora, Minas Gerais state, Brazil. Algae on the turtle’s shells were collected with a scalpel and ﬁxed in Transeau and formaline 4% solution. Biometric analysis suggested that six of these specimens were adults, with an average maximum carapace length of 150.60 ± 12.51 mm, and one was considered a juvenile, with a carapace length of 128.60 mm. All specimens had B. cf. chelonum adhering to their carapaces, on the vertebral plates, principally on the anterior and posterior regions, and on the marginal plates. The same distribution of algae was also noted for the chelids Phrynops geoffroanus and Hydromedusa tectifera from Brazil (Semir et al. 1988. Cienc. Cult. 40:885–888). The algae occurred mainly in the anterior and posterior regions of the carapace on the marginal scutes, there being little algae on the costal and central scutes. A possible correlation between the feeding habits of turtles and frequency of epizoophyte growth is supported by the observation that carnivorous species that ambush or actively hunt their food may be more often subject to algal growth. Examples include include the chelydrids Macrochelys and Chelydra, kinosternids Sternotherus and Kinosternon, and the emydids Deirochelys and Emys. all of which hunt active prey, such as frogs, ﬁsh and aquatic insects (Niel and Allen 1954. Ecology 35:581–584). H. maximiliani has similar predatory feeding habits (Souza and Abe 1995. Chel. Cons. Biol. 1:320–322). The presence of algae on the carapace of this species may serve as protection against predators but would also decrease detection by prey. This record of B. cf. chelonum colonizing H. maximiliani is apparently the ﬁrst report of the algae growing on this turtle species and supports a possible mutualistic relationship between the algae and the turtle host. This work was licensed by IBAMA (Process nº 02015.003546/0411), and was performed under the principles adopted by COBEA (Brazilian School of Animal Experimentation), which were approved by the Committee of Ethics in Animal Experimentation (Pro-Rectory of Research) of the Federal University of Juiz de Fora, in a meeting which took place on 04/12/2004 (Protocol nº 011/2005-CEA). Submited by IARA ALVES NOVELLI, Pós-Graduação em Biologia Animal, Universidade Federal Rural do Rio de Janeiro, 336 BR-465, Km 7, Seropédica, Rio de Janeiro, Brazil, 23890-000 (e-mail: iaranovelli27@gmail.com); BERNADETE MARIA DE SOUSA Laboratório de Herpetologia, Departamento de Zoologia, Instituto de Ciências Biológicas, Universidade Federal de Juiz de Fora, Campus Universitário, Bairro Martelos, Juiz de Fora, MG, Brazil, 36036-330 (e-mail: bernadete.sousa@ufjf.edu.br); and IZABEL CRISTINA ALVES DIAS, Laboratório de Ficologia, Museu Nacional, Universidade Federal do Rio de Janeiro, Quinta da Boa Vista s/n; CEP 20940-040. Rio de Janeiro, RJ, Brazil. MACROCHELYS TEMMINCKII (Alligator Snapping Turtle). AERIAL BASKING. Macrochelys temminckii is rarely observed out of the water. Terrestrial activity is typically limited to nesting females and hatchlings traveling from nest to water. Only two deadon- road individuals are known (Ewert et al. 2006. In Meylan [ed.], Biology and Conservation of Florida Turtles, pp. 58–71. Chelon. Res. Monogr. No. 3, 376 pp.). Apparently, only four instances of observed or inferred aerial basking have been reported: on a log in Texas (Ewert 1976. Herpetologica 32:150–156; also see Ewert et al., op. cit.), on a fallen tree in Texas (Farr et al. 2005. Herpetol, Rev. 36:168), on a basking trap in Mississippi (Shelby and Jenson 2002. Herpetol. Rev. 33:304), and an adult basking on a river bank, also in Mississippi (Selman et al. 2009. Herpetol. Rev. 40:79). Here I report aerial basking of M. temminckii on land in Florida. The observation was made as part of a long-term (since 2003) ecological study of Macrochelys temminckii in northern peninsular Florida. On 19 October 2008 at 1300 h, I found a juvenile (18.6 cm CL, 1.68 kg) M. temminckii on the northern bank of the Santa Fe River, Columbia Co. Florida, 1.7 km upstream from the US 27 bridge. This turtle was basking in partial sunlight at least 2 m from the water’s edge. The turtle was found on sandy soil and appeared completely dry. Air temperature and water temperature (10 cm depth) were 23.3 and 22.5°C, respectively. Upon capture, this turtle was found to have been originally marked with a passive integrated transponder (PIT) on 9 July 2008, and had traveled ca. 2.5 km from the initial capture site. Additionally, I counted over 100 leeches that were attached to the turtle. This is the ﬁrst report of aerial basking in Florida. Aerial basking in this species is certainly rare, but may occur sporadically in response to thermoregulatory needs of individual turtles. Submitted by TRAVIS M. THOMAS, Florida Museum of Natural History, Division of Herpetology, P.O. Box 117800, University of Florida, Gainesville, Florida 32611, USA; e-mail: tthomas@ﬂmnh.uﬂ.edu. PHRYNOPS HILARII (Hilaire’s Side-necked Turtle). FEEDING BEHAVIOR. The freshwater chelid turtle Phrynops hilarii is one of the most abundant chelonians in Rio Grande do Sul state, Brazil. Research by the Chelonia-RS Project in the lake at Moinhos de Vento Park, Porto Alegre city (30.0269444°S, 51.2011111°W) has been carried out intermittently since 2003. Between August and December 2008, 29 P. hilarii were captured and marked. Straightline carapace length ranged between 113 and 387.2 mm (mean = 280.21; SD = 81.2). In the lake, which is ca. 4675 m2 in area and has a maximum depth of 1.5 m, there are four other chelonian Herpetological Review 40(3), 2009 species (Trachemys dorbigni, T. scripta, Acanthochelys spixii, and Hydromedusa tectifera). This note provides additional data about the diet and feeding behavior of P. hilarii at this location. Predation of waterfowl and other birds by chelonians has been reported for the Snapping Turtles, Chelydra serpentina, and the Red-eared Slider, Trachemys scripta elegans (Ernst et al. 1994. Turtles of the United States and Canada. Smithsonian Inst. Press, Washington, D.C.; Ligon 2007. Herpetol. Rev. 38:201–202; Pryor 1996. Wilson Bull. 108:190–192). However, predation of nonaquatic birds by P. hilarii has not been previously described. Pigeons (Columba livia) are abundant in the park and capture of the species by P. hilarii was observed by Daniel Borba Rocha and co-workers in 2004. On 9 October 2007, I described (ﬁeld notes of the Chelonia-RS Project) the predation of a pigeon by a P. hilarii while the bird was drinking water at the margin of the lake. The turtle, without leaving the water, grabbed the bird by the neck with its jaws, and pulled it to the bottom of the lake, while other turtles (T. scripta and P. hilarii), also bit at the bird. On 22 January 2009, pictures and a report about pigeon predation by the Hilaire’s Side-necked Turtle were published in a local newspaper (Zero Hora newspaper 2009. 15849:42). In this situation, the capture occurred on land as pigeons foraging on the margins of the lake were ambushed by a P. hilarii. When the birds got close to the margin of the lake, the turtle left the water and quickly attacked one FIG. 1. Sequence of predation by Hilaire’s Side-necked Turtle on pigeons in Moinhos de Vento Park, Porto Alegre City, Rio Grande do Sul, Brazil. of the birds (Fig 1A). The prey, at different times, was captured by the legs (Fig. 1B, C) or by the base of the neck (Fig. 1D). After the capture, the turtle brought the prey into the water and submerged with it (Fig. 1E). In the water, other turtles, presumably attracted by the movement and by the chance of obtaining food, also attacked the prey (Fig. 1F, G). A video showing what was reported here is available at http://csbujes.blogspot.com/. The predation of small aquatic birds by turtles is apparently occasional, and/or opportunistic in nature, however, the behavior of leaving water displayed by P. hilarri suggests an intentional act of capture not previously described. I thank the photographer of Zero Hora, Ronaldo Bernardi, for making photographic material and videos available. Thanks to the administration of the Moinhos de Vento Park; to the biologist Soraya Ribeiro; to the wildlife management group of the Secretaria de Meio Ambiente de Porto Alegre; and to the researchers in the Chelonia-RS Project of the Departamento de Zoologia da Universidade Federal do Rio Grande do Sul. Submitted by CLOVIS S. BUJES, Laboratório de Herpetologia, Departamento de Zoologia, Instituto de Biociências, Universidade Federal do Rio Grande do Sul, Av. Bento Gonçalves, 9500, Bloco IV, prédio 43435, CEP 91450-000, Porto Alegre, Rio Grande do Sul, Brazil; e-mail: chelonia_rs@hotmail.com. PSEUDEMYS RUBRIVENTRIS (Northern Red-bellied Cooter). HATCHLING BEHAVIOR. We observed the locomotion and behavior of hatchling Pseudemys rubriventris that emerged and departed from a nest at Jug Bay Wetland Sanctuary, Anne Arundel County, Maryland, USA (38.7858333°N, 76.7136111°W). At Jug Bay, Red-bellied Cooters nest in open, sunny areas 100–225 m from a large freshwater tidal wetland bordering the Patuxent River (Swarth 2004. In Swarth et al. [eds.], Conservation and Ecology of Turtles of the Mid-Atlantic Region, pp. 73–84. Bibliomania!, Salt Lake City, Utah). Turtle hatchlings often disperse quickly from the nest and may move directly to water or to dense vegetation or under leaves where they are difﬁcult to observe. Tuttle and Carroll (2005. Northeast. Nat. 12:331–348) tracked hatchling Wood Turtles (Glyptemys insculpta) as they moved from nests to water in New Hampshire; researchers in New Jersey found that hatchling Wood Turtles did not travel directly to water, but remained in a ﬁeld for days to weeks (Castellano et al. 2008. Chelon. Conserv. Biol. 7:113–118). Here we describe the post-emergence movements and behavior of Red-bellied Cooter hatchlings under natural ﬁeld conditions. The hatchlings under observation emerged from a nest discovered at 0900 h on 27 June 2007 when the female was observed laying eggs. The nest was immediately covered with a screened predator exclusion cage and was checked daily after 1 September 2007 to determine the date of hatchling emergence. The nest was in welldrained, sandy substrate next to a tractor shed, about 90 m from a large tidal wetland. The nest was partially shaded by the shed canopy roof and there was no vegetation within several meters. Following a 98-day incubation period, eight hatchlings emerged from the nest at 1715 h on 3 October 2007. Air temperature was 27°C and there had been no rain for more than six days. The hatchlings were contained at the nest under the predator exclusion cage until the next morning. (Upon excavation, the nest held ﬁve dead, Herpetological Review 40(3), 2009 337 under-developed hatchlings and three intact, unfertilized eggs). On 4 October, we released four hatchlings at 1000 h and four at 1100 h. Air temperature during the observations varied from 21°C to 26°C. We glued a yellow, 10 cm long bristle in an upright position to the carapace of each hatchling to help us observe them as they moved through the grass. However, we removed the bristles when they became entangled in vegetation a short distance from the nest. We then afﬁxed a thin, 10 cm long strip of colored plastic ﬂagging to the rear of each shell. We observed the hatchlings from a distance of 25 m in order to minimize observer effect. We noted the hatchlings’ rate of movement and behavior as they crossed the ﬁeld heading for the shrub/sapling area where we lost sight of them in the leaf litter. The terrain between the nest and the wetland consisted of 30 m of open mowed ﬁeld; 30 m of level, shady shrub/sapling habitat with ample leaf litter; and 30 m of sloping forest-covered bluff extending downhill to the wetland. The hatchlings alternated between short bursts of rapid walking and periods of rest and concealment. As they walked across the 30 m of mowed ﬁeld most of the hatchlings moved steadily for 5–10 min., then rested in taller grass clumps before moving on. Prior to reaching the forest edge, two of the hatchlings burrowed into the ground for several hours before moving on. For six hatchlings, we measured path length, amount of time required to cover the distance from nest to forest edge, rate of movement, and maximum speed. All walking paths were N or NW away from the nest site directly toward the wetland. Mean path length across the mowed ﬁeld for each of the six hatchlings walking from nest to forest was 29.7 m (SD = 6.8 m, range: 21.9–40.8 m). Mean travel time to cross the ﬁeld was 134 min. (SD = 73 min., range: 40–244 min.). Mean rate of movement was 17.6 m/h (range 6.6–38.7 m/h). Maximum speed was 38.7 m/h. For aquatic turtle species that deposit eggs in upland areas, hatchlings are quite vulnerable to predation as they move from the nest toward water. Hatchlings minimize this risk by moving quickly across exposed ﬁelds or lawns. Upon emergence, Red-bellied Cooter hatchlings in our study area generally leave the nest site immediately and invariably walk directly toward the tidal wetlands. However, predators may also be alerted to the movement of a rapidly walking hatchling. Hatchlings may minimize the risk of predation by alternating between bursts of relatively rapid movement followed by periods of rest where they are immobile. Additionally, the variability in the pattern of movement and stasis that we observed, wherein some hatchlings crossed the ﬁeld quickly while others spent several hours making the journey, suggests that behavioral heterogeneity among nest mates might reduce predation risk. Each hatchling under observation walked in almost the same direction towards the wetlands. It is possible that hatchlings move towards bright light from the sky that is visible on the horizon and avoid moving toward areas that are dark or shady. The sky was visible and bright at ground level through the trees and shrubs to the northwest of the nest. A sky that is visible through vegetation may be a cue to hatchlings that the terrain drops off towards lower elevations. By walking in this direction hatchlings would travel downhill, eventually reaching the safety of a stream, pond, or wetland. 338 Submitted by RACHAEL DICKEY, SUSAN MATTHEWS, and CHRISTOPHER W. SWARTH, Jug Bay Wetlands Sanctuary, 1361 Wrighton Rd., Lothian, Maryland 20711, USA (e-mail: Cswarth@jugbay.org). TERRAPENE CAROLINA MAJOR (Gulf Coast Box Turtle). SWIMMING. With the exception of Terrapene coahuila (Cohuilan Box Turtle), members of the genus Terrapene have been considered to be primarily terrestrial (Ernst et al. 1994. Turtles of the United States and Canada. Smithsonian Institution Press, Washington, D.C., pp. 250–265). The occasional use of aquatic habitats, perhaps for drinking or thermoregulation, is often noted (Bonin et al. 2006. Turtles of the World. John Hopkins Univ. Press, Baltimore, Maryland. 416 pp; Dodd 2001. North American Box Turtles: A Natural History. Univ. Oklahoma Press, Norman, Oklahoma. 231 pp.), but instances of actual swimming are less common (McCauley 1945. The Reptiles of Maryland and the District of Columbia. Privately printed, Hagerstown, Maryland. 194 pp.; Stickel 1950. Ecol. Monogr. 20:361; Tyler 1979. Southwest. Nat. 24:189–190; McDowell et al. 2004. Herpetol. Rev. 35:265–266). Terrapene c. major occupies coastal wetland or near-wetland habitats in the southeastern USA and will walk and forage on the bottoms of ponds and canals (Bartlett and Bartlett 1999. A Field Guide to Florida Reptiles and Amphibians. Gulf Publ., Houston, Texas. 280 pp.). Here we report a long-distance swimming event for this subspecies. On 28 June 2008, at ca. 0900 h, we observed a male T. c. major swimming across the East Fowl River in Mobile Co., Alabama, USA. The turtle was sighted by boat in the center of the river, swimming in a northwesterly direction away from the south bank. The turtle was swimming at the surface with its head up and approximately two-thirds of its carapace above the water line. Upon approach, the turtle ceased swimming and began to ﬂoat. We collected the turtle to verify the identiﬁcation and sex. The turtle was photographed and then returned to the water. As we moved away, the turtle continued to swim towards the north bank. The width of the river at this location was measured, using ArcGIS, to be ca. 320 m. We observed the turtle for 15 minutes until it came within 20 m of the north bank, at which time we continued up the river. Swimming and river crossings have been previously reported for subspecies of T. carolina (references, op. cit. from above), and T. ornata is capable of swimming (Richtsmeier et al. 2008. Chelon. Conserv. Biol. 7:3–11), but the distances reported for these events are generally short (< 20 m), and large rivers have been considered as possible barriers to dispersal and gene ﬂow (Richtsmeier et al., op. cit.). In the instance described above, the circumstances resulting in this animal swimming such a long distance are unknown. However, the ability to swim across large bodies of water has implications for gene ﬂow between populations that would otherwise be considered geographically isolated. We thank James Lee, David H. Nelson, and Ashley B. Morris for their comments. Submitted by THOMAS G. JACKSON, JR. (e-mail: jacksontg@hotmail.com), and JUAN E. LOO (e-mail: juancho_0113@hotmail.com), Department of Biology, University of South Alabama, Mobile, Alabama 36688, USA. Herpetological Review 40(3), 2009 SQUAMATA — LIZARDS AMEIVA AMEIVA (Green Lizard). ATTEMPTED PREDATION OF AMPHISBAENIAN. Ameiva ameiva (Teiidae) is a moderate-sized (to 190 mm SVL), terrestrial lizard that occurs in nearly all tropical habitats of South America (Pianka and Vitt 2003. Lizards: Windows to the Evolution of Diversity. University of California Press, Berkeley, California. 333 pp.). It is an active forager with a generalist diet that includes plants and vertebrates, though dominated by arthropods (Vitt and Colli 1994. Can. J. Zool. 72[11]:1986–2008). On 19 October 2004 at 1438 h, in an area of Cerrado in the municipality of Bauru, São Paulo State, Brazil (22.340690°S 49.018190; datum: WGS84; elev. 580 m) we observed an adult A. ameiva (ca. 150 mm SVL) attacking a Spade-snouted Worm-Lizard (Leposternon microcephalum) ca. 300 mm total length (Fig. 1). During a period of 20 minutes, we observed the Ameiva repeatedly shake the L. microcephalum from side to side, occasionally releasing its grip and then biting again. Eventually, the Ameiva released the L. microcephalum and retreated to a hole in a shaded ravine. We examined the amphisbaenian and noted small bite marks but the skin was not torn; it seemed exhausted and moved slowly. The latter was collected and added to collection of the House of Reptiles of Bauru Zoo. Records of predation of L. microcephalum include a coralsnake (Micrurus corallinus) and a mammal (Nasua nasua) (Marques and Sazima 1997. Herpetol. Nat. Hist. 5:88–93; Oliveira et al. 2004. Herpetol. Rev. 35:170–171). This is the ﬁrst report of predation of L. microcephalum by A. ameiva and the ﬁrst occurrence of predation by this species on an amphisbaenian. Submitted by FLÁVIO KULAIF UBAID (e-mail: flavioubaid@yahoo.com.br), GERSON RODRIGUES DO NASCIMENTO, and FÁBIO MAFFEI, Departamento de Zoologia, Instituto de Biociências, Universidade Estadual Paulista, 18618-000, Botucatu, São Paulo, Brazil. ASPIDOSCELIS COSTATA (Western México Whiptail). CANNIBALISM. Cannibalism is common among reptiles (Mitchell 1986. SSAR Herpetol. Circ. No. 15, 37 pp.) and has been recorded in a number of lizard species (e.g., Castilla and Van Damme 1996. Copeia 1996:991–994; Moll and Koenig 2003. Sonoran Herpetologist. 16:90–91; Keren-Rotem et al. 2006. Behav. Ecol. Sociobiol. 59:723–731). The teiid genus Aspidoscelis includes insectivorous lizards (Anderson 1993. In Wright and Vitt [eds.], Biology of Whiptail Lizards (Genus Cnemidophorus), pp. 83–116. Oklahoma Mus. Nat. Hist., Norman; Vitt and Pianka 2004. In Pérez-Mellado et al. [eds.], The Biology of Lacertid Lizards, pp. 139–157. Evolutionary and Ecological Perspectives. Institut Menorquí d’Estudis, Recerca. Lesvos, Greece). Though the diet of Aspidoscelis spp. is often characterized as comprising mainly insects, some records exist of cannibalistic behavior, especially among temperate zone species (Etheridge and Wit 1982. Herpetol. Rev. 13:19; Mitchell 1979. Can. J. Zool. 57:1487–1499). However, as no reports of cannibalism exist for A. costata, we report an instance of cannibalism in this species. On 18 October 2008 (rainy season), MAGR captured a juvenile male A. costata (60 mm SVL, 6.6 g) at Mezcala, Guerrero (17.9101°N, 99.6148°W, datum: WGS84; 548 m elev.) in southern México. Dominant vegetation is tropical dry forest with a wet season from July to October and a dry season from November to June. Examination of its stomach contents revealed a conspeciﬁc hatchling (35 mm SVL, 1.0 g). The hatchling had been swallowed headﬁrst and showed little evidence of digestion, indicating that it had been consumed just before the juvenile’s capture. In shortlived animals such as small lizards, population dynamics appear to be particularly sensitive to ﬂuctuations in annual recruitment and survival of eggs and juveniles (Castilla and Vand Damme 1996. Copeia 1996:991–994). In that sense, cannibalism of hatchlings could be a mechanism to reduce intraspeciﬁc competition or a form of population regulation (Fox 1975. Ann. Rev. Ecol. Syst. 6:87–106). Both specimens (predator and prey) were deposited in Colección Nacional de Herpetología, Instituto de Biología, Universidad Nacional Autónoma de México (IBH 20202 and 20209, respectively). Submitted by MARTHA ANAHI GÜIZADO RODRIGUEZ (e-mail: anahigr@ibiologia.unam.mx), LORENA REYES VAQUERO, and GUSTAVO CASAS ANDREU, Laboratorio de Herpetología, Instituto de Biología, Universidad Nacional Autónoma de México, A.P. 70-153, México D.F. 04510, México. FIG. 1. Green Lizard (Ameiva ameiva) attacking a Spade-snouted WormLizard (Leposternon microcephalum). Photo by F. K. Ubaid. ASPIDOSCELIS MARMORATA (Western Marbled Whiptail). FACULTATIVE FRUGIVORY. Aspidoscelis marmorata inhabits semiarid environments in southern New Mexico and Texas, and in northern México in Coahuila, Chihuahua, and Herpetological Review 40(3), 2009 339 Durango (Stebbins 2003. A Field Guide to Western Reptiles and Amphibians. Houghton Mifﬂin, Boston, Massachusetts. 533 pp.). Previous studies of stomach contents have shown that insects comprise most of the diet, and that some plant material may be taken in very low quantity (Gadsden and PalaciosOrona 2000. Acta Zool. Mex. 79:61–76; Lemos-Espinal and Smith 2007. Amphibians and Reptiles of the State of Coahuila, Mexico. UNAM and CONABIO. 550 pp.) but frugivory has not been documented. This is the ﬁrst observation of intentional fruit ingestion in the species. On 2 November 2004 around 1300 h, we observed an adult A. marmorata at Mapimí Biosphere Reserve in central Chihuahuan Desert, Durango, México (26.6662°N, 103.7471°W, WGS84, elev. 1150 m) consuming a ﬂeshly, juicy, red fruit of Opuntia leptocaulis (Cactaceae). The lizard picked up the fruit from the ground in its mouth and consumed most of. Opuntia leptocaulis (Desert Christmas Cholla) occurs throughout the Chihuahuan Desert. Their ﬂeshy, highly attractive fruits are common and ripen during winter, and are frequently consumed by a variety of vertebrates, especially birds and small mammals (Goettsch and Hernández 2006. J. Arid Environ. 65:513–528). In xeric habitats, food availability is frequently unpredictable and scarce, so opportunistic feeding could shape the diets of many lizard species (Whiting and Greeff 1997. Copeia 1997:811–818). Fruiting O. leptocaulis are food resources that are widely available in late fall. Fruit consumption by Aspidoscelis lizards could be an interesting issue for the study of facultative frugivory. Submitted by CLAUDIA BALLESTEROS-BARRERA (e-mail: claudiab@ibiologia.unam.mx), CONSTANTINO GONZÁLEZ-SALAZAR, Laboratorio de Análisis Espaciales, Instituto de Biología, Universidad Nacional Autónoma de México, A.P. 70-153, México D.F. 04510; and HÉCTOR GADSDEN, Instituto de Ecología, A. C.-Centro Regional Chihuahua, Km. 33.3 Carretera, Chihuahua-Ojinaga, Ciudad Aldama, Chihuahua, México, C.P. 32900. ASPIDOSCELIS TESSELATA (Checkered Whiptail) × ASPIDOSCELIS SEXLINEATA VIRIDIS (Prairie Racerunner). REPRODUCTIVE POTENTIAL. Pattern class E (Zweifel 1965. Am. Mus. Novit. 2235:1–35) of diploid (Lowe et al. 1970. Syst. Zool. 19:114–127) parthenogenetic Aspidoscelis tesselata (sensu Reeder et al. 2002. Am. Mus. Novit. 3365:1–61), derived from hybridization between A. marmorata (= tigris) marmorata × A. gularis septemvittata (Parker and Selander 1976. Genetics 40:245–252; Cordes and Walker 2006. Copeia 2006:14–26), is syntopic with gonochoristic A. sexlineatus viridis and parthenogenetic A. neomexicana near the railroad depot in the city of Fort Sumner, De Baca Co., New Mexico, USA (34.47389°N, 104.23944°W; datum: WGS84; elev. 1250 m). On 12 July 2002, GJM collected a male lizard (University of Arkansas Department of Zoology, UADZ 7619) at this site possessing characters of scutellation and components of the dorsal and ventral color patterns that appeared to be phenotypically derived from hybridization between A. tesselata and A. sexlineata viridis. Analyses of morphological characters in the apparent hybrid and samples of the putative parental species from Fort Sumner (Manning and Walker 2006. Am. Midl. Nat. 340 2006. 155:411–416) conﬁrmed that UADZ 7619 resulted from a haploid sperm of A. sexlineata viridis (1n = 23) fertilizing an unreduced egg of A. tesselata (2n = 46). Though this is the ﬁrst reported example of this hybrid combination, A. tesselata E (= pattern class EC of Taylor et al. 2003. Am. Mus. Novit. 3424:1–40) has been identiﬁed as a participant in frequent hybridizations with its maternal progenitor A. marmorata marmorata at a site in New Mexico (Taylor at al. 2001. Am. Mus. Novit. 3345:1–65). Based on Walker et al. (1989. Copeia 1989:1059–1064; 2006. Herpetol. Rev. 37:344–345) and Taylor et al. (2001, op. cit.), JMW described the gross reproductive morphology of the preserved hybrid (UADZ 7619). Based on Goldberg and Beaman (2003. Herpetol. Rev. 34:143), SRG histologically examined the left testis and left epididymis of the hybrid and assessed its previous reproductive functionality. The hybrid was 70 mm in SVL. Compared to males of A. sexlineata viridis of similar size from Fort Sumner, its testes (mean dimensions of both = ca. 3.8 × 6 mm), epididymides, and hemipenes were of expected size and structure. Histology of the left testis indicated that UADZ 7619 was undergoing early spermiogenesis when collected. As reported for other teiid lizards (Lowe and Goldberg 1966. J. Morphol. 119:277–281); a circumtesticular tunic 1–3 Leydig cells thick surrounded the left testis. Clusters of metamorphosing spermatids were present and tails of spermatozoa projected into the lumina of the seminiferous tubules. Sperm were also present in the sectioned epididymis. This A. tesselata × A. sexlineata viridis hybrid would have been deemed fertile based on cytological characteristics of testes in species such as A. tigris (Lowe and Goldberg, op. cit.). However, though it seemed capable of producing sperm and inseminating females of A. tesselata, A. neomexicana, and A. sexlineata viridis at Fort Sumner, its ability to produce a stable haploid chromosome complement from a triploid number of 69 is dubious. Nevertheless, the hybrid likely would have interfered with normal reproduction in some females of these three species should any of them have been inseminated. Taylor et al. (2001, op. cit.) provided morphological and histological descriptions of gonads and ducts of both male and female hybrids of A. tesselata × A. marmorata collected at Macho Draw, Chaves Co., New Mexico, and provided indirect evidence that some hybrid males (N = 12) may have inseminated congeneric females at this site. Collection of UADZ 7619 was made under authority of Permit No. 3118 issued to GJM in 2002 by the New Mexico Department of Game and Fish. Submitted by GLENN J. MANNING, Department of Biological Sciences, University of Arkansas, Fayetteville, Arkansas 72701, USA (e-mail: gmannin@ uark.edu); JAMES M. WALKER, Department of Biological Sciences, University of Arkansas, Fayetteville, Arkansas 72701, USA (e-mail: jmwalker@uark. edu); and STEPHEN R. GOLDBERG, Department of Biology, Whittier College, Whittier, California 90608, USA (e-mail: sgoldberg@whittier.edu). CALOTES “VERSICOLOR” (Burmese Garden Lizard). GLIDING. During surveys of the Burmese herpetofauna (NSF DEB9971861, DEB-0451832), we periodically sampled the components Herpetological Review 40(3), 2009 of the Hlawga Wildlife Park fauna (17.043°N, 96.1271667°E; Yangon Division, Myanmar). On 28 February 2009, we focused on the lizard C. “versicolor” (Quotes indicate the population is a member of this species group; true C. versicolor does not occur in Myanmar.), and discovered that members of the Hlawga population glide. Hlawga C. “versicolor” are wary and typically take evasive action when a person approaches within 4–5 m of “their” tree. These lizards are sit-and-wait predators, and typically cling to the sides of trees, usually 1–2 m from the tree base. When a potential predator approaches, evasive action begins with a slight upward movement around the tree, apparently in an effort to escape detection. If pursued, even slowly, they continue to circle the tree trunk, constantly moving upward. Our observation concerns an adult on a tree isolated in a grassy patch. As we approached, the lizard immediately climbed to nearly 3 m. Being the dry season, the tree was leaﬂess. A park ranger decided to aid capture by climbing the tree. The lizard continued to the top (about 12 m); at about 10 m, the tree was too slender for the ranger to reach the lizard, so he chopped off the tree top and moved slowly and held the top horizontal. We expected the lizard to simply drop directly downward; instead it leaped outward and descended at roughly 45°, landing in a brush pile and eluding capture. The descent angle alone does not imply gliding, but in association with the lizard’s deliberate vertical outward leap and its posture of straight body and head with outstretched fore- and hindlimbs, dorsoventrally ﬂattened trunk, and tail held straight backward (rigidly so, i.e., no movement), the controlled descent can be deﬁned as gliding behavior. Gliding is certainly not an unknown agamid behavior owing to the spectacular gliding performance of the various species of Draco, but it has not been reported in other agamid taxa except Bronchocela cristatella (Allen 1957. Malayan Nat. J. 11:81; Reid 1958. Malayan Nat. J. 12:119). Both of these reports note that the lizard was observed to leap outward and descent vertically from its origin at a roughly 40–45° angle. We suggest that these and other species of lighter-bodied arboreal agamids may use this mode of escape, but that it would be rarely seen and uncommonly used because these lizards are often successful at eluding capture by climbing, hiding, or jumping short distances to other trees. Submitted by GEORGE R. ZUG, Department of Vertebrate Zoology, National Museum of Natural History, PO Box 37012, Washington, DC 20013-7012, USA (e-mail: zugg@si.edu); JENS V. VINDUM, Department of Herpetology, California Academy of Sciences, Golden Gate Park, San Francisco, California 98000, USA; JEREMY F. JACOBS, Department of Vertebrate Zoology, National Museum of Natural History, PO Box 37012, Washington, DC 20013-7012, USA; KYI SOE LWIN and YE MYO WIN, Nature and Wildlife Conservation Division, Forestry Department, Naypyidaw, Myanmar. CELESTUS CURTISSI (Curtis’ Galliwasp). REPRODUCTION. Celestus curtissi is restricted to Hispaniola where it is known from Haiti and the Dominican Republic (Schwartz and Henderson 1991. Amphibians and Reptiles of the West Indies. Descriptions, Distributions, and Natural History. Univ. Florida Press, Gainesville. 720 pp.). It is ovoviviparous with brood sizes of 2–5 (Schwartz and Henderson 1991, op. cit.). The purpose of this note is to report a new minimum brood size for C. curtissi. Two female C. curtissi from the Dominican Republic collected August 1976, were examined from the herpetology collection of the Natural History Museum of Los Angeles County (LACM). Both (LACM 125351, SVL 63 mm; LACM 125354, SVL 64 mm) were from Pedernales Province, 5 km NE of Oviedo (17.7833°N, 71.3666°W, datum: WGS 84; elev. 3 m). A mid-ventral incision was made in the lower abdomen and the ovaries were examined. One well-developed embryo was observed in LACM 125354 and one oviductal egg was in LACM 125351. Ober (1970. Herpetol. 26:275) reported one C. (as Diploglossus) curtissi gave birth to ﬁve young in September. One embryo represents a new minimum brood size for C. curtissi. I thank Christine Thacker (LACM) for permission to examine C. curtissi. Submitted by STEPHEN R. GOLDBERG, Whittier College, Department of Biology, Whittier, California 90608, USA; e-mail: sgoldberg@whittier.edu. CYCLURA CYCHLURA INORNATA (Allen Cays Rock Iguana). CAUSE OF MORTALITY. Jetsam, ﬂotsam, and other discarded material has been shown to be a source of mortality for reptiles (e.g., Laist 1997. In Coe and Rogers [eds.], Marine Debris: Sources, Impacts, and Solutions, pp. 99–139. Springer-Verlag, New York; Benedict and Billeter 2004. Southeast. Nat. 3:371–377; Mascarenhas et al. 2004. Mar. Poll. Bull. 49:354–355). We report on jetsam-related sources of mortality for the Cyclura cychlura inornata. The Allen Cays Rock Iguana is a CITES-listed species restricted to a few small cays in the Exuma Islands of the Bahamas (Iverson et al. 2005. Cat. Amer. Amphib. Rept. 810:1–9). It has few, if any, natural predators as adults, and relatively high adult and juvenile survivorship (Iverson et al. 2006. Biol. Conserv. 132:300–310; Iverson 2007. Copeia 2007:740–744). Documented sources of mortality include accidental death (Iverson et al. 2005. Herpetol. Rev. 36:175), and there is some concern about the possibility of poaching or other human- or tourism-related stressors (Iverson et al. 2006. Biol. Conserv. 132:300–310; Smith and Iverson 2006. Can. J. Zool. 84:1522–1527). We observed another possible cause of mortality related to oceanic refuse of jetsam causing the death of four juvenile Allen Cays Iguanas. On 15 May 2008, three juvenile iguanas were found dead inside a large 50-gallon (189 liter) plastic drum on a cay near the Allen Cays (exact location not provided for conservation purposes). The drum had apparently washed up on the coastline, probably within the previous year. There were two small openings on the top of the drum through which the juveniles apparently entered (Fig. 1A). All three iguanas were about the same size (SVL ≈ 14 cm, 20 cm, 21 cm) and at a similar state of decomposition (Figs. 1B, 1C), suggesting that they may have been using the drum as a communal retreat until wind or waves reoriented the drum and prevented the lizards’ escape. The three iguanas were all clearly juveniles based upon their size (Fig. 1C). One juvenile iguana was found dead in a small upright freezer half ﬁlled with rain water on Southwest Allen’s Cay (U Cay) (24.75°N, 76.84°W; see Iverson et al. 2004. Herpetol. Monogr. Herpetological Review 40(3), 2009 341 FIG. 2. A) The upright freezer partially ﬁlled with rain water; and B) the remains of one juvenile Allen Cays Iguana (Cyclura cychlura inornata) found in the water inside the freezer. Photos by Lynne Pieper. that the individual was a juvenile (Fig. 2B). Although the mortality events that we observed are likely rare, it does suggest that material washing up on the shores of these small cays can be an additional source of mortality, especially for smaller and younger iguanas, and potentially other native wildlife. We thank Bruce Dunham and Will McLean of the M/V Bahama Star for help opening the barrel. Funding for this project was provided by Earlham College, the Cope Museum Fund, and the Test Fund. Permits for research on the Allen Cays Iguana were provided by the Bahamas Department of Agriculture to JBI. FIG. 1. A) The 50-gallon barrel as originally discovered; B) the remains of the three juvenile Allen Cays Iguana (Cyclura cychlura inornata) as found inside the barrel; and C) the remains of the three iguanas after removal from the barrel. Photos by Kirsten Hines (A), Lynne Pieper (B), and Stesha Pasachnik (C). 18:1–36 for a detailed description) on 16 May 2008 (Fig. 2A). The iguana had apparently fallen into the water and drowned. Few remains were found, but based on their size, it is highly probable 342 Submitted by GEOFFREY R. SMITH, Department of Biology, Denison University, Granville, Ohio 43023, USA (e-mail: smithg@denison.edu); KIRSTEN HINES, The Institute for Regional Conservation, 22601 SW 152 Ave, Miami, Florida 33170, USA (e-mail: hines@regionalconservation.org); STESHA PASACHNIK, Department of Ecology and Evolutionary Biology, University of Tennessee, Knoxville, Tennesse 37996, USA (e-mail: spasachn@utk.edu); LYNNE PIEPER, College of Education, University of Illinois at Chicago, Chicago, Illinois 60607, USA (e-mail: lypieper@juno.com); ERIKA PHELPS and JOHN B. Herpetological Review 40(3), 2009 IVERSON, Department of Biology, Earlham College, Richmond, Indiana 47374, USA (e-mail: johni@earlham.edu) EUTROPIS CARINATA (Common Skink), EUTROPIS MACULARIA (Rock Skink). ENDOPARASITES. Eutropis carinata, a terrestrial, diurnal skink, occurs in a wide variety of habitats in Sri Lanka, India, Bangladesh, and Nepal (Das and De Silva 2005. A Photographic Guide to Snakes and Other Reptiles of Sri Lanka. Ralph Curtis Publ., Sanibel Island, Florida. 144 pp.). Eutropis macularia, a semi-fossorial skink, occurs in Sri Lanka, India, Pakistan, Nepal, Bangladesh, Bhutan, east to mainland southeast Asia (Das and de Silva, op. cit.). There are, to our knowledge, no published records of helminths from these species. The purpose of this note is to establish the initial helminth lists for E. carinata and E. macularia. One male E. carinata (SVL = 110 mm, Christopher C. Austin = CCA 2364, collected August 2002 at Buttala, Monaragala District, Central Province, Sri Lanka [6.6813°N, 81.2705°E, datum: WGS84], elev. 125 m) and one female E. macularia (SVL = 68 mm, CCA 1771, collected November 2002 at Batamdomba Cave, Rathapura District, Central Province, Sri Lanka [6.7797°N, 80.3969°E, WGS84], elev. 114 m) were examined for helminths. Lizards were sacriﬁced within 12 h of capture, preserved in 10% formalin and stored in 70% ethanol. The digestive tract was later removed, opened, and searched for helminths under a dissecting microscope. The three female nematodes found in their stomachs (1 in E. carinata and 2 in E. macularia), were removed, cleared in a drop of glycerol on a glass slide, cover-slipped, studied under a compound microscope and identiﬁed as Physalopteroides dactylurus. Nematodes were deposited in the United States National Parasite Collection, Bethesda, Maryland, USA as: E. carinata (USNPC 101175) and E. macularia (USNPC 101176). Lizards were deposited in the herpetology collection of the National Museum of Sri Lanka, Colombo, Sri Lanka. Physalopteroides dactylurus is a member of the Physalopertidae, which utilizes insects as intermediate hosts (Anderson 2000. Nematode Parasites of Vertebrates. Their Development and Transmission. CABI Publishing, Oxford, UK. 650 pp.). Eutropis likely become infected through diet. Eutropis carinata and E. macularia are new host records for P. dactylurus, which is also known from Calotes versicolor, Hemidactylus ﬂaviviridis, both collected in India, and Eumeces taeniolatus (currently Eurylepis taeniolatus) collected in Turkmenistan (Baker 1987. Synopsis of the Nematoda Parasitic in Amphibians and Reptiles. Memorial University of Newfoundland Occas. Pap. Biol., St. John’s Newfoundland, Canada. 325 pp.) We thank the Department of Wildlife Conservation, Sri Lanka, for scientiﬁc collecting permit (number WL/3/2/1/14/12). This research was funded in part by the People’s Trust for Endangered Species and National Science Foundation grant DEB 0445213 to CCA. Sarah Goldsberry assisted with dissections. Submitted by STEPHEN R. GOLDBERG, Department of Biology, Whittier College, Whittier, California 90608, USA (e-mail: sgoldberg@whittier.edu); CHARLES R. BURSEY, Department of Biology, Pennsylvania State University, Shenango Campus, Sharon, Pennsylvania 16146, USA (e-mail: cxb13@psu.edu); INDRANEIL DAS, Institute of Biodiversity and Environmetal Conservation, Universiti Malaysia Sarawak, 94300, Kota Samarahan, Sarawak, Malaysia (e-mail: idas@ibec.unimas.my); ANSELM DE SILVA, Amphibian and Reptile Research Organization of Sri Lanka, 15/1 Dolosbage Road, Gampola, Sri Lanka (e-mail: kald@sitnet.lk); and CHRISTOPHER C. AUSTIN, Department of Biological Sciences and Museum of Natural Science, Lousiana State University, Baton Rouge, Louisiana 70803, USA (e-mail: ccaustin@lsu.edu). HELODERMA SUSPECTUM (Gila Monster). DIET AND PREDATORY BEHAVIOR. Heloderma suspectum is a strict carnivore and nest-raiding specialist that shows geographic variation in its diet. Typical prey selection includes eggs of groundnesting birds and lizards, as well as small mammals, commonly the neonates of rodents (reviewed by Beck 2005. Biology of Gila Monsters and Beaded Lizards. University of California Press, Berkeley and Los Angeles. 247 pp.; Geiger and Tracy 2008. Herpetol. Rev. 39:225–226.). Here, we provide additional information on the diet and predatory behavior of adult H. suspectum from a population in the Sonoran Desert of south-central Arizona, USA. The individuals we discuss herein were subjects of a radio-telemetric study in which various features of their behavior, physiology, and spatial ecology have been investigated since March 2001 (e.g., Kwiatkowski et al. 2008. J. Zool. doi:10.1111/j.1469-7998.2008.00495.x). The present study site, located in Pinal County, is 40 km SSE of the city of Florence, 8 km W of State Route 79, and encompasses an area of ≈ 3 km2 at the extreme western edge of the Suizo Mountains (Iron Mine Hill). The region is ecologically designated as Arizona Upland Desertscrub subdivision (Brown 1994. Biotic Communities of the American Southwest—United States and Mexico. University of Utah Press, Salt Lake City. 342 pp.). Annual precipitation (chieﬂy rain) patterns are bimodal, with slight to moderate storms occurring during winter and early spring (December to March), and the strongest activity occurs from mid- to late summer (early July to mid-September), which is termed the North American monsoon (Brown 1994, op. cit.; Phillips and Comus 2000. A Natural History of the Sonoran Desert. The University of California Press, Berkeley and Los Angeles. 628 pp.). The area lacks permanent freestanding water, a common feature of the Sonoran Desert; at the present site, collection of rainwater in surface rocks and soil, as a rule, is transitory, and persists for only minutes to several hours. However, artiﬁcial alterations of the habitat (e.g., deep tire tracks in narrow dirt roads) occasionally provide a substantive reservoir for the collection and storage of rainwater, but even under these situations it is ephemeral, especially in summer. On 27 April 2003, at 0923 h, in an area interfacing bajada (Phillips and Comus 2000, op. cit.) and desert ﬂats, a female H. suspectum (HS-2: SVL = 320 mm; TL = 135 mm; body mass = 573 g) was radio-tracked and located. This individual was originally collected on 1 April 2001, and had been studied regularly since that time. She was found (Site 55, 813 m elev.) beside an active nest of Gambel’s Quail; 8 clustered, normal-sized eggs were present (Fig. 1A). The nest was a shallow soil depression, located centrally in a network of Engelmann’s Prickly Pear, Opuntia engelmannii (200 L × 100 W × 80 H cm). Nearby (3 m diam. Herpetological Review 40(3), 2009 343 from nest) vegetation was sparse and limited to Triangle Bursage and Pincushion. Her core body temperature (obtained via an implanted 11.0 g temperature-sensitive radio-transmitter; Holohil Systems Ltd., Ontario, Canada) was 30.5ºC. Ambient temperature (1 m above the ground in shade) was 29.5ºC. Cloud cover was ≈ 10%, relative humidity was ≈ 15%, and wind speed was slight (0–5 kph) and intermittent. The eight clustered eggs were inspected by HS-2 via nose-rubbing and tongue-ﬂicking. From 0923 h to 0937 h, seven eggs were consumed at a seemingly rapid rate (0.5 eggs/min), and from 0937 h to 0947 h, the last egg was consumed. From the time the ﬁrst egg was secured in her jaws, the eight eggs were consumed in 24 minutes. None of the eggs were broken prior to swallowing; each egg was slowly and gently secured in her jaws (Fig. 1A) without the aid of forelimbs. During ingestion, when the eggs were no longer visible in her mouth, cracking sounds were audible from a distance up to 3 m. When ingestion of the eggs was completed, she moved away from this site and was not re-located for 7 days. The previous day (26 April 2003), at 1800 h, HS-2 was located at Site 2 (elev. 826 m), a large triangular rock slab, which was located in the western-most area of the Suizo Mountain complex (Iron Mine Hill); she demonstrated ﬁdelity to Site 2 on multiple occasions since using it ﬁrst on 13 April 2001. The minimum distance traveled between the two sites (2 and 55) was 215 m and accomplished in < 15 h. On 11 May 2004, at 1831 h, in bajada, a male H. suspectum (HS-7: SVL = 334 mm; TL = 151 mm; body mass = 673 g) was radio-tracked and located (Site 19, elev. 821 m). This individual FIG. 1. Predation by adult wild-living Gila Monsters, Heloderma suspectum. A) An adult female (HS-2) at the nest of a Gambel’s Quail in the process of securing an egg to ingest. Eight eggs were observed at the nest and all were ingested. B–D) An adult female (HS-13) in various stages of consuming neonates of Desert Cottontails (Sylvilagus audubonii). B) Female HS-13 was ﬁrst located entering a burrow. C) Female HS-13 is next to the ﬁrst neonatal rabbit (within the yellow ellipse). Note the extensive torn skin in the area of the neck. D) The neonate in Fig. 1C is being consumed head-ﬁrst. All photographs by R. A. Repp. 344 Herpetological Review 40(3), 2009 was originally collected on 23 February 2003, and had been studied regularly since that time. He was found 30 cm W of an active Gambel’s Quail nest; 6 clustered, normal-sized eggs were present. He appeared to be moving away from the nest at our approach, likely as a result of our sudden presence. The nest was located on the west side of a stand of Engelmann’s Prickly Pear, which was circular-shaped (110 H cm × 200 cm in diam.). Nearby (3 m diam. from the nest) vegetation was Triangle Bursage, Buckhorn Cholla (Opuntia acanthocarpa), and Pincushion. His core body temperature (obtained via a 11.0 g temperature-sensitive radio-transmitter; Holohil Systems Ltd) was 31.3ºC. Ambient temperature was 30ºC, and ground temperature in the sun (“hot spot”) was 34.0ºC. Cloud cover was 0%, relative humidity was ≈ 10%, and wind speed was slight to moderate (5–20 kph) and intermittent. Because our presence seemed to distress HS-7, we promptly left this site. At 1947 h, we radio-tracked HS-7 a second time; he was relocated at a previously used location (Site 18, elev. 826 m) on the western edge of Iron Mine Hill, which was 37 m NW of Site 19. His core body temperature was 28.8ºC. Ambient temperature was 25ºC. Wind speed was slight (0-5 kph) and intermittent. At 1956 h, we returned to Site 19 and inspection revealed that all 6 eggs were gone; we thus presumed they had been consumed by HS7. Similar to our observations of HS-2, we found no signs that the eggs had been broken during feeding (e.g., no yolk or shell fragments). On 9 August 2008, at 0820 h, in bajada, a female H. suspectum (HS-13: SVL = 310 mm; TL = 146 mm; body mass = 466 g) was radio-tracked and located (Site 9, elev. 825 m). This individual was originally collected on 24 May 2008, and had been studied regularly since that time. She was found with her head and midbody in a north-facing burrow (Fig. 1B) that was 75 mm diameter. The burrow was situated between two clumps of Triangle Bursage (each ≈ 400 m height). Nearby plants included Pincushion. Her core body temperature (obtained via an implanted 11.0 g temperature-sensitive radio-transmitter; Holohil Systems Ltd) was 35.5ºC. Ambient temperature was 29.5ºC and the hot spot was 41.0ºC. Cloud cover was ≈ 10 %, relative humidity was ≈ 30%, and wind speed was slight (0–5 kph) and intermittent. Mufﬂed vocalizations (repetitious cat-like mews, 1 sec apart, presumed from a single individual) were heard emanating from the burrow; these calls were distinctive and could be heard several meters from the site. At 0821 h, her tail was grabbed, and she was quickly removed from the burrow and placed on the ground ≈ 1 m from the burrow. During this transfer, a neonatal Desert Cottontail (Sylvilagus audubonii), estimated to be 40–50 g, was gripped in her jaws but was dropped as she was placed on the ground. For about 1 min, the neonate was writhing and rapidly vocalizing while HS-13 re-located and inspected it; the neonate had severe lacerations with oozing ﬂuids (Fig. 1C). At 0822 h, HS-13 began ingesting the neonate headﬁrst (Fig. 1D), which was completed in 3 min. At 0825 h, HS-13 returned to and stayed at the nest; she entered to midbody. Mufﬂed vocalizations from a single neonate were heard; at 0830 h, the vocalizations stopped. Apparently, the neonate producing the vocalizations had been consumed. At 0831 h, with no further signs of predatory activity, HS-13 was gently removed from the burrow a second time, and it was noted that she had rabbit fur clinging to her jaws. The nest was gently and non-invasively inspected at 0832 h. It was shallow (120 mm), the cavity was larger than the entrance, and the substrate was lined with fur (presumably of the mother) and grasses (Bowers et al. 2004. Mammals of North America. Houghton Mifﬂin Co., New York. 352 pp.). During this assessment, no further neonates were detected. HS-13 re-entered the nest at 0833 h, inspected it for several minutes, then exited at 0837 h, traveling N of the burrow; she was not re-located until a later date (16 August 2008). We cannot determine whether HS-13 bit and envenomated the ﬁrst neonate prior to our grabbing and removing her from the burrow/nest (see Beck 2005, op. cit.). Clearly, she was startled by this procedure, and there is little doubt that the neonate was damaged (and possibly envenomated) by this intrusion. Nonetheless, based on the presence of vocalizations prior to our disturbances, it is possible that biting and envenomation occurred prior to our arrival. Similarly, the second neonate we heard vocalizing from the burrow/nest might have been bitten and envenomated. Furthermore, although we show that HS-13 consumed at least two neonates, it is possible that there were others eaten because litter size in S. audubonii is typically greater than two (Beck 2005, op. cit.). Support for our ﬁeld studies at the Suizo Mountains are from Arizona State University, Zoo Atlanta, Georgia State University, and David L. Hardy, Sr. Since 2001, many individuals provided assistance in radio-tracking, but most noteworthy are Hans-Werner Herrmann and Ryan Sawby. Also, Ryan Sawby provided invaluable assistance with photography and identiﬁcation of plants and invertebrates. This study was approved by the animal care committee (IACUC) of Arizona State University (98-429R), and appropriate scientiﬁc permits were obtained from the Arizona Game and Fish Department. Submitted by ROGER A. REPP, National Optical Astronomy Observatory, 950 North Cherry Avenue, Tucson, Arizona 85719, USA (e-mail: repp@noao.edu); and GORDON W. SCHUETT, Department of Biology and Center for Behavioral Neuroscience, Georgia State University, 33 Gilmer Street, Unit 8, Atlanta, Georgia 30303-3088, USA (e-mail: biogws@langate.gsu.edu, or gwschuett@yahoo.com). HEMIDACTYLUS FRENATUS (Common House Gecko). BROOD. On 9 February 2008, two gravid female Hemidactylus frenatus (SVL = 49.19 mm, tail length = 40.62 mm, mass = 2.365 g; SVL = 44.06 mm, tail length = 52.12 mm, mass = 2.360 g) were obtained from Poza Larga, Municipio Espinal Veracruz, Mexico (20.1608333°N, 97.5094444°W), and brought to the laboratory. Specimens were maintained in separate plastic boxes measuring 32 × 21 × 15 cm with newspaper as substrate, fresh water, and an ambient temperature of 28.5ºC. On the morning of 10 February 2009, each female produced a clutch of two eggs, which were placed in an incubator at 28–30ºC with 50–70% humidity in plastic boxes with agrolite substrate. The ﬁrst hatchling emerged on 17 March (37 days post oviposition [dpo]) and measured 19.6 mm SVL, 18.8 mm tail length, 0.19 g. The second hatchling emerged on 19 March (39 dpo) and measured 13.1 mm SVL, 14.1 mm tail length, 0.16 g. The third hatchling emerged on 25 March (45 dpo), and measured 16.1 mm SVL, 20.9 mm tail length, 0.21 g. The fourth hatchling emerged on 2 Herpetological Review 40(3), 2009 345 April (53 dpo), and measured 18.7 mm SVL, 19.6 mm tail length, 0.21g. Mean incubation period lasted 43.5 days, shorter than those presented in Krysko et al. (2003. Amphibia-Reptilia 24:390–396), who reported incubation times of 48–55 days at 28–29ºC. Submitted by BEATRIZ RUBIO MORALES, EDUARDO CID MENDEZ, and FELIPE CORREA SÁNCHEZ (e-mail: corsanfel@gmail.com), Laboratorio de Herpetología, FES Iztacala UNAM, Avenida de los Barrios 1, P.O. Box 54090 Tlalnepantla, Estado de México, México. IGUANA IGUANA (Green Iguana). AGGRESSIVE BEHAVIOR. Male green iguanas are reported to defend small territories in which variable numbers of females and sub-adults males reside (Muller 1972. Zool. Beitr. 18:109–131). Dominant males use varied displays to defend their territory. These displays involve vertical head and body movements (head bob) as well as tail slapping, often accompanied by dewlap extensions (Dugan 1982. Anim. Behav. 30:327–338). Reports in literature on display behavior of I. iguana are common, but descriptions of males engaging in contact ﬁghts are rare. We report here on aggressive behavior in which two males were observed ﬁghting violently and biting each other. At 1300 h on 28 May 2008, we observed two male I. iguana of approximately the same size (120 cm from snout to tip of tail) ﬁghting on a ranch near the Abobral River in the Pantanal of Mato Grosso do Sul, Brazil (56.9177111°W, 19.4762333°S, datum: WGS84, elev. 60 m). Both males were on the ground in an open grassland ca. 10 m away from trees bordering a marsh. The males were seen lunging, biting each other and thrashing on the ground (Figs. 1A–B). They interlocked for less than ﬁve minutes, until ﬁnally one male escaped. The remaining male displayed, using vertical head movements while extending the dewlap for ca. 5 min before retreating to the forested area near the marsh (Fig. 1C). Both males sustained superﬁcial wounds on their ﬂanks and one male’s front right leg was injured. Iguana iguana has an annual reproductive cycle and maximum testicular development in the Pantanal occurs between June and September with a peak in June (Ferreira et al. 2002. Brazil. J. Morphol. Sci. 19:23–28). Oviposition occurs between September and December (Campos 2004. Herpetol. Rev. 35:169). Hence, this behavior may be related territorial defense for access reproductive females. Submitted by ARNAUD L. J. DESBIEZ, Royal Zoological Society of Scotland Edinburgh, EH12 6TS, United Kingdom (e-mail: adesbiez@hotmail.com or adesbiez@cpap.embrapa.br); and ZILCA CAMPOS, Embrapa Pantanal CP 109 Corumbá MS 79320-900 Brazil (e-mail: zilca@cpap.embrapa.br). FIG. 1. A–B. Male Iguana iguana engaging in interlocking biting behaviors in the Brazilian Pantanal, May 2008. C) Victorious male iguana displaying after the ﬁght. Note the superﬁcial wound on the animal’s ﬂank. 346 Herpetological Review 40(3), 2009 LYGOSOMA PUNCTATA (Spotted Supple Skink). ENDOPARASITES. Lygosoma punctata is a diurnal, fossorial skink that is widepread in Sri Lanka, India, Bangladesh, and Pakistan (Das and de Silva 2005. A Photographic Guide to Snakes and Other Reptiles of Sri Lanka. Ralph Curtis Publ., Sanibel Island, Forida. 144 pp.). To our knowledge, the only helminth recorded from L. punctata is the nematode, Thelandros sp. (Lakshimi et al. 1985. Ind. J. Helminthol. 2:115–125). The purpose of this note is to add the nematode Parapharyngodon adamsoni to the helminth list of L. punctata. Two female L. punctata (SVL = 60 mm, Christopher C. Austin = CCA 2367 collected at Tampataya, Ampara District, Eastern Province [7.5996°N, 81.4272°E, WGS 84], elev, 20 m; and SVL = 49 mm, CCA = 2400 collected 2 km N central Puttalam, Puttalam District, North Western Province, Sri Lanka [8.0742°N, 79.7956°E, WGS 84], elev. 0 m) were examined for helminths. Lizards were sacriﬁced within 12 h of capture, preserved in 10% formalin, and stored in 70% ethanol. The digestive tract was removed, opened, and searched for helminths. The ﬁve nematodes found (CCA 2367 1 m, 2 f; CCA 2400 2 f) in their stomachs were removed, cleared in a drop of glycerol on a glass slide, cover-slipped, studied under a compound microscope, and identiﬁed as Parapharyngodon adamsoni. Nematodes were deposited in the United States National Parasite Collection, Bethesda, Maryland as (USNPC 101177). Lizards were deposited in the herpetology collection of the National Museum of Sri Lanka, Colombo, Sri Lanka. Parapharyngodon adamsoni was described from Chalcidoceps thwaitesi, Nessia smithi (currently Nessia bipes), and N. burtoni by Crusz and Daundasekera (1988. Ann. Parasitol. Hum. Comp. 63:439–447.). Parapharyngodon adamsoni is in the family Oxyuridae which has a direct life cycle with no intermediate host (Anderson 2000. Nematode Parasites of Vertebrates. Their Development and Transmission. CABI Publishing, Oxford, UK. 650 pp.). Infection likely occurs through ingestion of eggs. Lygosoma punctata represents a new host record for P. adamsoni. We thank the Department of Wildlife Conservation, Sri Lanka for permission (number WL/3/2/14/12). This research was funded in part by the People’s Trust for Endangered Species and National Science Foundation grant DEB 0445213 to CCA. Sarah Goldsberry assisted with dissections. MESOSCINCUS MANAGUAE (Managua Skink). ENDOPARASITES. Mesoscincus managuae is known from southern Honduras to Costa Rica (Köhler 1999. The Amphibians and Reptiles of Nicaragua. A Distributional Checklist with Keys. Cour. Forsch.-Inst. Senckenburg, Frankfurt a. M. 213:1–121). To our knowledge, there are no reports of helminths from M. managuae. The purpose of this note is to establish an initial helminth list for M. managuae. One female M. managuae from the herpetology collection of the Natural History Museum of Los Angeles County (LACM), Los Angeles, California collected in December 1964 in Nicaragua, Granada Department, near Isletas (11.7500°N, 85.3333°W, WGS84; elev. 10 m) was examined for endoparasites (LACM 37975; SVL = 85 mm). The body cavity was opened and the digestive tract examined under a dissecting microscope. Found in the large intestine were one fourth stage female and one male nematode which were cleared in a drop of glycerol on a glass slide, cover-slipped, and identiﬁed using a compound microscope as Parapharyngodon alvarengai. Nematodes were deposited in the United States National Parasite Collection, Beltsville, Maryland as Parapharyngodon alvarengai (USNPC 101743). Parapharyngodon alvarengai was originally described from Mabuya maculata from Brazil (Freitas 1957. Mem. Institut. Oswaldo Cruz 55:21–45) and later reported in Ameiva ameiva (Vicente et al. 1993. Rev. Brasil. Zool. 10:19–168) and Rhinella icterica (Luque et al. 2005. Acta Parasitol. 50:215–220), both hosts from Brazil. It was also reported from the lizards Anolis nebulosus, Phyllodactylus lanei, and Sceloporus nelsoni collected in Mexico (Moravec et al. 1997. J. Helminthol. Soc. Washington 64:240–247; Mayén-Peña and Salgado Maldonado 1998. J. Helminthol. Soc. Washington 65:108–111). Parapharyngdon alvarengai is a member of the Oxyuridae, which do not utilize intermediate hosts (Anderson 2000. Nematode Parasites of Vertebrates: Their Development and Transmission, 2nd ed. CABI Publishing, Oxfordshire, UK. 650 pp.). Infection of M. managuae presumably occurred via exposure to P. alvarengai eggs in fecally contaminated substrate. Mesoscincus managuae represents a new host record for P. alvarengai. Nicaragua is a new locality record. We thank Christine Thacker (LACM) for permission to examine M. managuae and Cecilia Nava (Whittier College) for assistance with dissections. Submitted by STEPHEN R. GOLDBERG, Department of Biology, Whittier College, Whittier, California 90608, USA (e-mail: sgoldberg@whittier.edu); CHARLES R. BURSEY, Department of Biology, Pennsylvania State University, Shenango Campus, Sharon, Pennsylvania 16146, USA (e-mail: cxb@ppsu. edu); INDRANEIL DAS, Institute of Biodiversity and Environmental Conservation, Universiti Malaysia Sarawak, 94300, Kota Samarahan, Sarawak, Malaysia (e-mail: idas@ibec.unimas.my); ANSELM DE SILVA, Amphibian and Reptile Research Organization of Sri Lanka, 15/1 Dolosbage Road, Gampola, Sri Lanka (e-mail: kalds@sitnet.lk); and CHRISTOPHER C. AUSTIN, Department of Biological Sciences and Museum of Natural Science, Louisiana State University, Baton Rouge, Louisiana 70803, USA (e-mail: ccaustin@lsu.edu). Submitted by STEPHEN R. GOLDBERG, Department of Biology, Whittier College, Whittier, California 90608, USA (email: sgoldberg@whittier.edu); and CHARLES R. BURSEY, Department of Biology, Pennsylvania State University, Shenango Campus, Sharon, Pennsylvania 16146, USA (e-mail: cxb13@psu. edu). PHRYNOSOMA ASIO (Giant Horned Lizard). DIET. Phrynosoma asio is the largest member of the genus, occurring along the Paciﬁc Coastal forests of Colima, Mexico south to northern Guatemala (Baur and Montanucci 1998. Krötenechsen. Herpeton, Offenbach. 158 pp.; Sherbrooke 2003. Introduction to Horned Lizards. Univ. California Press, Berkeley. 178 pp.). Little is known of the natural history of P. asio, but previous reports on the diet of this species show that it is more of a generalist and less myrmecophagous than congeners (Lemos-Espinal et al. 2004. Herpetological Review 40(3), 2009 347 Herpetol. Rev. 35:131–134; Montanucci 1989. Herpetologica 45:208–216; Pianka and Parker 1975. Copeia 1975:141–162). On 3 August 2008, Ian M. Recchio, Chris M. Rodriguez, Brett Baldwin, and Carlos Martinez collected 4 male and 2 female sub-adult (mean SVL = 50.0 mm) P. asio in tropical deciduous forest in Municipio de Ixtlahuacan, Colima, Mexico (19.03°N, 103.779°W). All six individuals were observed in late afternoon between 1400 and 1700 h, associated with secondary growth clearings adjacent to primary tropical deciduous forest, resting near or underneath bunchgrass. Fecal samples were obtained while the lizards were held prior to export. While being transported for export, the feces from the lizards were preserved for content analysis. Fecal samples were preserved in 75% isopropyl alcohol and examined for contents. The grouped fecal samples contained the remains of the following insect taxa: 56 Aphaenogaster ensifera (Formicidae, Myrmicinae), 23 larvae of an undetermined species of tortoise beetle (Chrysomelidae, Cassidinae), over 400 termites (Isoptera), three toad bugs (Gelastocoridae, Nerthra fuscipes), a single dung beetle (Scarabaeidae, Scarabaeinae, Canthon sp.), and several pieces of unidentiﬁable insect material. Our sample shows P. asio feeds on a variety of terrestrial insects including, but not limited to, ants and termites. This is consistent with previous studies. All research and collecting were done under the authority of SEMARNAT scientiﬁc research permit SGPA/DGVS/03804 issued to IR. Submitted by IAN M. RECCHIO, Reptile/Amphibian Department, Los Angeles Zoo and Botanical Gardens, 5333 Zoo Drive, Los Angeles, California 90027, USA (e-mail: ian. recchio@lacity.org); JAMES N. HOGUE, Department of Biology, California State University, Northridge, 18111 Nordhoff St., Northridge, California 91330-8303, USA (e-mail: james. n.hogue@csun.edu); and DAVID LAZCANO, Universidad Autónoma de Nuevo León, Facultad de Ciencias Biológicas, Laboratorio de Herpetología, Apartado Postal-513, San Nicolás de los Garza, Nuevo León, C.P. 66450, México (e-mail: dlazcanov@hotmail.com). PHRYNOSOMA CORNUTUM (Texas Horned Lizard). REPRODUCTION. We observed copulation between Phrynosoma cornutum on multiple occasions. On 3 May 2008 at 1250 h in Oklahoma Co., Oklahoma, USA, while conducting radiotelemetry observations on a male (61.0 mm SVL) P. cornutum, one of us (MTC) observed copulation with a radio-transmittered female (62.9 mm SVL) (Fig. 1). Both individuals were ﬁtted with Holohil model BD-2 transmitters (0.9 g, male; 1.5 g, female), with transmitters attached to the dorsal surface using silicone sealant and secured around the neck by an elastic collar. The same male was observed copulating with another female (70.0 mm SVL) on 16 May 2008 at 0715 h and with a third female (69.5 mm SVL) on 20 May 2008. The latter two females were not ﬁtted with transmitters. These observations are signiﬁcant in that they conﬁrm that our technique of transmitter attachment does not prevent coitus in our study animals, and also provides direct evidence for the polygynous mating strategy of male P. cornutum. 348 FIG. 1. Copulating pair of radio-transmittered Phrynosoma cornutum. Photograph by Megan T. Cook. Submitted by VICTOR BOGOSIAN III and MEGAN T. COOK, Cooperative Wildlife Research Laboratory, Southern Illinois University, Carbondale, Illinois 62901-6504, USA (e-mail: vicbogos@siu.edu); and RAYMOND W. MOODY, 7701 Arnold Street Suite 109, United States Air Force, Tinker Air Force Base, Oklahoma 73145-9100, USA. PODARCIS BOCAGEI (Bocage’s Wall Lizard). SPINAL FRACTURE. Pathologic spinal fractures can occur in reptiles as the result of metabolic bone disease (Bennett and Mehler 2006. In Mader [ed.], Reptile Medicine and Surgery, 2nd ed., pp. 244–245. Saunders Elsevier, St. Louis, Missouri), but in some cases they have also been reported to result from spinal osteopathies (Bennett and Mehler, op. cit.; Fitzgerald and Vera 2006. In Mader [ed.], op. cit., pp. 909–910 and periosteal newbone formation due to undetermined causes (Fitzgerald and Vera, op. cit.). Reptiles with spinal cord fractures may present paralysis (Mader 2006. In Mader [ed.], op. cit., p. 847) and reptiles with spinal cord injury generally have a loss of panniculus response caudally to the site of injury and a loss of tail or vent stimulation reﬂex (Bennett and Mehler, op. cit.). In some cases, hypertonia occurs cranial to the site of the injury (Bennett and Mehler, op. cit.). A reptile with paralysis and spinal fracture can loose its ability to urinate and defecate (Mader, op. cit.). Here, we report on a case of spinal fracture in a wild lacertid lizard from Portugal. During the radiographical examination of specimens of Podarcis undergoing experiments on locomotor performance, we detected an adult female P. bocagei (55.9 mm SVL) with a conspicuous spinal fracture. The lesion affected the thoracic vertebrae, next to the front limbs (Fig. 1) and we observed evidence of reossiﬁcation of the vertebrae cranially and caudally the fracture. This female had been collected 26 days earlier in São Mamede do Coronado, near Trofa, northwestern Portugal (41.2853°N, 8.5745°W; datum: WGS 1984 ; elev. 50 m), in a habitat consisting of agricultural ﬁelds separated by granite formations where lizards found refuge and attained high densities. The female displayed no obvious locomotor deﬁciencies before or after the radiographies. In fact, before being submitted to the x-ray revealing the malformation, this individual had participated in several Herpetological Review 40(3), 2009 experiments consisting of different locomotion trials. Nonetheless, during preliminary analysis of the data (video footage), a slight difference in this individual’s running pattern was noted, consisting of a “wriggling” or undulating movement different from all other specimens (N = 26) included in the study. Currently, detailed quantiﬁcation of the locomotion parameters is being carried out, which may provide more information on the exact impact of the injury on the locomotor capacities of the lizard. While the lizard was kept in the laboratory (for total period of 27 days), we detected no differences in drinking, feeding, defecating, or urinating habits between this individual and remaining lizards. Radiographs of the 26 other specimens (14 males, 12 females) from the same population revealed no fractures or anomalies of any kind. X-rays from a previous study that included 162 P. bocagei and 168 P. carbonelli (Kaliontzopoulou et al. 2008. Amphibia-Reptilia 29:288–292) also revealed no individuals with fractures or malformations. Though other kinds of vertebral malformations have frequently been observed in lizards, mostly are related to the metabolic bone disease suffered by many captive bred reptiles and usually result in either scoliosis/lordosis or bone demineralization (Grogan 1976. J. Herpetol. 10:262–263; Ahboucha and Gamrani 2001. Metabolic Brain Dis. 16:219–226; Mitchell and Georgel 2005. Herpetol. Rev. 36:183–184). The fracture we observed could be a case of spinal osteopathy or trauma, but we can not assess these hypotheses through radiography (all the individuals used were released to their place of capture after the experiment was completed). However, independent of its exact cause, this represents an exceptional case of recovery from a fracture that typically would have become lethal, either directly by causing a fracture in the spinal cord or indirectly by causing paralysis and consequently making the lizard unable to carry on its normal activities. The low frequency with which such a fracture was observed (only one out of more than 300 individuals of two closely related species of the genus) might indicate that such fractures typically are lethal and rarely seen in free-ranging animals. Special thanks to H. Fernandes from Park & Zoo Santo Inácio who provided X-rays, and W. Grogan for providing references related to vertebral anomalies in reptiles. Submitted by VICTOR BANDEIRA, Universidade de Aveiro, Departamento de Biologia, Campus Universitário de Santiago, 3810-193 Aveiro, Portugal (e-mail: victor.bandeira@ua.pt); ALEXANDRE AZEVEDO, Park & Zoo Santo Inácio, Rua 5 de Outubro nº 4503, 4430-809 Avintes, Portugal; ANTIGONI KALIONTZOPOULOU, CIBIO, Centro de Investigação em Biodiversidade e Recursos Genéticos, Campus Agrário de Vairão, 4485-661 Vairão, Portugal / Departament de Biologia Animal (Vertebrats), Facultat de Biologia, Universitat de Barcelona, Avgda. Diagonal 645, 08028 Barcelona, Spain; and MIGUEL A. CARRETERO, CIBIO, Centro de Investigação em Biodiversidade e Recursos Genéticos, Campus Agrário de Vairão, 4485-661 Vairão, Portugal. PRISTIDACTYLUS SCAPULATUS (NCN). SAUROPHAGY. Pristidactylus are generally considered to be insectivorous (Cei1993. Reptiles del Noroeste, Nordeste y Este de la Argentina. Mus. Reg. Sci. Nat. Torino Monogr. 14:1–949), but speciﬁc diet data are lacking for most species. Hence, here we report an observation of saurophagy in P. scapulatus. At 1213 h on 13 December 2007, we observed an adult male P. scapulatus (100 mm SVL; 26 mm mouth width) attack and start to eat an adult (51 mm SVL) male Liolaemus ruibali at the Reserva Uso Multiple Don Carmelo, Sierra de la Invernada, Departamento Ullum, San Juan, Argentina (30.5862°S, 69.0964°W; datum: WGS84; elev. 3100 m). The habitat where the observation was dominated for shrubs of Adesmia subterranea, Adesmia horrida and Licyum chañar, this is vegetation typical to the Puna (M. Almirón, pers. comm.). When captured, the P. scapulatus released the L. ruibali, which was unable to move and died soon thereafter. To our knowledge, this is the ﬁrst record of attempted saurophagy for any species of Pristidactlyus. We thank M. Jordán for permission to conduct research at the Reserva Don Carmelo (permit No 1204-2235-070), R. Espinoza and F. Lobo for assistance in the ﬁeld, and M. Almirón for identiﬁcation of the vegetation. Submitted by EDUARDO A. SANABRIA (e-mail: sanabria_ ea@yahoo.com.ar.), and LORENA QUIROGA, Departamento de Biología e Instituto y Museo de Ciencias Naturales, F.C.E.F. y N., Universidad Nacional de San Juan, Avenida España 400 (N) C.P. 5400, San Juan, Argentina. FIG. 1. X-ray of an injured female Podarcis bocagei. Detail of the fractured vertebral area can be seen in the round inset at upper right. The 36 Kv X-ray was taken with an exposure of 3mA/sec for 0.03 sec. SCELOPORUS MAGISTER (Desert Spiny Lizard). PREY. Members of the Sceloporus magister species group (Schulte et al. 2006. Mol. Phylogen. Evol. 39:873–880) are often reported to be generalized feeders (Parker and Pianka 1973. Herpetologica 29:143–152; Vitt and Ohmart 1974. Herpetologica 30:410–417). However, studies show that invertebrates are far more representative in the diet than vertebrates. Documented prey items include insects (especially Hymenoptera [mostly ants], Coleoptera, Hemiptera, and various larvae), spiders, and other invertebrates (Hotton 1955. Am. Midl. Nat. 53:88–114; Johnson 1966. Am. Midl. Nat. 76:504–509; Parker and Pianka 1973, op. cit.; Tanner and Krogh 1973. Great Basin Nat. 33:133–146; Vitt and Ohmart 1974, op. cit.). Vegetation is also reported in the diet in these studies, but it Herpetological Review 40(3), 2009 349 is not clear how much plant material is secondarily ingested along with invertebrate prey. Vertebrate prey, especially lizards, is widely reported in general literature, but is based on relatively few records in the primary literature (Cardwell 1994. Herpetol. Rev. 25:121–122). Parker and Pianka (1973, op. cit.) reported a single unidentiﬁed vertebrate in their sample of the stomach contents of 123 specimens in the S. magister complex (S. magister + S. bimaculosus + S. uniformis, ﬁde Schulte et al. 2006, op. cit.) and Vitt and Ohmart (1974, op. cit.) reported a single Aspidoscelis tigris from the sample of stomach contents of 66 specimens (S. magister + S. uniformis, ﬁde Schulte et al. 2006, op. cit.). Perkins et al. (1997. Herpetol. Rev. 28:89) reported Xantusia vigilis in the diet of these lizards in the Mojave Desert (hence, S. uniformis, ﬁde Schulte et al. 2006, op. cit.). Intraspeciﬁc predation upon neonates has also been reported (Tanner and Krogh 1973, op. cit.; Cardwell 1994, op. cit.). Jones and Schwalbe (2009. In Jones and Lovich [eds.], Lizards of the American Southwest: A Photographic Field Guide, pp. 226–229. Rio Nuevo Publishers, Tucson, Arizona) report “nestlings” in the diet of S. magister, but the details are not reported. However, we are aware that the source of this report comes from a reliable personal observation by R. Smith, Tucson, Arizona, of an adult S. m. magister feeding on a nestling Gambel’s Quail (Callipepla gambelii) in a Tucson suburb. To our knowledge, there are no records of S. magister (or any species of the complex) attempting to feed on mammalian prey. At 0900 h on 13 October 2008, one of us (WBG) heard squeals FIG. 1. Adult Sceloporus m. magister apparently attempting to prey on a bat, Pipistrellus hesperus. Photograph by W. B. Gillespie. 350 that were coming from the direction of an outside wall of a house in Tucson, Pima Co., Arizona. An adult S. m. magister had the left wing of a Western Pipistrelle Bat (Pipistrellus hesperus) in its mouth. The lizard and the bat were under a wooden false shutter next to a window on a south-facing wall (Fig. 1). The bat struggled for ca. 15 min before freeing itself and dropping to the ground. The bat remained on the ground for at least 15 more minutes, the duration of the observation. The lizard stayed in place and did not attempt to interact with the bat again during that time. The next morning the bat was gone, but the lizard was not. It seems likely this event was an attempt at predation, but other behavioral traits (e.g., territorial aggression) cannot be completely ruled out. The identity of the lizard was veriﬁed by Cecil Schwalbe (U.S. Geological Survey, Tucson, Arizona) and the bat’s identity was veriﬁed by Tim Snow and Angie McIntire (Arizona Game and Fish Department) and Ronnie Sidner (University of Arizona). Submitted by LAWRENCE L. C. JONES, Casa Araña, 3975 S. Wolf Spider Way, Tucson, Arizona 85735, USA (e-mail: gilaman@comcast.net); and WILLIAM B. GILLESPIE, 7413 N Paseo Ronceval, Tucson, Arizona 85704, USA. TROPIDURUS HISPIDUS (NCN). MINIMUM SIZE AT MATURITY; MAXIMUM BODY SIZE. Tropidurus hispidus has a broad distribution from central-eastern and northeastern Brazil to Venezuela (Rodrigues 1987. Arq. Zool. 31:105–230). The minimum size at sexual maturity is ≥ 70 and ≥ 71 mm, for females and males, respectively based on a series from northeastern Brazil (Vitt 1995. Occ. Pap. Oklahoma Mus. Nat. Hist. 1:1–29). Vitt (1995, op. cit.) also reported the maximum body size for adult female and male T. hispidus, respectively, as 107 and 129 mm. Herein, we report a new record for minimum size at maturity and for maximum body size for both sexes of this species in a caatinga at the Estação Ecológica do Seridó (ESEC Seridó), Serra Negra do Norte municipality, Estado do Rio Grande do Norte, Brazil. To study the reproductive ecology of Tropidurus in rocky habitats at ESEC Seridó (06.57672°S, 37.25575°W, datum: WGS84; elev. 192 m), we collected a large series of T. hispidus (nine juveniles, 75 adults). Of the 75 adult lizards (25 males, 50 females), three females (collected on 6 February 2007 and 23–24 January 2008, respectively) with SVLs of 67.8, 65.5, and 65.0 mm had vitellogenic follicles. Also, one male (SVL = 68 mm) collected on 7 February 2007 had testes and epididymides containing mature spermatozoa in the lumen, and showed secondary sexual characteristics (a black stripe on the ventral aspect of each thigh and a black compact spot in the pre-cloacal ﬂap.; Vanzolini et al. 1980. Répteis das Caatingas. Acad. Bras. de Ciênc., Rio de Janeiro, Brazil. 161 pp.). Thus, for the purpose of our study, we regarded the SVL of the smallest reproductive female as the minimum size for sexual maturity and all females ≥ 65 mm were considered adults. Also, all males with SVL ≥ 68 mm were regarded as adults. These sizes at sexual maturity precede the previous minima for females by 5 mm and for males by 3 mm. With relation to the maximum body size, we recorded a 113 mm SVL adult female on 21 November 2007, and a 139 mm SVL adult male on 13 December 2007. These sizes exceed the previous maxima for females by 6 mm and for males by 10 mm. Herpetological Review 40(3), 2009 Specimens of T. hispidus (CHBEZ 1589, 1592, 1961, 1974, 1983, 1987) were deposited in the herpetological collection of Universidade Federal do Rio Grande do Norte, Natal City. We thank the Programa PELD/CNPq – Caatinga: Estrutura e Funcionamento for logistic support, and R. Hansen for helpful comments on this note. This study was supported by a grant from the Conselho Nacional de Desenvolvimento Cientíﬁco e Tecnológico (CNPq) to L. Ribeiro (Process 141993/2006-5), and IBAMA provided a permit (Permit 206/2006 and Process 02001.004294/03-15). Submitted by LEONARDO B. RIBEIRO (e-mail: ribeiro.lb@gmail.com), and ELIZA M. X. FREIRE (e-mail: elizajuju@ufrnet.br), Programa de Pós-Graduação em Psicobiologia, Universidade Federal do Rio Grande do Norte, Centro de Biociências, Caixa Postal 1511, Campus Universitário Lagoa Nova, CEP 59078-970, Natal, Rio Grande do Norte, Brazil. TROPIDURUS MONTANUS (Calango-da-Montanha; Montane Collared Lizard). ESCAPE BEHAVIOR. Some knowledge of behavior is crucial for understanding an animal’s ecology, yet for many vertebrates, we lack even anecdotal information is lacking (Greene 1986. In Feder and Lauder [eds.], Predator-Prey Relationships, pp. 99–108. University of Chicago Press, Chicago, Illinois; Greene 2005. Trend. Ecol. Evol. 20:23–27). Lizards are known to present diverse defensive behaviors to avoid predation, including crypsis, immobility, mimicry, chemical defense, tail or skin shedding, social and aggressive displays, and use of retreats or escape routes (Pianka and Vitt 2003. Lizards: Windows to the Evolution of Diversity. University of California Press, Berkeley. 348 pp.). Tropidurus montanus is a small, diurnal, terrestrial lizard, endemic to the open, montane habitats of southern Espinhaço range (Rodrigues 1987. Arq. Zool., São Paulo 31:105–230). The defensive behavior of T. montanus was recently studied (Machado et al. 2004. S. Am. J. Herpetol. 2:136–140) and the primary strategy of defense is the predator avoidance through crypsis associated with immobility. When disturbed, locomotor escape was the tactic employed by this species (Machado et al., op. cit.). Here, we describe an unreported defensive behavior in T. montanus, and a novelty among Tropidurus lizards. At 1210 h on 16 March 2008, we observed an adult male T. montanus (80.6 mm SVL, 117.4 mm tail length) dive into a small stream after being chased by one of us. The observation occurred in Serra do Cabral, Augusto de Lima municipality, Espinhaço range, Minas Gerais State, Brazil (17.999500°S, 44.337028°W; datum: WGS84; elev. 1016 m). After having been disturbed, the lizard ﬂed on the rocks and dove to a depth of ca. 10 cm into a small stream and remained underwater over 3.5 min. Throughout this time, the animal kept its head partly oriented toward the surface and its eyes partially opened, presumably observing the surface and our movements. Three minutes after the observation began, the animal was visibly experiencing difﬁculties remaining under water, and bubbles escaped through its nostrils. Just over 3.5 min after the observation began, it tried to escape sideways on the rock where it was submerged, and was captured. Diving behavior would allow T. montanus opportunity to escape a predator during a risky chase. Lizards in many families are known to use water bodies as escape routes from potential predators (Pi- anka and Vitt, op. cit.). The majority of the reports on this defensive behavior in neotropical lizards is related to lowland rainforest or ﬂooded-ﬁeld species, especially semi-aquatic or river-associated lizards (Martins 1991. Faun. Environ. 26:179–190; Ávila-Pires 1995. Zool. Verh. Leiden 299:1–706; Pianka and Vitt, op. cit.). To date, the only known tropidurid for which water-escape has been described is the Amazonian Uranoscodon superciliosus. This semi-arboreal species relys largely on crypsis to avoid detection by predators, but when disturbed readily jumps to water, often running across the water surface, but in some cases, diving into the water, and remaining underwater for minutes before emerging (Howland et al. 1990. Can. J. Zool. 68:1366–1373). This represents of the ﬁrst report of water-based escaped behavior in the genus Tropidurus, and a second report for such behavior among tropidurid lizards. The specimen was deposited in the herpetological collection of the Museu de Zoologia da Universidade de São Paulo, São Paulo, Brasil (MZUSP 98259, José Cassimiro ﬁeld number JC 1358). We thank IBAMA for collection permits (10126-1). Fundação de Amparo à Pesquisa do Estado de São Paulo (FAPESP) provided research grants to JC (process 05/00283-6). Vanessa K. Verdade and Carolina Castro-Mello assisted in the MZUSP, and M. P. Hayes provided editorial suggestions. Submitted by JOSÉ CASSIMIRO (e-mail: geckoides@gmail. com), MAURO TEIXEIRA JR. (e-mail: mteixeirajr@usp.br), RENATO SOUSA RECODER (e-mail: renatorecoder@gmail. com), and MIGUEL TREFAUT RODRIGUES (e-mail: mturodri@usp.br), Departamento de Zoologia, Instituto de Biociências, Universidade de São Paulo, Caixa Postal 11.461, CEP 05422-970, São Paulo, SP, Brazil. XENOSAURUS GRANDIS (Knob-scaled Lizard). ENDOPARASITES. Xenosaurus grandis is known from the Mexican states of Chiapas, Oaxaca, and Veracruz (Flores Villela and Gerez 1994. Biodiversidad y Conservación en México: Vertebrados, Vegetación y Uso del Suelo. Comision Nacional para el Conocimiento y Uso de la Biodiversidad, Universidad Nacional Autónoma de México, Destricto Federal, 439 pp.). To our knowledge, there are no reports of endoparasites from X. grandis. The purpose of this note is to establish an initial helminth list for X. grandis. Eleven X. grandis (mean SVL = 110.0 mm ± 17.0 SD, range = 78–130 mm) from Cuautlapan, Veracruz, Mexico (18.8666°N, 97°0166°W, WGS84; elev. 1000–1200 m) collected in 1969, 1970–1972 and deposited in the herpetology collection of the Natural History Museum of Los Angeles County (LACM), Los Angeles, California were examined: LACM 75689, 75692–75694, 120051, 120060, 120065, 120071, 120077, 120083, 120086. The body cavity was opened and the digestive tract was removed and examined under a dissecting microscope. Only nematodes were found. These were cleared in a drop of glycerol on a glass slide, coverslipped, and identiﬁed using a compound microscope as Spauligodon xenosauri infection site: large intestine (prevalence = number infected/number examined × 100 = 91%; mean intensity = mean number helminths per infected individual =13.9 ± 12.7 SD. range = 1–36), Physaloptera sp. as 3rd stage larvae (the lack of adult specimens makes species identiﬁcation imprudent, however, P. retusa is commonly reported in lizards of Mexico), Herpetological Review 40(3), 2009 351 infection site: stomach, small intestine (prevalence = 9%, 5.0) and Seuratoidea gen. sp. (larvae in cysts), infection site: stomach wall (prevalence = 36%, mean intensity = 12.8 ± 12.6, range = 2–26). Voucher nematodes were deposited in the United States National Parasite Collection, (USNPC), Beltsville, Maryland as: Spauligodon xenosauri (USNPC 101562); Physaloptera sp. as 3rd stage larvae (USNPC 101563), and Seuratoidea gen. sp. (USNPC 101564). Spauligodon xenosauri was described from Xenosaurus platyceps from Tamaulipas, Mexico by Bursey et al. (2007. Zootaxa 1501:65–68). Xenosaurus grandis is the second host known to harbor it. Spauligodon xenosauri is a member of the Oxyuroidea, which infect hosts directly (Anderson 2000. Nematode Parasites of Vertebrates. Their Development and Transmission. CABI Publishing, Oxfordshire, UK, 650 pp.). Infection likely occurs when lizards lick substrate on which eggs are present (see Goldberg and Bursey 1992. J. Parasitol. 78:539–541). Physaloptera retusa is a common parasite in lizards of Mexico and North America and requires an insect intermediate host (Bursey et al. 2007. Comp. Parasitol. 74:108–140). Bursey and Goldberg (1991. J. Wildlife Dis. 27:710–715), in a study of P. retusa in the lizard Sceloporus jarrovii, suggested that approximately 10% of 3rd stage larvae ingested by a host survive to maturity and that those found in the small intestine were dead at time of host capture. Goldberg et al. (1993. Bull. South. California Acad. Sci. 92:43–51) reported hosts for physalopteran larvae; teiid and xenosaurid lizards were absent from that list which suggests that some lizards may not be suitable hosts for this nematode. In this case, these physalopteran larvae should be considered to be artifacts of diet. The life cycles of the Seuratoidea are largely unknown (Anderson 2000, op. cit.) but it is thought that insects serve as intermediate hosts. In this study, the cysts were in early granuloma formation perhaps indicating re-encystment of larvae in an inappropriate host. Xenosaurus grandis represents a new host record for Spauligodon xenosauri, Physaloptera sp. (3rd stage) and larvae of Seuratoidea gen. sp. We thank Christine Thacker (LACM) for permission to examine specimens and Cecilia Nava (Whittier College) for assistance with dissections. Submitted by STEPHEN R. GOLDBERG, Department of Biology, Whittier College, Whittier, California 90608, USA (email: sgoldberg@whittier.edu); and CHARLES R. BURSEY, Department of Biology, Pennsylvania State University, Shenango Campus, Sharon, Pennsylvania, USA (e-mail: cxb13@psu.edu). SQUAMATA — SNAKES BOTHROPS ASPER (Terciopelo). SCAVENGING BEHAVIOR. We ecountered a Bothrops asper as it scavenged a frog at La Selva Biological Station in Costa Rica (10.4333333ºN, 83.9833333ºW) on 20 April 2008 at dusk (1747 h local time). The B. asper (ca. 60 cm long and missing part of its tail) slid onto the cement trail to a dead, rotting frog. The snake grasped the frog in its mouth, dragged it to the leaf litter adjacent to the trail, let go of the frog, then re-approached and consumed it. The frog was decayed beyond identiﬁcation and attracting ﬂies (Fig. 1). Scavenging behavior has been documented in 12 species of Crotalinae under ﬁeld conditions (DeVault and Krochmal 2002, 352 FIG. 1. Bothrops asper consuming a scavenged frog at La Selva Biological Station, Costa Rica. Herpetologica 58:429-436). Sazima and Strüssman (1990. Rev. Brasil. Biol. 50:463–468) suggest that habitat and diet can inﬂuence carrion encounter rates. They expect terrestrial snakes with diverse diets to be “occasional” scavengers. As a dietary generalist, the terrestrial B. asper meets their prediction (Martins et al. 2002. In Schuett et al. [eds.], Biology of the Vipers, pp. 307–328. Eagle Mountain Publishing, Eagle Mountain, Utah). We speculate that scavenging is a prevalent opportunistic foraging strategy for many snakes (A. Solórzano, pers. comm.) that has gone unnoticed due to the difﬁculty of observing such behaviors under ﬁeld conditions. This research was sponsored by the Gates Cambridge Trust. Submitted by CORINA J. LOGAN, University of Cambridge, Department of Experimental Psychology, Cambridge, CB2 3EB, UK (e-mail: itsme@CorinaLogan.com); and CHRISTOPHER MONTERO, Ecosystem Services Section, Washington State Department of Natural Resources, Olympia, Washington 985047016, USA. CLELIA SCYTALINA (Mexican Snake Eater). DIET. Clelia scytalina is a large terrestrial snake whose distribution is apparently unclear and disjunct, occurring at low elevations from southern Mexico to Costa Rica (Savage 2002. The Amphibians and Reptiles of Costa Rica: A Herpetofauna Between Two Continents, Between Two Seas. Univ. Chicago Press. Chicago, Illinois. 934 pp.). Natural history of this species is poorly known. In the Los Tuxtlas region of Veracruz, Mexico diet records include frogs and lizards (Pérez-Higareda et al. 2007. Serpientes de la Región de Los Tuxtlas, Veracruz, México. Guía de Identiﬁcación Ilustrada. Universidad Nacional Autónoma de México. 189 pp.), but there are no previous records of snakes as prey in this area. On 17 May 2006 at 1700 h we observed a C. scytalina (ca. 120 cm total length) preying on a Parrot Snake (Leptophis ahaetulla) of similar length. The event occurred in a dry creek in the tropical rainforest reserve at Los Tuxtlas Field Station (Lote 67, Vigia Hill, 18.5851°N, 95.08055°W, 178 m elev., WGS84). The C. scytalina surprised the L. ahaetulla at rest, biting its head and starting to Herpetological Review 40(3), 2009 swallow without releasing. When approached to within 3 m, the C. scytalina detected our presence, released the prey, and ﬂed into nearby vegetation. The L. ahaetulla was still alive but died shortly after the attack. This is the ﬁrst report of a C. scytalina preying on a L. ahaetulla and it suggests that C. scytalina is an opportunistic and generalist feeder. This is consistent with other Clelia species (Campbell 1998. Amphibians and Reptiles of Northern Guatemala, the Yucatán, and Belize. Univ. Oklahoma Press. 380 pp.; da Costa Pinto and de Lema 2002. Iheringia, Ser. Zool. 92[2]:9–19; Vitt and Vangilder 1983. Amphibia-Reptilia 4:273–296). We thank Lindley McKay for improving the English language in this note. Submitted by ÁLVARO CAMPOS VILLANUEVA, Estación de Biología Los Tuxtlas, Instituto de Biología, Universidad Nacional Autónoma de México, Km 30 Carretera CatemacoMontepío, San Andrés Tuxtla, Veracruz, México, A.P. 94957001; and ELISA CABRERA-GUZMÁN, Tropical Ecology Research Facility, School of Biological Sciences, The University of Sydney, Middle Point, Northern Territory 0836, Australia (email: anﬁsbenido@yahoo.com). CROTALUS ATROX (Western Diamond-backed Rattlesnake). ADULT PREDATION ON LIZARDS. The prey of adult Crotalus atrox consists of a wide range of mammalian (endothermic) taxa, primarily rodents and lagomorphs. Occasionally, birds are subjugated and consumed (Klauber 1972. Rattlesnakes. Their Habits, Life Histories, and Inﬂuence on Mankind. Two vols., 2nd ed. University of California Press, Los Angeles and Berkeley. 1533 pp.; Cundall and Greene 2000. In K. Schwenk [ed.], Feeding, Form, Function, and Evolution in Tetrapod Vertebrates, pp. 293–333. Academic Press, San Diego, California; Campbell and Lamar 2004. The Venomous Reptiles of the Western Hemisphere. Cornell University Press, Ithaca, New York. 870 pp.). Here, we provide further information on the diet and predatory behavior of adult C. atrox from a population in the Sonoran Desert of south-central Arizona. Speciﬁcally, we show that adults of a large species of a North American temperate pitviper (Viperidae: Crotalinae) do not exclude relatively small, ectothermic prey from their diet (e.g., Sherbrooke 2003. Introduction to Horned Lizards of North America. University of California Press, Berkeley and Los Angeles. 178 pp.; Nowak et al. 2008. Biol. Rev. 2008: 83:601–620). The individuals of C. atrox we discuss were the subjects of a radio-telemetric study in which various aspects of their behavior, physiology, and spatial ecology have been investigated since March 2001 (e.g., Repp and Schuett 2008. Southwest. Nat. 53:108–114). The present study site, located in Pinal County, is 40 km SSE of Florence and encompasses an area ≈ 3 km2 at the western perimeter of the Suizo Mountains. This region is ecologically designated as Arizona Upland Desertscrub subdivision (Brown 1994. Biotic Communities of the American Southwest — United States and Mexico. University of Utah Press, Salt Lake City. 342 pp.). Annual precipitation (chieﬂy rain) patterns are bimodal, with slight to moderate storms occurring during winter and early spring (December–March); the strongest activity occurs from mid- to late summer (early July to mid-September), which is termed the North American monsoon (Brown 1994, op. cit.; Phillips and Comus 2000. A Natural History of the Sonoran Desert. The University of California Press, Berkeley and Los Angeles. 628 pp.). The area lacks permanent freestanding water, a common feature of the Sonoran Desert; at the present site, collection of rainwater in surface rocks and soil, as a rule, is transitory, and persists for only minutes to several hours. However, artiﬁcial alterations of the habitat (e.g., deep tire tracks in narrow dirt roads) occasionally provide a substantive reservoir for the collection and storage of rainwater, but even under these situations it is ephemeral, especially in summer. On 10 July 2004, at 2042 h, in an area interfacing bajada (Phillips and Comus 2000, op. cit.) and desert ﬂats, an adult male C. atrox (CA-32: SVL = 110 mm, TL = 9.5 mm, body mass = 745 g) was radio-tracked and located (Site 21, 822 m elev.) partially coiled beside an adult (SVL ≈100 mm; body mass ≈ 40 g) Regal Horned Lizard, Phrynosoma solare. The core body temperature of CA-32 (obtained via an implanted 11.0 g temperature-sensitive FIG. 1. Predation by adult wild-living Western Diamond-backed Rattlesnakes (Crotalus atrox). A) An adult male C. atrox (CA-32) beside an adult Regal Horned Lizard (Phrynosoma solare) that presumably was envenomated by CA-32. Note the two blood spots (within the yellow ellipses) that most likely resulted from fang punctures of the present snake. Photograph by R. Villa. B) An adult male C. atrox (CA-55) beside an adult Desert Spiny Lizard (Sceloporus magister) that it presumably subjugated; also, note that the lizard is gaping with its tongue protruding. This lizard was subsequently consumed by CA-55. Photograph by R. Repp. Herpetological Review 40(3), 2009 353 radio-transmitter; Holohil Systems Ltd, Ontario, Canada) was 32.7ºC. The ambient temperature (1 m above the ground) was 29.0ºC. Cloud cover was ≈ 10%, relative humidity was ≈ 20%, and wind speed was slight (0–5 kph) and intermittent. Upon closer inspection, the P. solare was sluggish and had been (presumably) bitten by the present snake; at least two presumptive fang punctures were present on both left and right left sides of the posterior body, and from these wounds oozing dark-colored blood (within yellow ellipses) was detected (Fig. 1A). From 2042 h to 2111 h, CA-32 was directly facing the lizard and periodically (N = 8 episodes) nudged it, presumably to determine whether it was dead or fully immobilized. During these nudging episodes, the lizard exhibited anti-predator behavior, primarily rearing and directing its head obliquely downward toward CA32 (see Sherbrooke 2003, op. cit.). At 2211 h, the lizard was still alive, and we departed to avoid possible interference. At 2300 h, we re-located CA-32; he was about 10 m from Site 21 and had a food bolus, undoubtedly the P. solare. On 24 July 2004, at 0643 h, 14 days following consumption of the P. solare, we radio-tracked male CA-32 and located him (Site 19, 821 m elev.) in bajada; he was coiled 1 m from of an active Neotoma albigula midden. An obvious large food bolus (documented via photography) was present; given his location, it was possibly an adult N. albigula. His core body temperature was 26.4ºC. The ambient temperature was 25.0ºC and ground temperature in direct sunlight (“hot spot”) was 28.0ºC. On 14 June 2008, from 0630 to 0733 h, in bajada, an adult male C. atrox (CA-55: SVL: 99 mm, TL: 80 mm, mass: 687 g) was radio-tracked and located (Site 52, 826 m elev.) beside an adult male (SVL ≈ 140 mm, body mass ≈ 40 g) Desert Spiny Lizard (Sceloporus magister). First inspection of the pair revealed that CA-55 was sprawled lengthwise, with the front one-third of his body positioned in a small area of shade in open ground, and his rear two-thirds was beneath a cluster of Triangle Bursage beneath the canopy of a 1.5 m tall Creosote Bush. The snake’s chin rested on the tail of the lizard, which was positioned directly in front of and facing away from CA-55. Neither subject was moving; the lizard’s eyes were slightly open. Minutes after locating CA-55, his core body temperature was 23.8ºC. The ambient temperature was 25ºC and hot spot was 32ºC. Cloud cover was 0%, relative humidity was ≈ 10%, and wind speed was slight (< 5 kph). Both ambient and hot spot temperatures changed rapidly as sunrise progressed. At 0648 h, CA-55 slowly crawled lengthwise beside the left side of the lizard, he occasionally rubbed the lizard with his mouth, and then started rubbing his snout on the snout of the lizard. At that point, the lizard ﬂinched and CA-55 slowly withdrew to resume his position behind the lizard. The core body temperature of CA-55 was 26.7ºC. The pair was temporarily left to radio-track another C. atrox subject; CA-55 was re-tracked at 0711 h. At that time, CA-55 was on the right side of the lizard, which gaped widely three times, at ≈ 30 sec intervals (Fig. 1B). Following the ﬁrst gape, the lizard shuddered violently; he also shuddered after the third gape. The third gape and subsequent shudder were the last obvious movements of the lizard. Both subjects were in full sunlight; the body temp of CA-55 was 31.3ºC. The ambient temperature was 29ºC and hot spot was 35ºC. Because CA-55 appeared disturbed, other subjects of C. atrox 354 were subsequently radio-tracked; ca. 20 min later, CA-55 was retracked at 0733 h and neither the snake nor the lizard were present at the original site (Site 52). The snake had moved 2 m N of Site 52 into a heavily vegetated arroyo. Inspection revealed that a minor food bolus was present, and it was assumed he had consumed the lizard. Again, CA-55 appeared startled by the observer and quickly coiled in 100% shade under a downed canopy of wolfberry (Lycium sp.) branches on the south side of the arroyo. His core body temperature at the conclusion of the observations was 35.5ºC. The ambient temperature was 32ºC and hotspot was 44.5ºC. In both of the above cases, body masses of the two species of lizards, estimated based on SVL, were ≈ 40 g; this represents 5% (CA-32, 745 g) and 6% (CA-55, 687 g) of the body masses of the respective snakes. Moreover, the resultant food boluses, as expected, were not highly discernible compared to larger prey such as commonly consumed mammals (e.g., ground squirrels, woodrats); owing to their size, it is likely that the present food items would have been overlooked under most situations. Hence, this suggests that predation attempts and feeding frequency in certain large rattlesnakes (and other vipers) are underestimated (Nowak et al. 2008, op. cit.). Also, we show that adult C. atrox can successfully feed on adult horned lizards (Phrynosoma spp.) without (apparent) subsequent injury (see comments by Sherbrooke 2003, op. cit.). Support for our ﬁeld studies at the Suizo Mountains is from Arizona State University, Zoo Atlanta, Georgia State University, and David L. Hardy, Sr. Since 2001, many individuals provided assistance in the ﬁeld, but most noteworthy are Hans-Werner Herrmann and Ryan Sawby. Also, Ryan Sawby provided invaluable assistance with photography and identiﬁcation of plant and invertebrate taxa. We thank Emory Schuett, Robert Villa, and Annamarie Saenger for their help in making observations on the present C. atrox and P. solare. This study was approved by the animal care and use committee (IACUC) of Arizona State University (98-429R), and appropriate scientiﬁc permits were obtained from the Arizona Game and Fish Department. Submitted by ROGER A. REPP, National Optical Astronomy Observatory, 950 N. Cherry Avenue, Tucson, Arizona 85719, USA (e-mail: repp@noao.edu); and GORDON W. SCHUETT, Department of Biology and Center for Behavioral Neuroscience, Georgia State University, 33 Gilmer Street, S. E., Unit 8, Atlanta, Georgia 30303-3088, USA (e-mail: biogws@langate.gsu.edu, or gwschuett@yahoo.com). EPICRATES CENCHRIA (Brazilian Rainbow Boa). DIET. On 27 February 2008, at 2350 h, during a radiotracking study at Floresta River (22.5275°S, 42.5516667°W), Reserva Botânica Águas Claras, Silva Jardim, Brazil, we located a female (280 g) radio-collared Neotropical Water Rat (Nectomys squamipes) inside an Epicrates cenchria (ca. 2 m total length). The snake was <1 m from the river’s edge and active. We observed it again two days later in a shelter at the base of a large tree (3.55 m perimeter at breast height), at 5.1 m above the river level and 0.5 m from the river. This shelter had three openings, the measures were: 16×14×66; 20×7×56 and 22×18×49 cm (biggest diameter, perpendicular diameter and depth, respectively). The radiotransmitter was found one month later in what appeared to be regurgitations of the snake Herpetological Review 40(3), 2009 in the same shelter. Epicrates cenchria is widely distributed in Central and South America, from Costa Rica to Argentina (Tolson 1987. Occas. Pap. Mus. Zool., Univ. Michigan 715:1–68; Vanzolini et al. 1980. Répteis das Caatingas. Academia Brasileira de Ciências, Rio de Janeiro. 161 pp). All Epicrates are nocturnal and most eat exclusively endotherms, with most diet records consisting of bats and rodents (Mus, Rattus) (Nellis et al. 1983. J. Herpetol. 17:413–417). To the best of our knowledge, this is the ﬁrst record of E. cenchria predation on N. squamipes, the largest and most adapted to semiaquatic life among sigmodontine rodents. We thank Idea Wild for equipment, Fundação o Boticário de Proteção à Natureza, CNPq and PIBIC/CNPq for ﬁnancial support. In addition, we thank our colleagues in the Laboratório de Ecologia e Conservação de Populações for assistance. Submitted by GABRIELA MEDEIROS DE PINHO (e-mail: gabriela.m.pinho@gmail.com) and DANIELA OLIVEIRA DE LIMA Departamento de Ecologia, Instituto de Biologia, Universidade Federal do Rio de Janeiro, CP 68020, Rio de Janeiro, RJ, 21941-902, Brazil; PAULO NOGUEIRA DA COSTA, Departamento de Zoologia, Instituto de Biologia, Universidade Federal do Rio de Janeiro, CP 68020, Rio de Janeiro, RJ, 21941-902, Brazil; and FERNANDO ANTONIO DOS SANTOS FERNANDEZ, Departamento de Ecologia, Instituto de Biologia, Universidade Federal do Rio de Janeiro, CP 68020, Rio de Janeiro, RJ, 21941902, Brazil. FARANCIA ERYTROGRAMMA (Rainbow Snake). HABITAT. Farancia erytrogramma is a secretive, nocturnal species occurring in southeastern USA (Gibbons and Dorcas 2005. Snakes of the Southeast. University of Georgia Press, Athens, Georgia. 253 pp.). In North Carolina, this species can be found throughout the coastal plain (Palmer and Braswell 1994. Reptiles of North Carolina. University of North Carolina Press, Chapel Hill, North Carolina. 412 pp.) and usually inhabits clear, ﬂuvial water systems with low turbidity, neutral pH, and standing vegetation for cover (Neill 1964. Amer. Midl. Nat. 71:257–295). However, past research revealed this species can occur in standing freshwater habitats, such as Carolina bays (Gibbons et al. 1977. Herpetologica 33:276–281). During a study of the reptile and amphibian community at Bull Neck Swamp (35.96667°N, 076.41667°W), one F. erytrogramma was captured with an aquatic funnel trap. Bull Neck Swamp (BNS) is a 2428-ha pocosin wetland located 28 km E of Plymouth, North Carolina, USA, and is managed by the Fisheries and Wildlife Sciences Program at North Carolina State University. The highly turbid canals of tannic ﬂows and thick bottom debris that meander throughout BNS were an unexpected habitat to discover F. erytrogramma, which prefer clear, moving water (Neill 1964, op. cit.). Neill (op. cit.) maintained that consistent temperatures of stream habitats were important for thermoregulation. Also, ﬂuvial habitats provide cover for American Eels (Anguilla rostrata) and Neill (op. cit.) suggested the predominant diet on A. rostrata further supported the constraint of F. erytrogramma to these habitats. However, steep, sandy canal banks prevented the growth of standing vegetation, which likely reduced refuge and staging habitats for nocturnal foraging. Further, Beaver (Castor canadensis) dams reduced what little ﬂow existed in some canals to standing quagmires more representative of the habitat selected by Eastern Mudsnakes (Farancia abacura; Neill 1964, op. cit.). Interestingly, one A. rostrata was observed near BNS, but none was captured within the swamp. It is possible that Rainbow Snakes leave bordering ﬂuvial habitats in pursuit of young eels that wandered into canals and swamp habitats. Capturing such a secretive and uncommon species as F. erytrogramma in unexpected habitat encourages consideration of their delicate ecological niche. Declining population indices for American Eels along the eastern United States are attributed to overﬁshing, parasitism, habitat loss, pollution, and changes in major currents related to climate change (Hightower and Nesnow 2006. Southeast. Nat. 5:693–710). Eel declines could negatively impact population sizes and distributions of Rainbow Snakes, especially in inland areas. We believe future studies based on conﬁrmed Rainbow Snake occurrences from museum records or North Carolina GAP data could better delineate the range within North Carolina. Additionally, sampling for American Eels to determine their population status and distribution in North Carolina could augment population and distribution data for Rainbow Snakes. We thank A. Braswell, J. Jensen, and P. Moler for comments on earlier drafts of this manuscript. S u b m i t t e d b y S TA N J . H U T C H E N S ( e - m a i l : stanhutchens@gmail.com) and CHRISTOPHER S. DEPERNO, (e-mail: chris_deperno@ncsu.edu), Fisheries and Wildlife Program, North Carolina State University, 110 Brooks Ave., Raleigh, North Carolina 27607, USA. HELMINTHOPHIS FRONTALIS (Northern Antsnake). FEEDING BEHAVIOR. Fossorial blind worm snakes are frequently found under debris in lowland moist and premontane forests in Costa Rica and Panama (Savage 2002. The Amphibians and Reptiles of Costa Rica: A Herpetofauna Between Two Continents, Between Two Seas. University of Chicago Press, Chicago, Illinois. 934 pp.). The habits of these snakes are nearly unknown because of their burrowing and secretive lifestyle, though it is known that the diet of Helminthophis frontalis (Anomalepididae) consists of ants, termites and their larvae (Solórzano 2004. Serpientes de Costa Rica. Instituto Nacional de Biodiversidad (INBio). Heredia, Costa Rica. 792 pp.). How blind worm snakes prey upon ants and termites without being attacked by these colonial insects has not yet been described. Leptotyphlops dulcis (Leptotyphlopidae), another blind burrowing snake that feeds on termites, preys on immature stages of colonial insects (e.g., ants and termites), apparently without being attacked. This blind snake covers itself with feces and a viscous odoriferous liquid that is secreted by the cloaca prior to crawling into the colony. This substance repels the ants and prevents their attack (Gehlbach et al. 1968. Bioscience 18:784–785). This information is missing for most fossorial blind worm snakes, though there are a few reports of some species using this strategy (Zug et al. 2001. Herpetology: An Introductory Biology of Amphibians and Reptiles. 2nd ed. Academic Press. California,. 630 pp.). On 21 July 2004, we observed a Helminthophis frontalis entering an ant colony in San José City, Costa Rica. As the snake attempted Herpetological Review 40(3), 2009 355 to enter, it was immediately attacked and expelled by the ants. The attack eventually ceased and the snake was able to enter the colony and remain inside, preying upon adult ants and their larvae. After 5 minutes, we removed the snake from the colony and observed that it was covered by an odoriferous viscous substance. Presumably such material repelled the ants and protected the snake from being attacked. A few minutes later, we placed the snake back into the colony and noted that the substance was produced from the cloaca and seemed to allow the snake to feed, undisturbed by ants. Despite the phylogenetic association of the families Leptotyphlopidae and Anomalepididae, this specialized feeding-related behavior has not been reported for species of the latter (Savage, op. cit.). Submitted by BRANKO HILJE, Department of Biology, University of Puerto Rico, Río Piedras P.O. Box 23360, San Juan, Puerto Rico, 00931-3360 (e-mail: bhilje@yahoo.com); and MARIEL YGLESIAS, Centro Agronómico Tropical de Investigación y Enseñanza (CATIE), Turrialba, Costa Rica (e-mail: mariel_yglesias@yahoo.es). LEPTODEIRA SEPTENTRIONALIS (Cat-eyed Snake). PREY. Many neotropical snakes feed primarily on anurans (Vitt 1983. Herpetologica 39:52–66) but information regarding their individual prey species is scarce. On 15 November 2004 at 21 h, in the Refugio de Vida Silvestre, Golﬁto (Costa Rica, Puntarenas Province, 8.65°N, 83.18°W, ca. 5 m elev.), I observed an adult Leptodeira septentrionalis swallowing a Leptodactylus bolivianus (Fig. 1). The frog was ingested head ﬁrst. L. bolivianus represents a hitherto unreported prey item for L. septentrionalis. LEPTOPHIS AHAETULLA PRAESTANS (Green Parrot Snake). PREDATION ATTEMPT. Leptophis ahaetulla is a large, diurnal, and arboreal species, occurring in different vegetation types from southern Mexico to Brazil and Argentina (Savage 2002. The Amphibians and Reptiles of Costa Rica: A Herpetofauna Between Two Continents, Between Two Seas. Univ. Chicago Press. Chicago, Illinois. 934 pp.). It has been reported that L. ahaetulla forages in trees and shrubs for sleeping or resting frogs, and to a lesser extent, arboreal lizards (Oliver 1948. Bull. Am. Mus. Nat. Hist. 92:157–280; Pérez-Higareda et al. 2007. Serpientes de la Región de Los Tuxtlas, Veracruz, México. Guía de Identiﬁcación Ilustrada. Universidad Nacional Autónoma de México. 189 pp.). There are no reports for prey items for L. ahaetulla in Mexico, so here we record a feeding attempt observed in Veracuz. On 18 April 2007 at ca. 1000 h on bare soil adjacent to buildings belonging to Los Tuxtlas Tropical Biology Station (18.5851°N, 95.0752°W, 119 m elev., WGS84), we observed a L. ahaetulla (ca. 150 cm total length) feeding on a Baudin’s Treefrog (Smilisca baudini, ca. 60 mm snout–vent length). The snake held the frog in its mouth for a minute but when we approached to a distance of 2 m, the snake released the frog and ﬂed. The frog was alive though motionless, and died some minutes later. Frogs (especially hylids) have been recorded as the main food items for L. ahaetulla elsewhere in its range (e.g., Campbell 1998. Amphibians and Reptiles of Northern Guatemala, the Yucatán, and Belize. Univ. Oklahoma Press. 380 pp.; De Albuquerque and DiBernardo 2005. Herpetol. Rev. 36:325; Lopez et al. 2003 Herpetol. Rev. 34:68–69). Other reported prey for L. ahaetulla are birds, bird’s eggs, and grasshoppers (Oliver 1948, op. cit; Lopez et al., op. cit). We thank Lindley McKay for improving the English language in this note. Submitted by ELISA CABRERA-GUZMÁN, Tropical Ecology Research Facility, School of Biological Sciences, The University of Sydney, Middle Point, Northern Territory 0836, Australia (e-mail: anﬁsbenido@yahoo.com); and ÁLVARO CAMPOS VILLANUEVA, Estación de Biología Los Tuxtlas, Instituto de Biología, Universidad Nacional Autónoma de México, Km 30 Carretera Catemaco-Montepío, San Andrés Tuxtla, Veracruz, México A.P. 94957001. FIG. 1. Leptodeira septentrionalis swallowing a Leptodactylus bolivianus. Submitted by DAVID MATTHIAS DEHLING, Department of Ecology, Animal Ecology, Faculty of Biology, Philipps-University Marburg, Karl-von-Frisch-Straße 8, D-35032 Marburg, Germany; e-mail: Dehling@students.uni-marburg.de 356 LIOPHIS VITTI (NCN). DIET. The colubrid snake Liophis vitti is known from the western Andean slopes in northern Ecuador at elevations between 1070–1650 m (Dixon 2000. Copeia 2000:482–490). A specimen collected “one meter high on leaf in forest at night” is all the natural history data available in the literature for this species (Dixon, op. cit.). On 24 February 2009, we collected a male L. vitti (QCAZ 8708, SVL = 368 mm, TL = 110 mm) in northwestern Ecuador, Carchi Province, Chilmá Bajo (0.8647222°N, 78.0497222°W, 2071 m elev.). This specimen had recently eaten, as indicated by an anterior swelling at about one fourth its SVL. The forced-out prey was a specimen of an undescribed species of the bufonid toad Osornophryne (QCAZ 40028, SVL = 30.4 mm). No records of Osornophryne are known from this locality. Moreover, this is the ﬁrst record of predation of Osornophryne by any vertebrate. Both specimens are deposited in the Herpetological Review 40(3), 2009 herpetological collection of Museo de Zoología QCAZ, Pontiﬁcia Universidad Católica del Ecuador. Submitted by OMAR TORRES-CARVAJAL (e-mail: omartorcar@gmail.com), JUAN M. GUAYASAMIN, ELICIO TAPIA, and SILVIA ALDÁS, Escuela de Biología, Pontiﬁcia Universidad Católica del Ecuador, Apartado Postal 17-01-2184, Quito, Ecuador. MASTIGODRYAS MELANOLOMUS (NCN). PREY. Information on the trophic ecology and individual prey species of certain tropical snakes is scarce. Mastigodryas melanolomus is a moderately-sized colubrid snake that forages on the ground and preys on lizards, small snakes, reptile eggs, nesting birds, and small mammals (Savage 2002. The Amphibians and Reptiles of Costa Rica: A Herpetofauna Between Two Continents, Between Two Seas. University of Chicago Press, Chicago, 994 pp.). On 14 October 2004 at 1530 h, in the Reserva Biológica Hitoy Cerere (Costa Rica, Limón Province, 9.66°N, 83.03°W, ca. 200 m elev.), I observed a M. melanolomus with a total length of about 1.2 m that had entered a hole in the ground up to about the anterior third of its body. When it reappeared, it was holding an adult Ameiva festiva in its mouth (Fig. 1). Although M. melanolomus is known to prey on other species of Ameiva (Seib 1984. J. Herpetol. 18:412–420), A. festiva has not yet been reported as a prey item of M. melanolomus. FIG. 1. Mastigodryas melanolomus preying on Ameiva festiva. Submitted by DAVID MATTHIAS DEHLING, Department of Ecology, Animal Ecology, Faculty of Biology, Philipps-University Marburg, Karl-von-Frisch-Straße 8, D-35032 Marburg, Germany; e-mail: Dehling@students.uni-marburg.de. OXYRHOPUS CLATHRATUS (NCN) PREY. Species of Oxyrhopus are known to prey on small lizards (Bernarde and Machado 2000. Herpetol. Rev. 31:247–248.). On 2 September 2004, during ﬁeldwork in Teresópolis municipality, Rio de Janeiro State, southeastern Brazil (22.448442°S, 42.983542°W), we collected a recently deceased juvenile Oxyrhopus clathratus (312 mm total length), that contained an adult Placosoma cordyline (153.3 mm total length). The cause of death of the snake is unknown. This is the ﬁrst report of predation of P. cordyline by O. clathratus. Both specimens are deposited in the herpetological collection of the Universidade Federal do Estado do Rio de Janeiro, Brazil, under the accession numbers ZUFRJ 1650 (Oxyrhopus clathratus) and 1651 (Placosoma cordyline). We thank Universidade Federal do Estado do Rio de Janeiro for transport logistics, and Henrique C. Costa and Paulo Sérgio Bernarde for suggestions on the manuscript. Submitted by VICTOR G. DILL ORRICO, Departamento de Zoologia, UNESP - Universidade Estadual Paulista Júlio de Mesquita Filho, Av. 24-A, num 744, Bela Vista, Departamento de Zoologia, I.B.; Cx. P. 199, CEP: 13506-900 - Rio Claro, SP – Brazil (e-mail: victordill@gmail.com); and PAULO NOGUEIRA DA COSTA,. Laboratório de Anfíbios e Répteis - Universidade Federal do Rio de Janeiro, Departamento de Zoologia, Caixa Postal 68.044, CEP 21944-970, Rio de Janeiro, RJ, Brazil (e-mail: nogpj@yahoo. com.br). OXYRHOPUS GUIBEI (False Coralsnake). DIET. Oxyrhopus guibei is a terrestrial, crepuscular-nocturnal pseudoboine snake in southeastern Brazil (Sazima and Abe 1991. Stud. Neotr. Fauna Environ. 26:159–164). Reported prey items include rodents, lizards (Andrade and Silvano 1996. Rev. Bras. Zool. 13[1]:143–150; França et al. 2008. Copeia 2008:23–38), and birds (Sazima and Abe 1991. Stud. Neotr. Fauna Environ. 26:159–164). We dissected 17 specimens of O. guibei from the vicinity of the Irapé Power Plant (16.75°S; 42.53°W), Minas Gerais state, Brazil, collected between February 2004 and July 2006 during a faunal monitoring program. We found 10 prey items in the stomachs of three males (total length: 404, 629, and 811 mm), two females (total length: 599 and 1016 mm), and three juveniles (total length: 289, 306, and 363 mm). Prey consisted of rodents (70%) and lizards (30%). Nonetheless, one adult female O. guibei (1016 mm total length) had digested parts of an individual Oxymycterus sp. (Cricetidae) in its stomach. This snake also contained 10 vitellogenic follicles (> 10 mm, following Pizzatto and Marques 2002. Amphibia-Reptilia 4:495–504). Rodents of the genus Oxymycterus have semi-fossorial habits and are endemic to South America (Hershkovitz 1994. Fieldiana Zool. 79:1–43; Câmara and Murta 2003. Mamíferos da Serra do Cipó. Ed. PUC Minas. Belo Horizonte. 129 pp.). The rodent body parts had a volume of 5478.7 mm3 and seemed derived from an adult individual. To our knowledge this is the ﬁrst record of the semi-fossorial rodent Oxymycterus as a food item of O. guibei. Furthermore, another two species of rodents belonging to the Cricetidae family were found: four Necromys lasiurus in three specimens and two Calomys tener in one. Three unidentiﬁed tails of lizards were found in three juvenile specimens. Representative specimens are deposited in the Herpetological Laboratory of Museu de Ciências Naturais of Pontifícia Universidade Católica de Minas, Minas Gerais, Brazil (MCNR 915, 1718, 1847, 1923, 2511). We thank Sônia A. Talamoni for identifying the rodents and FAPEMIG for ﬁnancial support to LBN. Herpetological Review 40(3), 2009 357 Submitted by LAURA RODRIGUES VIEIRA DE ALENCAR, Departamento de Ecologia, Instituto de Biociências, Universidade de São Paulo, Rua do Matão, Travessa 14, Cidade Universitária, São Paulo, SP, Brazil, CEP 05508-090 (e-mail: laura.alencar@globo.com); CONRADO A. B. GALDINO, present address: Departamento de Biologia, Universidade Federal do Ceará, Campus do Pici, Bloco 906, Fortaleza, Ceará, CEP 60455760 (e-mail: galdinoc@gmail.com); and LUCIANA BARRETO NASCIMENTO, Programa de Pós-graduação em Zoologia de Vertebrados, Laboratório de Herpetologia, Museu de Ciências Naturais, Pontifícia Universidade Católica de Minas Gerais, Avenida Dom José Gaspar, n. 290, Bairro Coração Eucarístico, Campus PUC Minas, Belo Horizonte, Minas Gerais, Brazil, CEP 30535-610 (e-mail: luna@pucminas.br). PHILODRYAS CHAMISSONIS (Long-tailed Snake) and LIOLAEMUS NITIDUS. PREDATION DETERMINED BY PIT TAG. As part of a multi-species capture-recapture study in central Chile (31.9861°S, 71.1696167°W, 820 m elev., datum Prov SudAm 56) during October 2007, we collected a Philodryas chamissonis (68 cm total length, 24 g) that contained a secondarily-ingested PIT tag in its stomach. The PIT tag originally had been placed in a juvenile Liolaemus nitidus (5.0 cm SVL, 4.6 g) in December 2006 (10 months earlier). Thus, the minimum prey/predator ratio is 19.2%, exceeding the mean value of 7.7 ± 3.1% reported by Greene and Jaksic (1992. Rev. Chil. Hist. Nat. Santiago, Chile 65:485–493) for museum specimens of P. chamissonis. Our observation suggests the potential utility of PIT tags in long-term studies of predation. Submitted by GABRIEL LOBOS, Centro de Estudio de Vida Silvestre, Facultad de Ciencias Veterinarias y Pecuarias, Universidad de Chile, Casilla 2 correo 15, Santiago, Chile (e-mail: galobos@yahoo.com); MARTÍN A. H. ESCOBAR, ROBERTO F. THOMSON (e-mail: rthomsonsaa@gmail.com), Laboratorio de Ecología de Vida Silvestre, Facultad de Ciencias Forestales, Universidad de Chile, Casilla 9206, Santiago, Chile; and ALEJANDRA ALZAMORA, Centro de Estudio de Vida Silvestre, Facultad de Ciencias Veterinarias y Pecuarias, Universidad de Chile, Casilla 2 correo 15, Santiago, Chile, Casilla 2 correo 15, Santiago, Chile. RHINOCHEILUS LECONTEI (Long-nosed Snake). CANNIBALISM. The diet of Rhinocheilus lecontei consists mainly of lizards, small mammals, and squamate eggs (Rodríguez-Robles et al. 1999. J. Herpetol. 33:87–92; Rodríguez-Robles and Greene 1999. J. Zool. Lond. 248:489–499). Although general references to predation on snakes have appeared (e.g., Degenhardt et al. 1996. Amphibians and Reptiles of New Mexico. Univ. New Mexico Press, Albuquerque. 431 pp.; Stebbins 2003. A Field Guide to Western Reptiles and Amphibians, 3rd ed., revised. Houghton Mifﬂin Co., Boston, Massachusetts. 533 pp.; Werler and Dixon 2000. Texas Snakes: Identiﬁcation, Distribution, and Natural History. Univ. Texas Press, Austin. 437 pp.), speciﬁc reports are lacking and thus it is unclear if R. lecontei preys on snakes in the wild. Klauber (1941. Trans. San Diego Soc. Nat. Hist. 9[29]:289–332) reported 358 FIG. 1. Adult Rhinocheilus lecontei ingesting a juvenile conspeciﬁc, Alameda Co., California. feeding on Chionactis occipitalis in captivity. Here I document an instance of predation on a juvenile conspeciﬁc. On 17 May 2009 at 2307 h, I observed an adult R. lecontei (estimated total length 90 cm) ingesting a juvenile conspeciﬁc (ca. 32 cm TL) (Fig. 1). Although ingestion had commenced before my arrival, the sequence I observed was completed in ca. 2 minutes. The smaller snake did not display any obvious injuries. This observation took place 24 km by air SE of Livermore, Alameda Co., California, USA. Submitted by CHAD M. LANE, 426 Swallow Ct, Manteca, California 95336, USA; e-mail: ChadMLane@aol.com. THAMNOPHIS ELEGANS VAGRANS (Wandering Gartersnake). DEFENSIVE BEHAVIOR. Several species of snakes have been observed to enlarge or triangulate the head as a defensive response to a predator (Greene 1988. In C. Gans and R. B. Huey [eds], Biology of the Reptilia, Volume 16, Ecology B, pp. 1–152. Alan R. Liss, New York). In June 2007, I observed this behavior as a defensive response in three specimens of Thamnophis elegans vagrans (ca. 45 cm total length) in a residential area of Springville, Utah, USA (40.1529°N, 111.5960°W). Sexes were not determined. Defensive responses in the snakes were elicited by a peck on the head by a nest-guarding American Robin (Turdus migratorius). The nest was located 1.75 m above ground in a cherry tree (Prunus cerasus). The robin attacked up to 4.6 m from its nest. The original observation occurred with no manipulation on my part. However, after the initial observation, I captured two other T. e. vagrans within 200 m of the nest site and placed them individually under the same robin nest. The bird responded in the same manner to these new snakes (i.e., ﬂying from nest to ground and pecking at the snake’s head). Upon being attacked, the snakes quickly coiled and exhibited head triangulation, accompanied by aggressive posturing, i.e. in a stance poised to strike, and repeated striking with mouth open. Once the snakes exhibited this defensive behavior, attacks by the bird ceased. Head triangulation as a defensive response has been documented for many colubrid species (Greene 1988, op. cit.), and has apparently evolved independently multiple times (Young et al. 1999. J. Zool., Lond. 248:169–177). However, to the best of my knowledge this is the ﬁrst report of head triangulation in Thamnophis. Werner Herpetological Review 40(3), 2009 (1983. Israel J. Zool. 32:205–228) postulated that this behavior can be classiﬁed as Batesian mimicry of sympatric venomous snakes. Crotalus viridis (Western Rattlesnake) is broadly sympatric with T. e. vagrans over most of its range (Stebbins 2003. A Field Guide to Western Reptiles and Amphibians. Houghton Mifﬂin Co., Boston, Massachusetts. 533 pp.). Special thanks to Caleb, Shiloh, Jubal, Micah, and Cyrus Rasmussen for help in catching and observing the snakes. Submitted by JOSH E. RASMUSSEN, Department of Biology, Brigham Young University, Provo, Utah 84663, USA; e-mail: josh_rasmussen@byu.edu. Cover Images for Herpetological Review We are looking for photographic images to appear on future covers of HR. To be considered, preferred images should have the following qualities: • • • • • Should be technically superior photographs (e.g., composition, lighting, etc., should be excellent; subject must be in focus). Should be taken in vertical format, or, if in landscape format, permit cropping to achieve a vertical orientation. Should be based on ﬁlm or digital media; if the latter, the native resolution must be sufﬁciently high to permit cropping and/or enlargement to print publication quality (many “point-and-shoot” digital cameras do not produce image ﬁles with sufﬁcient resolution for print publication). Preference will be given to images that depict poorly known species. These could include recently described taxa or species for which a color illustration has never been published. Preference will be given to images that communicate some aspect of the biology of the organism (e.g., predation, feeding, courtship, crypsis). Images for consideration should be submitted as low-resolution jpg or pdf ﬁles. Do not send full resolution images via email. All submissions or questions should be directed to the Editor (herpreview@gmail.com). GEOGRAPHIC DISTRIBUTION Instructions for contributors to Geographic Distribution appear in Volume 40, Number 1 (March 2009, p. 106). Please note that the responsibility for checking literature for previously documented range extensions lies with authors. Do not submit range extension reports unless a thorough literature review has been completed. CAUDATA – SALAMANDERS AMBYSTOMA MACULATUM (Spotted Salamander). USA: ARKANSAS: SEARCY CO.: Off AR 14, ca. 2 km down Ramblewood Trail in a small pool of water (36.0568°N, 92.6044°W; WGS 84). 20 February 2009. M. B. Connior. Veriﬁed by S. E. Trauth. Arkansas State University Museum of Zoology Herpetology Collection (ASUMZ 31298). This adult male is a new county record ﬁlling a distributional hiatus between previous records in Marion, Stone, Van Buren, and Newton counties in northern Arkansas (Trauth et al. 2004. The Amphibians and Reptiles of Arkansas. University of Arkansas Press, Fayetteville. 421 pp.). Submitted by MATTHEW B. CONNIOR, Department of Biological Sciences, Arkansas State University, P.O. Box 599, State University, Arkansas 72467, USA; e-mail: matthew. connior@smail.astate.edu. AMBYSTOMA MACULATUM (Spotted Salamander). USA: TENNESSEE: VAN BUREN CO.: Ephemeral pond near entrance to paved ﬁeld just prior to Fall Creek Falls Inn parking lot (35.6561ºN, 85.3711ºW: datum NAD 27). Matthew L. Brown. Veriﬁed by A. Floyd Scott. One adult located, photographed and catalogued at Austin Peay State University Center for Field Biology (APSU 18953). First county record (Redmond and Scott 1996. Atlas of Amphibians in Tennessee. Misc. Publ. No. 12. The Center for Field Biology, Austin Peay State University, Clarksville, Tennessee. 94 pp. Internet version [http://www.apsu.edu/amatlas] contains links to information regarding Tennessee distribution of amphibians recorded since 1996; accessed 05 February 2009). Extends range approximately 15 km SW of nearest recorded occurrence. Submitted by MATTHEW L. BROWN, Fall Creek Falls State Park, Pikeville, Tennessee 37367, USA; e-mail: matthew. brown@state.tn.us. AMBYSTOMA TIGRINUM (Eastern Tiger Salamander). USA: OKLAHOMA: MCCURTAIN CO.: 1.2 km E Pollard Community on Ferguson Road (33.47640°N, 94.42591°W; WGS84). 18 March 2009. J. Ferguson. Veriﬁed by S. E. Trauth. Arkansas State University Herpetological Museum (ASUMZ photographic voucher 31306). First record for the county (Oklahoma Biological Survey DOKARRS Search; http://www.biosurvey.ou.edu/dbsrch/dokalist. php); adult male with swollen vent found crossing gravel road in early morning. Numerous A. tigrinum from the state are in the collection of the Sam Noble Oklahoma Museum of Natural History of Oklahoma; one specimen (SNOMNH 25011) is from adjacent Pushmataha Co. The species occurs immediately southward across the Red River in Red River Co., Texas (Dixon 2000. Amphibians and Reptiles of Texas, 2nd ed. Texas A&M University Press, College Station, Texas. 421 pp.). However, A. t. tigrinum has not yet been reported Herpetological Review 40(3), 2009 359 north of the Red River from any counties along the southwestern tier in Arkansas (Trauth et al. 2004. Amphibians and Reptiles of Arkansas. Univ. Arkansas Press, Fayetteville. 421 pp.). Submitted by CHRIS T. MCALLISTER, RapidWrite, 102 Brown Street, Hot Springs National Park, Arkansas 71913, USA (e-mail: drctmcallister@aol.com); HENRY W. ROBISON, Department of Biology, Southern Arkansas University, Magnolia, Arkansas 71754, USA (e-mail: hwrobison@suddenlink.net); DAVID ARBOUR, De Queen, Arkansas 71832, USA (e-mail: arbour@windstream.net); JACKYE FERGUSON, United States Forest Service, HC 60, Box 430, Haworth, Oklahoma 74740, USA; and TERRY STUART, Oklahoma Department of Wildlife Conservation, HC 60, Box 160, Haworth, Oklahoma 74740, USA. AMBYSTOMA TIGRINUM (Eastern Tiger Salamander). USA: TEXAS: LEE CO.: 6.1 km W of the junction of Farm-Market Roads 116 and 696, on FM 696 (30.5725º N, 97.0888889ºW). 25 March 2009. Philip Ralidis. Veriﬁed by Heather Prestridge. Texas Cooperative Wildlife Collection (photo voucher) TCWC-PV001. The large adult female (total length 175.5 mm, tail 80 mm) was captured alive about 2100 h, crossing FM 696 during a heavy rain. New county record (Dixon 2000. Amphibians and Reptiles of Texas. 2nd ed. Texas A&M University Press, College Station, Texas). Submitted by PHILIP RALIDIS, 3003 Loveland Cove, Austin, Texas 78746, USA; JAMES R. DIXON (e-mail: jrdixon5@verizon. net) and TOBY J. HIBBITTS (e-mail: thibbitts@tamu.edu), Texas Cooperative Wildlife Collections, Department of Wildlife and Fisheries Sciences, Texas A&M University, 2258 Tamu, College Station, Texas 77843-2258, USA. CRYPTOBRANCHUS ALLEGANIENSIS (Hellbender). USA: TENNESSEE: RUTHERFORD CO.: J. Percy Priest Reservoir, Spring Creek branch (36.0353ºN, 86.4467ºW; NAD83). 20 April 1990. Mark Hodge. Veriﬁed by M. L. Niemiller. An adult female gravely injured by a ﬁsherman’s attempts to retrieve baited hook from stomach. Deposited in the Herpetology Collection at Middle Tennessee State University (MTSU 422C). First record for county and for the Stones River watershed, ca. 60 km SE of historical record for the Cumberland River in Davidson Co., Tennessee (Redmond and Scott 1996. Atlas of Amphibians in Tennessee. Misc. Publ. No. 12. The Center for Field Biology, Austin Peay State University, Clarksville, Tennessee. 94 pp. Internet version [http://www.apsu. edu/amatlas] contains links to information regarding Tennessee distribution of amphibians recorded since 1996; accessed 03 March 2009). Submitted by BRIAN T. MILLER, Department of Biology, Middle Tennessee State University, Murfreesboro, Tennessee 37132, USA (e-mail: bmiller@mtsu.edu); and JOYCE L. MILLER, MIMIC, Middle Tennessee State University, Murfreesboro, Tennessee 37132, USA (e-mail: jlmiller@mtsu.edu). NOTOPHTHALMUS VIRIDESCENS LOUISIANENSIS (Central Newt). USA: ARKANSAS: DALLAS CO.: 5.6 km S Sparkman off St. Hwy 7 at Brushy Creek (Sec. 15, T10S, R17W). 07 May 1986. Henry W. Robison. Veriﬁed by S. E. Trauth. Arkansas State University Herpetological Museum (ASUMZ 30829). New county record ﬁlling a distributional gap between Clark and Cleveland counties (Trauth et al. 2004. Amphibians and Reptiles of Arkan- 360 sas. Univ. Arkansas Press, Fayetteville. 421 pp.), and leaving only Lafayette Co. without documentation of N. v. louisianensis for all of south Arkansas. Submitted by HENRY W. ROBISON, Department of Biology, Southern Arkansas University, Magnolia, Arkansas 71754, USA (email: hwrobison@saumag.edu); and CHRIS T. MCALLISTER, Department of Physical and Life Sciences, Chadron State College, Chadron, Nebraska 69337, USA (e-mail: cmcallister@csc.edu). PSEUDOTRITON MONTANUS (Mud Salamander) USA: ALABAMA: BIBB CO.: 18.0 km WSW of Eoline (32.90679ºN, 87.39313ºW, WGS84). 14 November 2008. Heather Cunningham, Walter H. Smith, and Joseph J. Apodaca. Veriﬁed by Leslie J. Rissler. University of Alabama Herpetological Collection (UAHC 16175, 16178). Two adult Pseudotriton montanus were found under leaf litter along the margin of a beaver-impounded wetland in the Talladega National Forest (Oakmulgee District) of west-central Alabama. This wetland was located along an unnamed tributary of Fivemile Creek in a mixed bottomland hardwood/yellow pine forest, with extensive canopy cover. This site is located ca. 60 km outside of the currently accepted range for P. montanus and is 94 km from the nearest catalogued voucher for the species (Petranka 1998. Salamanders of the United States and Canada. Smithsonian Institution Press, Washington, DC). The presence of a disjunct portion of the range of P. montanus in eastern Mississippi (Petranka, op. cit.) presents several possible implications for this new locality. It is possible that the contiguous range of P. montanus extends westward at least to the Alabama locality described here. Because this Alabama locality is centered between the recognized contiguous range and the disjunct Mississippi population it may also be possible that the range of P. montanus extends westward across the Gulf Coastal Plain to encompass the Mississippi population. Submitted by HEATHER CUNNINGHAM, WALTER H. SMITH, and JOSEPH J. APODACA, Department of Biological Sciences, Ecology, Evolution, and Systematics Section, Box 870345, University of Alabama, Tuscaloosa, Alabama 35487, USA. TARICHA GRANULOSA (Rough-skinned Newt) USA: CALIFORNIA: BUTTE CO.: North Fork Feather River, ca. 1.4 mi. upstream from Poe Powerhouse, west bank (39.72273ºN, 121.47105ºW; datum: WGS 84); 286 m elev.; adult (California Academy of Sciences 241781) captured on 13 March 2008 at 1300 h underwater in a pool tail-out; water temperature 21.5ºC; K. D. Wiseman. Veriﬁed by Ricka Stoelting. A second specimen (CAS 234628) was found at 39.80197ºN, 121.44695ºW, 404 m elev. on 15 June 2006. Specimen veriﬁed by Jens Vindum. CAS 241781 extends the known distribution of T. granulosa in the Sierra Nevada of California ca. 7.7 miles SE, while CAS 234628 extends the distribution ca. 9.1 miles E of specimens previously regarded as the southern extent of their distribution in the Sierra Nevada (MVZ 56784–56798; Hayes and Cliff 1982. Herpetol. Rev. 13[3]:85–87; Koo and Vindum 1999. Amphibians and Reptiles of the Plumas National Forest: Historical Collections and California Academy of Sciences 1998 and 1999 Surveys; Koo et al. 2004. Results of 02-CS-11050650-029, the California Academy of Sciences Survey: Amphibians and Reptiles of the Lassen National Forest). These specimens are also signiﬁcant as they document T. Herpetological Review 40(3), 2009 granulosa occupying the North Fork Feather River watershed, a distinct watershed from the West Branch Feather River watershed, previously considered the southern extent of the species range in the Sierra Nevada of California (Hayes and Cliff, op. cit.). Taricha granulosa is found in sympatry with Taricha torosa at localities provided herein. Submitted by KEVIN D. WISEMAN, Garcia and Associates (GANDA), 2601 Mission Street, Suite 600, San Francisco, California 94110, USA; Department of Herpetology, California Academy of Sciences, 55 Music Concourse Drive, Golden Gate Park, San Francisco, California 94118, USA; e-mail: kwiseman@garciaan dassociates.com. Calls have been audible for at least two years at the Fall Creek Falls State Park Golf Course pond. Possible introduction, 62.2 km N of nearest recorded occurrence (Miller et al. 2007. Herpetol. Rev. 38:97). First county record (Redmond and Scott 1996. Atlas of Amphibians in Tennessee. Misc. Publ. No. 12. The Center for Field Biology, Austin Peay State University, Clarksville, Tennessee. 94 pp. Internet version [http://www.apsu.edu/amatlas] contains links to information regarding Tennessee distribution of amphibians recorded since 1996; accessed 17 January 2009). Submitted by MATTHEW L. BROWN, Fall Creek Falls State Park, Pikeville, Tennessee 37367, USA; e-mail: matthew. brown@state.tn.us. ANURA – FROGS GASTROPHRYNE CAROLINENSIS (Eastern Narrow-mouthed Toad). USA: ARKANSAS: BOONE CO.: 3.2 km SE of Harrison off US 65 (36.122014°N, 93.039479°N; NAD83). 10 June 1995. H. W. Robison. Veriﬁed by S. E. Trauth. Arkansas State University Herpetological Museum (ASUMZ 31305). New county record partially ﬁlling a distributional hiatus in northern Arkansas between Benton and Marion counties (Trauth et al. 2004. Amphibians and Reptiles of Arkansas. Univ. Arkansas Press, Fayetteville. 421 pp.). This frog has now been documented in 69 of 75 (92%) Arkansas counties. Submitted by CHRIS T. MCALLISTER, RapidWrite, 102 Brown Street, Hot Springs National Park, Arkansas 71913, USA (e-mail: drcmcallister@aol.com); and HENRY W. ROBISON, Department of Biology, Southern Arkansas University, Magnolia, Arkansas 71754, USA (e-mail: hwrobison@suddenlink.net). AGALYCHNIS SPURELLI (Gliding Treefrog). COSTA RICA: SAN JOSE: Distrito Carara, El Sur de Turrubares, (9.4339ºN, 84.34038ºW; WGS84), 385 m elev. 19 August 2006. A. Vega, J. M. Robertson, and G. Chaves. Veriﬁed by Federico Bolaños. Museum of Zoology, Universidad de Costa Rica, San Jose (UCR 19214). New province record and a 114 km NW range extension from the closest previous record at Palmar Norte, Puntarenas, expanding its distribution along the Paciﬁc versant on the Cordillera de Talamanca (Savage 2002. The Amphibians and Reptiles of Costa Rica: A Herpetofauna Between Two Continents, Between Two Seas. Univ. Chicago Press, Chicago, Illinois. xx + 934 pp.). The frog was calling from a tree-lined temporary pool in an agricultural ﬁeld once covered by lowland rainforest, along with a large breeding congregation of Agalychnis callidryas. Submitted by ANDRES VEGA, AMBICOR, 400 E., 75 S., 75 E. de la Municipalidad de Tibas, Tibas, Costa Rica (e-mail: avegak@hotmail.com); JEANNE M. ROBERTSON, Department of Ecology and Evolutionary Biology, Cornell University, Ithaca, New York 14853, USA (e-mail: jmr79@cornell.edu); and GERARDO CHAVES, Museum of Zoology, University of Costa Rica, San Jose (e-mail: cachi@biologia.ucr.ac.cr). ANAXYRUS DEBILIS DEBILIS (Eastern Green Toad). USA: TEXAS: STEPHENS CO.: N of Eolian (32.68165ºN, 99.03378ºW; 370 m elev.). 21 March 2009. J. Meik, J. Streicher, and S. Schaack. Veriﬁed by C. J. Franklin. University of Texas at Arlington, UTA A-59175. Collected under a rock adjacent to a gravel road in Post Oak savannah. First known county record of the species, which ﬁlls a distributional gap in north-central Texas (Dixon 2000. Amphibians and Reptiles of Texas. 2nd ed. Texas A&M Univ. Press, College Station). Collected under Texas Parks and Wildlife permit number SPR-0707-1387. Submitted by JEFFREY W. STREICHER (e-mail: e-mail: streicher@uta.edu.), JESSE M. MEIK, and SARAH SCHAACK, Department of Biology, The University of Texas at Arlington, Arlington, Texas 76019, USA. HYLA CINEREA (Green Treefrog). USA: TENNESSEE: VAN BUREN CO.: Fall Creek Falls State Park Golf Course Pond (35.65ºN, 85.3606ºW: datum NAD 27). 11 July 2008. Matthew L. Brown and Leigh Ann Brown. Veriﬁed by A. Floyd Scott. One adult photographed and catalogued at Austin Peay State University Center for Field Biology (APSU 19959). Only specimen found in park. KALOULA PULCHRA (Painted Bull Frog): MALAYSIA: SARAWAK: MIRI DIVISION: Kuala Lutong, nr. Lutong Town (4.4708333ºN, 114.0038889ºE). From beach forest, within a patch of Casuarina with blackwater pools. 25 October 2007. I. Das. USDZ 1.12153. Veriﬁed by Kelvin K. P. Lim. First record for Sarawak. Bornean records from Sinkawang, Kalimantan (Indonesia) by Inger (1966. Fieldiana Zool. 52:1–402), Sabah East Malaysia, without locality (Inger and Tan 1996. Rafﬂes Bull. Zool. 44:551–574), Crocker Range, Sabah (Das 2006. Amphibian and Reptile Conservation 4[1]:3–11), and Bandar Seri Begawan, in Brunei Darussalam (Charles 2008. Herpetol. Rev. 39:462–463). Sarawak locality is ca. 115 km W of Brunei record, and choruses of breeding males heard during late afternoons and nights of 25–26 October 2007 indicate species is established locally. Considered introduced by anthropogenic activities in Borneo and is currently spreading west along north coast, and widespread in Southeast Asian mainland (Inger 1999. In W. E. Duellman [ed.], Patterns of Distribution of Amphibians: A Global Perspective, pp. 445–482. The John Hopkins University Press, Baltimore and London). I thank Lee Nyanti, Andrew Alek Tuen, and Matteo Veronesi for supporting ﬁeld work in Lutong under Project Kuala for Shell Sarawak Berhad and Environmental Resources Management Sdn. Bhd. Submitted by INDRANEIL DAS, Institute of Biodiversity and Environmental Conservation, Universiti Malaysia Sarawak, 94300 Kota Samarahan, Sarawak, Malaysia; e-mail: idas@ibec.unimas. my. Herpetological Review 40(3), 2009 361 LITHOBATES GRYLIO (Pig Frog). USA: TEXAS: CHAMBERS CO.: Highway 124, 5 km N of High Island (29.61379ºN, 94.38073ºW; WGS 84). 25 June 2008. Collected by Paul Crump. Veriﬁed by Carl J. Franklin. University of Texas Arlington (UTADC 3691–3698), and tissue deposited at the MRJ Forstner Frozen Tissue Collection at Texas State University (MF 27033). Submitted by PAUL CRUMP (e-mail: pcrump@houstonzoo. org), RACHEL ROMMEL, Department of Conservation and Science, Houston Zoo, 1513 North Macgregor Way, Houston, Texas 77030, USA (e-mail: rrommel@houstonzoo.org); and JAIME GONZALEZ, Katy Prairie Conservancy, Katy Prairie Conservancy, Waller, Texas 77098, USA (e-mail: jaimegonzalez72@gmail. com). LITHOBATES PALUSTRIS (Pickerel Frog). USA: ARKANSAS: SEARCY CO.: Off AR 14, ca. 2.2 km down Ramblewood Trail in small drainage creek (UTM 15N 0536006E, 3990296N; WGS 84). 26 March 2009. M. B. Connior. Veriﬁed by S. E. Trauth. Arkansas State University Museum of Zoology Herpetology Collection (ASUMZ 31312). Adult female caught by hand. First county record ﬁlling a distributional gap among surrounding Stone, Marion, and Newton counties (Trauth et al. 2004. The Amphibians and Reptiles of Arkansas. Univeristy of Arkansas Press, Fayetteville. 421 pp.). Submitted by MATTHEW B. CONNIOR, Department of Biological Sciences, Arkansas State University, P.O. Box 599, State University, Arkansas 72467, USA; e-mail: matthew. connior@smail.astate.edu. LITHOBATES SYLVATICUS (Wood Frog). USA: TENNESSEE: VAN BUREN CO.: Flooded pit near Piney Falls at Fall Creek Falls State Park (35.67278ºN, 85.375ºW: datum NAD27). 06 January 2009. Matthew L. Brown and Thomas P. Solomon. Veriﬁed by A. Floyd Scott. Twenty-two adults and egg masses located, two adults photographed and catalogued at Austin Peay State University Center for Field Biology (APSU 18954).. First county record (Redmond and Scott 1996. Atlas of Amphibians in Tennessee. Misc. Publ. No. 12. The Center for Field Biology, Austin Peay State University, Clarksville, Tennessee. 94 pp. Internet version [http://www.apsu.edu/amatlas] contains links to information regarding Tennessee distribution of amphibians recorded since 1996; accessed 17 January 2009). Extends range from nearest documented occurrence by approximately 25 km west. Submitted by MATTHEW L. BROWN, Fall Creek Falls State Park, Pikeville, Tennessee, 37367, USA; e-mail: matthew. brown@state.tn.us PHYSALAEMUS KROYERI (Kroyer’s Dwarf Frog). BRAZIL: PIAUÍ: Municipalidade de Piracuruca: Parque Nacional de Sete Cidades (03.978ºS, 41.556ºW; datum SAD69; 110 m elev.). April 2007. B. B. Annunziata, I. S. Castro, and W. M. Fontenele. February 2009. B. B. Annunziata, M. R. A. Mendes and D. Cisne. Veriﬁed by D. Silvano and P. Valdujo. Herpetological Collection of Universidade Estadual do Piauí, Parnaíba (UESPI 0062, 0394–0404). This species ranges from the northern part of the State of Minas Gerais and from the central part of the State of Bahia, north to São José do Bonﬁm and Mamanguape in State of Paraíba, occurring up to 240 m elev. (Frost 2009 Amphibian Species of the World 362 http://research.amnh.org/herpetology/amphibia/index.php IUCN 2009 Global Amphibian Assessment). First state record, extends distribution ca. 588 SE from the city of São José do Bonﬁm, Paraíba state (Arzabe et al. 1998 Herpetol. J. 8:111–113), and a new elevation range of the species, up to 110 m. Submitted by BRUNO B. ANNUNZIATA, Departamento de Zoologia, Coleção Herpetológica, Universidade de Brasília (UnB), CEP 70.855-160, Brasília, Distrito Federal, Brazil; IRISMAR S. CASTRO, Departamento de Biologia, Universidade Estadual do Piauí (UESPI), CEP 64.202-220, Parnaíba, Piauí, Brazil; WOLNEY M. FONTENELE, Departamento de Biologia, Universidade Estadual do Piauí (UESPI), CEP 64.202-220, Parnaíba, Piauí, Brazil; and DANIEL CISNE, Departamento de Biologia, Universidade Estadual do Piauí (UESPI), CEP 64.202-220, Parnaíba, Piauí, Brazil; e-mail: barcellos.ba@gmail.com. PSEUDACRIS CRUCIFER (Spring Peeper). USA: ARKANSAS: CARROLL CO.: 3.2 km S Berryville off St. Hwy 21 (36.201993°N, 93.340094°W; NAD83). 23 March 1996. H. W. Robison. Veriﬁed by S. E. Trauth. Arkansas State University Herpetological Museum (ASUMZ 31320). New county record completely ﬁlling a distributional gap in northwestern Arkansas among Benton and Boone counties (Trauth et al. 2004. Amphibians and Reptiles of Arkansas. Univ. Arkansas Press, Fayetteville. 421 pp.). This frog has now been reported from all but six counties of the state (Trauth et al. op. cit.). Submitted by CHRIS T. MCALLISTER, RapidWrite, 102 Brown Street, Hot Springs National Park, Arkansas 71913, USA (e-mail: drcmcallister@aol.com); and HENRY W. ROBISON, Department of Biology, Southern Arkansas University, Magnolia, Arkansas 71754, USA (e-mail: hwrobison@suddenlink.net). RHACOPHORUS PSEUDOMALABARICUS (False Malabar Tree Frog). INDIA: KERALA STATE: WESTERN GHATS: Sakkulathumedu (9.8830556ºN, 77.2330556ºE), Idukki District, 1080 m elev., 10 km N of Periyar Tiger Reserve. 2100 h, 19 April 2008. G. Srinivas and Suganthan R. Sakthivel. Digital image voucher, USDZ (IMG) 1.27. Veriﬁed by Karthikeyan Vasudevan. Metamorphs observed 12 June 2008. Including type locality, currently known from two sites in Andiparai shola, 1190 m elev., Indira Gandhi National Park, Tamil Nadu, India (Vasudevan and Dutta 1998. Hamadryad 25[1]:21–28). Appears to be distributed in narrow elevational range. First report for Kerala State, 60 km S of type locality. Submitted by G. SRINIVAS (e-mail: sriniherp@gmail.com), S. BHUPATHY (e-mail: bhupathy.s@gmail.com), Salim Ali Centre for Ornithology and Natural History, Anaikatty, Coimbatore 641 108, Tamil Nadu, India; and SUGANTHAN R. SAKTHIVEL, Kerala Forest Research Institute, Peechi 680 653, Kerala, India. SPHAENORHYNCHUS PRASINUS: BRAZIL: PERNAMBUCO: MUNICIPALITY OF SÃO LOURENÇO DA MATA: Estação Ecológica do Tapacurá (08.03333 oS, 35.18333 oW, WGS 84, elev. 140 m). 10 January 2000. E. Maranhão dos Santos. Coleção Herpetologica da Universidade Federal Rural de Pernambuco, Unidade Acadêmica de Serra Talhada, Serra Talhada, Pernambuco, Brazil (CHUFRPE 966, adult specimen SVL 26.8 mm). Veriﬁed by M. Trefault Rodrigues. Previously known from states of Bahia (Centro de Pesquisa Herpetological Review 40(3), 2009 do Cacau, Ilheus municipality (type locality); Teixeira de Freitas municipality, and Mata de São João municipality), and Minas Gerais (Parque Estadual do Rio Doce), Brazil (Bokermann 1973. Brasil. Biol. 33[4a]:589–594; Silvano and Pimenta 2003. CD-ROM, Ilhéus, IESB/CI/CABS/UFMG/UNICAMP; Juncá 2006. Biota Neotropica May/Aug 2006 6[2], http://www.biotaneotropica.org.br 2006; Feio et al. 1999. Herpetol. Rev. 30:56–57). First state record, extends the known distribution 860 km NE from type locality and 600 km NE from Mata de São João municipality. Submitted by EDNILZA MARANHÃO DOS SANTOS, Unidade Acadêmica de Serra Talhada, Universidade Federal Rural de Pernambuco, 56900-000, Serra Talhada, PE, Brazil; and GERALDO JORGE BARBOSA DE MOURA, Post-graduate Program in Biological Sciences, Universidade Federal da Paraíba, 58059-900, Cidade Universitária, João Pessoa, PB, Brazil. TRACHYCEPHALUS NIGROMACULATUS (Black-spotted Casque-headed Treefrog). BRAZIL: BAHIA: MUNICIPALITY OF POTIRAGUÁ: Cedro Farm (0414551ºS, 8265887ºW), 188 m elev. 20 January 2008. M. A. Freitas, T. O. Lima, and T. F. Miranda. Museu de Zoologia da Universidade Federal da Bahia, Salvador, Brazil (MZUFBA 7825, collected at night below terrestrial bromeliads). Veriﬁed by I. Nunes. Previously known from coastal regions of southern Brazil. First state record extends distribution ca. 380 km N from Fundão Municipality, Espírito Santo State (Ramos and Gasparini 2004. Anfíbios do Goiapaba-Açu, Fundão, estado do Espírito Santo. Fundão. 75 pp.). Submitted by MARCO ANTÔNIO DE FREITAS, Programa de pós-graduação em zoologia, UESC (Universidade Estadual de Santa Cruz), CEP 46.500-000 Rodovia Ilhéus/Itabuna, Ilhéus, Bahia, Brazil (e-mail: philodryas@hotmail.com); and TIAGO DE OLIVEIRA LIMA, Rua São Joaquim n 11, Sagrada Família, CEP 31-700-000, Belo Horizonte, Minas Gerais, Brazil (e- mail: tiagoboidae@yahoo.com.br). TESTUDINES – TURTLES TERRAPENE CAROLINA CAROLINA (Eastern Box Turtle). USA: OHIO: SUMMIT CO.: Bath Township, corner of Kemery and Glengary Roads (41.18437ºN, 81.60213ºW). 10 July 2006. Scott and Alice Mason. Veriﬁed by Jeffrey G. Davis. Cincinnati Museum Center (CMC Herpetology Photodocumentation Collection HP 245–247). New county record (Wynn and Moody 2006. Ohio Turtle, Lizard, and Snake Atlas. Ohio Biological Survey Misc. Contrib. No. 10, Columbus). Submitted by JOHN W. FERNER, Department of Biology, Thomas More College, Crestview Hills, Kentucky 41017, USA (e–mail: john.ferner@thomasmore.edu); and VERNA VANDER KOOI, 1903 Glengary Rd., Akron, Ohio 44333, USA. TRACHEMYS SCRIPTA ELEGANS (Red-eared Slider) USA: TEXAS: EASTLAND CO.: 1.7 miles S of Cisco on TX 183, (32.35350˚N, 98.96579˚W; datum WGS84). 05 April 2009. Collected by D. J. Leavitt. Veriﬁed by T. J. Hibbitts. Texas Cooperative Wildlife Collection (TCWC 93636). One individual found dead on the road. New county record ﬁlls gap within adjacent counties (Dixon 2000. Reptiles and Amphibians of Texas. Texas A&M Univ. Press, College Station. 500 pp.). Submitted by DANIEL J. LEAVITT, Texas Cooperative Wildlife Collection, Department of Wildlife and Fisheries Sciences, Texas A&M University, College Station, Texas 77843, USA. TRACHEMYS SCRIPTA SCRIPTA (Yellow-bellied Slider) × T. S. ELEGANS (Red-eared Slider). USA: VIRGINIA: CHESTERFIELD C O .: City of Petersburg (Virginia State University, Randolph Farm), 4415 River Road (3722881°N, 77.43800°W; WGS84). 22 September 2008. Christian A. d’Orgeix and Edward N. Sismour. Veriﬁed by Jeffrey C. Beane. North Carolina State Museum (NCSM 74672–74675). Intergrades between native T. s. scripta and introduced T. s. elegans occur in southeastern Virginia (Mitchell 1994. The Reptiles of Virginia. Smithsonian Institution Press, Washingon, DC. 352 pp.). This new county record is ca. 75 km NW from the nearest known intergrade population in James City Co., Virginia (Mitchell 1994, op. cit.). An adult female, adult male, and two juveniles were caught in ﬂoating (1.2 m L × 1.2 m W × 1.2 m H), 0.6 cm2 mesh, black polyethylene ﬁsh aquaculture cages in two 0.1 ha artiﬁcial aquaculture ponds located ca. 430 m N of the Appomattox River. These individuals represent a range of markings on the head and plastron (i.e., amount and pattern of red and yellow pigment on the head and black spots on the plastron) between the two parental species. Backcrossing of intergrades with parental subspecies or among themselves would explain the range of observed phenotypic variation in the intergrades. We hypothesize that original colonization of these ponds occurred through immigration of the parental subspecies and possibly intergrades from the Appomattox River. Field work supported through S. Newton (Evans-Allen program, USDA/CSREES, P. L. 95-113, Sec. 1445) and through Virginia General Assembly appropriations for the Virginia State University Agricultural Research Station. Submitted by CHRISTIAN A. D’ORGEIX, National Evolutionary Synthesis Center, 2024 West Main Street, Durham, North Carolina 27701, USA; Department of Biology, Virginia State University, Petersburg, Virginia 23806, USA (e-mail: cdorgeix@vsu.edu); and EDWARD N. SISMOUR, Agricultural Research Station, School of Agriculture, Virginia State University, Petersburg, Virginia 23806, USA (e-mail: esismour@vsu.edu). SQUAMATA – LIZARDS ANOLIS CHLOROCYANUS (Hispaniolan Green Anole). USA: FLORIDA: PALM BEACH CO.: 1301 Summit Boulevard, West Palm Beach, The Palm Beach Zoo (26.667731°N, 80.068944°W, WGS84; elev. < 1m). 16 February 2009. Brian J. Camposano and Mark D. Halvorsen. Veriﬁed by Joseph Burgess. Florida Museum of Natural History (UF 154683–84). New county record and extends the range ca. 42 km NE of the closest known locality in Parkland, Broward Co. (Butterﬁeld et al. 1994. Herpetol. Rev. 25:77–78). Submitted by BRIAN J. CAMPOSANO, Division of Herpetology, Florida Museum of Natural History, Dickinson Hall, University of Florida, Gainesville, Florida 32611, USA (e-mail: biscuit1@uﬂ.edu); MARK D. HALVORSEN, Curator of Herpetology, The Palm Beach Zoo, 1301 Summit Boulevard, West Palm Beach, Florida 33405, USA (e-mail: mhalvorsen@palmbeachzoo. Herpetological Review 40(3), 2009 363 org); and KENNETH L. KRYSKO, Division of Herpetology, Florida Museum of Natural History, Dickinson Hall, University of Florida, Gainesville, Florida 32611, USA (e-mail: kenneyk@ﬂmnh. uﬂ.edu). ANOLIS DISTICHUS (Bark Anole). USA: FLORIDA: COLLIER CO.: 722 16th Avenue South, Naples (26.12797°N, 81.79576°W; WGS84; elev. < 1 m). 12 April 2008. Steve A. Johnson and Sam Logue. Veriﬁed by Kenneth L. Krysko. Florida Museum of Natural History (UF 152789). New county record and extends the range ca. 150 km NW of the closest known locality at SR 997 & SW 300 Street, Homestead, Miami-Dade Co. (UF 147676, Meshaka et al. 2004. The Exotic Amphibians and Reptiles of Florida. Krieger Publ. Co., Malabar, Florida. 155 pp.). Submitted by BRIAN J. CAMPOSANO, Division of Herpetology, Florida Museum of Natural History, Dickinson Hall, University of Florida, Gainesville, Florida 32611, USA (e-mail: biscuit1@uﬂ.edu); and STEVE A. JOHNSON, Wildlife Ecology and Conservation, UF/IFAS Plant City Campus, 1200 North Park Road, University of Florida, Plant City, Florida 33563, USA (email: tadpole@uﬂ.edu). PHELSUMA MADAGASCARIENSIS (Madagascar Day Gecko). USA: FLORIDA: MONROE CO.: Sugarloaf Key, 19580 Mayan Street (24.6682797ºN, 81.5277097ºW; WGS84; elev. < 1 m). 31 March 2009. Andrew P. Borgia. Veriﬁed by Kevin M. Enge. Florida Museum of Natural History (Photographic voucher UF 155282). First voucher specimen and seventh known island in the Florida Keys to which this species has been introduced (Krysko and Sheehy 2005. Caribbean J. Sci. 41:169–172, Krysko and Hooper 2007. Gekko 5:33–38). Submitted by KENNETH L. KRYSKO, Division of Herpetology, Florida Museum of Natural History, Dickinson Hall, University of Florida, Gainesville, Florida 32611, USA (e-mail: kenneyk@ﬂmnh.uﬂ.edu); and ANDREW P. BORGIA, P.O. Box 4346, Key West, Florida 33041, USA. PSEUDOCALOTES MICROLEPIS (Small-scaled Montane Forest Lizard). THAILAND: CHIANG RAI PROVINCE: Wieng Pa Pao District, Tambon (= Subdistrict) Pagnew, Moo. 7.6 km S of Ban Pa Miang Mae Hang (19.3505556ºN, 99.5341667ºE). 18 November 2007. K. Kunya. Korat Zoo Museum (KZM 002) and Queen Saovabha Memorial Institute, Bangkok (QSMI 1001). Veriﬁed by M. Sumontha (Ranong Marine Fisheries Station). First provincial record based on the rejection by Pauwels et al. (2003. Nat. Hist. J. Chulalongkorn Univ. 3[1]:23–53) of a record from Phetchaburi Province in west Thailand. Species known in Thailand from northern provinces of Chiang Mai and Chiang Rai and northeastern provinces of Chaiyaphum and Loei (Nabhitabhata et al. “2000” 2004. Checklist of Amphibians and Reptiles in Thailand. Ofﬁce of Environmental Policy and Planning, Bangkok. 152 pp.). We thank L. Chanhome (QSMI) for providing working facilities. Submitted by OLIVIER S. G. PAUWELS, Département des Vertébrés Récents, Institut Royal des Sciences naturelles de Belgique, Rue Vautier 29, 1000 Brussels, Belgium (email: osgpauwels@yahoo.fr); and KIRATI KUNYA, Korat Zoo, Muang District, Nakhon Ratchasima, Thailand (e-mail: kkunya2006@yahoo.com). 364 SQUAMATA – SNAKES BOA CONSTRICTOR (Boa Constrictor). MÉXICO: HIDALGO. Municipality of Huehuetla, Barrio Aztlán (98.0836°N, 20.43125°W; WSG 84), 496 m elev. 28 August 2008. Froylán Ramírez, Alejandro Ramírez, Adriana López, Nallely Morales, Arely Penguilly, and Arely Flores. Veriﬁed by Norma Manríquez-Morán. Colección Herpetológica, Centro de Investigaciones Biológicas, Universidad Autónoma del Estado de Hidalgo (CIB-UAEH 1623). First record for the municipality, extending the range in Hidalgo ca. 99 km S from Tepehuacán de Guerrero (Mendoza et al. 2006. In A. Ramírez-Bautista et al. [eds.], Inventarios Herpetofaunísticos de México: Avances en el Conocimiento de su Biodiversidad, pp. 99–109. Publ. Soc. Herpetol. Mex. No. 3. México, D.F.). The snake was found in tropical deciduous forest. Fieldwork was funded by CONACyT- 43761. Submitted by IRENE GOYENECHEA (e-mail: ireneg@uaeh. edu.mx), JESÚS M. CASTILLO, FROYLÁN RAMÍREZRESÉNDIZ, and ALEJANDRO RAMÍREZ-PÉREZ, Centro de Investigaciones Biológicas (CIB), Universidad Autónoma del Estado de Hidalgo, A.P. 1-69 Plaza Juárez, Pachuca, Hidalgo, México. BUNGARUS SLOWINSKII (Red River Krait). VIETNAM: QUANG NAM PROVINCE: Tay Giang District: B Ha Lee Commune (16.0086111ºN, 107.521666ºE), 750 m elev. 27 April 2006. Doan Van Kien. IEBR 2978. Veriﬁed by David Kizirian. Adult male. Previously known from Lao Cai and Yen Bai provinces (Kuch et al. 2005. Copeia 2005:818–833). Third and southernmost record from Vietnam, and ca. 750 km from type locality in Yen Bai Province. Submitted by DOAN VAN KIEN (e-mail: doanvankien@yahoo. fr) and NGUYEN QUANG TRUONG (e-mail: NQT2@YAHOO. COM), Institute of Ecology and Biological Resources, 18 Hoang Quoc Viet, Hanoi, Vietnam. COLUBER (= MASTICOPHIS) FLAGELLUM FLAGELLUM (Eastern Coachwhip). USA: GEORGIA: CHARLTON CO.: ~16 km SSW of St. George (straight distance from town center) (30.39450°N, 82.11297°W; WGS84). 16 June 2004. Nathan H. Nazdrowicz. Veriﬁed by John Jensen. Georgia Museum of Natural History photographic voucher (GMNH 50108). Photographed and released. New county record (Jensen et al. [eds.] 2008. Amphibians and Reptiles of Georgia. Univ. Georgia Press, Athens, 575 pp.). Submitted by NATHAN H. NAZDROWICZ, Department of Entomology and Wildlife Ecology, University of Delaware, Newark, Delaware 19716, USA; e-mail: spinifer@aol.com. COLUBER (= MASTICOPHIS) FLAGELLUM (Coachwhip). USA: GEORGIA: EMANUEL CO.: Ohoopee Dunes Natural Area, Hall’s Bridge Tract, just south of Hall’s Bridge Rd., 2.33 km NW of intersection with Frank Wimberly Rd. (32.530277°N, 82.4516861°W; WGS 84). 13 December 2007. Kerry T. Nelson and Andrew M. Durso. Veriﬁed by Kenneth L. Krysko. FLMNH 154856. New county record (Jensen et al. [eds.] 2008. Amphibians and Reptiles of Georgia. Univ. Georgia Press, Athens, 575 pp.). A juvenile M. ﬂagellum was found on a sandhill under a log at ca. 1230 h. This record is ca. 27.6 km NW of a 2008 record from Herpetological Review 40(3), 2009 Candler Co., Georgia (FLMNH 154400), and ca. 32 km S of a 1994 record from Jefferson Co., Georgia (SSM 13101). Submitted by KERRY T. NELSON, University of Georgia, Warnell School of Forestry and Natural Resources, Athens, Georgia 30602, USA (e-mail: kerrytnelson@gmail.com); and ANDREW M. DURSO, University of Georgia, Odum School of Ecology, Athens, Georgia 30602, USA (e-mail: amdurso@gmail.com). FARANCIA ERYTROGRAMMA ERYTROGRAMMA (Common Rainbow Snake). USA: GEORGIA: CHARLTON CO.: west bank of St. Marys River at Earnest A. Bell Bridge, Moniac Rd. (Hwy 94) near St. George (30.52438°N, 82.01898°W; WGS84). 31 March 2006. Nathan H. Nazdrowicz and Nigel Watson. Veriﬁed by John Jensen. Georgia Museum of Natural History photographic voucher (GMNH 50107). Found under rock on steep bank of river. New county record (Jensen et al. [eds.] 2008. Amphibians and Reptiles of Georgia. Univ. Georgia Press, Athens, 575 pp.). S u b m i t t e d b y N AT H A N H . N A Z D R O W I C Z ( email: spinifer@aol.com) and NIGEL WATSON (e-mail: nigelgwatson@yahoo.com), Department of Entomology and Wildlife Ecology, University of Delaware, Newark, Delaware 19716, USA. HYDRODYNASTES GIGAS (False Water Cobra). BRAZIL: PIAUÍ: MUNICIPALITY OF ILHA GRANDE: 2.8219°S, 41.8245°W; datum WGS84, 4 m elev. 23 December 2008. Pedro da Costa Silva. Instituto Butantan, São Paulo (IBSP 77245). Veriﬁed by V. J. Germano. Species widely distributed in South America with recognized records from Suriname, French Guyana, Venezuela, Peru, Bolivia, Paraguay, Argentina, and Brazil (Giraudo and Scrocchi. 2002. Smithson. Herpetol. Inf. Serv. 132:1–53). In Brazil, the species has been recorded mainly in the northern, southeastern, and central western states (Franco et al. 2007. Zootaxa. 1613:57–65), however, there is a unique record for northeastern Brazil in Paraíba State (Pereira Filho and Montingelli. 2006. Herpetol. Rev. 37:497). This is the ﬁrst record for the state of Piauí, extending the distribution previously known as follow: ca. 900 km NW from the municipality of João Pessoa, Paraíba, Brazil (Pereira Filho and Montingelli, op. cit.); ca. 1000 km NE from the municipality of Mateiros, Tocantins, Brazil (Vitt et al. 2005. Occas. Pap. Sam Noble Oklahoma Mus. Nat. Hist. [2]:1–24); and ca. 340 km E from the municipality of Perim-Mirim, Maranhão, Brazil (Franco et al. 2007, op. cit.). Submitted by ROBERTA ROCHA SILVA-LEITE, Universidade Federal do Piauí, Campus Ministro Reis Velloso, Av. São Sebastião nº 2819, Parnaíba, Piauí, Brazil, CEP 64202-020 (e-mail: roberta.ufpi@gmail.com); DANIEL LOEBMANN, Departamento de Zoologia, Instituto de Biociências, Universidade Estadual Paulista, Rio Claro, São Paulo, Brazil, Caixa Postal 199, CEP 13506-970; PEDRO DA COSTA SILVA, Associação de Guias e Condutores de Turismo de Ilha Grande, Centro, Ilha Grande, Piauí, Brazil, CEP 64224-970. LAMPROPELTIS TRIANGULUM (Milksnake). USA: TEXAS: NACOGDOCHES CO.: Stephen F. Austin Experimental Forest (31.49726°N, 094.78005°W; WGS 84). Stephen F. Austin State University Vertebrate Collection 4916. 01 April 2008 by T. B. Cotten. Veriﬁed by T. J. Hibbitts. New county record (Dixon 2000. Amphibians and Reptiles of Texas. Texas A&M Univ. Press, Col- lege Station. 129 pp.). Submitted by TAYLOR B. COTTEN, Department of Biology, Stephen F. Austin State University, P.O. Box 13003, S.F.A. Station, Nacogdoches, Texas 75962-3003, USA. MICRURUS FULVIUS (Harlequin Coralsnake). USA: ALABAMA: COOSA CO.: Cahaba and Columbiana Forever Wild Tract, Coosa Wildlife Management Area (32.8°N, 86.4°W; WGS84). 20 April 2009. Nicholas W. Sharp and Josh D. Landrum. Veriﬁed by Robert Mount. Alabama Herpetological Atlas Project AHAPD-203. Photographed and released. Previously believed to be extirpated in the county. Although range maps typically depict a disjunct population in the Alabama Piedmont north of the Fall Line (Mount 1975. Reptiles and Amphibians of Alabama. Agricultural Experiment Station, Auburn University. 347 pp.), this is the only record of such in 40 years. Auburn University holds only one specimen from Coosa Co., lacking a date and locality; collected sometime before 1975 near Strickland Crossroads, ca. 13 miles N of the current specimen (R. Mount, pers. comm.). Adult captured traversing open ground at 1040 h after heavy rains the previous evening. Habitat was mature Longleaf Pine over well-drained, sandy loam ridges and steep slopes. Submitted by NICHOLAS W. SHARP, Alabama Department of Conservation and Natural Resources, State Lands Division, 64 North Union Street, Montgomery, Alabama 36104, USA; e-mail: nicholas.w.sharp@gmail.com. MICRURUS FULVIUS (Harlequin Coralsnake). USA: GEORGIA: TALBOT CO.: Manchester, ca. 10.8 km E (32.865223ºN, 84.493107ºW; NAD83), 252 m elev. 13 October 2008. Nathan A. Klaus. UF 153883. Veriﬁed by Kenneth Krysko. First county record and only the ﬁfth documented specimen from the Pine Mountain/Piedmont physiographic region (Stevenson and Moulis 2008. In Jensen et al. [eds.], Amphibians and Reptiles of Georgia, pp. 421–423. Univ. Georgia Press, Athens. 575 pp.). Not seen in this area since 1977 (Pike Co. specimen, AUM 32733). Found on surface along a ridgetop in an area of old growth montane Longleaf Pine (Pinus palustris) and Chestnut Oak (Quercus montana) forest with rocky/gravelly soil. Submitted by NATHAN A. KLAUS (e-mail: nathan. klaus@gadnr.org), and JOHN B. JENSEN, Georgia Department of Natural Resources, Nongame Conservation Section, 116 Rum Creek Drive, Forsyth, Georgia 31029, USA. MICRURUS FULVIUS (Harlequin Coralsnake). USA: GEORGIA: T ELFAIR CO.: Horse River Wildlife Management Area (31.83007°N, 82.85688°W; WGS84). 29 March 2008. Nigel Watson and Nathan H. Nazdrowicz. Veriﬁed by John Jensen. Georgia Museum of Natural History photographic voucher (GMNH 50106). One adult observed crossing a dirt road at night. New county record (Jensen et al. [eds.] 2008. Amphibians and Reptiles of Georgia. Univ. Georgia Press, Athens, 575 pp.). S u b m i t t e d b y N AT H A N H . N A Z D R O W I C Z ( email: spinifer@aol.com) and NIGEL WATSON (e-mail: nigelgwatson@yahoo.com), Department of Entomology and Wildlife Ecology, University of Delaware, Newark, Delaware 19716, USA. Herpetological Review 40(3), 2009 365 MICRURUS LEMNISCATUS LEMNISCATUS (Guiana’s Ribbon Coralsnake). BRAZIL: CEARÁ: MUNICIPALITY OF UBAJARA: Ubajara National Park (03.830417°S, 40.900278°W; datum WGS84), 445 m elev. 13 November 2008. D. Loebmann. Veriﬁed by F. L. Franco. Coleção Instituto Butantan, São Paulo, Brazil (IBSP 77079). Species recorded for the Amazon Rainforest Biome with distribution in Guyana, Suriname, French Guyana, Colombia, Bolivia, Ecuador, Peru, Venezuela, and Brazilian states of Acre, Amapá, Amazonas, Pará, Maranhão, Rondônia, and Roraima (Cunha and Nascimento 1993. Bol. Mus. Para. Emílio Goeldi, sér. Zool. 9[1]:1–191; Roze 1996. Coral Snakes of the Americas: Biology, Identiﬁcation, and Venoms. Krieger Publishing Company, Malabar, Florida. 328 pp.; Campbell and Lamar 2004. The Venomous Reptiles of the Western Hemisphere. Cornell University Press, Ithaca, New York. 1032 pp.; Feitosa 2006. Unpubl. thesis. Universidade Federal do Pará, Belém, Pará. 170 pp.). First record for Ceará state and the Caatinga biome, extending distribution ca. 350 km E from state of Maranhão, in surroundings of municipality of Coroatá, Brazil (Campbell and Lamar 2004, op. cit.). Submitted by DANIEL LOEBMANN, Departamento de Zoologia, Instituto de Biociências, Universidade Estadual Paulista, Rio Claro, São Paulo, Brazil, Caixa Postal 199, CEP 13506-970; e-mail: contato@danielloebmann.com. OXYRHOPUS MELANOGENYS ORIENTALIS (Black-headed Calico Snake). BRAZIL: CEARÁ: MUNICIPALITY OF UBAJARA: 3.8450°S, 40.9344°W (datum WGS84), 857 m elev. 10 February 2008. D. Loebmann. Coleção Instituto Butantan, São Paulo, Brazil (IBSP 77061); MUNICIPALITY OF GUARAMIRANGA: 04.2723°S, 38.9492°W, 844 m elev. 10 December 2006. T. Pinto, C. Albano, and I. J. Roberto (IBSP 76979); MUNICIPALITY OF PACOTI: 04.2208°S, 38.9259°W, 758 m elev. 15 April 2006. C. Albano and I. J. Roberto. (IBSP 77980). All veriﬁed by F. L. Franco. Subspecies distributed in the Brazilian states of Pará and Maranhão (Cunha and Nascimento 1993. Bol. Mus. Para. Emílio Goeldi, sér. Zool. 9[1]:1–191). First state records represented by two isolated populations in the relict rain forests of the plateau of Ibiapaba and Baturité Hills. Also the ﬁrst records in the Caatinga biome, extending distribution ca. 480 km E (Plateau of Ibiapaba) and ca. 710 km E (Baturité Hills) from municipality of Santa Inês, state of Maranhão, Brazil (Cunha and Nascimento 1983. Bol. Mus. Para. Emílio Goeldi, sér. Zool. [122]:1–42). Submitted by DANIEL LOEBMANN, Departamento de Zoologia, Instituto de Biociências, Universidade Estadual Paulista, Rio Claro, São Paulo, Brazil, Caixa Postal 199, CEP 13506-970 (e-mail: contato@danielloebmann.com); IGOR JOVENTINO ROBERTO, Aquasis - Associação de Pesquisa e Preservação de Ecossistemas Aquáticos, Programa Biodiversidade. Praia de Iparana s/n, SESC Iparana, Caucaia, Ceará, Brazil, CEP 61627-010. PANTHEROPHIS GUTTATUS (Red Cornsnake). USA: GEORGIA: EMANUEL CO.: DOR adult male on Hall’s Bridge Rd. (CR 160), ~10.75 km SW of Swainsboro (straight distance from town center) (32.54500°N, 82.43683°W; WGS84). 02 April 2002. Nathan H. Nazdrowicz and Shawn T. Dash. Veriﬁed by Elizabeth McGhee. Georgia Museum of Natural History (GMNH 50109). New county record (Jensen et al. [eds.] 2008. Amphibians and Reptiles of Georgia. Univ. Georgia Press, Athens, 575 pp.). 366 Submitted by NATHAN H. NAZDROWICZ, Department of Entomology and Wildlife Ecology, University of Delaware, Newark, Delaware 19716, USA; e-mail: spinifer@aol.com. RAMPHOTYPHLOPS BRAMINUS (Brahminy Blindsnake). USA: TEXAS: HIDALGO CO.: McAllen (26.261983°N, 98.210933W; WGS 84). 15 March 2009. John Merino. Veriﬁed by Frank W. Judd. University of Texas – Pan American Vertebrate Museum (UTPA) 03092. Female (11.3 cm total length and 0.5879 g) found under ﬂat landscaping stones alongside ant colonies in a residential area. This is the ﬁrst county record and third known locality in the state. Previously reported in adjacent Cameron Co. (Dixon 2000. Amphibians and Reptiles of Texas. 2nd ed. Univ. of Texas A&M Press, College Station. viii + 421 pp.; Godwin et al. 2007. Herpetol. Rev. 38:356; Wallach. 2008. Bull. Chicago Herpetol. Soc. 43[5]:80–82.). Our record lies ca. 88 km WNW of the nearest (= Cameron Co.) published record. On the same day, another was captured at a residence in Edinburg, also in Hidalgo Co. (26.287050°N, 98.175717°W; WGS 84). This individual was found underneath an established plant in a ﬂowerbed. On 19 March 2009 a third individual was captured by G. Martinez and J. M. Cantu in Brownsville, Cameron Co. in a different locality (University of Texas – Brownsville campus) from the previous Cameron Co. record. From 20–30 March 2009 an additional 13 specimens were captured at the two Hidalgo Co. locations. After its ﬁrst mention in the area over 20 years ago, it appears that R. braminus is now well established in the Lower Rio Grande Valley. Submitted by JOHN MERINO, Department of Biology, University of Texas – Pan American, Edinburg, Texas 78539, USA; LEE BAINES, Department of Biology, South Texas College, 3201 W. Pecan Blvd., McAllen, Texas 78501, USA; and FREDERIC ZAIDAN III, Department of Biology, University of Texas – Pan American, Edinburg, Texas 78539, USA (e-mail: fzaidan@utpa. edu). SCOLECOPHIS ATROCINCTUS (Black-Banded Snake). HONDURAS: CORTÉS: Sierra de Omoa, Parque Nacional El Cusuco, Buenos Aires Village (15.499722ºN, 88.176667ºW; WGS84), 1120 m elev. 27 July 2007. Jonathan E. Kolby and Douglas Fraser. Veriﬁed by Larry D. Wilson. (USNM Herp Images 2685–2688). First record for Cortés, extending the range ca. 111 km N of Lempira, Gracias (FMNH 28559). It is also a 420 m upward extension in elevation from the highest previous record of 700 m reported by Kohler (2003. Reptiles of Central America. Herpeton Verlag Elke Kohler. 367 pp.). The snake was found DOR in a deforested region converted to coffee cultivation. Submitted by JONATHAN E. KOLBY, The Conservation Agency, 6 Swinburne Street, Jamestown, Rhode Island 02835, USA (e-mail: J_Kolby@hotmail.com); and DOUGLAS FRASER, Operation Wallacea, Wallace House, Old Bolingbroke, Lincolnshire PE23 4EX, United Kingdom. SIBON NEBULATUS (Cloudy Snail-eating Snake). MÉXICO: JALISCO: MUNICIPALITY OF SAN SEBASTIÁN DEL OESTE: on the road from Mascota to Ixtapa (20.78017°N, 104.92650°W; WGS84), 917 m elev. 14 June 2007. U. O. García-Vázquez and R. C. Jadin. Both veriﬁed by E. N. Smith. Museo de Zoología, Facultad de Ciencias UNAM (MZFC 22051). First veriﬁed municipality record that Herpetological Review 40(3), 2009 bridges a distribution gap between Tepic, Nayarit (80 km N) and Chamela, Jalisco (140 km S) (Zweifel 1959. Amer. Mus. Novit. 1953:1–13; Ramirez-Bautista 1994. Manual y Claves Ilustradas de los Anﬁbios y Reptiles de la Región de Chamela, Jalisco. UNAM, Cuadernos del Inst. de Biol. 23:1–127). Another individual was found nearby on the same road and date (20.81104°N, 104.95510°W, 710 m elev.; UTA R-55780). The vegetation at the capture site was a patch of pristine tropical deciduous forest, although much of what remained was in the process of being burned for agriculture and other developmental purposes. Submitted by ROBERT C. JADIN, Amphibian and Reptile Diversity Research Center and Department of Biology, University of Texas at Arlington, Arlington, Texas 76019, USA (e-mail: snakeman1982@hotmail.com); and URI OMAR GARCÍAVÁZQUEZ, Museo de Zoología, Facultad de Ciencias, Universidad Nacional Autónoma de México, A.P. 70-399, México D.F. (e-mail: urigarcia@gmail.com). SPILOTES PULLATUS (Tropical Rat Snake). MÉXICO: HIDALGO: MUNICIPALITY OF HUEHUETLA: El Paraíso (98.13972°N, 20.43125°W, WGS 84), 498 m elev. 28 August 2008. Adán Martínez Sosa. Veriﬁed by Norma Manríquez-Morán. Colección Herpetológica, Centro de Investigaciones Biológicas, Universidad Autónoma del Estado de Hidalgo (CIB-UAEH 1622). First municipality record, extending the range in Hidalgo ca. 92 km S from 2.3 km E of San Felipe Orizatlán, Orizatlán (Mendoza-Quijano and Hernández 2001. Herpetol. Rev. 32:279). The snake was found in a coffee ﬁeld surrounded by tropical deciduous forest. Fieldwork was funded by CONACyT- 43761. Submitted by IRENE GOYENECHEA (e-mail: ireneg@uaeh. edu.mx), JESÚS M. CASTILLO, FROYLÁN RAMÍREZRESÉNDIZ, and ALEJANDRO RAMÍREZ-PÉREZ, Centro de Investigaciones Biológicas (CIB), Universidad Autónoma del Estado de Hidalgo, A.P. 1-69 Plaza Juárez, Pachuca, Hidalgo, México. STORERIA DEKAYI TEXANA (Texas Brownsnake). USA: TEXAS: STEPHENS CO.: S of State Highway 180 and E of Ranch Road 717 near Caddo (32.64668ºN, 98.61677ºW). 05 April 2008. Collected by J. W. Streicher. Veriﬁed by Carl J. Franklin. Deposited in the herpetological collection of the University of Texas at Arlington (UTA R-56675, ﬁeld ID: JWS 053). Adult (SVL = 225 mm, TL = 33 mm) found under a roadside log represents ﬁrst county record. This account is a local range extension W (ca. 4 km) and ﬁlls in a longitudinal gap (ca. 50 km) within the known distribution of S. d. texana (Dixon 2000. Amphibians and Reptiles of Texas, 2nd ed. Texas A&M University Press, College Station. 421 pp.; Werler and Dixon 2000. Texas Snakes. University of Texas Press, Austin. 437 pp.). Collected under Texas Parks and Wildlife scientiﬁc permit number SPR-0707-1387. Submitted by JEFFREY W. STREICHER, Amphibian and Reptile Diversity Research Center, Department of Biology, The University of Texas at Arlington, Arlington, Texas 76019, USA; e-mail: streicher@uta.edu. STORERIA DEKAYI WRIGHTORUM (Midland Brownsnake). USA: ARKANSAS: LINCOLN CO.: 1.6 km E Palmyra (33.552712°N, 91.563533°W; NAD83). 05 May 2001. M. B. Gannet. Veriﬁed by S. E. Trauth. Arkansas State University Herpetological Museum (ASUMZ 31321). New county record ﬁlling a distributional hiatus among Arkansas and Cleveland counties (Trauth et al. 2004. Amphibians and Reptiles of Arkansas. Univ. Arkansas Press, Fayetteville. 421 pp.). Since the publication of Trauth et al. (op. cit.), no fewer than seven new county records in various parts of the state have been documented for this snake. Submitted by CHRIS T. MCALLISTER, RapidWrite, 102 Brown Street, Hot Springs National Park, Arkansas 71913, USA (e-mail: drctmcallister@aol.com); and HENRY W. ROBISON, Department of Biology, Southern Arkansas University, Magnolia, Arkansas 71754, USA (e-mail: hwrobison@suddenlink.net). VIRGINIA VALERIAE ELEGANS (Western Smooth Earthsnake). USA: ARKANSAS: HOWARD CO.: 9.7 km NW Center Point off US 278 (34.025766°N, 93.571243°W; NAD83). 10 June 2004. S. Fowler. Veriﬁed by S. E. Trauth. Arkansas State University Herpetological Museum (ASUMZ 31322). New county record ﬁlling a distributional hiatus in southwestern Arkansas south of previous records in Polk Co. (Trauth et al. 2004. Amphibians and Reptiles of Arkansas. Univ. Arkansas Press, Fayetteville. 421 pp.). Submitted by CHRIS T. MCALLISTER, RapidWrite, 102 Brown Street, Hot Springs National Park, Arkansas 71913, USA (e-mail: drctmcallister@aol.com); and HENRY W. ROBISON, Department of Biology, Southern Arkansas University, Magnolia, Arkansas 71754, USA (e-mail: hwrobison@suddenlink.net). Herpetological Review, 2009, 40(3), 367–371. © 2009 by Society for the Study of Amphibians and Reptiles One Hundred and One New County Records for Alabama Amphibians and Reptiles SEAN P. GRAHAM Department of Biological Sciences, Auburn University 331 Funchess Hall Auburn, Alabama 36849, USA e-mail: grahasp@auburn.edu SHANNON K. HOSS Department of Biology, San Diego State University, 5500 Campanile Drive San Diego, California 92182-4614, USA ROGER D. BIRKHEAD Auburn University Science In Motion, 206 Allison Lab, Auburn University Auburn, Alabama 36849, USA CHELSEA K. WARD Department of Biological Sciences, Auburn University of Montgomery P.O. Box 244023, Montgomery, Alabama 36124, USA DAVID A. STEEN Department of Biological Sciences, Auburn University 331 Funchess Hall Auburn, Alabama 36849, USA KATHERINE M. GRAY Department of Biological Sciences, Auburn University 331 Funchess Hall Auburn, Alabama 36849, USA and ROBERT H. MOUNT Department of Biological Sciences, Auburn University 331 Funchess Hall Auburn, Alabama 36849, USA The distribution of Alabama’s herpetofauna was treated thoroughly in Mount’s (1975) regional ﬁeld guide. Since its publication, few additional distribution contributions for Alabama Herpetological Review 40(3), 2009 367 (mostly in the form of single-taxon notes) have appeared. Although most counties in Alabama have numerous recorded taxa, a few counties have not been sampled extensively and have fewer than 18 species recorded. To address this situation, we began road cruise surveys on rainy nights in counties with few records. In addition, we have included new records discovered during class ﬁeld trips and other related activities. Here we report 101 new county records for Alabama. These new records include at least one new county record for 48 species. All new records were veriﬁed by Craig Guyer. Caudata – Salamanders Ambystoma opacum (Marbled Salamander). DALE CO.: Found AOR 0.3 km W of Judy Creek on Dale County Rd. 36 (31.462594°N, 85.574776°W; WGS 84). 29 December 2007. S. Graham and S. Hoss. AHAP-D 77. LOWNDES CO.: Found AOR on Lowndes County Rd. 40 (32.324877°N, 86.676382°W; WGS 84). 18 October 2007. S. Graham. AHAP-D 64. Ambystoma talpoideum (Mole Salamander). LOWNDES CO.: Lowndes WMA. Captured by dipnet in shallow marsh (32.357364°N, 86.731320°W; WGS 84). 20 September 2008. S. Graham and K. Gray. AHAP-D 154–155. Ambystoma tigrinum (Eastern Tiger Salamander). BULLOCK CO.: Collected AOR in Union Springs, crossing US Hwy 29 in historic district at intersection with Hunter St. (32.15068°N, 85.713527°W; WGS 84). 10 December 2008. S. Graham, R. Birkhead, and K. Gray. AHAP-D 176. Cryptobranchus alleganiensis (Hellbender). LIMESTONE CO.: Caught on ﬁshing line in Sugar Creek at Buck Island Road (State Rt. 99) bridge (34.944524°N, 87.155586°W; WGS 84). 09 March 1990. H. Bloodworth. Photograph of H. Bloodworth and specimen appeared in Athens News Courier 25 March 1990; locality and additional information veriﬁed by M. Bailey and R. Mount. Newspaper clipping with photo and other information entered into AUM as AHAP-P182. This represents a new county record and one of the most recent veriﬁed records for a hellbender in Alabama. Eurycea cirrigera. (Southern Two-lined Salamander). HALE CO.: Payne Lake Recreational Area, Talladega National Forest. In spring W of Lake. (32.886059°N, 87.445250°W; WGS 84). 20 September 2008. S. Graham and K. Gray. AHAP-D 167. Eurycea guttolineata (Three-lined Salamander). CRENSHAW CO.: In Threemile Cr. ﬂoodplain ~ 25 m E of US Hwy 29 (31.520517°N, 86.335433°W; WGS 84). 18 March 2008. S. Graham. AHAP-D 105. HALE CO.: Payne Lake Recreational Area, Talladega National Forest. In swampy area S. of dam. (32.878206°N, 87.443716°W; WGS 84). 20 September 2008. S. Graham and K. Gray. AHAP-D 157. Hemidactylium scutatum (Four-toed Salamander). LAMAR CO.: Found AOR Lamar County Rd. 12 .64 km W of Mud Creek Rd. intersection (33.577378°N, 88.144967°W; WGS 84). 29 November 2008. S. Graham and K. Gray. AHAP-D 163. MARENGO CO.: 5.8 km (air km) NE of Dixon’s Mill. 1 January 1977. J. Autery. AUM 29664. Notophthalmus viridescens (Eastern Newt). BULLOCK CO.: Found DOR on Bullock County Road 102 (32.22618°N, 85.4796052°W; 368 WGS 84). 29 December 2007. S. Graham and S. Hoss. AUM 37612. CRENSHAW CO.: Found AOR Crenshaw County Rd. 59 NE of Luverne (31.751883°N, 86.232050°W; WGS 84). 10 December 2008. S. Graham, R. Birkhead, and K. Gray. AHAP-D 168. LAMAR CO.: Found AOR on State Rt. 17 N of Millport 10m N of bridge over Luxapalilla Cr. (33.575406°N, 88.083639°W; WGS 84). 29 November 2008. S. Graham and K. Gray. AHAP-D 162. LOWNDES CO.: Collected AOR on Brownshill Rd. ~ 200m W of Lowndes County Rd. 40 (32.314128°N, 86.584818°W; WGS 84). 21 September 2007. S. Graham, S. Hoss, D. Steen, V. Johnson. AHAP-D 97. WILCOX CO.: Near dam of large beaver marsh along Howard Steen Rd. 3.86 km SW of Wilcox County Rd. 4 (31.884343°N, 86.913055°W; WGS 84). 5 May 2008. S. Graham and R. Birkhead. AHAP-D 99. Plethodon grobmani (Southeastern Slimy Salamander). CRENCO.: In Floodplain of Threemile Cr. ~ 25 m E of US Hwy 29 (31.520517°N, 86.335433°W; WGS 84). 18 March 2008. S. Graham. AHAP-D 106. LOWNDES CO.: Collected AOR on Lowndes County Rd. 40 (32.325749°N, 86.731768°W; WGS 84). 18 November 2007. S. Graham and K. Gray. AHAP-D 68. These specimens were not assigned to this taxon using molecular methods; they were assumed to be P. grobmani based on existing range maps (Lannoo, 2005) and in any event represent new county records for a species of the P. glutinosus complex. SHAW Pseudotriton ruber (Southern Red Salamander). CRENSHAW CO.: In seepage S of Threemile Cr. ﬂoodplain ~ 25 m E of US Hwy 29 (31.520517°N, 86.335433°W; WGS 84). 18 March 2008. S. Graham. AHAP-D 104. LOWNDES CO.: Collected in spring just S of Lowndes County Rd. 40 (32.325966°N, 86.681629°W; WGS 84). 10 February 2008. S. Graham and S. Hoss. AUM 37643. Anura – Frogs Acris crepitans (Northern Cricket Frog). BULLOCK CO.: Collected AOR Bullock County Road 14 at the Conecuh River (32.010724°N, 85.740373°W; WGS 84). 29 December 2007. S. Graham and S. Hoss. AUM 37613. LAMAR CO.: Collected DOR on Lamar County Rd. 12 (33.579901°N, 88.0989721°W; WGS 84). 12 February 2008. S. Graham and S. Hoss. AUM 37644. Acris gryllus (Southern Cricket Frog). CRENSHAW CO.: Collected DOR on Crenshaw County Rd. 1200 m NW Crenshaw County Rd. 7 (31.700153°N, 86.375061°W; WGS 84). At least one male also heard calling at this locality on this night (large beaver marsh N of road). 10 December 2008. S.Graham, R. Birkhead, and K. Gray. AHAP-D 180–181. Anaxyrus fowleri (Fowler’s Toad). CRENSHAW CO.: Found at US Hwy 331 at the Conecuh River (31.574056°N, 86.253057°W; WGS 84). 02 October 2008. S. Graham, D. Steen, and K. Gray. AHAP-D 149. Anaxyrus terrestris (Southern Toad). CRENSHAW CO.: Collected DOR on State Rt. 106 (31.604150°N, 86.414700°W; WGS 84). 10 December 2008. S. Graham, R. Birkhead, and K. Gray. AHAPD 173. MONROE CO.: Collected AOR on Monroe County Rd. 47 (31.815335°N, 86.959945°W; WGS 84). 22 September 2007. S. Graham and S. Hoss. AHAP-D 80 Gastrophryne carolinensis (Eastern Narrow-mouthed Toad). Herpetological Review 40(3), 2009 CONECUH CO.: In seepage 5m N of State Rt. 106 (31.707310°N, 86.929000°W; WGS 84). 5 May 2008. R. Birkhead and S. Graham. AHAP-D 190. CRENSHAW CO.: Found at US Hwy 331 at the Conecuh River (31.574056°N, 86.253057°W; WGS 84). 2 October 2008. S. Graham, D. Steen, and K. Gray. AHAP-D 150. LOWNDES CO.: Collected AOR Lowndes County Rd. 40 (32.327441°N, 86.743206 °W; WGS 84). 03 September 2007. S. Graham AHAP-D 75. Hyla avivoca (Bird-voiced Treefrog). LOWNDES CO.: Collected DOR on Lowndes County Rd. 40, 0.64 km E of Cypress Cr. bridge (32.331069°N, 86.669781°W; WGS 84). 18 October 2007. S. Graham. AUM 37570. Hyla chrysoscelis (Cope’s Gray Treefrog). BULLOCK CO.: Collected AOR on Bullock County Rd. 23, 0.8 km NW of US Hwy 82 (32.17664122°N, 85.7088937°W; WGS 84). 10 July 2008. S. Graham. AHAP-D 130–131. CRENSHAW CO.: Collected DOR State Rt. 59 N of Patsburg (31.790033°N, 86.229567°W; WGS 84). 10 December 2008. R. Birkhead, S. Graham, and K. Gray. AHAP-D 179. CULLMAN CO.: Found along millpond at ClarksonLegg covered bridge (34.207298°N, 86.990100°W; WGS 84). 11 July 2008. S. Graham, M. Connell, D. McMoran, K. Gray. AHAP-D 141. LAMAR CO.: Collected DOR on Lamar County Rd. 12 (33.572793°N, 88.212234°W; WGS 84). 29 November 2008. S. Graham and K. Gray. AHAP-D 165. Hyla cinerea (Green Treefrog). BUTLER CO.: Collected DOR on State Rt. 106, 0.8 km W of Butler County Rd. 7 (31.665940°N, 86.856180°W; WGS 84). 18 October 2007. S. Hoss and A. Liner. AUM 37581. CRENSHAW CO.: Collected AOR on State Rt. 106 (31.589400°N, 86.396317°W; WGS 84). 10 December 2008. S. Graham, R. Birkhead, and K. Gray. AHAP-D 172. CULLMAN CO.: Found along millpond at Clarkson-Legg covered bridge (34.207298°N, 86.990100°W; WGS 84). 11 July 2008. S. Graham, M. Connell, D. McMoran, K. Gray. AHAP-D 140. LOWNDES CO.: Collected DOR Lowndes County Rd. 29/40 intersection (32.33260°N, 86.646066°W; WGS). 3 September 2007. S. Graham. AUM 37476. LAMAR CO.: Collected AOR Lamar County Rd. 12 (33.578240°N, 88.111256°W; WGS 84). 10 January 2008. S. Graham. AHAP-D 101. Hyla femoralis (Pine Woods Treefrog). MARENGO CO.: Collected on Marengo County Rd. 30 200m W of railroad at Magnolia Terminal. 1 August 1976. J. Autery. AUM 29665. Hyla gratiosa (Barking Treefrog). CRENSHAW CO.: Collected AOR on US Hwy 29 SW Brantly (31.577350°N, 86.281700°W; WGS 84). 10 December 2008. S. Graham, R. Birkhead, and K. Gray. AHAP-D 173. LOWNDES CO.: Collected AOR on Lowndes County Rd. 40 (32.326992°N, 86.702447°W; WGS 84). 18 October 2007. S. Graham AHAP-D 66. Hyla squirella (Squirrel Treefrog). LOWNDES CO.: Collected AOR on Lowndes County Rd. 40 (32.331811°N, 86.657170°W; WGS 84). 21 September 2007. S. Graham, S. Hoss, D. Steen, V. Johnson. AHAP-D 79. MONROE CO.: Collected DOR on State Rt. 21 6.12 km N of State Rt. 47 bypass (31.570260°N, 87.285750°W; WGS 84). 18 October 2007. S. Hoss and A. Liner. AUM 37580. Lithobates catesbeianus (American Bullfrog). LOWNDES CO.: Collected AOR on Lowndes County Rd. 40 (32.331925°N, 86.649499°W; WGS 84). 21 September 2007. S. Graham, S. Hoss, D. Steen, and V. Johnson. AHAP-D 78. Lithobates clamitans (Green Frog). COLBERT CO.: Along Cane Creek in Cane Creek Canyon Nature Preserve (34.626323°N, 87.805596°W; WGS 84). 13 September 2008. D. Steen, S. Graham, C. Guyer, and K. Bakkegard. AHAP-D 144. CULLMAN CO.: Found along millpond at Clarkson-Legg covered bridge (34.207298°N, 86.990100°W; WGS 84). 11 July 2008. S. Graham, M. Connell, D. McMoran, K. Gray. AHAP-D 188. DALE CO.: Collected DOR on Dale County Road 19 100m from State Rt. 123 (31.5711208°N, 85.6722654°W; WGS 84). 29 December 2007. S. Graham and S. Hoss. AHAP-D 185–186. HALE CO.: Found AOR on Hale County Rd. 50 ~ 2km E of intersection with Hale County Rd. 44 (32. 966308°N 87.546789°W; WGS 84). 20 September 2008. S. Graham and K. Gray. AHAP-D 161. LAMAR CO.: Collected DOR on Lamar County Rd. 12 adjacent to Fernbank Baptist Ch. (33.578645°N, 88.140127°W; WGS 84). 12 February 2008. S. Graham and S. Hoss. AUM 37645. LOWNDES CO.: Collected AOR on Lowndes County Rd. 40 (32.331842°N, 86.648872°W; WGS 84). 3 September 2007. S. Graham. AHAP-D 76. Lithobates palustris (Pickerel Frog). CULLMAN CO.: Found under Clarkson-Legg covered bridge (34.207298°N, 86.990100°W; WGS 84). 11 July 2008. S. Graham, M. Connell, D. McMoran, K. Gray. AHAP-D 143. Lithobates sphenocephalus (Southern Leopard Frog). BULLOCK CO.: Found DOR Bullock County Road 14 at the Conecuh River (32.010724°N, 85.740373°W; WGS 84). 29 December 2007. S. Graham and S. Hoss. AUM 37610. CRENSHAW CO.: Collected along west bank of Conecuh River at Crenshaw County Rd. 77 (31.487593°N, 86.361388°W; WGS 84). 4 October 2008. S. Graham, D. Steen, and K. Gray. AHAP-D 152. LAMAR CO. Collected AOR St. Route 96 SW of Belk, AL (33.616678°N, 87.954633°W; WGS 84). 10 January 2008. S. Graham. AHAP-D 102. LOWNDES CO.: Found AOR on Lowndes County Rd. 40 (32.327348°N, 86.710297°W; WGS 84). 18 October 2007. S. Graham. AHAP-D 65. PIKE CO.: Collected DOR on US Hwy 231, 0.16km N of intersection with State Rt. 125 (31.681467°N, 85.824700°W; WGS 84). 10 December 2008 S. Graham, R. Birkhead, and K. Gray. AHAP-D 182.s Pseudacris crucifer (Spring Peeper). BULLOCK CO.: Found AOR Bullock County Road 35 (32.029348°N, 85.588407°W; WGS 84). 29 December 2007. S. Graham and S. Hoss. AHAP-D 183. COFFEE CO.: Collected AOR on State Rt. 189 0.5 km E of Crenshaw County Line (31.524867°N, 86.189500°W; WGS 84). 10 December 2008. S. Graham, R. Birkhead, and K. Gray. AHAP-D 174. CONECUH CO.: Collected DOR on State Rt. 106 0.1 km E of Conecuh County Rd. 37 and State Rt. 106 intersection (31.706196°N, 86.921304°W; WGS 84 ). 17 October 2008. S. Graham and K. Gray. AUM 37828. DALE CO.: Found AOR Dale County Road 36 E of Ozark, AL (31.461183°N, 85.579572°W; WGS 84). 29 December 2007. S. Graham and S. Hoss. AHAP-D 184. FAYETTE CO.: Collected DOR Fayette County Rd. 12 at the Sipsey River (33.543883°N, 87.784320°W; WGS 84). 12 February 2008. S. Graham and S. Hoss. AUM 37646. LAMAR CO. Collected AOR St. Route 96 SW of Belk, AL (33.616678°N, 87.954633°W; WGS 84). 10 January 2008. S. Graham. AHAP-D 100. LOWNDES CO.: Collected AOR on Lowndes County Rd. 40 (32.326603°N, 86.680310 °W; WGS 84). 18 November 2007. S. Graham and K. Gray. AHAP-D 67. PICKENS CO.: Collected DOR State Rt. 17 N of Reform (33.511498°N, Herpetological Review 40(3), 2009 369 87.980888°W; WGS 84). 29 November 2008. S. Graham and K. Gray. AHAP-D 166. Pseudacris feriarum (Upland Chorus Frog). CRENSHAW CO.: Collected in roadside ditch along Bradleyton Rd. (31.880455°N, 86.250700°W; WGS 84). Several males heard chorusing. 10 December 2008. R. Birkhead, S. Graham, and K. Gray. AHAP-D 171. DALE CO.: Collected in roadside ditch off Dale County Rd. 59 0.6 km NW of Choctawhatchee River (31.478963°N, 85.537040°W; WGS 84). Several males heard chorusing. 29 December 2007. S. Graham and S. Hoss. AHAP-D 187. LAMAR CO.: Collected AOR on Lamar County Rd. 12 .80 km E of Mississippi State Line (33.574091°N, 88.262090°W; WGS 84). 29 November 2008. S. Graham and K. Gray. AHAP-D 164. Scaphiopus holbrookii (Eastern Spadefoot). BULLOCK CO.: Collected AOR on Bullock County Rd. 40 3.22 km E of Union Springs (32.167561°N, 85.612993°W; WGS 84). 10 July 2008. S. Graham. AHAP-D 132. DALE CO.: Found DOR Dale County Road 36 (31.462133°N, 85.576611°W; WGS 84). 29 December 2007. S. Graham and S. Hoss. AUM 37611. Testudines – Turtles Chelydra serpentina (Snapping Turtle). COVINGTON CO.: Found AOR on US Hwy 29 (31.455568°N, 86.433406°W; WGS 84) 3.56 km SW of Covington County Line. 18 March 2008. S. Graham. AHAP-D 107. LAUDERDALE CO.: Found in Little Cypress Cr. at Lauderdale County Rd. 8 (34.949325°N, 87.694589°W; WGS 84). 12 July 2008. S. Graham, M. Connell, D. McMoran, and K. Gray. AHAP-D 143. MACON CO.: Captured in hoop trap in large beaver marsh in Tuskegee National Forest (32.428926°N, 85.647345°W; WGS 84). 19 June 2008. S. Graham and M. Connell. AHAP-D 129. Terrapene carolina (Eastern Box Turtle). CULLMAN CO.: Found under Clarkson-Legg covered bridge (34.207298°N, 86.990100°W; WGS 84). 11 July 2008. S. Graham, M. Connell, D. McMoran, K. Gray. AHAP-D 142. Squamata – Lizards Anolis carolinensis (Green Anole). CRENSHAW CO.: Collected DOR on Crenshaw County Rd. 59 (31.751883°N, 86.232050°W; WGS 84). 10 December 2008. R. Birkhead, S. Graham, and K. Gray. AHAP-D 177–178. Plestiodon laticeps (Broad-headed Skink). CRENSHAW CO.: Found at US Hwy 331 at the Conecuh River (31.574056°N, 86.253057°W; WGS 84). 2 October 2008. S. Graham, D. Steen, and K. Gray. AHAP-D 146-148. Scincella lateralis (Little Brown Skink). HALE CO.: Payne Lake Recreational Area, Talladega National Forest. Near spring W of Lake. (32.886059°N, 87.445250°W; WGS 84). 20 September 2008. S. Graham and K. Gray. AHAP-D 160. Squamata – Snakes Agkistrodon contortrix (Copperhead). CRENSHAW CO.: In Threemile Creek ﬂoodplain ~ 25 m E of US Hwy 29 (31.520517°N, 370 86.335433°W; WGS 84). 04 October, 2008. S. Graham, K. Gray, and D. Steen. AHAP-D 153. CULLMAN CO.: Found DOR on Cullman County Rd. 1043 100 m E of Winston County Line (34.228753°N, 87.109102°W; WGS 84). 12 April 2008. S. Graham. AHAP-D 121. Carphophis amoenus (Eastern Wormsnake). CONECUH CO.: In seepage 5 m N of State Rt. 106 (31.707310°N, 86.929000°W; WGS 84). 05 May 2008. R. Birkhead and S. Graham. Coluber constrictor (North American Racer). LOWNDES CO.: Collected DOR Lowndes County Rd. 40 (32.327227°N, 86.762491°W; WGS 84). 03 September 2007. S. Graham. AUM 37480. WILCOX CO.: Collected in small branch 5 m S of Howard Steen Rd. (31.884617°N 96.922506°W; WGS 84). 5 May 2008. R. Birkhead and S. Graham. AHAP-D 98. Crotalus horridus (Timber Rattlesnake). LOWNDES CO.: Collected DOR Lowndes Wildlife Management Area, 200 m W of Check Station turnoff (32.364884°N, 86.740857°W; WGS 84). 03 September 2007. S. Graham. AUM 37478. Diadophis punctatus (Ring-necked snake). HALE CO.: Payne Lake Recreational Area, Talladega National Forest. In swampy area S of dam (32.878206°N, 87.443716°W; WGS 84). 20 September 2008. S. Graham and K. Gray. AHAP-D 158. LOWNDES CO.: Collected AOR Lowndes County Rd. 40 (32.326305°N, 86.680024°W; WGS 84). 21 November 2007. S. Graham. AHAP-D 71. Lampropeltis getula niger (Eastern Black Kingsnake). LIMESTONE CO.: Found along Limestone Cr. 5 m S of Interstate Hwy 565 (34.648517°N, 86.822358°W; WGS 84). 12 July 2008. S. Graham, M. Connell, D. McMoran, and K. Gray. AHAP-D 189. Masticophis ﬂagellum (Eastern Coachwhip). ELMORE CO.: DOR State Rt. 229, 4.83 km S of Tallassee (32. 493624°N, 85.892708°W; WGS 84). 05 May 2009. R. Birkhead. AHAP-D 204. Nerodia erythrogaster (Plain-bellied Watersnake). LOWNDES CO.: Collected DOR Lowndes County Rd. 40 (32.326305°N, 86.680024°W; WGS 84). 21 November 2007. S. Graham. AHAPD 70. Opheodrys aestivus (Rough Greensnake). LOWNDES CO.: Collected DOR 8.85 km W of Montgomery County line at Tallawassee Creek bridge over Lowndes County Rd. 40 (32.329826°N, 86.557229°W; WGS 84). 18 October 2007. S. Graham. AUM 37572. Pantherophis guttatus (Red Cornsnake). COVINGTON CO.: Found DOR on US Hwy 331 at mile marker 10. 2 October 2008. S. Graham, D. Steen, and K. Gray. AHAP-D 151. ELMORE CO.: Found DOR Elmore County Road 4 (32.451117°N, 86.075117°W; WGS 84). 17 September 2008. S. Graham and R. Birkhead. AHAP-D 191. Pantherophis spiloides (Gray Ratsnake). CHILTON CO.: Found DOR on US Hwy 82 3.22 km N of Autauga County Line (32.683793°N, 86.791322°W; WGS 84). 30 November 2008. S. Graham. AHAPD 164. LOWNDES CO.: Collected DOR in Lowndes WMA 0.8 km E of Check Station (32.350679°N, 86.725180°W; WGS 84). 21 September 2007. S. Graham, V. Johnson, D. Steen and S. Hoss. AUM 37491. Herpetological Review 40(3), 2009 Storeria dekayi (DeKay’s Brownsnake). BULLOCK CO.: Found AOR on US Hwy 82 N of Union Springs (32.175567°N, 85.709517°W; WGS 84). 10 December 2008. S. Graham, R. Birkhead, and K. Gray. AHAP-D 175. CONECUH CO.: Collected DOR on State Rt. 106 1.45 km E of County Rd. 29 intersection (31.707533°N, 86.933592°W; WGS 84). 17 February 2008. S. Hoss. AUM 37650. HALE CO.: Payne Lake Recreational Area, Talladega National Forest. In spring W of lake. (32.878206°N, 87.443716°W; WGS 84). 20 September 2008. S. Graham and K. Gray. AHAPD 159. LOWNDES CO.: Collected AOR Lowndes County Rd. 40 (32.327649°N, 86.743344°W; WGS 84). 03 September 2007. S. Graham. AHAP-D 74. The Conecuh and Lowndes county records ﬁll a substantial distribution gap from the nearest documented populations to the N (Bibb County) and NE (Montgomery County) to those to the SW (Washington County; Mount 1975). Storeria occipitomaculata (Redbelly Snake). HENRY CO.: Collected in swamp S of Hutto Pond, 4.61 air km W of Laurenceville (31.660815°N, 85.317152°W; WGS 84). 13 May 2009. S. Graham and D. Laurencio. AUM 34915. Thamnophis sauritus (Eastern Ribbonsnake). CRENSHAW CO.: Collected DOR on Crenshaw County Rd. 1, 200m NW Crenshaw County Rd. 7 (31.700153°N, 86.375061°W; WGS 84). 10 December 2008. S.Graham, R. Birkhead, and K. Gray. AHAP-D 169. LOWNDES CO.: Collected DOR on Lowndes County Rd. 40 (32.331976°N, 86.594544 °W; WGS 84). 21 September 2007. S. Graham, S. Hoss, D. Steen, V. Johnson. AUM 37490. ST. CLAIR CO.: Found under rock next to small creek along US Hwy 231 ~ 10 km S of Ashville (33.754353°N, 86.275546°W; WGS 84). 2 March 2008. S. Graham and S. Hoss. AHAP-D 103. Thamnophis sirtalis (Common Gartersnake). LOWNDES CO.: Collected DOR on Brownshill Rd. 100 m W of Lowndes County Rd. 40 (32.318552 °N, 86.577542 °W; WGS 84). 3 September 2007. S. Graham AUM 37477. Acknowledgments.—We thank Craig Guyer for verifying these records. We thank V. Johnson, D. J. McMoran, M. Connell, K. Bakkegard, and A. Liner for their assistance during these ﬁeld trips, and J. Autery for his contributions. These collection trips were partially supported by NIH grant R01-A149724 to T. Unnasch under ADCNR Permit 4268. LITERATURE CITED MOUNT, R. H. 1975. The Reptiles and Amphibians of Alabama. Auburn University Agricultural Experiment Station, Auburn, Alabama. vii +347 pp. LANNOO, M. 2005. Amphibian Declines: The Conservation Status of United States Species. University of California Press. Berkeley, California. xxi +1093 pp. BOOK REVIEWS Herpetological Review, 2009, 40(3), 371–374. © 2009 by Society for the Study of Amphibians and Reptiles Lizard Ecology: The Evolutionary Consequences of Foraging Mode, edited by Stephen M. Reilly, Lance B. McBrayer, and Donald B. Miles. 2007. Cambridge University Press, Cambridge, New York. xiv + 531 pp. Hardcover. US $142.00. ISBN 978-0521-83358-5. BRETT A. GOODMAN School of Marine & Tropical Biology James Cook University, Townsville, 4811,Queensland, Australia e-mail: Brett.Goodman@jcu.edu.au Lizards have become a model group for research spanning a wide range of biological disciplines, including: morphology, physiology, ecology, behavior, and evolution. Over the past 40 years, many of the advances in lizard ecology have been periodically summarized in books beginning with the title Lizard Ecology. These include: Lizard Ecology: A Symposium (Milstead 1967), Lizard Ecology: Studies of a Model Organism (Huey et al. 1983), and Lizard Ecology: Historical and Experimental Perspectives (Pianka and Vitt 1994). The most recent addition, Lizard Ecology: The Evolutionary Consequences of Foraging Mode adds to this lineage and provides further evidence of the importance of lizards in understanding the evolutionary complexity and trade-offs inherent in the way animals forage for their food. At its core, Lizard Ecology focuses on the sit-and-wait (SW)/ambush mode of foraging versus the active, widely-foraging (WF) dichotomy (analogous to the r-K dichotomy of life history theory), where species are lumped into one or other foraging mode (FM). In the latest release, researchers have tested an immense range of hypotheses from the ﬁelds of ecology, evolutionary biology, and animal behavior using species with an equally diverse range of natural histories. The goal of the book is to review research on FM and assess its inﬂuence on the biology of squamates that has accumulated over the past 40 years. The book is divided into two parts: I. Organismal patterns of variation in FM, and II. Environmental inﬂuences of FM. The ﬁrst 11 chapters make up Part I and include relationships between FM and various aspects of squamate biology, such as physiology, morphology, anatomy, performance, behavior, diet and life history. Part II centers around the inﬂuence of nocturnality (geckos) on FM, plasticity in FM in response to environmental variation, and habitat use and its relationship to food acquisition. To begin, Ray Huey and Eric Pianka provide interesting insight into the emergence of the term FM (Pianka 1966). Following on in one of the most important chapters (Chapter 1), particularly Herpetological Review 40(3), 2009 371 in terms of data collection and quality, Gad Perry examines and identiﬁes the all-too-often neglected issues of methodology and terminology of foraging behavior. This is followed by theoretical predictions of foraging theory. Measuring foraging behavior, and the qualitative versus quantitative measures of speed, moves per minute and percentage time spent moving has often been problematic and there has rarely been consensus on how such foraging behaviors should be measured. Perry examines various taxonomic groups to determine the number of observations necessary to reliably estimate foraging behavior. In general, we ﬁnd that, with the exception of gekkonid lizards, relatively few samples (≥15) are needed to reliably estimate a species’ movement rates. Early predictions of foraging mode suggested “SW” foragers should have high sprint speed and low endurance, highlighting an evolutionary trade-off. In the longest chapter of the book (Chapter 2; 45 pages), Miles et al. use published data on sprint speed, endurance and FM to examine these hypotheses, and provide a macroevolutionary perspective on whether morphology and locomotor performance correlates with FM, using size-corrected data for the majority of lizard groups. Finally, they examine data within Anolis to provide a microevolutionary perspective using ﬁeld data on foraging behavior and locomotor performance in the ﬁeld. Nonetheless, their ancestral reconstruction reveals that shifts in FM from SW-WF occurred early in the evolution of squamate reptiles, probably at the node of scleroglossans, as other studies have suggested (e.g., Vitt and Pianka 2005). One result to emerge is that SW species have greater sprint speeds than WF species. In closing, the authors warn against categorizing SW-WF as end points of a continuum, but instead encourage the collection of additional data to develop a clearer picture of the differences in foraging behavior. In Chapter 3, correlates of lizard foraging mode, Kevin Bonine highlights known and potential correlations between physiology and performance traits that relate primarily to FM. Using Arnold’s (1983) paradigm (morphology → performance → behavior → ﬁtness) he presents results within a phylogenetic context to explore the link between morphology and performance. In addition, Bonine provides a thorough introduction to FM and physiology, and whole-animal performance measures, including energy balance, sprint speed, endurance, temperature, water loss, aerobic capacity, anaerobic scope, and the sub-organismal physiological traits of respiration and muscle physiology. Chapter 4 (Brown and Nagy) addresses the fundamental ecological question: do WF species have greater energetic costs than SW species? The authors use published data on ﬁeld metabolic rates (FMR), and doubly labeled water (DLW) for 46 lizard species, develop allometric equations for members of each FM, which they use to assess the position of new species, and to predict, develop and test bioenergetic hypotheses. They begin by reviewing older studies that examined distantly related SW-FW species, and move onto more recent studies which focus on more closely related species. Next, they examine DLW studies in an attempt to detect general patterns among SW and FW species, before examining the phylogenetic effects of energy use among lacertilians. While their results for the FMR data support the SW-FW dichotomy they suggest that as more foraging behavior data become available, it may be possible to use a continuous predictor (e.g., MPM) to investigate patterns in FM and energy use. 372 In Chapter 5 the modern-day fathers of lizard ecology, Laurie Vitt and Eric Pianka begin with the admission that… “the foraging mode paradigm is more complex than originally envisioned” (page 141). Fittingly, their chapter sets out to highlight major evolutionary and non-evolutionary factors that affect the prey types consumed by lizards, using diet data for 184 lizard species from Africa, Australia, North America and the New World Tropics. They begin by presenting factors likely to inﬂuence the prey consumed by lizards in addition to foraging mode, including; body size, biomechanics of feeding structures, thermoregulatory behavior, times of activity, sensory capabilities, physiological constraints, and resource availability. The results of their phylogenetic analyses revealed that variation among lizard diet is reduced by 80%, suggesting many of the differences are nested deep within the evolutionary history of lizards. The emergence of chemical prey discrimination, jaw prehension, and the use of a WF mode of locomotion to ﬁnd prey, no doubt led to an increase in prey types that were unavailable previous to iguanians. The sheer number of extant scleroglossans (snakes and lizards; 6000) to iguanians (1230) provides ample evidence for the success of these evolutionary innovations. In Chapter 6 Shine and Wall examine the reasons for the dramatic degree of intraspeciﬁc niche divergence in body size and sex observed in snakes compared to lizards. To begin, the authors highlight a series of mechanisms that may drive observed patterns of: size-dependent shifts in prey consumption, and intersexual niche divergence, including other factors that may cause variation in foraging traits within a species. In the ﬁnal part of their chapter, the authors highlight the difference between lizards and snakes (e.g., snakes consume a wider range of prey sizes), and follow this with a series of hypothesis that test these ideas. The authors suggest that the functional basis for the intraspeciﬁc shift in dietary niche frequently observed in snakes relates mainly to differences in the anatomy, physiology, ecology and behavior observed between snakes and lizards. Anthony Herrel (Chapter 7) aims to identify those traits (ecological, morphological and performance) in lizards that are typically associated with the two foraging modes, and compares these with the traits typically associated with an herbivorous lifestyle. One of the highlights of this chapter is the ﬁnding that ancestry appears not to have constrained the dietary mode of lizards. Regardless of FM the shift to herbivory has led to the evolution of ﬂat, bladelike teeth, high bite force, large size and a longer colon. Moreover, given that these traits are present within omnivorous species, it suggests that the evolutionary shift to herbivory has occurred via an omnivorous diet. Given this, Herrel poses the question of why are their so few scleroglossan herbivores? Clearly, this is one question requiring additional data! In Chapter 8, William Cooper describes the morphology and physiology of lizard chemosensory systems, the evidence for prey chemical discrimination, and how lizard FMs have inﬂuenced these relationships. Evidence is presented in the form of correlated evolution between the lingual-vomeronasal system, food chemical discrimination, and FM. We ﬁnd that differences in FM have inﬂuenced the evolution of diet, and this has then affected the responsiveness of species to the different chemicals of speciﬁc food types. In closing, Cooper discusses the role of the lingual-vomeronasal system and foraging mode, for prey chemical discrimination, and Herpetological Review 40(3), 2009 in driving the evolutionary diversiﬁcation of lizards. In Chapter 9 McBrayer and Corbin examine patterns of head shape variation in response to FM, in order to establish the existence of trade-offs between head shape, biting force, and FM using 22 species of lizard representing 12 families. Here we ﬁnd that head length and width has evolved in concert with FM, but that much remains to be learned. In particular, it is unclear whether head shape is changing in response to foraging mode alone, or whether other forces play a part. Following on, Reilly and McBrayer (Chapter 10) examine the convergence and divergence in prey capture and processing behavior and the evolution of lingual and sensory traits in lizards. In general, three distinct patterns of FM emerge: two distinct SW predators, one mixed forager, and thee distinct WFs. They suggest that despite the similar prey processing behaviors of the Iguania and Gekkota, it is achieved via fundamentally different forms of tongue use when capturing prey. The mixed FM and retention of primitive Autarchoglossan features and tongue prehension place the Scincoidea midway. Indeed, the evolution of a tongue with chemosensory abilities appears to have been a pre-determinate of a widely-foraging strategy and is probably a key component of the WF vs. SW FM dichotomy. Beaupre and Montgomery (Chapter 11) change gear by focusing on the FM of snakes, with reference to interspeciﬁc and broad-scale patterns among snakes. They point out that the foraging modes of snakes differ from that of lizards by being strongly determined by phylogeny: species within the same family tend to forage in a similar way, with relatively few exceptions. The authors consider an impressive range of factors likely to inﬂuence snake foraging mode. There is evidence, however, that snake foraging modes conform to the “syndrome hypothesis” by being sufﬁciently variable. Results from a bioenergtic model suggest that within snakes the dichotomous classes of FM may represent adaptive peaks, with intermediate FMs favored only under certain conditions. In closing, it becomes clear that a suitable deﬁnition of FM for snakes is lacking. In Chapter 12 Aaron Bauer examines a group that appears to be an exception to the SW–WF dichotomy, the Gekkota. The Gekkota contain some 1100+ species representing 106 genera from essentially three families. A large summary table (7 pages) provides details of the genera, number of species and FM. The Gekkota display a mixed foraging strategy, and provide some evidence that WF evolved with the Scleroglossa, and was probably facilitated by the development of chemosensory abilities. Thus, despite similar chemosensory abilities as autarchoglossans, the Gekkotans moved along an alternative evolutionary pathway for foraging, having retained visual predation and SW foraging. However, apart from this generality little more can be concluded. Thus, while geckos do not fall neatly into the dichotomous foraging paradigm, Bauer concedes that more data (movement patterns, additional lineages, etc.) are necessary to provide greater conﬁrmation on this placement. In Chapter 13 Martin Whiting examines plasticity in FM using the lizard Platysaurus broadleyi – a member of a clade of SW foragers which shows considerable variation in FM. For instance, juveniles move more in order to hunt insects, while adults move less often making short movements, but switch to active “herbivory” in order to increase the likelihood of encountering ﬁgs when available. This provides a clear ﬁtness beneﬁt in situations where a high quality resource becomes available, but which requires a different FM then that which is typically employed. Indeed, whether lizards experience a trade-off between the amount of time spent foraging and courtship/reproducetive behavior, or simply increase energy intake, is unknown—and a question in need of study. In closing, Whiting posses two questions highlighted by the Platysaurus system: 1. how widespread is FM plasticity and what is the effect of a spatially and temporally variable high energy resource on FM? and 2. what effect on FM does having traits common to both ambush and active foragers? Clearly, there awaits considerable research potential for such a system! In Chapter 14 Vanhooydonck et al. provide insight into the relationship between locomotor performance, bite performance and head morphology of lacertid lizards. They investigate how endurance and sprint performance affect an organism’s feeding ecology, and whether there is a trade-off between species that rely on sprint performance to acquire prey versus species that are reliant on increased stamina for prey capture. The results suggest endurance and the proportion of soft-bodied prey consumed is co-evolved in lacertid lizards. However, speed is not correlated with evasive prey, but rather with the proportion of intermediate prey in the diet. In males, body ﬂattening and climbing trade-off is due to a reduction in head height, which is likely to be beneﬁcial for maintaining the center of mass close to the substrate (Aerts et al. 2003), and reducing the possibility of the animal lifting off the substrate. Conversely, there iss no trade-off between body ﬂattening and climbing in females, possibly because of differential selection on head shape in males versus females. In Chapter 15 Roger Anderson examines FM from the perspective of the principal ecological features required by any organism to survive: ﬁnding food, avoiding becoming food, avoiding abiotic extremes (e.g., temperature), and reproducing. Anderson examines how food acquisition mode (FAM), as opposed to FM per se, varies among lizards, and among habitats. Unfortunately, the message from this chapter is that we still know too little to conclude much regarding the role of FAM on the evolution of lizard FM. Nonetheless, Anderson proposes promising possibilities for future research, such as the use of laboratory microcosms and semi-natural mesocosms for conducting experimental tests of FAM, prey types, competitors and predators. In the shortest chapter (Chapter 16) Vitousek et al. examine the Galapagos Marine Iguana, whose short, intense bouts spent foraging on macrophytic marine algae are unique among reptiles, and more reminiscent of a SW foraging style. They the marine iguana as a model system to test how natural and sexual selection drive morphological and behavioral adaptation, including the physiological and environmental constrains that act as intense selective pressures on this species. We ﬁnd that in response to the strong selective pressures of their energetically costly grazing bouts, marine iguanas have evolved a blunt head, salt glands, and dark coloration allow for maximal energy intake. In Chapter 17 McBrayer et al. summarize by highlighting those studies to emerge since Huey and Pianka (1981). They point out that while many traits relate to foraging mode, there remain many areas in need of additional research attention. The authors suggest that because FM span both physiological and morphological parameters there is a need for more integrative approaches and thinking; only in this way can both general patterns and variation in lizard foraging biology be understood. Herpetological Review 40(3), 2009 373 In closing, Lizard Ecology brings together a diverse range of information, offering a great starting point for anyone contemplating lizard studies likely to incorporate aspects of food acquisition and foraging behavior. Presumably, this is because it achieves its main goal by determining the inﬂuence of FM on the biology of squamate reptiles. Overall, the book contains few errors, a complement to the three editors. While some may lament the lack of color images, the text is well complemented with numerous tables (37), and ﬁgures (90). On the downside, however, the book is highly priced (US $142), and may be out of reach for students or people with just a general interest in lizard ecology. Price not withstanding, this work is a worthy acquisition for anyone interested in lizard ecology and behavior. As stated by Kevin Bonine in Chapter 3, the success of books of the Lizard Ecology series (Milstead 1967; Huey et al. 1983; Pianka and Vitt 1994), and the recent Lizards: Windows to the Evolution of Diversity (Pianka and Vitt 2003) highlight the appeal of lizard biology to scientists and a wider audience alike. LITERATURE CITED AERTS, P., R. VAN DAMME, K. D’AOUT, AND B. VANHOOYDONCK. 2003. Bipedalism in lizards: whole-body modeling reveals a possible spandrel. Proc. Roy. Soc. Lond. B 358:1525–1533. ARNOLD, S. J. 1983. Morphology, performance and fitness. Am. Zool. 23:347–361. HUEY, R. B., AND E. R. PIANKA. 1981. Ecological consequences of foraging mode. Ecology 62:991–999. ________ ________ , , AND T. W. SCHOENER. 1983 Lizard Ecology: Studies of a Model Organism. Harvard University Press, Cambridge, Massachusetts. vi + 501 pp. MILSTEAD, W. W. 1967. Lizard Ecology: A Symposium. University of Missouri Press, Columbia, Missouri. ix + 300 pp. PIANKA, E. R. 1966. Convexity, desert lizards, and spatial heterogeneity. Ecology 47:1055–1059. ________ , AND L. J. VITT. 1994. Lizard Ecology: Historical and Experimental Perspectives. Princeton University Press, Princeton. xii + 403 pp. ________ , AND ________. 2003. Lizards: Windows to the Evolution of Diversity. University of California Press, Berkeley. xiii + 333 pp. VITT, L. J., AND E. R. PIANKA. 2005. Deep history impacts present-day ecology and biodiversity. Proc. Natl. Acad. Sci. USA 102:7877–7881. Erratum In two recently published geographic distribution notes, the name of one of the coauthors (Paulo Nogueira da Costa) was printed incorrectly. The citations, with correct author names, are: SILVA-SOARES, THIAGO, PAULO NOGUEIRA DA COSTA, AND RODRIGO B. FERREIRA. 2009. Geographic distribution: Chiasmocleis carvalhoi (Central Humming Frog). Herpetological Review 40:107. SILVA-SOARES, THIAGO, RODRIGO B. FERREIRA, AND PAULO NOGUEIRA DA COSTA. 2009. Geographic distribution: Ischnocnema oea (Espírito Santo Robber frog). Herpetological Review 40:108. 374 Herpetological Review, 2009, 40(3), 374–376. © 2009 by Society for the Study of Amphibians and Reptiles Ecological and Environmental Physiology of Amphibians, by Stanley S. Hilman, Philip C. Withers, Robert C. Drewes, and Stanley D. Hilyard. 2009. Oxford University Press (www.oup.com). xii + 469 pp. Softcover. US $65.00. ISBN 978–0–19–857032–5. DAVID C. BLACKBURN Natural History Museum and Biodiversity Research Center University of Kansas, Lawrence, Kansas, 66045, USA e-mail: david.c.blackburn@gmail.com The new book Ecological and Environmental Physiology of Amphibians provides an up-todate and relatively concise summary of comparative research on amphibian physiology. This is the ﬁrst volume in the Ecological and Environmental Physiology Series published by Oxford University Press; similar volumes are in the works for reptiles, ﬁshes, birds, insects, and crustaceans (http://www.eeps-oxford.com). One may wonder whether this review is necessary following the relatively recent synthesis edited by Feder and Burrgren (1992). However, this new book ﬁlls a different niche and is more appropriate for those looking for a concise review, especially students. This multi-authored book provides a relatively well-integrated and cohesive view of the topic that should prove accessible to researchers, graduate students, and those teaching courses in physiology or amphibian biology. While Hillman and co-authors touch on many aspects of amphibian physiology, the book places a particular emphasis on water balance, an obviously important topic for amphibians and a dominant area of expertise of the authors. This book will serve as a useful and up-to-date addition to the much larger volume by Feder and Burggren (1992). Most importantly, this book will serve as an entry point for students and researchers interested in integrating a physiological component into their comparative and phylogenetic studies. While not as all encompassing as Feder and Burggren (1992), Hillman et al. succeed in producing a work is perhaps a more engaging introduction to amphibian physiology. This book provides references to and short summaries of relevant literature published since 1992, especially for some topics such as metabolic depression. It also provides gems for young physiologists and comparative biologists interested in unusual features of amphibians, such as the sections dealing with cutaneous water exchange (including “waterproof” frogs in Chiromantis or Phyllomedusa), the physiology of the “pelvic” or “seat” patch, cocoon formation, dehydration tolerance, and hypoxia. The book is organized into six chapters: an introduction with a discussion of diversity, phylogeny, and basic physiological challenges faced by amphibians; two chapters summarizing both basic and specialized aspects of amphibian physiology; a chapter on the Herpetological Review 40(3), 2009 physiology of species living in “extreme” environments; and two ﬁnal short chapters, one detailing approaches and techniques and the other on future directions. I found the separation of chapters into basic, specialized, and “extreme” physiology to be a useful division that highlights the physiology of unusual taxa living in deserts or regions with freezing soils. The section concerning techniques in amphibian physiology provides a quick introduction to oft-used methods and the inclusion of some historical background for some methods is a nice touch. The section on cardiovascular oxygen and carbon dioxide exchange (2.6) is probably my favorite. In many ways, it is representative of the many positive aspects of this book and its utility for those requiring an informed yet brief introduction. This section includes reviews of cardiovascular anatomy and basic physiology, and has a number of short, readable passages on topics such as arterial pressure, mixing of pulmocutaneous and systemic ﬂows, and cardiac output. This basic summary and introduction to the relationship between physiology and ecology will be of interest to audiences other than physiologists, such as those interested in ecological niche modeling. The chapter on physiological adaptations to extreme environments provides interesting food for thought for those considering how environmental variables contribute to determining species distributions. There is a general lack of knowledge about where amphibians actually spend their time (e.g., in trees, under ground, and high up or far down), but this book engenders an appreciation that this likely has important physiological consequences. There is also a fair amount of physiological variation between species, for instance in the ability to “supercool” which would effect species distributions. Basic data on the relationship between physiology and ecology is also important to evolutionary studies of the relationship between morphology and the environment, which, for example, has been relatively little explored in frogs, especially the Neobatrachia. In a relatively short review of a topic as large and varied as amphibian physiology, there are inevitably many areas that receive little attention; examples include endocrinology, development, or locomotory and behavioral energetics. Despite the focus on the relationship between physiology and the environment, the mention of phenotypic plasticity is limited to an exceedingly brief discussion of developmental rates. Studies on plasticity and reaction norms can form a beneﬁcial basis for many research programs (e.g., Schlichting and Pigliucci, 1998; West-Eberhard, 2003), including those on amphibian physiology. There are several short sections on larval physiology (e.g., energy budgets, skin morphology), but, in general, there is little attention paid to larvae or metamorphosis. For those interested in these topics, I suggest turning to the thorough review by Burggren and Just (1992). I appreciated that many of the ﬂow charts summarizing available data contain references pointing to appropriate literature. Summary tables, such as Table 2.2, are very useful and help to drive home the lack of knowledge for most species. Many of the plots and ﬂow charts will prove useful as teaching aids, though some drawings and photographs could be of higher quality (e.g., Figs. 2.37, 3.41, 4.6A, 5.10). The ﬁrst chapter of this work lays out a comparative framework that aims to encapsulate all living amphibian diversity, including a lengthy section titled “Habitats and Morphotypes.” While some discussion does appear in the literature (e.g., Emerson, 1988), to my knowledge, such a detailed and explicit discussion of amphibian morphotypes has not been published previously in a form that is easily accessible to students. The authors state that the discussion of morphotypes is intended to reﬁne the traditional categories of aquatic, fossorial, terrestrial, and arboreal. However, the categories proposed by the authors will need further scrutiny as some are separated by few substantial differences. For example, imagine trying to teach the difference between “semi-aquatic” and “semiterrestrial (mesic)”; the only character state given that differs between these morphotypes is the amount of webbing (fully webbed vs. not fully webbed). This is a good start, but further quantitative studies of morphotypes will be necessary. One drawback is that the relationship between morphotypes and environment is not adequately explored using comparative methods. Morphotypes are summarized by family in Table 1.1 (with the number of species examined in parentheses; R. Drewes, pers. comm.), but it is only much later in the book (p. 377) that the often extensive variation within families and genera is clearly pointed out. However, this review of morphotypes still provides a concise summary for students and is a satisfactory start at dissecting ecomorphological diversity within the three orders of living amphibians. At the time of the last thorough synthesis of environmental physiology of amphibians (i.e., 1992), the described diversity of amphibians was considerably less (~ 70% of the diversity recognized at the time of writing this review; D. Frost, pers. comm.) and a central problem confounding comparative research was the lack of phylogenetic data. Yet descriptions of new species and the accumulation of phylogenetic data now far outpace comparative morphological and physiological studies, which can be labor-intensive for obtaining data for a few, or even just one, species. While limited, comparative data do exist for some topics and could be analyzed using comparative methods to investigate correlations between physiological features or examine character evolution over the phylogeny. In fact, the authors do make some claims regarding the change or conservation of traits across phylogeny. For example, the authors state that the energy source underlying metamorphic climax varies “interspeciﬁcally with little phylogenetic inertia” (p. 110), though this appears to be based on only four species from three families. It seems that the authors intended to use comparative analyses to make use of recent phylogenetic data. However, the only example that I could ﬁnd of using comparative data in a phylogenetic context is the analysis of standard metabolic rate that appears in the sections on phylogeny and environmental physiology and on comparative methods. The authors might have missed an opportunity by not combining available data from recent phylogenetics research with that from either comparative physiology or the morphotypes that they describe. The text is peppered with comparisons between “aquatic” and “terrestrial” species, though usually with little mention of phylogenetic relationships. By couching these arguments in a phylogenetic context, the authors could have made an even stronger case concerning the environmental correlates of observed physiological diversity. Perhaps most importantly, this review by Hillman and co-authors underscores that the vast majority of living amphibians still remains to be studied in physiological research. We know little of the physiology of most amphibian larvae, exceedingly little about caecilians, not much about the relationship of physiological Herpetological Review 40(3), 2009 375 phenotypes with genotype × environment interactions, and without better studies of interspeciﬁc variation we will lack the insight into the “hows and whys” underlying physiological variation. With appropriate estimates of the amphibian tree of life in hand, we now need to integrate these with ecological and physiological data to understand “how environmental differences have shaped intra- and interspeciﬁc variation in amphibians’ physiological performance” (p. 376). A book such as this one is poised to inspire students to do exactly that. LITERATURE CITED BURGGREN, W. W., AND J. J. JUST. 1992. Developmental changes in physiological systems. Pp. 467–530 in M.E. Feder and W.W. Burggren (Eds). Environmental Physiology of the Amphibians. University of Chicago Press, Chicago. EMERSON, S. B. 1988. Convergence and morphological constraint in frogs: variation in postcranial morphology. Fieldiana (Zoology) 43:1–19. FEDER, M. E., AND W. W. BURGGREN (Eds.). 1992. Environmental Physiology of the Amphibians. University of Chicago Press, Chicago. vii + 646 pp. SCHLICHTING, C. D., AND M. PIGLUICCI. 1998. Phenotypic Evolution: a Reaction Norm Perspective. Sinauer Associates, Inc., Sunderland, Massachusetts. xii + 387 pp. WEST-EBERHARD, M. J. 2003. Developmental Plasticity and Evolution. Oxford University Press, Oxford. xx + 794 pp. Herpetological Review, 2009, 40(3), 376–377. © 2009 by Society for the Study of Amphibians and Reptiles Frogs and Toads of the Southeast, by Mike Dorcas and Whit Gibbons. 2008. The University of Georgia Press (www.ugapress. uga.edu). US $22.95. viii + 238 pp. Flexible cover. ISBN-978-08203-2922-2. JEFFREY C. BEANE North Carolina State Museum of Natural Sciences, Research Laboratory MSC # 1626, Raleigh, North Carolina 27699-1626, USA e-mail: jeff.beane@ncdenr.gov This book is the third in a series on amphibians and reptiles of the Southeast published by the University of Georgia Press. It closely follows the format of the two previous volumes on snakes (Gibbons and Dorcas 2005) and turtles (Buhlmann et al. 2008), and covers the same nine-state region (Virginia, Tennessee, the Carolinas, Georgia, Florida, Alabama, Mississippi, and Louisiana) deﬁned as the Southeast in the other volumes. Like the others in this series, it was written with a broad and general readership in mind, but it will also be highly useful to professional herpetologists and seasoned naturalists. 376 Species accounts are included for all 42 (38 native and four introduced) species known from the region. Each account is divided into sections on “Description,” “What do the tadpoles look like?,” “Similar species,” “Distribution and habitat,” “Behavior and activity,” “Food and feeding,” Description of call,” “Reproduction,” “Predators and defense,” “Conservation,” and “Comments.” Each also contains several photographs, shaded distribution maps of the species’ range within the region as well as its overall range, a sidebar with quick identiﬁcation tips, and a bar chart with peak calling months indicated by shading. In addition, the book includes a thorough introductory section, a glossary, a table of calling months for all species, a quick-reference table of species found in each state, a list of further reading, indices of scientiﬁc and common names, and a closing section on human-anuran interactions. The pages throughout are punctuated with “Did you know?” sidebars featuring interesting facts on frogs and toads. The book carries a strong conservation message throughout, and I have only praise for this conservation-oriented approach. The more than 300 color photographs range in quality from adequate to excellent. Virtually all serve their purpose of showcasing anuran beauty and diversity, pointing out important identiﬁcation features, and illustrating intraspeciﬁc variation. This last is an especially important feature — too many works allow only a single photograph or illustration of each species covered and fail to adequately cover the often extensive range of variation exhibited by most species. In a few cases, the choices of included photographs could have been improved upon. For example, all the included images of Hyla squirella show similarly-colored bright green individuals. As this frog is, color-wise, one of the most variable species in the Southeast, it would have been helpful to have included more images of individuals in gray, brown, and heavily spotted liveries. Several photographs are included more than once. While there is certainly nothing wrong with that per se, it may present an illusion that the book has more photos than it actually does, and considering the cost of color printing and the great array of quality anuran photographs available, any such repetition seems unnecessary. Most of the errors I noticed are ones I would regard as minor. To cite a few examples, on pages 10 and 78 (one of the aforementioned instances of an image being included more than once) a ribbon snake is misidentiﬁed as a common garter snake. I also believe the metamorphs identiﬁed as green treefrogs at the bottom of page 13 are probably squirrel or possibly pine woods treefrogs. “Brumation” is misspelled in the glossary and “Pine Barrens” is misspelled twice (“Pine Barren” and “Pine barrens”) on page 82. The green treefrog account (page 76) states that “no other southeastern treefrog has small, bright orange or gold spots” (in fact, the barking treefrog also frequently has such spots). Page 142 shows “a river frog transforming from a tadpole into an adult” (“. . . into a juvenile” would be more accurate). There are a number of errors in the index. For example, the index lists a nonexistent reference to squirrel treefrogs on page 211, and page 207 and 212 feature photographs of a southern toad and a barking treefrog, respectively, but these are not listed in the index. I also found the variable fonts and type sizes, and inconsistent use of italics among the listed page numbers in the index to be slightly confusing. Finally, I am inclined to question the statement (page 44): “Southern and northern cricket frogs occasionally interbreed in areas where Herpetological Review 40(3), 2009 both occur and can produce hybrids that are difﬁcult to identify.” While Mount (1975) and Jensen (2005) mentioned possible hybrids between these two species, and Mecham (1964) was able to produce laboratory hybrids, there is apparently scant, if any, solid evidence to support natural hybridization. Mette (2008) and Micancin and Mette (2009) suggested that specimens thought to be hybrids may be merely indicative of the difﬁculty in identifying these two species morphologically. I was pleased to see each species account include a brief description of the tadpole, as well as photographs of tadpoles for some species; this is another feature sorely lacking in many anuran ﬁeld guides. I did feel that some of the tadpole descriptions could have been a bit more accurate, thorough, or clear. While some might argue that tadpoles are often so variable that short verbal descriptions, or even photographs, could potentially be more misleading than useful, I view them as generally helpful, especially for species with particularly distinctive tadpoles (river frog, eastern narrowmouth toad, cricket frogs). At 7¾” x 10¼”, the book’s size may hinder its use somewhat as a ﬁeld guide, but the ﬂexible cover may compensate enough for it to ﬁt into a backpack or otherwise be more easily transported than would a hardback of the same dimensions. Overall, this volume is an excellent and welcome addition to the series. As the only published work covering exclusively the anurans occurring in the nine-state Southeast region, it will be a valuable addition to the library of anyone interested in the natural history of that region. LITERATURE CITED BUHLMANN, K., T. TUBERVILLE, and W. GIBBONS. 2008. Turtles of the Southeast. The University of Georgia Press, Athens. 252 pp. GIBBONS, W., and M. DORCAS. 2005. Snakes of the Southeast. The University of Georgia Press, Athens. 253 pp. JENSEN, J. B. 2005. Acris gryllus (LeConte, 1825). Pp. 443–445 in M. J. Lannoo (Ed.), Amphibian Declines: the Conservation Status of United States Species. University of California Press, Berkeley. MECHAM, J.S. 1964. Ecological and genetic relationships of the two cricket frogs, genus Acris, in Alabama. Herpetologica 20:84−91. METTE, J.T. 2008. The distribution of the cricket frogs Acris crepitans and Acris gryllus in a zone of sympatry. Unpublished undergraduate research report, University of North Carolina, Chapel Hill. 16 pp. MICANCIN, J. P., and J. T. METTE. 2009. Acoustic and morphological identiﬁcation of the sympatric cricket frogs Acris crepitans and A. gryllus and the disappearance of A. gryllus near the edge of its range. Zootaxa 2076:1–36. MOUNT, R.H. 1975. The Reptiles and Amphibians of Alabama. Auburn University Agricultural Experiment Station, Auburn, Alabama. 347 pp. Herpetological Review, 2009, 40(3), 377–379. © 2009 by Society for the Study of Amphibians and Reptiles Urban Herpetology (Herpetological Conservation, Number 3), edited by Joseph C. Mitchell, Robin E. Jung Brown, and Breck Bartholomew. 2008. Society for the Study of Amphibians and Reptiles, Salt Lake City (www.SSARherps.org). Hardcover. xvii + 586 pp. US $75.00. ISBN 978–0–916984–79–3. TRENTON W. J. GARNER Institute of Zoology, Zoological Society of London Regents Park, London NW1 4RY, United Kingdom e-mail: trent.garner@ioz.ac.uk Urban Herpetology is a brick. At 40 chapters and 13 case studies, this is not the type of tome one moves casually between ofﬁce and home. My 20 minute morning walk to work with this bad boy in my rucksack buckled my knees several times and cracked the casing of my lap top. I’ve successfully blocked ravaging graduate students from entering my ofﬁce by leaning it against my closed ofﬁce door. Be forewarned. Also be forewarned that there’s useful information in this book. Although I’ve had some experience with its effects…..the type locality for the Italian agile frog, Rana latastei, with which I have worked for several years, is located somewhere in the heart of downtown Milano, where the species is no longer to be found…..I had little familiarity with the speciﬁc topic of urban herpetology before agreeing to review this book. I was laboring under the delusion that the study of herps in cities and related areas fell under the old standards of habitat loss and modiﬁcation, pollution, invasive species and other IUCN-type threats (although even the IUCN has modiﬁed their categories: anyone ever heard of transportation and service corridors?). I was pleased to see some familiar words in the section headings and my readings made me realize more clearly how urbanization can bring speciﬁc, new and worrying takes on the old IUCN standards. Reading this book has persuaded me that there is a need for me to reconsider how urbanization does and will impact amphibian and reptile populations. I can envision how I may use the information contained within it to some effect, and the chapters I gave short shrift to will be subject to a more thorough reading in the very near future. In the age of narrow focus topical books or superﬁcial undergraduate text books, it is a pleasure to hold a substantial book that covers a fairly broad bit of ground with some depth on a topic that is moving rapidly up the research priority list. The book opens with a list of contributors, a forward and a preface, followed by an opening overview chapter. This chapter I found exceptionally useful, as it provided me with a tidy historical and geographic overview of urban herpetology. Mitchell and Jung Brown have spent some time collating a valuable list of relevant Herpetological Review 40(3), 2009 377 references, restricted to English language reference sources, and synthesized their contents neatly. I found their deﬁnitions of urban environments most useful, although there is some scope for further reﬁnement. Multiple deﬁnitions for exurban are provided, summarized as 1 house per 0.4–16 ha in a matrix that may include native landscapes. This does suggest that Caiapó settlements could be viewed as urbanized, while most would treat the Brazilian forest within which they live as natural. The rest of the book is broken into sections dealing with various types of habitat alteration (Section I: Direct and Indirect Effects of Habitat Loss and Alteration; Section II: Effects of Roads, Trails and Railroad Tracks; Section IV: Stormwater Ponds, Swimming Pools, Urban Lakes and Golf Courses), pollution (Section III: Chemical and Light Pollution), introduced species (Section V: Introduced Species, Urbanophiles, and Urbanophobes), reviews (Section VI: State, Regional, and Country Reviews), policy (Section VII: Management and Regulations) and outreach (Section VIII: Education and Citizen Involvement). Michael Lannoo provides the afterword. Each section contains up to eight chapters and case studies. Most of the papers specialized on a single or a few species focus on amphibians or chelonians, with a few snake, lizard and crocodilian papers. There are also several papers that attempt to cover both amphibians and reptiles. I found the papers varied widely in content. Ignoring reviews, quantitative studies ranged from excellent examples of population ecological studies that examined how human-driven habitat change affected population dynamics over time (Chapter 7, Plummer and Mills) to a two-page ‘case study’ consisting of a collection of anecdotes containing no new empirical data (Case Study 2, Vandeman). In addition, I am unsure that the weight afforded each section was justiﬁed. While the number of chapters and case studies in Section I was certainly warranted, why were there only ﬁve state, regional, and country reviews? Was it really necessary to have eight chapters and case studies in Section IV, or three chapters outlining the welfare issues of roads? Having said that, I applaud Andrews and Gibbons (Chapter 10) for presenting some rare empirical evidence of possible ecological costs of road mortality to snake populations. As discussed by Andrews, Gibbons and Jochimsen (Chapter 9), most studies of road mortality simply present counts of animals found dead on roads and extrapolating this animal welfare issue to actual population regulation and conservation is fraught with assumptions. While counting dozens or even hundreds of dead reptiles and amphibians on roads is appalling from a welfare point of view, if such a level of slaughter is maintained over time, this suggests that road mortality may not be increasing the likelihood of population extirpation. I encourage other researchers to go and determine if road mortality actually has the ability to regulate populations. The organization and classiﬁcation of papers is, on occasion, confusing. Hayes et al.’s study (Chapter 31), arguably one of the best of the book, reviews the ecology of Rana aurora in depth and puts it in context of existing legislation, making recommendations as to where legislation fails this species. Even the authors consider it a case study for the species however the paper has been classiﬁed as a chapter and put in the section on state, regional and country reviews, even though it is a species review. This chapter is an exception: overall, papers are correctly categorized. The book does have a large number of unreplicated studies that rely 378 on comparisons with previously published data to make conclusions as to the ‘health’ status of urbanized populations. Not ideal, and any such conclusion must be taken with a grain of salt, even though I do acknowledge the manpower and funding limitations imposed on research groups that dictate how much new data can be collected at any one time. Perhaps this may serve as a call for different research teams studying the same species in urban versus rural versus ‘natural’ habitats to join forces and develop broader, more comparative studies? I have three major issues with this book. First, the editors claim that Urban Herpetology encapsulates a global perspective, stating their hope that the book will “....set the stage for future research and conservation efforts around the world….” Clearly overblown: Mitchell and Jung Brown cite numerous recent and empirical studies from Africa, Europe, Canada and Australia in their opening chapter, along with some additional studies from Asia and Latin America, yet most chapters and case studies are based on studies of herps in the United State. There is a smattering of data chapters from Europe, Canada and Australia, along with two review articles representing Russia and the West Indies, but these come across as sops when compared to the number of U.S. studies included in the book versus the spectrum of available published literature tabulated and cited in Chapter 1. This bias narrows the insight into urban herpetology afforded readers of the book because the contents do not reﬂect the distribution of research effort on the topic. As stated by Mitchell and Jung Brown “….much of the available herpetofaunal literature on the impacts of urbanization……is based on work done in Europe. The only serious long-term studies have been conducted (t)here.” Further: “A growing body of literature on urban amphibians and reptiles continues to be produced in Australia.” The bias extends to the selection of section headings and their contents, to the detriment of herpetologists in other parts of the world. I ﬁnd it unlikely that researchers in much of Africa, Asia and Latin America will ﬁnd the section on stormwater ponds, swimming pools, urban lakes and golf courses valuable. Researchers studying how urbanization affects herps in the U.S.A. would certainly beneﬁt from understanding how advanced and long-term urbanization in areas such as Europe, Africa, the Fertile Crescent and some parts of Asia has impacted herps in those regions. Imagine studying the ecology of urbanized herps in Tokyo, the focus of Imperial Japanese power since the shogun of Edo won a civil war in the 16th century and subject to increasing levels of urbanization ever since! Are there any herps to be found in Greater Tokyo? Urban Herpetology doesn’t answer this question, but a quick Google hunt returns a study by Kusano and Inoue (2009) on the breeding phenology of three Japanese amphibian species in the suburbs of Tokyo, rather ironically published in the Journal of Herpetology. Of course, SSAR is an organization based in the United States, but once claims of a global perspective are made, a lack of attention to it grates. My second issue is with the complete lack of a genetics perspective. Habitat alteration, fragmentation and pollution, to name a few relevant urbanization processes, are known to inﬂuence genetic structure, and there is a wealth of genetic studies on urbanized herps to support this (e.g., Madsen et al. 1996, 1999; Hitchings and Beebee 1997; Andersen et al. 2004; Johansson et al. 2005; Lesbarrères et al. 2006; Ficetola et al. 2007; Noël et al. 2007; see Beebee 2005 for a review of genetics applications that are applica- Herpetological Review 40(3), 2009 ble to studies of urbanized herps). These studies consistently show how habitat fragmentation due to urbanization decrease genetic variability and gene ﬂow among populations, often with associated population-averaged or individual ﬁtness costs. Quantitative genetics have been used to explore amphibian responses to acid stress, providing insight into the potential for a species to adapt in the face of increasing chemical pollution (Merilä et al. 2004). Recent developments in Bayesian population genetics approaches have been successfully used to reveal demographic processes that were not detected using standard ﬁeld ecology methods (e.g., Jehle et al. 2005). Many of the studies included in Urban Herpetology describe demographic processes that should elevate genetic drift, such as low migration rates, and populations exposed to factors that should impose selection, yet the editors did not think the topic worthy of a single chapter or case study. Why the omission? My third issue is the decidedly mechanistic perspective evinced throughout the book. Urbanization is set to increase and more species will be forced to exist in human-modiﬁed habitat, thus identifying the circumstances that allow species to persist over long time scales is crucial for herpetological conservation. Where is the evolutionary perspective that would facilitate managing urbanized populations? In fact, where is the section on adaptive management, brieﬂy discussed by Hayes et al in Chapter 31? Important questions that can only be addressed using evolutionary concepts are not addressed in Urban Herpetology. How can we expect urbanization and climate change to interact and what are the consequences for amphibians and reptiles? Some chapters do take a more dynamic approach to urban herpetology and herpetological conservation, but by and large species and populations are treated as static units and conservation applications as one-off responses to interpretations of current data and trends. In the end, if you can stump up US $75, the book is worth buying after you determine if the range of topics, taxa, and geographic regions are relevant to your research program. My hope is that this ﬁrst book on urban herpetology will generate follow-ups, along the lines of Seigel and Collins’ snake ecology books or, better yet, Krebs and Davies’ series on Behavioral Ecology, that take this decent, but limited, ﬁrst effort into as yet unexplored aspects of urban herpetology. LITERATURE CITED ANDERSEN, L. W., K. FOG, AND C. DAMGAARD. 2004. Habitat fragmentation causes bottlenecks and inbreeding in the European tree frog (Hyla arborea). Proc. Roy. Soc. London B 271:1293–1302. BEEBEE, T. J. C. 2005. Conservation genetics of amphibians. Heredity 95:423-427. FICETOLA, G. F., T. W. J. GARNER, AND F. DE BERNARDI. 2007. Genetic diversity and ﬁtness in the threatened frog, Rana latastei, are inﬂuenced by the joint effect of post glacial colonization and isolation. Mol. Ecol. 16:1787–1797. HITCHINGS, S. P., AND T. J. C. BEEBEE. 1997. Genetics substructuring as a result of barriers to gene ﬂow in urban common frog (Rana temporaria) populations: implications for biodiversity conservation. Heredity 79:117–127. JEHLE, R., G. A. WILSON, J. W. ARNTZEN, AND T. BURKE. 2005. Contemporary gene ﬂow and the spatio-temporal genetic structure of subdivided newt populations (Triturus cristatus, T. marmoratus). J. Evol. Biol. 18:619–628. JOHANSSON, M., C. R. PRIMMER, J. SAHLSTEN, AND J. MERILÄ. 2005. The inﬂuence of landscape structure on occurrence, abundance and genetic diversity of the common frog, Rana temporaria. Global Change Biol. 11:1664–1679. KUSANO, T., AND M. INOUE. 2009. Long-term trends toward earlier breeding of Japanese amphibians. J. Herpetol. 42:608–614. LESBARRÈRES, D., C. R. PRIMMER, T. LODÉ, AND J. MERILÄ. 2006. The effects of 20 years of highway presence on the genetic structure of Rana dalmatina populations. Ecoscience 14:311–323. MADSEN, T., M. OLSSON, R. SHINE, AND H. WITTZELL. 1999. Restoration of an inbred population of adder (Vipera berus). Nature 402:34–35. ________ , B. STILLE, AND R. SHINE. 1996. Inbreeding depression in an isolated population of adders, Vipera berus. Biol. Conserv. 75:113–118. MERILÄ, J., F. SÖDERMAN, R. O’HARA, K. RÄSÄNEN, AND A. LAURILA. 2004. Local adaptation and genetics of acid-stress tolerance in the moor frog, Rana arvalis. Conserv. Genet. 5:513–527. NOËL, S., M. OUELLET, P. GALOIS, AND F.-J. LAPOINTE. 2007. Impact of urban fragmentation on the genetic structure of the eastern red-backed salamander. Conserv. Genet. 8:599–606. Happy 100th Birthday to Dr. Henry S. Fitch (December 25, 1909 – ) from The Herpetological Community Your many years of dedication to ﬁeld research, keen eye, perseverance, deep understanding and appreciation of your subjects, and timeless contributions will continue to inspire those of us who study reptiles and amphibians for many, many years to come. Herpetological Review 40(3), 2009 379 380 Herpetological Review 40(3), 2009 SSAR COMMITTEE CHAIRS COORDINATORS AND CHAIRPERSONS Standard English and Scientiﬁc Names BRIAN I. CROTHER Department of Biological Sciences Southeastern Louisiana University Hammond, Louisiana 70402, USA Conservation BETSIE ROTHERMEL Archbold Biological Station PO Box 2057 Lake Placid, Florida 33862, USA JOSEPH R. MENDELSON, III Zoo Atlanta 800 Cherokee Ave., SE Atlanta, Georgia 30315-1440, USA Grants-In-Herpetology ERIK R. WILD Department of Biology University of Wisconsin-Stevens Point Stevens Point, Wisconsin 54481-3897, USA JOSHUA M. KAPFER Natural Resources Consulting, Inc. 119 South Main Street, PO Box 128 Cottage Gove, Wisconsin 53527, USA Kennedy Student Award LYNNETTE SIEVERT Department of Biological Sciences Emporia State University Emporia, Kansas 66801, USA Metter Memorial Award JOSEPH J. BEATTY Department of Zoology Oregon State University Corvallis, Oregon 97331-2914, USA Nominating GREGORY WATKINS-COLWELL Yale Peabody Museum of Natural History New Haven, Connecticut 06520-8118, USA Resolutions RICHARD WASSERSUG Anatomy Department Dalhousie University Halifax, NS B3H 4H7 Canada Seibert Awards PATRICK OWEN Department of EEO Biology The Ohio State University at Lima Lima, Ohio 45804, USA Student Travel Awards MATTHEW VENESKY Department of Biology The University of Memphis Memphis, Tennessee 38152, USA CARI HICKERSON Biological, Geological & Environmental Science Cleveland State University Cleveland, Ohio 44115, USA Webmaster RAUL E. DIAZ University of Kansas Medical Center Lawrence, Kansas 66160, USA e-mail: lissamphibia@gmail.com COORDINATORS Elector DANIEL NOBLE Department of Integrative Biology University of Guelph Guelph, Ontario N1G 2W1, Canada Symposium Coordinator RICHARD D. DURTSCHE Department of Biological Sciences Northern Kentucky University Highland Heights, Kentucky 41099, USA INFORMATION FOR CONTRIBUTORS Herpetological Review is a peer-reviewed quarterly that publishes, in English, articles and notes of a semi-technical or non-technical nature, as well as book reviews, institutional features, commentaries, regional and international herpetological society news, and letters from readers directed to the ﬁeld of herpetology. Articles reporting the results of experimental research, descriptions of new taxa, or taxonomic revisions are not published in HR, but should be submitted to the Journal of Herpetology (see inside front cover for Editor’s address). To submit a manuscript to HR, please consult the SSAR webpage at: <http://www.ssarherps.org/pages/HRinfo.php> Reprints and Page Proofs Reprints of notes or articles published in HR may be ordered through EZReprints (EzReprint@odysseypress.com), an online reprint odering system. Authors will receive email notiﬁcation as the issue goes to press. Page proofs are sent electronically to authors of all articles, book reviews, and obituaries, but are not sent for natural history notes or geographic distribution notes. Proofs are sent as pdf ﬁles approximately 2–4 weeks prior to publication. Advertising Herpetological Review accepts commercial advertising. Rates and copy information are available from the SSAR web page (http://www.ssarherps.org/pages/HRinfo.php). Herpetological Review (ISSN: 0018-084X) is published quarterly (March, June, September, and December) by the Society for the Study of Amphibians and Reptiles at Central Michigan University, Department of Biology, 217 Brooks Hall, Mt. Pleasant, MI 48859, USA. Periodicals postage paid at Mt. Pleasant, MI 48859 and at additional mailing ofﬁces. All rights reserved. No part of this periodical may be reproduced without written permission of the Editor, except that authors and their agents are permitted to reproduce and distribute their own articles and notes. POSTMASTER: Send address changes to Herpetological Review, Allen Press, Inc., P.O. Box 1897, Lawrence, KS 66044, USA. Regarding missing or damaged issues, please notify Herpetological Review, Allen Press, Inc., P.O. Box 1897, Lawrence, KS 66044, USA (tel. 785/843-1234 or e-mail: herps@allenpress.com). SSAR Publications SSAR publishes seven different series: Journal of Herpetology, Herpetological Review, Facsimile Reprints in Herpetology, Contributions to Herpetology, Herpetological Circulars, Herpetological Conservation, and the Catalogue of American Amphibians and Reptiles (see below for CAAR details). SSAR members receive pre-publication discounts on all Facsimiles and Circulars and on volumes in the Contributions and Herpetological Conservation series. A complete pricelist of Society publications is available at: http://www.ssarbooks.com/. Catalogue of American Amphibians and Reptiles The Catalogue consists of loose-leaf accounts of taxa prepared by specialists, including synonymy, deﬁnition, description, distribution map, and comprehensive list of literature for each taxon. Covers amphibians and reptiles of the entire Western Hemisphere. Available for purchase from the SSAR Bookstore (http://www.ssarbooks. com/). Use the prices below to order back issues. COMPLETE SET: NUMBERS 1 – 840 US $460 INDEX TO ACCOUNTS 1 – 400: Cross-referenced, 64 pages $6 INDEX TO ACCOUNTS 401 – 600: Cross-referenced, 32 pages $6 SYSTEMATIC TABS (Ten tabs to ﬁt binder: “Class Amphibia,” “Order Caudata,” etc.) $6 IMPRINTED POST BINDER (Note: one binder holds about 200 accounts) $35 INCOMPLETE SET: NUMBERS 1 – 190 $75 191 – 410 $85 411 – 840 $320 To order: make checks payable to “SSAR” and mail to Breck Bartholomew, SSAR Publications Secretary, P.O. Box 58517, Salt Lake City, Utah 84158, USA (fax 801/4530489). e-mail: ssar@herplit.com. Online orders at: http://www.ssarbooks.com/. ISSN 0018-084X The Ofﬁcial News-Journal of the Society for the Study of Amphibians and Reptiles Herpetological Review Volume 40, Number 3 September 2009 ARTICLES The Taxonomic Status of the Inornate (Unstriped) and Ornate (Striped) Whiptail Lizards (Aspidoscelis inornata [Baird]) from Coahuila and Nuevo León, México ................................................................................by J. M. WALKER, J. R. DIXON, R. W. AXTELL, AND J. E. CORDES 276 Discovery of a Surviving Population of the Montane Streamside Frog Craugastor milesi (Schmidt) ..... by J. E. KOLBY AND J. R. MCCRANIE 282 Observations on the Ecology of Trachemys adiutrix and Kinosternon scorpioides on Curupu Island, Brazil ....................................................................................................................................................... by L. BARRETO, L. C. LIMA, AND S. BARBOSA 283 Foraging Ecology of Spotted Turtles (Clemmys guttata) in Ontario, Canada ........... by M. L. RASMUSSEN, J. E. PATERSON, AND J. D. LITZGUS 286 Herpetofaunal Conservation in the Rainforest: Perceptions of Ecotourists................................................................................ by T. M. DOAN 289 Distribution and Natural History Notes on Some Poorly Known Frogs and Snakes from Peninsular Malaysia .........................................................................................................................................................................by CHAN K. O AND NORHAYATI A. 294 On the Distribution and Status of Plectrohyla hazelae (Taylor, 1940) (Amphibia: Hylidae) from Oaxaca, Mexico ........................................................................................................................................................... by J. R. MENDELSON III AND E. K. KABAY 301 One Hundred and One New County Records for Alabama Amphibians and Reptiles ................................ by S. P. GRAHAM AND COLLEAGUES 367 TECHNIQUES A Method for Constructing an Adjustable Platform to Obtain Lateral Photographs of Larval Anurans .............................................................................................................................................................by M. C. SCHACHT AND L. D. MCBRAYER 303 Baiting Differentially Inﬂuences Capture Rates of Large Aquatic Salamanders, Siren and Amphiuma .................................................................................................................................... by C. P. SMITH, D. R. GREGOIRE, AND M. S. GUNZBURGER 304 AMPHIBIAN DISEASES Reassessment of the Historical Timeline for Batrachochytrium dendrobatidis Presence in Honduras and Conservation Implications for Plectrohyla dasypus .......................................................................................................................... by J. E. KOLBY AND G. E. PADGETT-FLOHR 307 Local and Regional Patterns of Amphibian Chytrid Prevalence on the Osa Peninsula, Costa Rica .............................................................................................................................................by C. S. GOLDBERG, T. J. HAWLEY, AND L. P. WAITS 309 Occurrence of Batrachochytrium dendrobatidis in an Anuran Community in the Southeastern Talamanca Region of Costa Rica ............................................................................................................................... by D. SAENZ, C. K. ADAMS, J. B. PIERCE, AND D. LAURENCIO 311 Detecting Batrachochytrium dendrobatidis in the Wild When Amphibians Are Absent ........................... by J. G. WIXSON AND K. B. ROGERS 313 First Report of Ranavirus Infecting Lungless Salamanders ............................................... by M. J. GRAY, D. L. MILLER, AND J. T. HOVERMAN 316 Amphibian Chytrid Fungus in Western Toads (Anaxyrus boreas) in British Columbia and Yukon, Canada .......................... by B. G. SLOUGH 319 Chytridiomycosis-Associated Mortality in Rana palustris Collected in Great Smoky Mountains National Park, Tennessee, USA ...................................................................................................................by M. TODD-THOMPSON, D. L. MILLER, P. E. SUPER, AND M. J. GRAY 321 HERPETOLOGICAL HUSBANDRY Nutrient Composition of Whole Crayﬁsh (Orconectes and Procambarus Species) Consumed by Hellbender (Cryptobranchus alleganiensis) .......................................... by E. S. DIERENFELD, K. J. MCGRAW, K. FIRTSCHE, J. T. BRIGGLER, AND J. ETTLING 324 BOOK REVIEWS Lizard Ecology: The Evolutionary Consequences of Foraging Mode.................................................................. reviewed by B. A. GOODMAN 371 Ecological and Environmental Physiology of Amphibians ............................................................................... reviewed by D. C. BLACKBURN 374 Frogs and Toads of the Southeast ............................................................................................................................... reviewed by J. C. BEANE 376 Urban Herpetology............................................................................................................................................... reviewed by T. W. J. GARNER 377 SSAR BUSINESS ......................................... 257 NEWSNOTES ............................................ 259 MEETINGS....................................................... 259 CURRENT RESEARCH ............................ 260 ZOO VIEW ................................................. 263 LETTERS TO THE EDITOR ...................... 273 NATURAL HISTORY NOTES ................... 330 GEOGRAPHIC DISTRIBUTION ................ 359

Log In

Geographic Distribution: Oxyrhopus melanogenys orientalis

Geographic Distribution: Oxyrhopus melanogenys orientalis

Related Papers

RELATED PAPERS