Published in the United States of America 2013 • VOLUME 7 • NUMBER 1 AMPHIBIAN & REPTILE CONSERWION SPECIAL MEX CO ISSUE amphibian-reptile-conservation.org ISSN: 1083-446X elSSN: 1525-9153 Editor Craig Hassapakis Berkeley, California, USA Associate Editors Raul E. Diaz Howard O. Clark, Jr. Erik R. Wild University of Kansas, USA Garcia and Associates, USA University of Wisconsin-Stevens Point, USA Assistant Editors Alison R. Davis University of California, Berkeley, USA Daniel D. Fogell Southeastern Community College, USA Editorial Review Board David C. Blackburn California Academy of Sciences, USA C. Kenneth Dodd, Jr. University of Florida, USA Harvey B. Lillywhite University of Florida, USA Peter V. Lindeman Edinboro University of Pennsylvania, USA Jaime E. Pefaur Universidad de Los Andes, VENEZUELA Jodi J. L. Rowley Australian Museum, AUSTRALIA Bill Branch Port Elizabeth Museum, SOUTH AFRICA Lee A. Fitzgerald Texas A&M University, USA Julian C. Lee Taos, New Mexico, USA Henry R. Mushinsky University of South Florida, USA Rohan Pethiyagoda Australian Museum, AUSTRALIA Peter Uetz Virginia Commonwealth University, USA Jelka Crnobrnja-Isailovc IBISS University of Belgrade, SERBIA Adel A. Ibrahim Ha’il University, SAUDIA ARABIA Rafaqat Masroor Pakistan Museum of Natural History, PAKISTAN Elnaz Najafimajd Ege University, TURKEY Nasrullah Rastegar-Pouyani Razi University, IRAN Larry David Wilson Institute Regional de Biodiversidad, USA Allison C. Alberts Zoological Society of San Diego, USA Michael B. Eisen Public Library of Science, USA Advisory Board Aaron M. Bauer Villanova University, USA James Hanken Harvard University, USA Walter R. Erdelen UNESCO, FRANCE RoyW. McDiarmid USGS Patuxent Wildlife Research Center, USA Russell A. Mittermeier Conservation International, USA Robert W. Murphy Royal Ontario Museum, CANADA Eric R. Pianka University of Texas, Austin, USA Antonio W. Salas Environment and Sustainable Development, PERU Dawn S. Wilson AMNH Southwestern Research Station, USA Honorary Members Carl C. Gans Joseph T. Collins ( 1923 - 2009 ) ( 1939 - 2012 ) Cover : Upper left: Bolitoglossa franklini. Photo by Sean Rovito. Upper right: Diaglena spatulata. Photo by Oscar Medina Aguilar. Center left: Agkistrodon bilineatus. Photo by Chris Mattison. Center right: Trachemys gaigeae. Photo by Vicente Mata-Silva. Lower left: Heloderma horridum. Photo by Tim Burkhardt. Lower right: Cerro Mariana, Balsas-Tepalcatepec Depression, ca. 12 km NW of Caracuaro, Michoacan. Photo by Javier Alvar ado -Diaz. Amphibian & Reptile Conservation — Worldwide Community-Supported Herpetological Conservation (ISSN: 1083-446X; elSSN: 1525-9153) is published by Craig Hassapakis/Amphibian & Reptile Conservation as full issues at least twice yearly (semi-annually or more often depending on needs) and papers are immediately released as they are finished on our website; http://amphibian-reptile-conservation.org; email: arc.publisher@gmail.com Amphibian & Reptile Conservation is published as an open access journal. Please visit the official journal website at: http://amphibian-reptile-conservation.org Instructions to Authors : Amphibian & Reptile Conservation accepts manuscripts on the biology of amphibians and reptiles, with emphasis on conservation, sustainable management, and biodiversity. Topics in these areas can include: taxonomy and phylogeny, species inventories, distri- bution, conservation, species profiles, ecology, natural history, sustainable management, conservation breeding, citizen science, social network- ing, and any other topic that lends to the conservation of amphibians and reptiles worldwide. Prior consultation with editors is suggested and important if you have any questions and/or concerns about submissions. Further details on the submission of a manuscript can best be obtained by consulting a current published paper from the journal and/or by accessing Instructions for Authors at the Amphibian and Reptile Conservation website: http://amphibian-reptile-conservation.org/submissions.html © Craig Hassapakis! Amphibian & Reptile Conservation Copyright: © 2013 Wilson. This is an open-access article distributed under the terms of the Creative Commons At- tribution-NonCommercial-NoDerivs 3.0 Unported License, which permits unrestricted use for non-commercial and education purposes only provided the original author and source are credited. Amphibian & Reptile Conservation 7(1): /-//. PREFACE AMPHIBIAN & REPTILE CONSERVATION SPECIAL MEXICO ISSUE Citation: Wilson LD. 2013. Preface ( Amphibian & Reptile Conservation Special Mexico Issue). Amphibian & Reptile Conservation 7(1): The allure of Mexico first beckoned me in 1957, but only from across the border, as along with my parents and sister I was visiting family members in Mission, Texas. Mission is a bit west of McAllen, just north of the interna- tional border, with Reynosa located on the southern bank of the Rio Bravo directly across from McAllen. We went to Reynosa just to say we had been in Mexico. My first herpetological trip to Mexico occurred in 1966, when Ernest A. Liner kindly took me on one of his many journeys. We traveled as far south as Chiapas, and saw much of the country and plenty of amphibians and reptiles. In the ensuing years, I traveled south of the border on several occasions, and ultimately visited all but one of Mexico’s 31 states. Among several others, 1 took one of those trips with Louis Porras, the senior author of the pa- per on cantils in this issue. I made another extensive trip with my father, Ward Wendell Wilson, and visited many of the ancient ruins for which the country is well known. During my career I have always been interested in Mexico, although in recent years I spent much of my time in Central America. Nevertheless, I was delighted at the opportunity to work on the book Conservation of Mesoamerican Amphibians and Reptiles (2010), which dealt with all of Mexico and Central America. This mas- sive undertaking presented me with the chance to work closely with two long-time friends, Jerry Johnson, one of my co-editors, and Louis Porras, the proprietor of Eagle Mountain Publishing, LC, and both are involved in this Special Mexico Issue. The herpetofauna of Mexico is impressive from a number of perspectives. At 1,227 species, it is almost twice the size of that of its northern neighbor (presently, the United States is known to contain 628 native species, according to the Center for North American Herpetol- ogy [naherpetology.org]; data accessed 17 March 2013); Mexico, however, is only about one-fifth the size of the United States. Mexico’s herpetofauna also is larger than that of the seven Central American nations combined (1,024 native species, according to Wilson and Johnson [2010], and my updating since), although the disparity be- tween Mexico and its southern neighbors is much smaller. Notably, Central America’s land area is slightly over one- fourth that of Mexico. amphibian-reptile-conservation.org The level of endemicity in Mexico also is spectacu- lar. In this Special Mexico Issue, Wilson, Mata-Silva, and Johnson report that 482 species of reptiles (excluding the marine species) of a total of 849 (56.8%) are Mexi- can endemics; Wilson, Johnson, and Mata-Silva indicate that 253 species of amphibians of a total of 378 (66.9%) are not found outside of Mexico. The combined figure is 736 endemics out of 1,227 species (60.0%), a percent- age substantially higher than that for Central America. In Central America, 367 endemic species have been re- corded to date (Wilson and Johnson [2010], and my up- dating since), which equates to 35.8%. According to the accounting at the Center for North American Herpetol- ogy website (www.cnah.org), however, compared to the figures for Mexico (see the two Wilson et al. papers in- dicated below), Canada (www.carcnet.ca) and the West Indies (Powell and Henderson 2012), of the 628 species listed, 335 are endemic to the United States, for which the resulting percentage (53.3%) is much closer to that of Mexico than for Central America. Because the United States is about five times the size of Mexico, when one compares the degree of endemism in these two countries with their respective land areas (area/number of endem- ics), the resulting figures (areas from the CIA World Fact- book; www.cia.gov) are as follows: Mexico (1,943,945 km 2 /736 = 2,641); and the United States (9,161,966 km 2 /335 = 25,808). Thus, the area/endemism ratio for the United States is almost 10 times that of Mexico, indicat- ing that endemism in Mexico is that much greater than that of its neighbor to the north. The comparable figure for Central America is 507,966 km 2 /367 = 1,384, which is even lower than that for Mexico, and this region already is regarded as a major source of herpetofaunal diversity (Wilson et al. 2010). The Mexican herpetofauna also is of immense impor- tance and interest from a conservation standpoint. In both of the Wilson et al. papers indicated below, the authors applied the Environmental Vulnerability Score (EVS) measure to Mexico’s herpetofauna and found that 222 of 378 amphibian species (58.7%) and 470 of 841 rep- tile species in (55.9%) were assigned an EVS that falls into the high vulnerability category. In total, 692 species (56.8%) fall into the highest category of susceptibility to environmental deterioration. The relatively small portion / June 2013 I Volume 7 | Number 1 | e62 Preface of humanity that recognizes the value and critical neces- sity of biodiversity is fighting an uphill battle to salvage as much biodiversity as possible before it disappears into extinction (Wilson 2006). Given the rate of human popu- lation growth and the commensurate rate of loss of natu- ral habitats, populations of these unique components of the Mexican patrimony likely will decline steadily, as is happening over the remainder of the planet (Raven et al. 2011 ). One of the most important imperatives we face, there- fore, is to take appropriate steps to conserve the Mexican herpetofauna. Toward this end, five papers collectively written by 10 contributors are expected to appear in this Special Mexico Issue of Amphibian & Reptile Conserva- tion. These papers are as follows: A conservation reassessment of the reptiles of Mexico based on the EVS measure by Larry David Wilson, Vicente Mata-Silva, and Jerry D. Johnson. A taxonomic reevaluation and conservation assess- ment of the common cantil, Agkistrodon bilinea- tus (Squamata: Viperidae): a race against time by Louis W. Porras, Larry David Wilson, Gordon W. Schuett, and Randall S. Reiserer. Patterns of physiographic distribution and conserva- tion status of the herpetofauna of Michoac an, Mex- ico by Javier Alvarado-Diaz, Ireri Suazo-Ortuno, Larry David Wilson, and Oscar Medina- Aguilar. Taxonomic reevaluation and conservation of beaded lizards, Heloderma horridum (Squamata: Helo- dermatidae) by Randall S. Reiserer, Gordon W. Schuett, and Daniel D. Beck. A conservation reassessment of the amphibians of Mexico based on the EVS measure by Larry David Wilson, Jerry D. Johnson, and Vicente Mata-Silva. All of these papers deal with issues of herpetofaunal con- servation, and range in coverage from the entire country of Mexico, through a single Mexican state, to what have been regarded as single species. Each study provides a set of recommendations. These five papers are gathered under this Preface and an issue cover. The concept behind the cover is to draw the papers into a coherent whole that reinforces the mis- sion of the journal, which is to “support the sustainable management of amphibian and reptile biodiversity.” Thus, the photograph of Cerro Mariana, located in the Balsas-Tepalcatepec Depression between Huetamo and Morelia, in Michoacan, is intended to illustrate dry forest, the type of vegetation most heavily damaged in Meso- america (Janzen 1988), one of the major features of the state’s environment and in which a significant portion of the herpetofauna is found. This type of environment is amphibian-reptile-conservation.org inhabited by two of the reptiles featured in this issue, the common cantil (Agkistrodon bilineatus ) and the beaded lizard (Heloderma horridum ), as well as the shovel-head- ed treefrog (Diaglena spatulataf all three of these species are relatively broadly distributed in subhumid environ- ments along the Pacific coastal region of Mexico, as well as in the extensive valley of the Balsas and Tepalcatepec rivers, of which the western portion lies in the state of Michoacan. Finally, our aim is to examine the conservation status of the amphibians and reptiles of Mexico, in general, and to focus more closely on a state herpetofauna (of Micho- acan) and on two prominent and threatened Mexican flag- ship species, the common cantil and the beaded lizard. Thus, we hope to contribute to the ongoing effort to pro- vide for a sustainable future for the world’s amphibians (Stuart et al. 2010) and reptiles (Bohm et al. 2013). Literature Cited Bohm M et al. 2013. The conservation status of the world’s reptiles. Biological Conservation 157: 372- 385. Janzen DH. 1988. Tropical dry forests: the most en- dangered major tropical ecosystem. Pp. 130-137 In: Biodiversity. Editor, Wilson EO. National Academy Press, Washington, DC, USA. Powell R, Henderson RW (Editors). 2012. Island lists of West Indian amphibians and reptiles. Florida Museum of Natural History Bulletin 51 : 85—166. Raven PH, Hassenzahl DM, Berg LR. 2011. Environ- ment (8 th edition). John Wiley & Sons, Inc., Hoboken, New Jersey, USA. Stuart SN, Chanson JS, Cox NA, Young BE. 2010. The global decline of amphibians: current trends and fu- ture prospects. Pp. 2-15 In: Conservation of Meso- american Amphibians and Reptiles. Editors, Wilson LD, Townsend JH, Johnson JD. Eagle Mountain Pub- lishing, LC, Eagle Mountain, Utah, USA. Wilson, EO. 2006. The Creation: An Appeal to Save Life on Earth. W. W. Norton & Company, New York, New York, USA. Wilson LD, Johnson JD. 2010. Distributional patterns of the herpetofauna of Mesoamerica, a biodiversity hotspot. Pp. 30-235 In: Conservation of Mesoameri- can Amphibians and Reptiles. Editors, Wilson LD, Townsend JH, Johnson JD. Eagle Mountain Publish- ing, LC, Eagle Mountain, Utah, USA. Wilson LD, Townsend JH, Johnson JD. 2010. Conserva- tion of Mesoamerican Amphibians and Reptiles. Ea- gle Mountain Publishing, LC, Eagle Mountain, Utah, USA. Larry David Wilson 2 May 2013 ii June 2013 | Volume 7 | Number 1 | e62 Copyright: © 2013 Johnson et al. This is an open-access article distributed under the terms of the Creative Com- mons Attribution-NonCommercial-NoDerivs 3.0 Unported License, which permits unrestricted use for non-com- mercial and education purposes only provided the original author and source are credited. Amphibian & Reptile Conservation 7(1): iii-vi. DEDICATIONS Citation: Johnson JD, Porras LW, Schuett GW, Mata-Silva V, Wilson LD. 2013. Dedications ( Amphibian & Reptile Conservation Special Mexico Issue). Amphibian & Reptile Conservation 7(1): iii-vi. With the publication of this Special Mexico Issue (SMI), the contributing authors were provided with an opportu- nity to dedicate it to herpetologists who have played a sig- nificant role in their lives, as well as the lives of other her- petologists past and present. Each of the 10 contributors was asked to identify the person who was most influential in their respective careers, especially with respect to what each of them has contributed to SMI. The dedicatees are: Miguel Alvarez del Toro. Miguel Alvarez del Toro (August 23, 1917-August 2, 1996) was bom in the city of Colima, Colima, Mexico, according to an obituary in Herpetological Review by Os- car Flores-Villela and Wendy Hodges in 1999. He moved to Mexico City in 1932, where he attended and later grad- uated from high school. Although his formal education was limited, his repute as an avid naturalist spread rapidly and at the age of 21, while still in Mexico City, he began a long career devoted to a multitude of zoological and conservation related disciplines. He moved to Chiapas in 1942, and after a short stint as keeper and curator became the Director of what then was known as the Instituto de Historia Natural located near downtown Tuxtla Gutierrez. His reputation grew exponentially because of his tireless work at the Zoological Park and Natural History Muse- um, his publication record, including books and papers on numerous vertebrate and invertebrate groups, and his sol- emn activism on conservation issues. One of his greatest legacies was convincing several generations of politicians in Chiapas to help develop a system of natural protected areas, and also to expand the Zoological Park and move it to “El Zapotal,” a relatively pristine site on the southern edge of the city. That new and remarkable facility was s . named “Zoologico Regional Miguel Alvarez del Toro, or ZOOMAT as it is popularly called today. Because of his lifetime efforts, “Don Miguel,” as he was called respect- fully, was justly awarded honorary doctoral degrees from the Universidad de Chapingo, in 1992, and from the Uni- versidad Autonomo de Chiapas, in 1993. Over his long career he received a plethora of other awards, and also was involved in numerous conservation projects in con- junction with various local, state, national, and interna- tional organizations. Jerry D. Johnson, an avid “herper” since grade school and recently discharged from the Marine Corps after a stint in Viet Nam, enrolled in the 1971 wintermester course at Fort Hays State University (Kansas), and ac- companied Dr. Charles A. Ely to Chiapas on a migratory bird study. Dr. Ely, after recognizing Johnson’s eagerness to search for amphibians and reptiles through all sorts of tropical and highland environments, included him on many return trips during the next several years. On that initial 1971 trip, Johnson briefly met Don Miguel at the old Zoological Park. In 1974, Dr. Ely arranged for he and Johnson to pitch tents in Don Miguel’s back yard, located near the Zoo. This initiated an opportunity to mingle with s all sorts of interesting people, including the Alvarez del Toro family, their friends, and a continuous flow of trav- eling naturalists who were visiting the Zoo. During those times Johnson realized just how influential Don Miguel’s scientific and conservation work had become, in Chiapas and elsewhere. On a typical day, Don Miguel often would walk among the Zoological Park’s animal enclosures, and during those walks Jerry came to know him while dis- cussing the status of herpetology in Chiapas, how con- servation efforts were in dire straits, and pondering his doubts about the possibility that anything resembling a natural Chiapas would persist into the future. In 1985, Don Miguel published a book entitled \Asi Era Chiapasl that described how Chiapas had changed in the 40 years since he had arrived in the state. Even today, Johnson of- ten thinks about how habitat destruction had altered the Chiapan environment since he began investigations there in 1971, as a college sophomore. He now realizes that his life and professional experiences have passed rather quickly, but sadly, environmental decay is accelerating at an even greater pace. Johnson now concentrates much of his professional efforts on conservation issues, hoping that humankind can avoid total environmental devasta- tion. Jerry also is reasonably sure that Don Miguel really didn’t expect preservation efforts to be very successful, amphibian-reptile-conservation.org Hi June 2013 I Volume 7 | Number 1 | e64 Dedications but he didn’t give up his dream of a more conservation- oriented populace by continually teaching people why preserving natural habitats is important to their own well- being, which probably is the only way conservation will ever succeed. With great pleasure, Johnson dedicates his contributions to this special Mexico edition of Amphib- s ian and Reptile Conservation to Miguel Alvarez del Toro, who in his opinion was the leading advocate and pioneer of biodiversity conservation in 20 th century Mexico. Roger Conant in his early 20s. Roger Conant (May 6, 1909-December 19, 2003) was born in Mamaroneck, New York, USA. As a child he de- veloped a passion for reptiles, especially snakes, and at the age of 19 became the Curator of Reptiles at the Tole- do Zoo. After assembling a sizeable collection of reptiles for public display, he was promoted to General Curator. Because of the close proximity of Toledo to Ann Arbor, he occasionally would visit herpetologists at the Univer- sity of Michigan and became close friends with a then- graduate student, Howard K. Gloyd. Eventually, Roger left Toledo to become the Curator of Herpetology at the Philadelphia Zoo, and in time became the zoo’s Director. Throughout his 38-year career at Philadelphia he partici- pated in weekly radio shows, edited the zoo’s publica- tions, and made frequent television appearances. During this time he also helped establish the Philadelphia Herpe- tological Society, served as President of the Association of Zoological Parks and Aquariums, and as President of the American Association of Ichthyologists and Herpe- tologists. In 1947 Roger married Isabelle Hunt Conant, an accomplished photographer and illustrator who had been working at the zoo for several years, and during the following two decades the couple made several collecting trips to Mexico. Roger’s first of 240 scientific publica- tions (including 12 books) came at the age of 19; about a decade later he authored The Reptiles of Ohio, a landmark amphibian-reptile-conservation.org book that set the standard for state herpetological publi- cations. Roger perhaps is best known as the author of the best selling book in herpetological history, A Field Guide to the Reptiles and Amphibians of Eastern North Ameri- ca, which was illustrated by Isabelle. The book was pub- lished in 1958, and expanded versions followed in 1975, 1991, and 1998. For the majority of amphibian and reptile enthusiasts and herpetologists living in the eastern part of the United States during those years, this book became their bible. In 1973, Roger retired early from the Philadel- phia Zoo, after Isabelle had become ill. The Conants then moved to Albuquerque, where Roger became an adjunct professor at the University of New Mexico and devoted much of his time to herpetology. Isabelle passed away in 1976, and soon after Roger discovered that his close friend, Howard K. Gloyd, was terminally ill. Howard had been busy working on a project that he and Roger started in 1932, and because of Howard’s deteriorating condi- tion Roger made an enormous commitment and assured Howard that the project would be completed. This hugely important contribution, entitled Snakes of the Agkistro- don Complex: a Monographic Review, was published by the Society for the Study of Amphibians and Reptiles (SSAR) in 1990. During this time Roger also was busy writing his memoirs, A Field Guide to the Life and Times of Roger Conant, which was published in 1997 by Selva, and details his remarkable life and illustrious career. Roger Conant in Santa Rosa National Park, Costa Rica (1982). Louis W. Porras and Gordon W. Schuett, two very close friends of Roger’s, were involved at several levels with the Agkistrodon monograph and Roger’s autobiog- raphy. Because of their mutual interest in Agkistrodon, in January of 1982 the trio traveled to Costa Rica in search of cantils and although no individuals were found in the iv June 2013 | Volume 7 I Number 1 | e64 Dedications field, they managed to secure preserved specimens for study. In July of that year, Porras returned to Costa Rica with John Rmdfleish and collected what became the holo- type of Agkistrodon bilineatus howardgloydi. Additional information on the life of Roger Conant appears in an obituary published in the June 2004 issue of Herpetologi- cal Review. Among several solicited tributes indicating how Roger had affected his colleague’s lives and careers, Porras wrote the following summary: As a giant in herpetology, no doubt many will be writing about Roger Conant ’s amazing organizational skills, at- tention to detail, literary contributions, lifelong produc- tivity, and so on. From a personal perspective, however, Roger was my friend, mentor, and father figure. He en- riched my life in so many ways, and it would warm his heart to know that by simply following his example, he will continue to do so. Schuett summarized his tribute as follows: In reflection, I have no doubt that Roger Conant pos- sessed genius. His was not displayed in eccentric man- nerisms and arrogant actions, but in a subtle and quiet ability to collect, organize, and process information for large-scale projects. In his research, each and every de- tail was painstakingly considered. Roger’s vast achieve- ments are even more remarkable knowing that he was largely self-educated. If genius is measured by the degree to which one’s ideas and work influence others, Roger stands among the giants of knowledge. . . Cheers to you, Roger, to your remarkable and enviable life. Yes, Indeed! Aurelio Ramfrez-Bautista was bom in Xalapa, Vera- cruz, Mexico, and today is a professor and biological in- vestigator at the Universidad Autonoma del Estado de Hi- dalgo. Dr. Ramfrez-Bautista has authored or co-authored more than 100 publications, including five books and 40 book chapters, made numerous presentations on the ecology and conservation of the Mexican herpetofauna, and has become one of the leading herpetologists in the country. During his many years as an educator and re- searcher, Dr. Ramfrez-Bautista advised numerous bache- lor, master, and doctoral students. Vicente Mata-Silva met Dr. Ramfrez-Bautista in the summer of 1998, as an un- dergraduate student working on his thesis on the herpeto- fauna of a portion of the state of Puebla. They developed a friendship, and through Dr. Ramirez-Bautista’s mentor- ing Vicente developed a passion for Mexican herpetolo- gy, especially Chihuahuan Desert reptiles, that continued throughout his undergraduate studies and later through master’s, doctoral, and post-doctoral work in the Ecology and Evolutionary Biology program at the University of Texas at El Paso. They have continued to work on sig- nificant research projects on the conservation and ecology of the Mexican herpetofauna. Vicente is extremely grate- amphibian-reptile-conservation. org Aurelio Ramfrez-Bautista in Chamela, Jalisco (2011). ful to Dr. Ramfrez-Bautista for his farsighted and life-al- tering introduction to herpetology. Their association has led to a lifetime friendship, and a road of excitement and opportunities that Vicente never envisioned possible. Dr. Ramfrez-Bautista is the epitome of what an educator and mentor should be, providing students the opportunity to become professional scientists working in a world sorely in need of commitment to environmental sustainability. Hobart M. Smith in Mexico (1930). Hobart Muir Smith (September 26, 1912-March 4, 2013) was bom Frederick William Stouffer in Stanwood, Iowa, USA. At the age of four, he was adopted by Charles and Frances Smith; both of his adoptive parents died, however, before Dr. Smith finished college at Kansas State University (KSU). In the engaging “historical per- spective” written by David Chiszar, Edwin McConkey, i/ June 2013 | Volume 7 | Number 1 | e64 Dedications and Margaret M. Stewart and published in the 2004(2) issue of Copeia, the authors recount an amazing story in- dicating that when Dr. Smith (HMS) was in his senior year in high school he was plagued by tachycardia and an allergy to caffeine, which ended his interest in running and led to youthful resolution that they reported as fol- lows: “If I’m gonna do anything worthwhile, I had better get to it, because I not gonna live very long” (!). Upon completing high school, he headed for KSU with expecta- tions of a major in entomology. A fortunate meeting with Howard K. Gloyd, a somewhat older student who was majoring in herpetology, brought HMS a change of heart, however, and he became determined to study amphibians and reptiles. He made this decision after having traveled to the American West on collecting trips with Dr. Gloyd, whose association with Dr. Conant is discussed above. Gloyd and his major professor at the University of Michi- gan, Dr. Frank Blanchard, suggested that HMS contact Edward H. Taylor at the University of Kansas (KU). As noted by Chiszar et al. (2004: 419), “this was probably the act that cinched HMS to a herpetological orientation and kiboshed entomology.” In fact, these authors also claim that “HMS literally collected his BA and moments later hopped into Taylor’s car bound for Mexico,” and that “the rest is history.” Hobart M. Smith and Rozella B. Smith at the University of Wyoming (1960). In 1940 (Wilson’s birth year), at age 26, he married Rozella Pearl Beverly Blood, who he met while both were graduate students at KU. Their marriage endured until Rozella’s death in 1987. Dr. Smith began working in Mexico in 1932, before any of the SMI contributors was born, and those early collecting trips instilled a life- long dedication for studying the Mexican herpetofauna. Other collecting ventures followed during the remainder of the decade. The material assembled during these trips allowed him to begin a life-long journey to record the composition, distribution, and systematics of the amazing Mexican herpetofauna. During his long life he authored more than 1,600 publications, including 29 books — the greatest output in the history of herpetology. Chiszar et al. (2004: 421-422) indicated that HMS was most proud of the three Mexican checklists, the Sceloporus monograph, the Handbook of Lizards, the comparative anatomy text- book (which Wilson used when he took the course under HMS), the Synopsis of the Herpetofauna of Mexico, the Pliocercus book, and the Candoia monograph. In 1947, HMS became a professor of zoology at the University of Illinois at Urbana-Champaign, and remained there until 1968. During this period in his career, one of the SMI contributors came under his influence. In 1958, Larry Da- vid Wilson graduated from Stephen Decatur High School in Decatur, Illinois, and the following year enrolled at Millikin University in that city. After two years and hav- ing exhausted the coursework offered by the biology department at Millikin, Wilson decided to move to the U of I, which became a turning point in his life. There, he met HMS and managed to survive a number of his courses, including comparative anatomy. During the two years that led to his graduation, Wilson cemented his in- terest in zoology and, due to Smith’s influence, decided to attend graduate school and major in herpetology. Also, due to Smith’s interest in Mesoamerican amphibians and reptiles, Wilson was determined to specialize in studying these creatures, and in 1962 ventured south and never re- turned to live in the flatlands of the “Great Corn Desert.” In 1983, Wilson had the opportunity to acknowledge his gratitude to the Smiths by organizing a symposium on the Mexican herpetofauna in their honor, which was held in connection with the annual SSAR meeting in Salt Lake City, Utah. Although much of Wilson’s overall work has focused on the Honduran herpetofauna, this special issue on the Mexican herpetofauna provided him with an op- portunity to reawaken his love for the country where his fieldwork outside the US began in 1966, and to again ac- knowledge his debt to Dr. Hobart Muir Smith, one of the most important people in the history of herpetology. As Wilson stated in a tribute to HMS on his centenary pub- lished last year in Herpetological Review, “I know I am only one of many people who are indebted to Dr. Smith in ways small and large. For me, however, his influence determined the direction of my career and, in a significant way, the nature of the contributions I have made to our field.” Acknowledgments. — The authors of the papers com- prising the Special Mexico Issue are very grateful to Sally Nadvornik, who kindly supplied the photographs we used of her father, Hobart M. Smith, and Uriel Hemandez-Sali- nas, who helpfully provided the image we used of Aurelio Ramfrez-Bautista. Louis Porras provided the photographs / of Roger Conant. The image of Miguel Alvarez del Toro was taken from the 3 rd edition of his book, Los Reptiles de Chiapas. amphibian-reptile-conservation.org vi June 2013 I Volume 7 | Number 1 | e64 Xenosaurus tzacualtipantecus. The Zacualtipan knob-scaled lizard is endemic to the Sierra Madre Oriental of eastern Mexico. This medium-large lizard (female holotype measures 188 mm in total length) is known only from the vicinity of the type locality in eastern Hidalgo, at an elevation of 1,900 m in pine-oak forest, and a nearby locality at 2,000 m in northern Veracruz (Woolrich- Pina and Smith 2012). Xenosaurus tzacualtipantecus is thought to belong to the northern clade of the genus, which also contains X. newmanorum and X. platyceps (Bhullar 2011). As with its congeners, X. tzacualtipantecus is an inhabitant of crevices in limestone rocks. This species consumes beetles and lepidopteran larvae and gives birth to living young. The habitat of this lizard in the vicinity of the type locality is being deforested, and people in nearby towns have created an open garbage dump in this area. We determined its EVS as 17, in the middle of the high vulnerability category (see text for explanation), and its status by the IUCN and SEMAR- NAT presently are undetermined. This newly described endemic species is one of nine known species in the monogeneric family Xenosauridae, which is endemic to northern Mesoamerica (Mexico from Tamaulipas to Chiapas and into the montane portions of Alta Verapaz, Guatemala). All but one of these nine species is endemic to Mexico. Photo by Christian Berriozabal-Islas. amphibian-reptile-conservation.org 01 June 2013 I Volume 7 | Number 1 | e61 Copyright: © 2013 Wilson et al. This is an open-access article distributed under the terms of the Creative Com- mons Attribution-NonCommercial-NoDerivs 3.0 Unported License, which permits unrestricted use for non-com- mercial and education purposes only provided the original author and source are credited. Amphibian & Reptile Conservation 7(1): 1-47. A conservation reassessment of the reptiles of Mexico based on the EVS measure ^arry David Wilson, 2 Vicente Mata-Silva, and 3 Jerry D. Johnson 1 Centro Zamomno de Biodiversidad, Escuela Agricola Panamericana Zamorano, Departamento de Francisco Morazdn, HONDURAS 2 3 Depart- ment of Biological Sciences, The University of Texas at El Paso, El Paso, Texas 79968-0500, USA Abstract. — Mexico is the country with the most significant herpetofaunal diversity and endemism in Mesoamerica. Anthropogenic threats to Mexico’s reptiles are growing exponentially, commensu- rate with the rate of human population growth and unsustainable resource use. In a broad-based multi-authored book published in 2010 ( Conservation of Mesoamerican Amphibians and Reptiles ; CMAR), conservation assessment results differed widely from those compiled in 2005 by IUCN for a segment of the Mexican reptile fauna. In light of this disparity, we reassessed the conservation status of reptiles in Mexico by using the Environmental Vulnerability Score (EVS), a measure previ- ously used in certain Central American countries that we revised for use in Mexico. We updated the total number of species for the Mexican reptile fauna from that reported in CMAR, which brought the new number to 849 (three crocodilians, 48 turtles, and 798 squamates). The 2005 assessment categorized a small percentage of species in the IUCN threat categories (Critically Endangered, En- dangered, and Vulnerable), and a large number of species in the category of Least Concern. In view of the results published in CMAR, we considered their approach overoptimistic and reevaluated the conservation status of the Mexican reptile fauna based on the EVS measure. Our results show an inverse (rather than a concordant) relationship between the 2005 IUCN categorizations and the EVS assessment. In contrast to the 2005 IUCN categorization results, the EVS provided a conservation assessment consistent with the threats imposed on the Mexican herpetofauna by anthropogenic en- vironmental degradation. Although we lack corroborative evidence to explain this inconsistency, we express our preference for use of the EVS measure. Based on the results of our analysis, we provide eight recommendations and conclusions of fundamental importance to individuals committed to reversing the trends of biodiversity decline and environmental degradation in the country of Mexico. Key words. EVS, lizards, snakes, crocodilians, turtles, IUCN categories, IUCN 2005 Mexican Reptile Assessment Resumen. — Mexico es el pais que contiene la diversidad y endemismo de herpetofauna mas signifi- cative en Mesoamerica. Las amenazas antropogenicas a los reptiles de Mexico crecen exponencial- mente acorde con la tasa de crecimiento de la poblacion humana y el uso insostenible de los recur- sos. Un libro publicado por varios autores en 2010 ( Conservation of Mesoamerican Amphibians and Reptiles; CMAR) produjo resultados sobre conservacion ampliamente contrarios a los resultados de una evaluacion de un segmento de los reptiles mexicanos conducida en 2005 por la UICN. A la luz de esta disparidad, se realizo una nueva evaluacion del estado de conservacion de los reptiles mexicanos utilizando una medida llamada el Calculo de Vulnerabilidad Ambiental (EVS), revisado para su uso en Mexico. Se actualizo el numero de especies de reptiles mexicanos mas alia del es- tudio de CMAR, por lo que el numero total de especies se incremento a 849 (tres cocodrilidos, 48 tortugas, y 798 lagartijas y serpientes). La evaluacion de 2005 de la UICN clasifico una proporcion inesperadamente pequena de especies en las categories para especies amenazadas (En Peligro Critico, En Peligro, y Vulnerable) y un porcentaje respectivamente grande en la categoria de Preo- cupacion Menor. En vista de los resultados publicados en CMAR, consideramos que los resultados de este enfoque son demasiado optimistas, y reevaluamos el estado de conservacion de todos los reptiles mexicanos basandonos en la medida de EVS. Nuestros resultados muestran una relacion inversa (mas que concordante) entre las categorizaciones de la UICN 2005 y EVS. Contrario a los resultados de las categorizaciones de la UICN 2005, la medida de EVS proporciono una evaluacion para la conservacion de reptiles mexicanos que es coherente con las amenazas impuestas por la degradacion antropogenica del medio ambiente. No tenemos la evidencia necesaria para propor- cionar una explicacion para esta inconsistencia, pero expresamos las razones de nuestra prefer- ence por el uso de los resultados del EVS. A la luz de los resultados de nuestro analisis, hemos Correspondence. Emails: 'bufodoc@aol.com (Corresponding author), 2 vmata@ utep.edu, 3 jjohnson@ utep.edu amphibian-reptile-conservation.org 02 June 2013 I Volume 7 | Number 1 | e61 Wilson et al. construido ocho recomendaciones y conclusiones de importancia fundamental para las personas comprometidas en revertir las tendencias asociadas con la perdida de biodiversidad y la degra- dation del medio ambiente. Palabras claves. EVS, lagartijas, culebras, cocodrflidos, tortugas, categories de UICN, 2005 UICN valoracion de reptiles mexicanos Citation: Wilson LD, Mata-Silva V, Johnson JD. 2013. A conservation reassessment of the reptiles of Mexico based on the EVS measure. Amphibian & Reptile Conservation 7(1 ): 1-47 (e61 ). The history of civilization is a history of human beings as they become increasingly knowledgeable about biologi- cal diversity. Beattie and Ehrlich 2004: 1. Introduction From a herpetofaunal standpoint, Mexico is the most significant center of diversity in the biodiversity hotspot of Mesoamerica (Mexico and Central America; sensu Wilson and Johnson [2010]). Of the 1,879 species of amphibians and reptiles listed by Wilson and Johnson (2010) for all of Mesoamerica, 1,203 (64.0%) occur in Mexico; reptiles are especially diverse in this country, with 830 species (72.3%) of the 1,148 species distributed throughout Mesoamerica. Wilson and Johnson (2010) also reported that the highest level of herpetofaunal endemism in Mesoamerica is found in Mexico (66.8% for amphibians, 57.2% for reptiles [60.2% combined]), with the next highest level in Honduras (36.2% for amphibians, 19.2% for reptiles [25.3% combined]). The reported level of herpetofaunal diversity and endemism in Mexico has continued to in- crease, and below we discuss the changes that have oc- curred since the publication of Wilson et al. (2010). Interest in herpetofaunal diversity and endemicity in Mexico dates back nearly four centuries (Johnson 2009). Herpetologists, however, only have become aware of the many threats to the survival of amphibian and reptile populations in the country relatively recently. The prin- cipal driver of these threats is human population growth (Wilson and Johnson 2010), which is well documented as exponential. “Any quantity that grows by a fixed percent at regular intervals is said to possess exponential growth” (www.regentsprep.org). This characteristic predicts that any population will double in size depending on the percentage growth rate. Mexico is the 11 th most popu- lated country in the world (2011 Population Reference Bureau World Population Data Sheet), with an estimated mid-2011 total of 114.8 million people. The population of Mexico is growing at a more rapid rate (1.4% rate of natural increase) than the global average (1.2%), and at a 1 .4% rate of natural increase this converts to a doubling time of 50 years (70/1.4 = 50). Thus, by the year 2061 the population of Mexico is projected to reach about 230 amphibian-reptile-conservation.org 03 million, and the population density will increase from 59 to 118/km 2 (2011 PBR World Population Data Sheet). Given the widely documented threats to biodiversity posed by human population growth and its consequences (Chiras 2009; Raven et al. 2011), as well as the increas- ing reports of amphibian population declines in the late 1980s and the 1990s (Blaustein and Wake 1990; Wake 1991), the concept of a Global Amphibian Assessment (GAA) originated and was described as “a first attempt to assess all amphibians against the IUCN Red List Cat- egories and Criteria” (Stuart et al. 2010). The results of this assessment were startling, and given broad press coverage (Conservation International 2004; Stuart et al. 2004). Stuart et al. (2010) reported that of the 5,743 spe- cies evaluated, 1,856 were globally threatened (32.3%), i.e., determined to have an IUCN threat status of Criti- cally Endangered (CR), Endangered (EN), or Vulnerable (VU). An additional 1,290 (22.5%) were judged as Data Deficient (DD), i.e., too poorly known for another deter- minable status. Given the nature of the Data Deficient category, eventually these species likely will be judged in one of the threat categories (CR, EN, or VU). Thus, by adding the Data Deficient species to those determined as globally threatened, the total comes to 3,146 species (54.8% of the world’s amphibian fauna known at the time of the GAA). Our knowledge of the global amphib- ian fauna has grown since the GAA was conducted, and a website (AmphibiaWeb) arose in response to the real- ization that more than one-half of the known amphibian fauna is threatened globally or too poorly known to con- duct an evaluation. One of the functions of this website is to track the increasing number of amphibian species on a global basis. On 8 April 2013 we accessed this website, and found the number of amphibian species at 7,116, an increase of 23.9% over the number reported in Stuart et al. (2010). As a partial response to the burgeoning reports of global amphibian population decline, interest in the con- servation status of the world’s reptiles began to grow (Gibbons et al. 2000). Some of this interest was due to the recognition that reptiles constitute “an integral part of natural ecosystems and [...] heralds of environmental quality,” just like amphibians (Gibbons et al. 2000: 653). Unfortunately, Gibbons et al. (2000: 653) concluded that, “reptile species are declining on a global scale,” and fur- ther (p. 662) that, “the declines of many reptile popula- tions are si mil ar to those experienced by amphibians in June 2013 I Volume 7 | Number 1 | e61 Conservation reassessment of Mexican reptiles Dermatemys mawii. The Central American river turtle is known from large river systems in Mexico, from central Veracruz south- ward into Tabasco and Chiapas and northeastward into southwestern Campeche and southern Quintana Roo, avoiding the northern portion of the Yucatan Peninsula. In Central America, it occurs in northern Guatemala and most of Belize. The EVS of this single member of the Mesoamerican endemic family Dermatemyidae has been calculated as 17, placing it in the middle of the high vulner- ability category, and the IUCN has assessed this turtle as Critically Endangered. This image is of an individual emerging from its egg, with its egg tooth prominently displayed. The hatching took place at the Zoologico Miguel Alvarez del Toro in Tuxtla Gutier- rez, Chiapas, as part of a captive breeding program for this highly threatened turtle. The parents of this hatchling came from the hydrologic system of the Rio Usumacinta and Playas de Catazaja. Photo by Antonio Ramirez Velazquez. Terrapene mexicana. The endemic Mexican box turtle is distributed from southern Tamaulipas southward to central Veracruz and westward to southeastern San Luis Potosf. Its EVS has been determined as 19, placing it in the upper portion of the high vulnerabil- ity category, but this turtle has not been evaluated by IUCN. This individual is from Gomez Farias, Tamaulipas, within the Reserva de la Biosfera El Cielo. Photo by Eli Garda Padilla. amphibian-reptile-conservation.org 04 June 2013 | Volume 7 | Number 1 | e61 Wilson et al. terms of taxonomic breath, geographic scope, and sever- ity.” They also identified the following significant threats to reptile populations: habitat loss and degradation, intro- duced invasive species, environmental pollution, disease [and parasitism], unsustainable use, and global climate change. Essentially, these are the same threats identified by Vitt and Caldwell (2009) in the Conservation Biology chapter of their textbook Herpetology. In the closing chapter of Conservation of Mesoameri- can Amphibians and Reptiles, Wilson and Townsend (2010: 774-777) provided six detailed and intensely critical recommendations for the conservation of the herpetofauna of this region, based on the premise that “problems created by humans ... are not solved by treat- ing only their symptoms.” Because of the nature of these recommendations, we consider it important to note that the IUCN conducted a conservation assessment of the Mexican reptiles in 2005, for which the results were made available in 2007 (see NatureServe Press Release, 12 September 2007 at www.natureserve.org). The contents of this press release were startling and unexpected, how- ever, as indicated by its title, “New Assessment of North American Reptiles Finds Rare Good News,” and contrast the conclusions of Wilson and Townsend (2010), which were based on the entire herpetofauna of Mesoamerica. The principal conclusion of the press release was that “a newly completed assessment of the conservation status of North American reptiles shows that most of the group is faring better than expected, with relatively few spe- cies at severe risk of extinction.” Wilson and Townsend (2010: 773) commented, however, that “conserving the Mesoamerican herpetofauna will be a major challenge for conservation biologists, in part, because of the large number of species involved and the considerable number that are endemic to individual countries, physiographic regions, and vegetation zones.” Given the contrast in the conclusions of these two sources, and because the 2005 Mexican reptile assess- ment was based on the IUCN categories and criteria without considering other measures of conservation sta- tus, herein we undertake an independent reassessment of the reptile fauna of Mexico based on the Environmen- tal Vulnerability Score (EVS), a measure developed by Wilson and McCranie (2004) for use in Honduras, which was applied to the herpetofauna of certain Central Amer- ican countries in Wilson et al. (2010), and modified in this paper for use in Mexico. The IUCN System of Conservation Status Categorization The 2005 Mexican reptile assessment was conducted using the IUCN system of conservation status categori- zation. This system is used widely in conservation biol- ogy and applied globally, and particulars are found at the IUCN Red List of Threatened Species website (www. iucnredlist.org). Specifically, the system is elaborated in amphibian-reptile-conservation.org the online document entitled “IUCN Red List of Catego- ries and Criteria” (2010), and consists of nine categories, identified and briefly defined as follows (p. 9): Extinct (EX): ‘ ‘A taxon is Extinct when there is no rea- sonable doubt that the last individual has died.” Extinct in the Wild (EW): ‘ ‘A taxon is Extinct in the Wild when it is known only to survive in cultivation, in captivity or as a naturalized population (or popula- tions) well outside the past range.” Critically Endangered (CR): ‘ ‘A taxon is Critically En- dangered when the best available evidence indicates that it meets any of the criteria A to E for Critically Endangered, and it is therefore considered to be fac- ing an extremely high risk of extinction in the wild.” Endangered (EN): “A taxon is Endangered when the best available evidence indicated that it meets any of the criteria A to E for Endangered, and is therefore considered to be facing a very high risk of extinction in the wild.” Vulnerable (VU): ‘ ‘A taxon is Vulnerable when the best available evidence indicates that it meets any of the criteria A to E for Vulnerable, and it is therefore con- sidered to be facing a high risk of extinction in the wild.” Near Threatened (NT): “A taxon is Near Threatened when it has been evaluated against the criteria but does not quality for Critically Endangered, Endan- gered, or Vulnerable now, but is close to qualifying for or is likely to qualify for a threatened category in the near future. Least Concern (LC): “A taxon is Least Concern when it has been evaluated against the criteria and does not qualify for Critically Endangered, Endangered, Vul- nerable or Near Threatened. Widespread and abun- dant taxa are included in this category.” Data Deficient (DD): “A taxon is Data Deficient when there is inadequate information to make a direct, or indirect, assessment of its risk of extinction based on its distribution and/or population status.” Not Evaluated (NE): “A taxon is Not Evaluated when it is has not yet been evaluated against the criteria.” As noted in the definition of the Near Threatened catego- ry, the Critically Endangered, Endangered, and Vulner- able categories are those with a threat of extinction in the wild. A lengthy discussion of criteria A to E mentioned in the definitions above is available in the 2010 IUCN document. A Revised EVS for Mexico In this paper, we revised the design of the EVS for Mex- ico, which differs from previous schemes in the compo- nents of geographic distribution and human persecution. Initially, the EVS was designed for use in instances where the details of a species’ population status (upon June 2013 I Volume 7 | Number 1 | e61 05 Conservation reassessment of Mexican reptiles Trachemys gaigeae. The Big Bend slider is distributed along the Rio Grande Valley in south-central New Mexico and Texas, as well as in the Rio Conchos system in Chihuahua. Its EVS has been calculated as 18, placing it in the upper portion of the high vulner- ability category, and the IUCN has assessed this turtle as Vulnerable. This individual is from the Rio Grande about 184 straight kilo- meters SE of Ciudad Juarez, Chihuahua. Although the picture was taken on the US side (about 44 km SSW of Van Horn, Hudspeth County, Texas), it was originally in the water. Photo by Vicente Mata-Silva. Kinosternon oaxacae. The endemic Oaxaca mud turtle occurs in southern Oaxaca and adjacent eastern Guerrero. Its EVS has been estimated as 15, placing it in the lower portion of the high vulnerability category, and the IUCN considers this kinosternid as Data Deficient. This individual was found in riparian vegetation along the edge of a pond in La Soledad, Tututepec, Oaxaca. Photo by Vicente Mata-Silva. amphibian-reptile-conservation.org 06 June 2013 | Volume 7 | Number 1 | e61 Wilson et al. which many of the criteria for the IUCN status catego- rizations depend) are not available, so as to estimate its susceptibility to future environmental threats. In this regard, the EVS usually can be calculated as soon as a species is described, as it depends on information gen- erally available when the species is discovered. Use of the EVS, therefore, does not depend on population as- sessments, which often are costly and time consuming. Nonetheless, its use does not preclude the implementa- tion of other measures for assessing the conservation sta- tus of a species, when these measures can be employed. After all, conservation assessment measures are only a guide for designing conservation strategies, and consti- tute an initial step in our effort to protect wildlife. The version of the EVS algorithm we developed for use in Mexico consists of three scales, for which the val- ues are added to produce the Environmental Vulnerabil- ity Score. The first scale deals with geographic distribu- tion, as follows: 1 = distribution broadly represented both inside and outside Mexico (large portions of range are both inside and outside Mexico) 2 = distribution prevalent inside Mexico, but limited outside Mexico (most of range is inside Mexico) 3 = distribution limited inside Mexico, but preva- lent outside Mexico (most of range is outside Mexico) 4 = distribution limited both inside and outside Mexico (most of range is marginal to areas near border of Mexico and the United States or Central America) 5 = distribution only within Mexico, but not re- stricted to vicinity of type locality 6 = distribution limited to Mexico in the vicinity of type locality The second scale deals with ecological distribution based on the number of vegetation formations occupied, as follows: 1 = occurs in eight or more formations 2 = occurs in seven formations 3 = occurs in six formations 4 = occurs in five formations 5 = occurs in four formations 6 = occurs in three formations 7 = occurs in two formations 8 = occurs in one formation The third scale relates to the degree of human persecution (a different measure is used for amphibians), as follows: 1 = fossorial, usually escape human notice 2 = semifossorial, or nocturnal arboreal or aquatic, nonvenomous and usually non-mimicking, sometimes escape human notice 3 = terrestrial and/or arboreal or aquatic, generally ignored by humans 4 = terrestrial and/or arboreal or aquatic, thought to be harmful, might be killed on sight 5 = venomous species or mim ics thereof, killed on sight 6 = commercially or non-commercially exploited for hides, meat, eggs and/or the pet trade The score for each of these three components is added to obtain the Environmental Vulnerability Score, which can range from 3 to 20. Wilson and McCranie (2004) divided the range of scores for Honduran reptiles into three cat- egories of vulnerability to environmental degradation, as follows: low (3-9); medium (10-13); and high (14-19). We use a similar categorization here, with the high cat- egory ranging from 14-20. For convenience, we utilized the traditional classifica- tion of reptiles, so as to include turtles and crocodilians, as well as lizards and snakes (which in a modern context comprise a group). Recent Changes to the Mexican Reptile Fauna Our knowledge of the composition of the Mexican rep- tile fauna keeps changing due to the discovery of new species and the systematic adjustment of certain known species, which adds or subtracts from the list of taxa that appeared in Wilson et al. (2010). Since that time, the fol- lowing nine species have been described: Gopherus morafkai : Murphy et al. (2011). ZooKeys 113:39-71. Anolis unilobatus : Kohler and Vesely (2010). Herpe- tologica 66: 186-207. Gerrhonotus f cirri: Bryson and Graham (2010). Her- petologica 66: 92-98. Scincella kikaapoda : Garcia- Vasquez et al. (2010). Copeia 2010: 373-381. Lepidophyma cuicateca: Canseco-Marquez et al. (2008). Zootaxa 1750: 59-67. Lepidophyma zongolica : Garcia- Vasquez et al. (2010). Zootaxa 2657: 47-54. Xenosaurus tzacualtipantecus : Woolrich-Pina and Smith (2012). Herpetologica 68: 551-559. Coniophanes michoacanensis : Flores- Villela and Smith (2009). Herpetologica 65: 404-412. Geophis occabus : Pavon-Vazquez et al. (2011). Her- petologica 67: 332-343. amphibian-reptile-conservation.org 07 June 2013 I Volume 7 | Number 1 | e61 Conservation reassessment of Mexican reptiles Abronia smithi. Smith’s arboreal alligator lizard is endemic to the Sierra Madre de Chiapas, in the southeastern portion of this state. Its EVS has been determined as 17, placing it in the middle of the high vulnerability category; the IUCN, however, lists this lizard as of Least Concern. This individual was found in cloud forest in the Reserva de la Biosfera El Triunfo, Chiapas. Photo by Eli Garcia- Pad ilia. amphibian-reptile-conservation.org 08 June 2013 | Volume 7 | Number 1 | e61 Wilson et al. The following 1 8 taxa either have been resurrected from the synonymy of other taxa or placed in the synonymy of other taxa, and thus also change the number of species in the CMAR list: Phyllodactylus nocticolus : Blair et al. (2009). Zoo- taxa 2027 : 28-42. Resurrected as a distinct species from P. xanti. Sceloporus albiventris : Lemos-Espinal et al. (2004). Bulletin of the Chicago Herpetological Society 39: 164-168. Resurrected as a distinct species from S. horridus. Sceloporus bimaculatus : Leache and Mulcahy (2007). Molecular Ecology 16: 5216-5233. Returned to the synonymy of S. magister. Plestiodon bilineatus : Feria-Ortiz et al. (2011). Her- petological Monographs 25: 25-51. Elevated to full species from P brevirostris. Plestiodon dicei: Feria-Ortiz et al. (2011). Herpeto- logical Monographs 25: 25-51. Elevated to full species from P. brevirostris. Plestiodon indubitus : Feria-Ortiz et al. (2011). Herpe- tological Monographs 25: 25-51. Elevated to full species from P. brevirostris. Plestiodon nietoi: Feria-Ortiz and Garcia- Vazquez (2012). Zootaxa 3339: 57-68. Elevated to full spe- cies from P brevirostris. Aspidoscelis sticto gramma: Walker and Cordes (2011). Herpetological Review 42: 33-39. Elevat- ed to full species from A. burti. Xenosaurus agrenon: Bhullar (2011). Bulletin of the Museum of Comparative Zoology 160: 65-181. El- evated to full species from X. grandis. Xenosaurus rackhami : Bhullar (2011). Bulletin of the Museum of Comparative Zoology 160: 65-181. El- evated to full species from X. grandis. Lampropeltis californiae: Pyron and Burbrink (2009). Zootaxa 2241: 22-32. Elevated to full species from L. getula. Lampropeltis holbrooki: Pyron and Burbrink (2009). Zootaxa 2241: 22-32. Elevated to full species from L. getula. Lampropeltis splendida: Pyron and Burbrink (2009). Zootaxa 2241: 22-32. Elevated to full species from L. getula. Sonora aequalis: Cox et al. (2012). Systematic s and Biodiversity 10: 93-108. Placed in synonymy of S. mutabilis. Coniophanes taylori: Flores-Villela and Smith (2009). Herpetologica 65: 404-412. Resurrected as a dis- tinct species from C. piceivittis. Leptodeira maculata: Daza et al. (2009). Molecular Phylogenetics and Evolution 53: 653-667. Synon- ymized with L. cussiliris. The correct name of the taxon, however, contrary to the decision of Daza et al. (2009), is L. maculata , inasmuch as this name was originated by Hallowell in 1861, and thus has priority. Leptodeira cussiliris, conversely, origi- nally was named as a subspecies of L. annulata by Duellman (1958), and thus becomes a junior syn- onym of L. maculata. Crotalus ornatus: Anderson and Greenbaum (2012). Herpetological Monographs 26: 19-57. Resur- rected as a distinct species from the synonymy of C. molossus. Mixcoatlus browni: Jadin et al. (2011). Zoological Journal of the Linnean Society 163: 943-958. Res- urrected as a distinct species from M. barbouri. The following species have undergone status changes, including some taxa discussed in the addendum to Wil- son and Johnson (2010): Anolis beckeri: Kohler (2010). Zootaxa 2354: 1-18. Resurrected as a distinct species from A. pentapri- on, which thus no longer occurs in Mexico. Marisora brachypoda: Hedges and Conn (2012). Zoo- taxa 3288: 1-244. Generic name originated for a group of species formerly allocated to Mabuya. Sphaerodactylus continentalis: McCranie and Hedges (2012). Zootaxa 3492: 65-76. Resurrection from synonymy of S. millepunctatus, which thus no lon- ger occurs in Mexico. Holcosus chaitzami, H. festivus, and H. undulatus: Harvey et al. (2012). Zootaxa 3459: 1-156. Gener- ic name originated for a group of species formerly allocated to Ameiva. Lampropeltis knoblochi: Burbrink et al. (2011). Mo- lecular and Phylogenetic Evolution. 60: 445-454. Elevated to full species from L. pyromelana, which thus no longer is considered to occur in Mexico. Leptodeira cussiliris: Mulcahy. 2007. Biological Journal of the Linnean Society 92: 483-500. Re- moved from synonymy of L. annulata, which thus no longer occurs in Mexico. See Leptodeira macu- lata entry above. Leptodeira uribei: Reyes- Velasco and Mulcahy (2010). Herpetologica 66: 99-110. Removed from the genus Pseudoleptodeira. Rhadinella godmani: Myers. 2011. American Muse- um Novitates 3715: 1-33. Species placed in new genus from Rhadinaea. Rhadinella hannsteini: Myers (2011). American Mu- seum Novitates 3715: 1-33. Species placed in new genus from Rhadinaea. Rhadinella kanalchutchan: Myers (2011). American Museum Novitates 3715: 1-33. Species placed in new genus from Rhadinaea. Rhadinella kinkelini: Myers (2011). American Mu- seum Novitates 3715: 1-33. Species placed in new genus from Rhadinaea. amphibian-reptile-conservation.org 09 June 2013 I Volume 7 | Number 1 | e61 Conservation reassessment of Mexican reptiles Barisia ciliaris. The widespread Sierra alligator lizard is endemic to Mexico, and is part of a complex that still is undergoing system- atic study. Its distribution extends along the Sierra Madre Occidental from southern Chihuahua southward through western Durango and into central Jalisco, and thence into northern Guanajuato and central Queretaro and northward in the Sierra Madre Oriental to central Nuevo Leon. Its EVS has been calculated as 15, placing it in the lower portion of the high vulnerability category. The IUCN does not recognize this taxon at the species level, so it has to be considered as Not Evaluated. This individual is from 10. 1 km WNW of La Congoja, Aguascalientes. Photo by Louis W. Porras. Lampropeltis mexicana. The endemic Mexican gray-banded kingsnake is distributed from the Sierra Madre Occidental in southern Durango and the Siena Madre Oriental in extreme southeastern Coahuila southward to northern Guanajuato. Its EVS has been gauged as 15, placing it in the lower portion of the high vulnerability category, but its IUCN status, however, was determined as of Least Concern. This individual was found at Banderas de Aguila (N of Coyotes), Durango. Photo by Ed Cassano. amphibian-reptile-conservation.org 010 June 2013 I Volume 7 | Number 1 | e61 Wilson et al. RhadineUa lachrymans : Myers (2011). American Mu- seum Novitates 3715: 1-33. Species placed in new genus from Rhadinaea. RhadineUa posadasi: Myers (2011). American Mu- seum Novitates 3715: 1-33. Species placed in new genus from Rhadinaea. RhadineUa schistosa : Myers (2011). American Mu- seum Novitates 3715: 1-33. Species placed in new genus from Rhadinaea. Sonora aemula: Cox et al. (2012). Systematic s and Biodiversity 10: 93-108. Generic name changed from Procinura, which thus becomes a synonym of Sonora. Epictia goudotii: Adalsteinsson et al. (2009). Zootaxa 2244: 1-50. Species placed in a new genus from Leptotyphlops. Rena boettgeri: Adalsteinsson et al. (2009). Zootaxa 2244: 1-50. Species placed in a new genus from Leptotyphlops. Rena bressoni: Adalsteinsson et al. (2009). Zootaxa 2244: 1-50. Species placed in a new genus from Leptotyphlops. Rena dissecta: Adalsteinsson et al. (2009). Zootaxa 2244: 1-50. Species placed in a new genus from Leptotyphlops. Rena dulcis: Adalsteinsson et al. (2009). Zootaxa 2244: 1-50. Species placed in a new genus from Leptotyphlops. Rena humilis: Adalsteinsson et al. (2009). Zootaxa 2244: 1-50. Species placed in a new genus from Leptotyphlops. Rena maxima : Adalsteinsson et al. (2009). Zootaxa 2244: 1-50. Species placed in a new genus from Leptotyphlops. Rena myopica: Adalsteinsson et al. (2009). Zootaxa 2244: 1-50. Species placed in a new genus from Leptotyphlops. Mixcoatlus barbouri: Jadin et al. (2011). Zoological Journal of the Linnean Society 163: 943-958. New genus for species removed from Cerrophidion. Mixcoatlus melanurus: Jadin et al. (2011). Zoological Journal of the Linnean Society 163: 943-958. New genus for species removed from Ophryacus. Results of the 2005 Mexican Reptile Assessment The 2005 Mexican Reptile Assessment “was carried out by zoologists from the non-profit conservation group NatureServe, working in partnership with reptile ex- perts from universities, the World Conservation Union (IUCN), and Conservation International” (NatureServe Press Release; available at natureserve.org/aboutUS/ PressReleases). This study dealt with “721 species of lizards and snakes found in Mexico, the United States, and Canada.” Turtles and crocodilians previously were assessed. The press release indicated that, “about one amphibian-reptile-conservation.org in eight lizards and snakes (84 species) were found to be threatened with extinction [i.e., judged as Critically Endangered, Endangered, or Vulnerable], with another 23 species labeled Near Threatened. For 121 lizards and snakes, the data are insufficient to allow a confident es- timate of their extinction risk [i.e., judged as Data Defi- cient], while 493 species (about two-thirds of the total) are at present relatively secure [i.e., judged as Least Con- cern].” Thus, the percentages of species that fall into the standard IUCN assessment categories are as follows: CR, EN, and VU (11.7); NT (3.2); DD (16.8); and LC (68.4). Inasmuch as the above results include species that occur in the United States, Canada, and also those not evaluated in the survey, we extracted information from the IUCN Red List website on the ratings provided for Mexican species alone, and also used the “NE” designa- tion for species not included in the 2005 assessment. We list these ratings in Appendix 1 . Critique of the 2005 Results Our primary reason for writing this paper is to critique the results of the Mexican reptile assessment, as reported in the above press release, and to reassess the conserva- tion status of these organisms using another conserva- tion assessment tool. We begin our critique with the data placed in Appendix 1, which we accessed at the IUCN Red List website up until 26 May 2012. The taxa listed in this appendix are current to the present, based on the changes to the Mexican reptile fauna indicated above. The data on the IUCN ratings are summarized by family in Table 1 and discussed below. We based our examination on the understanding that the word “critique” does not necessarily imply an unfa- vorable evaluation of the results of the Mexican reptile assessment, as conducted using the IUCN categories and criteria. “Critique,” in the strict sense, implies neither praise nor censure, and is neutral in context. We under- stand, however, that the word sometimes is used in a neg- ative sense, as noted in the 3 rd edition of The American Heritage Dictionary (1992: 443). Nonetheless, our usage simply means to render a careful analysis of the results. Presently, we recognize 849 species of reptiles in Mexico, including three crocodilians, 48 turtles, 413 liz- ards and amphisbaenians, and 385 snakes, arrayed in 42 families. This total represents an increase of 19 species (14 lizards, five snakes) over the totals listed by Wilson and Johnson (2010). The number and percentage of each of these 849 species allocated to the IUCN categories, or not evaluated, are as follows: CR = 9 (1.1%); EN = 38 (4.5%); VU = 45 (5.3%); NT = 26 (3.1%); LC = 424 (49.9%); DD = 118 (13.9%); and NE (not evaluated) = 189 (22.2%). The number and percentage of species col- lectively allocated to the three threat categories (CR, EN, and VU) are 92 and 10.8%, respectively. This number is exceeded by the 118 species placed in the DD category, and is slightly less than one-half of the 189 species not June 2013 I Volume 7 | Number 1 | e61 011 Conservation reassessment of Mexican reptiles Anolis dollfusianus . The coffee anole is distributed on the Pacific versant from southern Chiapas to western Guatemala. Its EVS has been determined as 13, placing it at the upper end of the medium vulnerability category, and its IUCN status is undetermined. This individual was found in cloud forest in Reserva de la Biosfera El Triunfo, Chiapas. Photo by Eli Garcia-Padilla. amphibian-reptile-conservation.org 012 June 2013 I Volume 7 | Number 1 | e61 Table 1 . IUCN Red List categorizations for the Mexican reptile families (including crocodilians, turtles, lizards, and snakes). Families Number of species IUCN Red List categorizations Critically Endangered Endangered Vulnerable Near Threatened Least Concern Data Deficient Not Evaluated Alligatoridae 1 — — — — 1 — — Crocodylidae 2 — — 1 — 1 — — Subtotals 3 — — 1 — 2 — — Cheloniidae 5 2 2 1 — — — — Chelydridae 1 — — 1 — — — — Dermatemydidae 1 1 — — — — — — Dermochelyidae 1 1 — — — — — — Emydidae 15 — 2 4 2 2 1 4 Geoemydidae 3 — — — 2 — — 1 Kinosternidae 17 — — — 6 6 3 2 Testudinidae 3 — — 1 — 1 — 1 Trionychidae 2 — — — — 1 — 1 Subtotals 48 4 4 7 10 10 4 9 Biporidae 3 — — — — 3 — — Anguidae 48 — 10 4 1 17 10 6 Anniellidae 2 — 1 — — 1 — — Corytophanidae 6 — — — — 1 — 5 Crotaphytidae 10 — 1 1 8 Dactyloidae 50 — 3 2 — 16 12 17 Dibamidae 1 — — — — 1 — — Eublepharidae 7 — — — — 6 — 1 Gymnophthalmi- dae 1 — — — — — — 1 Helodermatidae 2 — — — 1 1 — — Iguanidae 19 1 — 2 2 3 — 11 Mabuyidae 1 — — — — — — 1 Phrynosomatidae 135 1 5 8 6 89 6 20 Phyllodactylidae 15 — — — 1 10 1 3 Scincidae 23 — — 1 — 12 5 5 Sphaerodactylidae 4 — — — — — — 4 Sphenomorphidae 6 — — — — 3 — 3 Teiidae 46 — — 3 1 35 2 5 Xantusiidae 25 — 1 2 — 6 8 8 Xenosauridae 9 — 2 1 — 2 1 3 Subtotals 413 2 23 24 12 214 45 93 Boidae 2 — — — — 1 — 1 Colubridae 136 2 3 1 3 77 18 32 Dipsadidae 115 — 3 3 — 44 38 27 Elapidae 19 — — 1 — 13 4 1 Leptotyphlopidae 8 — — — — 5 1 2 Loxocemidae 1 — — — — — — 1 Natricidae 33 — 2 3 — 20 3 5 Typhlopidae 2 — — — — 2 — — Ungaliophiidae 2 — — 1 — — — 1 Viperidae 59 1 3 4 1 33 4 13 Xenodontidae 8 — — — — 3 1 4 Subtotals 385 3 11 13 4 198 69 87 Totals 849 9 38 45 26 424 118 189 amphibian-reptile-conservation.org 013 June 2013 I Volume 7 I Number 1 I e61 Conservation reassessment of Mexican reptiles Mastigodrycis cliftoni. The endemic Clifton’s lizard eater is found along the Pacific versant from extreme southeastern Sonora southward to Jalisco. Its EVS has been determined as 14, placing it at the lower end of the high vulnerability category, and its IUCN status has not been assessed. This individual is from El Carrizo, Sinaloa. Photo by Ed Cassano. Geophis dugesi. The endemic Duges’ earthsnake occurs from extreme southwestern Chihuahua along the length of the Sierra Madre Occidental southward to Michoacan. Its EVS has been assessed as 13, placing it at the upper end of the medium vulner- ability category, and its IUCN status has been determined as of Least Concern. This individual was found at El Carrizo, Sinaloa. Photo by Ed Cassano. amphibian-reptile-conservation.org 014 June 2013 | Volume 7 | Number 1 | e61 Wilson et al. evaluated on the website. Thus, of the total of 849 spe- cies, 307 (36.2%) are categorized either as DD or NE. As a consequence, only 542 (63.8%) of the total number are allocated to one of the other five categories (CR, EN, VU, NT, or LC). These results provided us with a substantially in- complete picture of the conservation status of reptiles in Mexico, which sharply contrasts the picture offered for Central American reptiles (the other major portion of Mesoamerica), as recorded in Wilson et al. (2010). This situation is underscored by the relatively low spe- cies numbers of Mexican reptiles placed in any of the three IUCN threat categories. In addition, a substantial proportion (13.9%) of the Mexican species are assessed as DD, indicating that insufficient information exists for the IUCN rating system to be employed. Finally, 189 species (22.3%) are not evaluated, largely because they also occur in Central America (and in some cases, also in South America) and will be assessed presumably in future workshops, which was the case for most of these species when they were assessed in a Central American workshop held on May 6-10, 2012; as yet, the results of that assessment are not available. Given that only 10.8% of the Mexican species were allocated to one of the three IUCN threat categories and that about six in 10 species in the country are en- demic, we examined the IUCN ratings reported for spe- cies inhabiting five of the countries in Central America (see Wilson et al. 2010). For Guatemala, Acevedo et al. (2010) reported that 56 reptile species (23.0%) of a total of 244 then recognized were assigned to one of the three threat categories. Of 237 Honduran reptiles assessed by Townsend and Wilson (2010), 74 (31.2%) were placed in one of the threat categories. Sunyer and Kohler (2010) listed 165 reptile species from Nicaragua, a country with only three endemic reptiles known at the time, but judged 10 of them (6.1%) as threatened. Of 231 reptile species assessed by Sasa et al. (2010) for Costa Rica, 36 (15.6%) were placed in a threat category. Finally, Jaramillo et al. (2010) placed 22 of 248 Panamanian reptile species (8.9%) in the threat categories. Collectively, 17% of the reptile species in these countries were assessed in one of the three threat categories. The number of species in Central America placed into one of the threat categories apparently is related to the number allocated to the DD category. Although the DD category is stated explicitly as a non-threat category (IUCN Red Fist Categories and Criteria 2010), its use highlights species so poorly known that one of the other IUCN categories cannot be applied. The percentage of DD species in the reptile faunas of each of the five Cen- tral American countries discussed above ranges from 0.9 in Honduras to 40.3 in Panama. Intermediate figures are as follows: Nicaragua = 1.2; Guatemala = 5.3; Costa Rica = 34.2. These data apparently indicate that the conser- vation status of the Costa Rican and Panamanian reptile faunas are by far more poorly understood than those of Guatemala, Honduras, and Nicaragua. The length of time for placing these DD species into another category is unknown, but a reassessment must await targeted surveys for the species involved. Given the uncertainty implied by the use of this category sup- plemented by that of NE species in Mexico, we believe there is ample reason to reassess the conservation status of the Mexican reptiles using the Environmental Vulner- ability Score (EVS). EVS for Mexican Reptiles The EVS provides several advantages for assessing the conservation status of amphibians and reptiles. First, this measure can be applied as soon as a species is described, because the information necessary for its application generally is known at that point. Second, the calculation of the EVS is an economical undertaking and does not require expensive, grant-supported workshops, such as those held in connection with the Global Reptile Assess- ment sponsored by the IUCN. Third, the EVS is predic- tive, because it provides a measure of susceptibility to anthropogenic pressure, and can pinpoint taxa in need of immediate attention and continuing scrutiny. Finally, this measure is simple to calculate and does not “penalize” species that are poorly known. One disadvantage of the EVS, however, is that it was not designed for use with marine species. So, the six species of marine turtles and two of marine snakes occurring on the shores of Mexico could not be assessed. Nevertheless, given the increas- ing rates of human population growth and environmental deterioration, an important consideration for a given spe- cies is to have a conservation assessment measure that can be applied simply, quickly, and economically. We calculated the EVS for each of the 841 species of terrestrial reptiles occurring in Mexico (Wilson and Johnson 2010, and updated herein; see Appendix 1). In this appendix, we listed the scores alongside the IUCN categorizations from the 2005 Mexican Reptile Assess- ment, as available on the IUCN Red List website (www. iucnredlist.org) and as otherwise determined by us (i.e., as NE species). Theoretically, the EVS can range from 3 to 20. A score of 3 is indicative of a species that ranges widely both within and outside of Mexico, occupies eight or more forest formations, and is fossorial and usually escapes human notice. Only one such species (the leptotyphlo- pid snake Epictia goudotii) is found in Mexico. At the other extreme, a score of 20 relates to a species known only from the vicinity of the type locality, occupies a single forest formation, and is exploited commercially or non-commercially for hides, meat, eggs and/or the pet trade. Also, only one such species (the trionychid turtle Ap alone atra ) occurs in Mexico. All of the other scores fall within the range of 4-19. We summarized the EVS for reptile species in Mexico by family in Table 2. June 2013 I Volume 7 | Number 1 | e61 amphibian-reptile-conservation.org 015 Conservation reassessment of Mexican reptiles Rhadinaea laureata. The endemic crowned graceful brownsnake is distributed along the Sierra Madre Occidental from west-central Durango southward into the Tran verse Volcanic Axis as far as central Michoacan, Morelos, and the Distrito Federal. Its EVS has been calculated as 12, placing it in the upper portion of the medium vulnerability category, and its IUCN status has been determined as Least Concern. This individual is from Rancho Las Canoas, Durango. Photo by Louis W. Porras. Thamnophis mendax. The endemic Tamaulipan montane gartersnake is restricted to a small range in the Sierra Madre Oriental in southwestern Tamaulipas. Its EVS has been determined as 14, placing it at the lower end of the high vulnerability category, and its IUCN status has been assessed as Endangered. This individual came from La Gloria, in the Gomez Farias region of Tamaulipas. Photo by Ed Cassano. amphibian-reptile-conservation.org 016 June 2013 I Volume 7 | Number 1 | e61 Wilson et al. Table 2. Environmental Vulnerability Scores for the Mexican reptile species (including crocodilians, turtles, lizards, and snakes, but excluding the marine species), arranged by family. Shaded area to the left encompasses low vulnerability scores, and to the right high vulnerability scores. Families Number of species Environmental Vulnerability Scores 8 10 11 12 13 14 15 16 17 18 19 20 Alligatoridae 1 Crocodylidae Subtotals Subtotal % 33.3 33.3 33.3 Chelydridae Dermatemydi- dae Emydidae 15 Geoemydidae Kinosternidae 17 Testudinidae Trionychidae Subtotals 42 Subtotal % 2.4 7.1 2.4 2.4 7.1 19.0 14.3 9.5 7.1 11.9 14.3 2.4 Bipedidae Anguidae 48 11 Anniellidae Corytophani- dae Crotaphyti- dae 10 Dactyloidae 50 15 Dibamidae Eublephari- dae Gymnoph- thalmidae Heloderma- tidae Iguanidae 19 Mabuyidae Phrynosoma- tidae 135 11 18 22 16 23 23 11 Phyllodactyli- dae 15 Scincidae 23 Sphaerodac- tylidae Sphenomor- phidae Teiidae 46 14 Xantusiidae 25 Xenosauridae Subtotals 413 11 13 14 28 39 49 54 67 78 38 10 Subtotal % Boidae 0.2 0.7 1.5 2.7 3.1 3.4 6.8 9.4 11.9 13.1 16.2 18.9 9.2 2.4 0.5 amphibian-reptile-conservation.org 017 June 2013 I Volume 7 I Number 1 I e61 Conservation reassessment of Mexican reptiles Table 2. Continued. Colubridae 136 — — 4 7 3 6 10 15 8 8 18 22 14 16 5 — — — Dipsadidae 115 — 1 3 3 3 8 4 7 6 13 14 13 19 15 6 — — — Elapidae 17 — — — — — 2 — — 2 — 2 2 3 — 2 3 1 — Leptotyphlo- pidae 8 1 — — — — 1 — — 2 — 2 2 — — — — — — Loxocemidae 1 1 Natricidae 33 — — — — 3 1 — 2 2 2 3 6 7 4 2 1 — — Typhlopidae 2 1 1 Ungaliophi- idae 2 — — — — — — — 1 — — — — 1 — — — — — Viperidae 59 — — — — — 1 2 1 3 7 5 6 6 9 8 5 6 — Xenodontidae 8 — — — — — — 1 1 1 — 3 1 — — 1 — — — Subtotals 383 1 1 7 10 9 19 17 30 25 31 47 52 50 44 24 9 7 — Subtotal % — 0.3 0.3 1.8 2.6 2.3 5.0 4.4 7.8 6.5 8.1 12.3 13.6 13.1 11.5 6.3 2.3 1.8 — Totals 841 1 1 8 13 15 31 30 47 54 71 100 115 123 127 65 24 15 1 Total % — 0.1 0.1 1.0 1.5 1.8 3.7 3.6 5.6 6.4 8.4 11.9 13.7 14.6 15.1 7.7 2.9 1.8 0.1 The range and average EVS for the major reptile groups are as follows: crocodilians = 13-16 (14.3); tur- tles = 8-20 (15.3); lizards = 5-19 (13.8); and snakes = 3-19 (12.8). On average, turtles are most susceptible and snakes least susceptible to environmental degradation, with lizards and crocodilians falling in between. The av- erage scores either are at the upper end of the medium category, in the case of snakes and lizards, or at the lower end of the high category, in the case of crocodilians and turtles. The average EVS for all the reptile species is 13.4, a value close to the lower end of the high range of vulnerability. Nineteen percent of the turtle species were assigned an EVS of 14, at the lower end of the high vulnerability category. For lizards, the respective figures are 18.9% and 16, about midway through the range for the high vul- nerability category; for snakes, the values are 13.6% and 14. The total EVS values generally increase from the low end of the scale (3) to about midway through the high end (16), with a single exception (a decrease from 31 to 29 species at scores 8 and 9), then decrease thereafter to the highest score (20). The peak number of taxa (127) was assigned an EVS of 16, a score that falls well within the range of high vulnerability. Of the 841 total taxa that could be scored, 99 (11.8%) fall into the low vulnerability category, 272 (32.3%) in the medium category, and 470 (55.9%) in the high cat- egory. Thus, more than one-half of the reptile species in Mexico were judged as having the highest degree of vulnerability to environmental degradation, and slightly more than one-tenth of the species the lowest degree. This increase in absolute and relative numbers from the low portion, through the medium portion, to the high portion varies somewhat with the results published for both the amphibians and reptiles for some Central Amer- ican countries (see Wilson et al. 2010). Acevedo et al. (2010) reported 89 species (23.2%) with low scores, 179 (46.7%) with medium scores, and 115 (30.0%) with high scores for Guatemala. The same trend is seen in Hon- duras, where Townsend and Wilson (2010) indicated the following absolute and relative figures in the same order, again for both amphibians and reptiles: 71 (19.7%); 169 (46.8%); and 121 (33.5%). Comparable figures for the Panamanian herpetofauna (Jaramillo et al. 2010) are: 143 (33.3%); 165 (38.4%); and 122 (28.4%). The principal reason that EVS values are relatively high in Mexico is because of the high level of endemism and the relatively narrow range of habitat occurrence. Of the 485 endemic species in Mexico (18 turtles, 264 lizards, 203 snakes), 124 (25.6%) were assigned a geo- graphic distribution score of 6, signifying that these crea- tures are known only from the vicinity of their respective type localities; the remainder of the endemic species (361 [74.4%]) are more broadly distributed within the country (Appendix 1). Of the 841 terrestrial Mexican reptile spe- cies, 212 (25.2%) are limited in ecological distribution to one formation (Appendix 1). These features of geograph- ic and ecological distribution are of tremendous signifi- cance for efforts at conserving the immensely important Mexican reptile fauna. Comparison of IUCN Categorizations and EVS Values Since the IUCN categorizations and EVS values both measure the degree of environmental threat impinging on a given species, a certain degree of correlation between the results of these two measures is expected. Townsend and Wilson (2010) demonstrated this relationship with reference to the Honduran herpetofauna, by comparing the IUCN and EVS values for 362 species of amphibians and terrestrial reptiles in their table 4. Perusal of the data in this table indicates, in a general way, that an increase in amphibian-reptile-conservation.org 018 June 2013 I Volume 7 | Number 1 | e61 Wilson et al Crotalus catalinensis . The endemic Catalina Island rattlesnake is restricted in distribution to Santa Catalina Island in the Gulf of California. Its EVS has been determined as 19, placing it in the upper portion of the high vulnerability category, and its IUCN status as Critically Endangered. Photo by Louis W. Porras. Crotalus stejnegeri. The endemic Sinaloan long-tailed rattlesnake is restricted to a relatively small range in western Mexico, where it is found in the western portion of the Sierra Madre Occidental in western Durango and southeastern Sinaloa. Its EVS has been determined as 17, placing it in the middle of the high vulnerability category, and its IUCN status as Vulnerable. This individual came from Plomosas, Sinaloa. Photo by Louis W. Porras. amphibian-reptile-conservation.org 019 June 2013 | Volume 7 | Number 1 | e61 Conservation reassessment of Mexican reptiles Table 3. Comparison of the Environmental Vulnerability Scores (EVS) and IUCN categorizations for terrestrial Mexican reptiles. Shaded area on top encompasses the low vulnerability category scores, and at the bottom high vulnerability category scores. EVS IUCN categories Totals Critically Endangered Endangered Vulnerable Near Threatened Least Concern Data Deficient Not Evaluated 3 — — — — — — 1 1 4 — — — — 1 — — 1 5 — — — — 3 — 5 8 6 — — — — 5 — 8 13 7 — — — — 5 — 10 15 8 — — — — 20 — 11 31 9 — — 1 — 16 — 13 30 10 — — — — 25 1 21 47 11 — — 1 1 36 2 14 54 12 — 1 1 — 49 4 16 71 13 — 2 5 3 66 5 19 100 14 — 5 6 8 65 15 16 115 15 — 13 11 7 54 25 13 123 16 — 8 3 6 48 38 24 127 17 4 3 11 1 21 14 11 65 18 — 2 2 — 4 12 4 24 19 2 2 3 — 4 2 2 15 20 — — — — — — 1 1 Totals 6 36 44 26 422 118 189 841 EVS values is associated with a corresponding increase in the degree of threat, as measured by the IUCN catego- rizations. If average EVS values are determined for the IUCN categories in ascending degrees of threat, the fol- lowing figures result: LC (206 spp.) =10.5; NT (16 spp.) = 12.9; VU (18 spp.) = 12.5; EN (64 spp.) = 14.1; CR (50 spp.) = 15.1; and EX (2 spp.) = 16.0. The broad cor- respondence between the two measures is evident. Also of interest is that the average EVS score for the six DD species listed in the table is 13.7, a figure closest to that for the EN category (14.1), which suggests that if and when these species are better known, they likely will be judged as EN or CR. In order to assess whether such a correspondence ex- ists between these two conservation measures for the Mexican reptiles, we constructed a table (Table 3) simi- lar to table 4 in Townsend and Wilson (2010). Important similarities and differences exist between these tables. The most important similarity is in general appearance, i.e., an apparent general trend of decreasing EVS values with a decrease in the degree of threat, as indicated by the IUCN categorizations. This similarity, however, is more apparent than real. Our Table 3 deals only with Mexi- can reptiles, excludes the IUCN category EX (because presently this category does not apply to any Mexican species), and includes a NE category that we appended to the standard set of IUCN categories. Apart from these obvious differences, however, a closer examination of the data distribution in our Table 3 reveals more signifi- cant differences in the overall picture of the conserva- tion status of the Mexican reptiles when using the IUCN categorizations, as opposed to the EVS, especially when viewed against the backdrop of results in Townsend and Wilson (2010: table 4). 1. Nature of the IUCN categorizations in Table 3 Unlike the Townsend and Wilson (2010) study, we in- troduced another category to encompass the Mexican reptile species that were not evaluated in the 2005 IUCN study. The category is termed “Not Evaluated” (IUCN 2010) and a large proportion of the species (189 of 841 Mexican terrestrial reptiles [22.5%]) are placed in this category. Thus, in the 2005 study more than one-fifth of the species were not placed in one of the standard IUCN categories, leaving their conservation status as undeter- mined. In addition, a sizable proportion of species (118 [14.0%]) were placed in the DD category, meaning their conservation status also remains undetermined. When the species falling into these two categories are added, evidently 307 (36.5%) of the 841 Mexican terrestrial rep- tiles were not placed in one of the IUCN threat assess- ment categories in the 2005 study. This situation leaves less than two-thirds of the species as evaluated. amphibian-reptile-conservation.org 020 June 2013 I Volume 7 | Number 1 | e61 Wilson et al. Xantusia sanchezi ■ The endemic Sanchez’s night lizard is known only from extreme southwestern Zacatecas to central Jalisco. This lizard’s EVS has been assessed as 16, placing it in the middle of the high vulnerability category, but its IUCN status has been deter- mined as Least Concern. This individual was discovered at Huaxtla, Jalisco. Photo by Daniel Cruz-Sdenz. amphibian-reptile-conservation.org 021 June 2013 I Volume 7 | Number 1 | e61 Conservation reassessment of Mexican reptiles 2. Pattern of mean EVS vs. IUCN categoriza- tions In order to more precisely determine the relationship be- tween the IUCN categorizations and the EVS, we calcu- lated the mean EVS for each of the IUCN columns in Ta- ble 3, including the NE species and the total species. The results are as follows: CR (6 spp.) = 17.7 (range 17-19); EN (36 spp.) = 15.4 (12-19); VU (44 spp.) = 15.3 (10- 19); NT (26 spp.) = 14.6 (11-17); LC (422 spp.) = 13.0 (4-19); DD (118 spp.) = 15.5 (10-19); and NE (189 spp.) = 12.0 (3-20); and Total (841 spp.) = 13.3 (3-20). The results of these data show that the mean EVS decreases from the CR category (17.7) through the EN category (15.4) to the VU category (15.3), but only slightly be- tween the EN and VU categories. They also continue to decrease from the NT category (14.6) to the LC category (13.0). This pattern of decreasing values was expected. In addition, as with the Townsend and Wilson (2010) Honduran study, the mean value for the DD species (15.5) is closest to that for the EN species (15.4). To us, this indicates what we generally have suspected about the DD category, i.e., that the species placed in this category likely will fall either into the EN or the CR categories when (and if) their conservation status is better under- stood. Placing species in this category is of little benefit to determining their conservation status, however, since once sequestered with this designation their significance tends to be downplayed. This situation prevailed once the results of the 2005 assessment were reported, given that the 1 1 8 species evaluated as DD were ignored in favor of the glowing report that emerged in the NatureServe press release (see above). If the data in Table 3 for the DD spe- cies is conflated with that for the 86 species placed in one of the three threat categories, the range of EVS values represented remains the same as for the threat categories alone, i.e., 10-19, and the mean becomes 15.5; the same as that indicated above for the DD species alone and only one-tenth of a point from the mean score for EN species (15.4). On the basis of this analysis, we predict that if a concerted effort to locate and assess the 118 DD spe- cies were undertaken, that most or all of them would be shown to qualify for inclusion in one of the three IUCN threat categories. If that result were obtained, then the number of Mexican reptile species falling into the IUCN threat categories would increase from 86 to 204, which would represent 24.3% of the reptile fauna. Based on the range and mean of the EVS values, the pattern for the LC species appears similar to that of the NE species, as the ranges are 4-19 and 3-20 and the means are 13.0 and 12.0, respectively. If these score dis- tributions are conflated, then the EVS range becomes the broadest possible (3-20) and the mean becomes 12.7, which lies close to the upper end of the medium vulner- ability category. While we cannot predict what would happen to the NE species once they are evaluated (pre- sumably most species were evaluated during the Central amphibian-reptile-conservation.org American reptile assessment of May, 2012), because they were evaluated mostly by a different group of herpetolo- gists from those present at the 2005 Mexican assessment, we suspect that many (if not most) would be judged as LC species. A more discerning look at both the LC and NE species might demonstrate that many should be par- titioned into other IUCN categories, rather than the LC. Our reasoning is that LC and NE species exhibit a range of EVS values that extend broadly across low, medium, and high categories of environmental vulnerability. The number and percentage of LC species that fall into these three EVS categories are as follows: Low (range 3-9) = 50 spp. (11.8%); Medium (10-13) = 176 (41.7); and High (14-20) = 196 (46.5). For the NE species, the fol- lowing figures were obtained: Low = 48 (25.8); Medium = 68 (36.6); and High = 70 (37.6). The percentage values are reasonably similar to one another, certainly increas- ing in the same direction from low through medium to high. Considering the total number of species, 99 (11.8%) fall into the low vulnerability category, 272 (32.3%) into the medium vulnerability category, and 470 (55.9%) into the high vulnerability category. These results differ sig- nificantly from those from the 2005 study. If the three IUCN threat categories can be considered most compa- rable to the high vulnerability EVS category, then 86 spe- cies fall into these three threat categories, which is 16.1% of the total 534 species in the CR, EN, VU, NT, and LC categories. If the NT category can be compared with the medium vulnerability EVS category, then 26 species fall into this IUCN category (4.9% of the 534 species). Fi- nally, if the LC category is comparable to the low vul- nerability EVS category, then 422 species (79.0%) fall into this IUCN category. Clearly, the results of the EVS analysis are nearly the reverse of those obtained from the IUCN categorizations discussed above. Discussion In the Introduction we indicated that our interest in con- ducting this study began after the publication of Wilson et al. (2010), when we compared the data resulting from that publication with a summary of the results of a 2005 Mexican reptile assessment conducted under the aus- pices of the IUCN, and later referenced in a 2007 press release by NatureServe, a supporter of the undertaking. Our intention was not to critique the IUCN system of conservation assessment (i.e., the well-known IUCN cat- egorizations), but rather to critique the results of the 2005 assessment. We based our critique on the use of the En- vironmental Vulnerability Score (EVS), a measure used by Wilson and McCranie (2004) and in several Central American chapters in Wilson et al. (2010). Since the IUCN assessment system uses a different set of criteria than the EVS measure, we hypothesized that the latter could be used to test the results of the for- mer. On this basis, we reassessed the conservation status June 2013 I Volume 7 | Number 1 | e61 022 Wilson et al. of the reptiles of Mexico, including, by our definition of convenience, crocodilians, turtles, lizards, and snakes, by determining the EVS value for each terrestrial spe- cies (since the measure was not designed for use with marine species). Based on our updating of the species list in Wilson and Johnson (2010), our species list for this study consisted of 841 species. We then developed an EVS measure applicable to Mexico, and employed it to calculate the scores indicated in Appendix 1 . Our analysis of the EVS values demonstrated that when the scores are arranged in low, medium, and high vulnerability categories, the number and percentage of species increases markedly from the low category, through the medium category, to the high category (Ta- ble 2). When these scores (Table 3) are compared to the IUCN categorizations documented in Table 1, however, an inverse correlation essentially exists between the re- sults obtained from using the two methods. Since both methods are designed to render conservation status as- sessments, the results would be expected to corroborate one another. We are not in a position to speculate on the reason(s) for this lack of accord, and simply are offering a reassess- ment of the conservation status of Mexico’s reptile spe- cies based on another measure (EVS) that has been used in a series of studies since it was introduced by Wilson and McCranie (1992), and later employed by McCranie and Wilson (2002), Wilson and McCranie (2004), and several chapters in Wilson et al. (2010). Nonetheless, we believe our results provide a significantly better assess- ment of the conservation status Mexico’s reptiles than those obtained in the 2005 IUCN assessment. We con- sider our results more consonant with the high degree of reptile endemism in the country, and the restricted geo- graphic and ecological ranges of a sizable proportion of these species. We also believe that our measure is more predictive, and reflective of impact expected from con- tinued habitat fragmentation and destruction in the face of continuing and unregulated human population growth. Conclusions and Recommendations Our conclusions and conservation recommendations draw substantially from those promulgated by Wilson and Townsend (2010), which were provided for the en- tire Mesoamerican herpetofauna; thus, we refined them specifically for the Mexican reptile fauna, as follows: 1. In the introduction we noted the human population size and density expected for Mexico in half a cen- tury, and no indication is available to suggest that this trend will be ameliorated. Nonetheless, although 66% of married women in Mexico (ages 15-49) use modern methods of contraception, the current fertility rate (2.3) remains above the replacement level (2.0) and 29% of the population is below the age of 15, 1% above the average for Latin America and the Carib- bean (2011 PRB World Population Data Sheet). 2. Human population growth is not attuned purposefully to resource availability, and the rate of regeneration depends on the interaction of such societal factors as the level of urbanization; in Mexico, the current level is 78%, and much of it centered in the Distrito Fed- eral (2011 PRB World Population Data Sheet). This statistic is comparable to that of the United States (79%) and Canada (80%), and indicates that 22% of Mexico’s population lives in rural areas. Given that the level of imports and exports are about equal (in 2011, imports = 350.8 billion US dollars, exports = 349.7 billion; CIA World Factbook 2012), the urban population depends on the basic foodstuffs that the rural population produces. An increase in human pop- ulation demands greater agricultural production and / or efficiency, as well as a greater conversion of wild lands to farmlands. This scenario leads to habitat loss and degradation, and signals an increase in biodiver- sity decline. 3. Although the rate of conversion of natural habitats to agricultural and urban lands varies based on the meth- ods and assumptions used for garnering this determi- nation, most estimates generally are in agreement. The Secretarfa del Medio Ambiente y Recursos Na- turales (SEMARNAT; Secretariat of Environment and Natural Resources; semarnat.gob.mx) has attempted to measure the rate of deforestation from 1978 to 2005, with estimates ranging from about 200,000 to 1.500.000 ha/yr. Most estimates, however, range from about 200,000 to 400,000 ha/yr. A study conducted for the years 2000 to 2005 reported an average rate of 260.000 ha/yr. Another source of information (www. rainforests.mongabay.com) reports that from 1990 to 2010 Mexico lost an average of 274,450 ha (0.39% of the total 64,802,000 ha of forest in the country), and during that period lost 7.8% of its forest cover (ca. 5.489.000 ha). No matter the precise figures for for- est loss, this alarming situation signifies considerable endangerment for organismic populations, including those of reptiles. About one-third of Mexico is (or was) covered by forest, and assuming a constant rate of loss all forests would be lost in about 256 years (starting from 1990), or in the year 2246. Forest loss in Mexico, therefore, contributes significantly to the global problem of deforestation. 4. As a consequence, no permanent solution to the prob- lem of biodiversity decline (including herpetofaunal decline) will be found in Mexico (or elsewhere in the world) until humans recognize overpopulation as the major cause of degradation and loss of humankind’s fellow organisms. Although this problem is beyond the scope of this investigation, solutions will not be June 2013 I Volume 7 | Number 1 | e61 amphibian-reptile-conservation.org 023 Conservation reassessment of Mexican reptiles available until humanity begins to realize the origin, nature, and consequences of the mismatch between human worldviews and how our planet functions. Wil- son (1988) labeled this problem “the mismanagement of the human min d.” Unfortunately, such realignment is only envisioned by a small cadre of humans, so crafting provisional solutions to problems like biodi- versity decline must proceed while realizing the ul- timate solution is not available, and might never be. 5. Mexico is the headquarters of herpetofaunal diver- sity and endemism in Mesoamerica, which supports the conclusions of Ochoa-Ochoa and Flores-Villela (2006), Wilson and Johnson (2010), and the authors of four chapters on the Mexican herpetofauna in Wil- son et al. (2010). Furthermore, field research and sys- tematic inquiry in Mexico will continue to augment the levels of diversity and endemicity, which are of immense conservation significance because reptiles are significant contributors to the proper functioning of terrestrial and aquatic ecosystems (Gibbons et al. 2000). From a political and economic perspective, diversity and endemism are important components of Mexico’s patrimony, as well as a potential source of income from ecotourism and related activities. In- vesting in such income sources should appeal to local stakeholders, as it provides an incentive for preserv- ing natural habitats (Wilson 2011). 6. Given that the ultimate solutions to biodiversity de- cline are unlikely to be implemented in any pertinent time frame, less effective solutions must be found. Vitt and Caldwell (2009) discussed a suite of ap- proaches for preserving and managing threatened species, including the use of reserves and corridors to save habitats, undertaking captive management initiatives, and intentionally releasing individuals to establish or enlarge populations of target species. Un- questionably, preserving critical habitat is the most effective and least costly means of attempting to res- cue threatened species. Captive management is less effective, and has been described as a last-ditch effort to extract a given species from the extinction vortex (Campbell et al. 2006). Efforts currently are under- way in segments of the herpetological community to develop programs for ex situ and in situ captive man- agement of some of the most seriously threatened her- petofaunal species, but such efforts will succeed only if these species can be reproduced in captivity and reintroduced into their native habitats. In the case of Mexico, Ochoa-Ochoa, et al. (2011: 2710) comment- ed that, “given the current speed of land use change, we cannot expect to save all species from extinction, and so must decide how to focus limited resources to prevent the greatest number of extinctions,” and for amphibians proposed “a simple conservation triage method that: evaluates the threat status for 145 micro- endemic Mexican amphibian species; assesses current potential threat abatement responses derived from existing policy instruments and social initiatives; and combines both indicators to provide broad-scale con- servation strategies that would best suit amphibian micro-endemic buffered areas (AMBAs) in Mexico.” These authors concluded, however, that a quarter of the micro-endemic amphibians “urgently need field- based verification to confirm their persistence due to the small percentage of remnant natural vegetation within the AMBAs, before we may sensibly recom- mend” a conservation strategy. Their tool also should apply to Mexican reptiles, and likely would produce similar results. 7. The preferred method for preserving threatened spe- cies is to protect habitats by establishing protected areas, both in the public and private sectors. Habitat protection allows for a nearly incalculable array of re- lationships among organisms. Like most countries in the world, Mexico, has developed a governmentally established system of protected areas. Fortunately, studies have identified “critical conservation zones” (Ceballos et al. 2009), as well as gaps in their cover- age (Koleff et al. 2009). The five reserves of great- est conservation importance for reptiles are the Los Tuxtlas Biosphere Reserve, the islands of the Gulf of California in the UNESCO World Heritage Site, the Sierra Gorda Biosphere Reserve, the Tehuacan-Cui- catlan Biosphere Reserve, and the Chamela-Cuixmala Biosphere Reserve. Significantly, all of these areas are part of the UNESCO World Network of Biosphere Reserves, but attainment of this status does not guar- antee that these reserves will remain free from anthro- pogenic damage. Ceballos et al. (2009, citing Dirzo and Garcia 1992) indicated that although the Los Tuxtlas is the most important reserve in Mexico for amphibians and reptiles, a large part of its natural veg- etation has been lost. This example of deforestation is only one of many, but led Ceballos et al. (2009: 597) to conclude (our translation of the original Spanish) that, “The determination of high risk critical zones has diverse implications for conservation in Mexico. The distribution of critical zones in the entire country con- firms the problem of the loss of biological diversity is severe at the present time, and everything indicates it will become yet more serious in future decades. On the other hand, the precise identification of these zones is a useful tool to guide political decisions con- cerning development and conservation in the country, and to maximize the effects of conservation action. Clearly, a fundamental linchpin for the national con- servation strategy is to direct resources and efforts to protect the high-risk critical zones. Finally, it also is evident that tools for management of production and development, such as the land-use planning and en- vironmental impact, should be reinforced in order to June 2013 I Volume 7 | Number 1 | e61 amphibian-reptile-conservation.org 024 Wilson et al. fully comply with their function to reconcile develop- ment and conservation.” We fully support this recom- mendation. 8. Humans have developed an amazing propensity for living in an unsustainable world. Organisms only can persist on Earth when they live within their environ- mental limiting factors, and their strategy is sustain- ability, i.e., in human terms, living over the long term within one’s means, a process made allowable by or- ganic evolution. Homo sapiens is the only extant spe- cies with the capacity for devising another means for securing its place on the planet, i.e., a strategy of un- sustainability over the short term, which eventually is calculated to fail. Conservation biology exists because humans have devised this unworkable living strategy. What success it will have in curbing biodiversity loss remains to be seen. Acknowledgments. — We are grateful to the follow- ing individuals for improving the quality of this contri- bution: Irene Goyenechea, Pablo Lavm-Murcio, Julio Lemos-Espinal, and Aurelio Ramirez-Bautista. We are especially thankful to Louis W. Porras, who applied his remarkable editorial skills to our work. 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Pavon-Vazquez CJ, Garcia- Vazquez UO, Blancas- Hernandez JC, Nieto-Montes A. 2011. A new species of the Geophis sieboldi group (Squamata: Colubridae) exhibiting color pattern polymorphism from Guerre- ro, Mexico. Herpetologica 67: 332-343. Population Reference Bureau. 2011. World Population Data Sheet. Available: www.prb.org [Accessed: 27 January 2012]. Pyron RA, Burbrink FT. 2009. Systematics of the com- mon kingsnake ( Lampropeltis getula; Serpentes: Col- ubridae) and the burden of heritage in taxonomy. Zoo- taxa 2241: 22-32. Raven PH, Hassenzahl DM, Berg LR. 2009. Environ- ment (8 th edition). John Wiley & Sons, Inc. Hoboken, New Jersey, USA. Reyes- Velasco J, Mulcahy DG. 2010. Additional taxo- nomic remarks on the genus Pseudoleptodeira (Ser- pentes: Colubridae) and the phylogenetic placement of “P. uribeiC Herpetologica 66: 99-110. Sasa M, Chaves G, Porras LW. 2010. The Costa Rican herpetofauna: conservation status and future perspec- tives. Pp. 510-603 In: Conservation of Mesoameri- can Amphibians and Reptiles. Editors, Wilson LD, Townsend JH, Johnson JD. Eagle Mountain Publish- ing, LC, Eagle Mountain, Utah, USA. Stuart SN, Chanson JS, Cox NA, Young BE, Rodrigues ASL, Fischman DL, Waller RW. 2004. Status and trends of amphibian declines and extinctions world- wide. Science 306(5702): 1783-1786. Stuart SN, Chanson JS, Cox NA, Young BE. 2010. The global decline of amphibians: Current trends and fu- ture prospects. Pp. 2-15 In: Conservation of Meso- american Amphibians and Reptiles. Editors, Wilson LD, Townsend JH, Johnson JD. Eagle Mountain Pub- lishing, LC, Eagle Mountain, Utah, USA. Sunyer J, Kohler G. 2010. Conservation status of the herpetofauna of Nicaragua. Pp. 488-509 In: Conser- vation of Mesoamerican Amphibians and Reptiles. Editors, Wilson LD, Townsend JH, Johnson JD. Eagle Mountain Publishing, LC, Eagle Mountain, Utah, USA. Townsend JH, Wilson LD. 2010. Conservation of the Honduran herpetofauna: issues and imperatives. Pp. 460-487 In: Conservation of Mesoamerican Amphib- ians and Reptiles. Editors, Wilson LD, Townsend JH, Johnson JD. Eagle Mountain Publishing, LC, Eagle Mountain, Utah, USA. Vitt LJ, Caldwell JP. 2009. Herpetology (3 rd edition). Academic Press, Burlington, Maine, USA. Wake DB. 1991. Declining amphibian populations. Sci- ence 253(5022): 860. Walker JM, Cordes JE. 2011. Taxonomic implications of color pattern and meristic variation in Aspidoscelis burti burti, a Mexican whiptail lizard. Herpetological Review 42: 33-39. Wilson EO (Editor). 1988. Biodiversity. National Acad- emy Press, Washington, DC, USA. Wilson LD. 2011. Imperatives and opportunities: refor- mation of herpetology in the age of amphibian de- cline. Herpetological Review 42: 146-150. Wilson LD, Johnson JD. 2010. Distributional patterns of the herpetofauna of Mesoamerica, a biodiversity hotspot. Pp. 30-235 In: Conservation of Mesoameri- can Amphibians and Reptiles. Editors, Wilson LD, Townsend JH, Johnson JD. Eagle Mountain Publish- ing, LC, Eagle Mountain, Utah, USA. Wilson LD, McCranie JR. 1992. Status of amphibian populations in Honduras. Unpublished Report to the Task Force on Declining Amphibian Population, 15 August 1992. 14 p. Wilson LD, McCranie JR. 2004. The conservation status of the herpetofauna of Honduras. Amphibian & Rep- tile Conservation 3: 6-33. Wilson LD, Townsend JH. 2010. The herpetofauna of Mesoamerica: biodiversity significance, conservation status, and future challenges. Pp. 760-812 In: Con- servation of Mesoamerican Amph ibians and Reptiles. Editors, Wilson LD, Townsend JH, Johnson JD. Eagle Mountain Publishing, LC, Eagle Mountain, Utah, USA. Wilson LD, Townsend JH, Johnson JD. 2010. Conserva- tion of Mesoamerican Amphibians and Reptiles. Ea- gle Mountain Publishing, LC, Eagle Mountain, Utah, USA. Woolrich-Pina, GA, Smith GR. 2012. A new species of Xenosaurus from the Sierra Madre Oriental, Mexico. Herpetologica 68: 551-559. World Wildlife Fund. 2012. Living Planet Report 2012. WWF International, Gland, Switzerland. Received: 18 Feb 2013 Accepted: 24 April 2013 Published: 09 June 2013 amphibian-reptile-conservation.org 027 June 2013 I Volume 7 | Number 1 | e61 Conservation reassessment of Mexican reptiles Larry David Wilson is a herpetologist with lengthy experience in Mesoamerica, totaling six collective years (combined over the past 47). Larry is the senior editor of the recently published Conservation ofMeso- american Amphibians and Reptiles and a co-author of seven of its chapters. He retired after 35 years of service as Professor of Biology at Miami-Dade College in Miami, Florida. Larry is the author or co-author of more than 290 peer-reviewed papers and books on herpetology, including the 2004 Amphibian & Rep- tile Conservation paper entitled “The conservation status of the herpetofauna of Honduras.” His other books include The Snakes of Honduras, Middle American Herpetology, The Amphibians of Honduras, Amphibians & Reptiles of the Bay Islands and Cay os Cochinos, Honduras, The Amphibians and Reptiles of the Honduran Mosquitia, and Guide to the Amph ibians & Reptiles ofCusuco National Park, Honduras. He also served as the Snake Section Editor for the Catalogue of American Amphibians and Reptiles for 33 years. Over his career, Larry has authored or co-authored the descriptions of 69 currently recognized herpetofaunal species and six species have been named in his honor, including the anuran Craugastor lauraster and the snakes Cerrophidion wilsoni, Myriopholis wilsoni, and Oxybelis wilsoni. Vicente Mata-Silva is a herpetologist interested in ecology, conservation, and the monitoring of amphibians and reptiles in Mexico and the southwestern United States. His bachelor’s thesis compared herpetofaunal richness in Puebla, Mexico, in habitats with different degrees of human related disturbance. Vicente’s master’s thesis focused primarily on the diet of two syntopic whiptail species of lizards, one unisexual and the other bisexual, in the Trans-Pecos region of the Chihuahuan Desert. Currently, he is a postdoctoral research fellow at the University of Texas at El Paso, where his work focuses on rattlesnake populations in their natural habitat. His dissertation was on the ecology of the rock rattlesnake, Crotalus lepidus, in the northern Chihuahuan Desert. To date, Vicente has authored or co-authored 34 peer-reviewed scientific publications. Jerry D. Johnson is Professor of Biological Sciences at The University of Texas at El Paso, and has exten- sive experience studying the herpetofauna of Mesoamerica. He is the Director of the 40,000 acre “Indio Mountains Research Station,” was a co-editor on the recently published Conservation of Mesoamerican Amphibians and Reptiles, and is Mesoamerica/Caribbean editor for the Geographic Distribution section of Herpetological Review. Johnson has authored or co-authored over 80 peer-reviewed papers, includ- ing two 2010 articles, “Geographic distribution and conservation of the herpetofauna of southeastern Mexico” and “Distributional patterns of the herpetofauna of Mesoamerica, a biodiversity hotspot.” amphibian-reptile-conservation.org 028 June 2013 I Volume 7 | Number 1 | e61 Wilson et al. Appendix 1 . Comparison of the IUCN Ratings from the 2005 Mexican Assessment (updated to the present time) and the Environmental Vulner- ability Scores for 849 Mexican reptile species (crocodilians, turtles, lizards, and snakes). See text for explanation of the IUCN and EVS rating sys- tems. * = species endemic to Mexico. 1 = IUCN status needs updating. 2 = Not rated because not recognized as a distinct species. 3 = not described at the time of assessment. Species IUCN Ratings Environmental Vulnerability Scores Geographic Distribution Ecological Distribution Degree of Human Persecution Total Score Order Crocodylia (3 species) Family Alligatoridae (1 species) Caiman crocodilus LC 1 3 7 6 16 Family Crocodylidae (2 species) Crocodylus acutus VU 3 5 6 14 Crocodylus moreletii LC 2 5 6 13 Order Testudines (48 species) Family Cheloniidae (5 species) Caretta caretta EN — — — — Chelonia mydas EN — — — — Eretmochelys imbricata CR — — — — Lepidochelys kempii CR — — — — Lepidochelys olivacea VU — — — — Family Chelydridae (1 species) Chelydra rossignonii VU 4 7 6 17 Family Dermatemydidae (1 species) Dermatemys mawii CR 4 7 6 17 Family Dermochelyidae (1 species) Dermochelys coriacea CR — — — — Family Emydidae (15 species) Actinemys marmorata VU 3 8 6 17 Chrysemys picta LC 3 8 3 14 Pseudemys gorzugi NT 4 6 6 16 Terrapene coahuila* EN 5 8 6 19 Terrapene mexicana* NE 5 8 6 19 Terrapene nelsoni* DD 5 7 6 18 Terrapene ornata NT 3 6 6 15 Terrapene yucatana* NE 5 7 6 18 Trachemys gaigeae VU 4 8 6 18 Trachemys nebulosa* NE 5 7 6 18 Trachemys ornata* VU 5 8 6 19 Trachemys scripta LC 3 7 6 16 Trachemys taylori* EN 5 8 6 19 Trachemys venusta NE 3 4 6 13 Trachemys yaquia* VU 5 8 6 19 Family Geoemydidae (3 species) Rhinoclemmys areolata NT 4 6 3 13 Rhinoclemmys pulcherrima NE 1 4 3 8 Rhinoclemmys rubida* NT 5 6 3 14 Family Kinosternidae (17 species) Claudius angustatus NT 1 4 7 3 14 Kinosternon acutum NT 1 4 7 3 14 amphibian-reptile-conservation.org 029 June 2013 I Volume 7 I Number 1 I e61 Conservation reassessment of Mexican reptiles Kinosternon alamosae* DD 5 6 3 14 Kinosternon arizonense LC 4 8 3 15 Kinosternon chimalhuaca* LC 5 8 3 16 Kinosternon creaseri* LC 5 7 3 15 Kinosternon durangoense* DD 5 8 3 16 Kinosternon flavescens LC 3 6 3 12 Kinosternon herrerai* NT 5 6 3 14 Kinosternon hirtipes LC 2 5 3 10 Kinosternon integrum* LC 5 3 3 11 Kinosternon leucostomum NE 3 4 3 10 Kinosternon oaxacae* DD 5 7 3 15 Kinosternon scorpioides NE 3 4 3 10 Kinosternon sonoriense NT 4 7 3 14 Staurotypus salvinii NT 1 4 6 3 13 Staurotypus triporcatus NT 1 4 7 3 14 Family Testudinidae (3 species) Gopherus berlandieri LC 1 4 8 6 18 Gopherus flavomarginatus* VU 5 8 6 19 Gopherus morafkai NE 3 4 5 6 15 Family Trionychidae (2 species) Apalone atra* NE 6 8 6 20 Apalone spinifera LC 3 6 6 15 Order Squamata (798 species) Family Bipedidae (3 species) Bipes biporus* LC 5 8 1 14 Bipes canaliculatus* LC 5 6 1 12 Bipes tridactyl us* LC 5 8 1 14 Family Anguidae (48 species) Abronia bogerti* DD 6 8 4 18 Abronia chiszari* EN 6 7 4 17 Abronia deppii* EN 6 6 4 16 Abronia fuscolabialis* EN 6 8 4 18 Abronia graminea* EN 5 6 4 15 Abronia leurolepis* DD 6 8 4 18 Abronia lythrochila* LC 6 7 4 17 Abronia martindelcampoi* EN 5 6 4 15 Abronia matudai EN 4 7 4 15 Abronia mitchelli* DD 6 8 4 18 Abronia mixteca* VU 6 8 4 18 Abronia oaxacae* VU 6 7 4 17 Abronia ochoterenai DD 4 8 4 16 Abronia ornelasi* DD 6 8 4 18 Abronia ramirezi* DD 6 8 4 18 Abronia reidi* DD 6 8 4 18 Abronia smithi* LC 6 7 4 17 Abronia taeniata* VU 5 6 4 15 Anguis ceroni* NE 5 7 2 14 Anguis incomptus* NE 5 8 2 15 Baris ia ci Haris* NE 5 7 3 15 amphibian-reptile-conservation.org 030 June 2013 I Volume 7 I Number 1 I e61 Wilson et al. Baris ia herrerae* EN 5 7 3 15 Barisia imbricata* LC 5 6 3 14 Barisia jonesi* NE 2 6 7 3 16 Barisia levicollis* DD 5 7 3 15 Barisia planifrons* NE 2 5 7 3 15 Barisia rudicollis* EN 5 7 3 15 Celestus enneagrammus* LC 5 6 3 14 Celestus ingridae* DD 6 8 3 17 Celestus iegnotus* LC 5 6 3 14 Celestus rozellae NT 4 6 3 13 Elgaria cedrosensis* LC 5 8 3 16 Elgaria kingii LC 2 5 3 10 Elgaria multicarinata LC 3 4 3 10 Elgaria nana* LC 5 8 3 16 Elgaria paucicarinata* VU 5 5 3 13 Elgaria velazquezi* LC 5 6 3 14 Gerrhonotus farri* NE 3 6 8 3 17 Gerrhonotus infernal is* LC 5 5 3 13 Gerrhonotus liocephalus LC 2 1 3 6 Gerrhonotus lugoi* LC 6 8 3 17 Gerrhonotus ophiurus* LC 5 4 3 12 Gerrhonotus parvus* EN 6 8 3 17 Mesaspis antauges* DD 6 7 3 16 Mesaspis gadovii* LC 5 6 3 14 Mesaspis juarezi* EN 5 7 3 15 Mesaspis moreleti LC 3 3 3 9 Mesaspis viridiflava* LC 5 8 3 16 Family Anniellidae (2 species) Anniella geronimensis* EN 5 7 1 13 Anniella pulchra LC 3 8 1 12 Family Corytophanidae (6 species) Basiliscus vittatus NE 1 3 3 7 Corytophanes cristatus NE 3 5 3 11 Corytophanes hernandesii NE 4 6 3 13 Corytophanes percarinatus NE 4 4 3 11 Laemanctus longipes NE 1 5 3 9 Laemanctus serratus LC 2 3 3 8 Family Crotaphytidae (10 species) Crotaphytus antiquus* EN 5 8 3 16 Crotaphytus collaris LC 3 7 3 13 Crotaphytus dickersonae* LC 5 8 3 16 Crotaphytus grismeri* LC 5 8 3 16 Crotaphytus i ns u laris* LC 6 7 3 16 Crotaphytus nebrius LC 2 7 3 12 Crotaphytus reticulatus VU 4 5 3 12 Crotaphytus vestigium LC 3 3 3 9 Gambelia copeii LC 2 6 3 11 Gambelia wislizenii LC 3 7 3 13 amphibian-reptile-conservation.org 031 June 2013 I Volume 7 I Number 1 I e61 Conservation reassessment of Mexican reptiles Family Dactyloidae (50 species) Anolis allisoni NE 3 7 3 13 Anolis alvarezdeltoroi* DD 6 8 3 17 Anolis anisolepis* LC 5 7 3 15 Anolis barkeri* VU 5 7 3 15 Anolis beckeri NE 3 3 6 3 12 Anolis biporcatus NE 3 4 3 10 Anolis breedlovei* EN 6 7 3 16 Anolis capito NE 3 6 3 13 Anolis compressicauda* LC 5 7 3 15 Anolis crassulus NE 3 4 3 10 Anolis cristifer DD 4 6 3 13 Anolis cuprinus* LC 6 7 3 16 Anolis cymbops* DD 6 8 3 17 Anolis dollfusianus NE 4 6 3 13 Anolis duellmani* DD 6 8 3 17 Anolis dunni* LC 5 8 3 16 Anolis forbesi* DD 6 7 3 16 Anolis gadovi* LC 5 8 3 16 Anolis hobartsmithi* EN 6 6 3 15 Anolis isthmicus* DD 5 8 3 16 Anolis laeviventris NE 3 3 3 9 Anolis lemurinus NE 3 2 3 8 Anolis liogaster* LC 5 6 3 14 Anolis macrinii* LC 5 8 3 16 Anolis matudai NE 4 6 3 13 Anolis megapholidotus* LC 5 8 3 16 Anolis microlepidotus* LC 5 7 3 15 Anolis milleri* DD 5 6 3 14 Anolis naufragus* VU 5 5 3 13 Anolis nebuloides* LC 5 6 3 14 Anolis nebulosus* LC 5 5 3 13 Anolis omiltemanus* LC 5 7 3 15 Anolis parvicirculatus* LC 6 7 3 16 Anolis petersii NE 2 4 3 9 Anolis polyrhachis* DD 5 8 3 16 Anolis pygmaeus* EN 5 8 3 16 Anolis quercorum* LC 5 8 3 16 Anolis rodriguezii NE 4 3 3 10 Anolis sagrei NE 2 7 3 12 Anolis schiedii* DD 5 8 3 16 Anolis schmidti* LC 5 8 3 16 Anolis sericeus NE 2 3 3 8 Anolis serranoi NE 4 5 3 12 Anolis simmonsi* DD 5 7 3 15 Anolis subocu laris* DD 5 7 3 15 Anolis taylori* LC 5 8 3 16 Anolis tropidonotus NE 4 2 3 9 Anolis uniformis NE 4 6 3 13 amphibian-reptile-conservation.org 032 June 2013 I Volume 7 I Number 1 I e61 Wilson et al. Anolis unilobatus NE 3 1 3 3 7 Anolis utowanae* DD 6 8 3 17 Family Dibamidae (1 species) Anelytropsis papillosus* LC 5 4 1 10 Family Eublepharidae (7 species) Coleonyx brevis LC 4 6 4 14 Coleonyx elegans NE 2 3 4 9 Coleonyx fasciatus* LC 5 8 4 17 Coleonyx gypsicolus * LC 6 8 4 18 Coleonyx reticulatus LC 4 7 4 15 Coleonyx switaki LC 4 6 4 14 Coleonyx variegatus LC 4 3 4 11 Family Gymnophthalmidae (1 species) Gymnophthalmus speciosus NE 3 3 3 9 Family Helodermatidae (2 species) Heloderma horridum LC 2 4 5 11 Heloderma suspectum NT 4 6 5 15 Family Iguanidae (19 species) Ctenosaura acanthura NE 2 4 6 12 Ctenosaura alfredschmidti NT 4 8 3 15 Ctenosaura dark!* VU 5 7 3 15 Ctenosaura conspicuosa* NE 5 8 3 16 Ctenosaura defensor* VU 5 7 3 15 Ctenosaura hemilopha* NE 5 7 6 18 Ctenosaura macrolopha* NE 5 8 6 19 Ctenosaura nolascensis* NE 6 8 3 17 Ctenosaura oaxacana* CR 5 8 6 19 Ctenosaura pectinata* NE 5 4 6 15 Ctenosaura similis LC 1 4 3 8 Dipsosaurus catalinensis* NE 6 8 3 17 Dipsosaurus dorsalis LC 4 4 3 11 Iguana iguana NE 3 3 6 12 Sauromalus ater LC 4 6 3 13 Sauromalus hispid us* NT 5 6 3 14 Sauromalus klauberi* NE 6 7 3 16 Sauromalus slevini* NE 5 8 3 16 Sauromalus varius* NE 5 8 3 16 Family Mabuyidae (1 species) Marisora brachypoda NE 1 2 3 6 Family Phrynosomatidae (135 species) Callisaurus draconoides LC 4 5 3 12 Cophosaurus texanus LC 4 7 3 14 Holbrookia approximans* NE 5 6 3 14 Holbrookia elegans LC 4 6 3 13 Holbrookia lacerata NT 4 7 3 14 Holbrookia maculata LC 1 6 3 10 Holbrookia propinqua LC 4 8 3 15 Petrosaurus mearnsi LC 4 5 3 12 Petrosaurus repens* LC 5 5 3 13 amphibian-reptile-conservation.org 033 June 2013 I Volume 7 I Number 1 I e61 Conservation reassessment of Mexican reptiles Petrosaurus slevini* LC 5 8 3 16 Petrosaurus thalassinus* LC 5 5 3 13 Phrynosoma asio NE 2 6 3 11 Phrynosoma blainvillii NE 3 7 3 13 Phrynosoma braconnieri* LC 5 7 3 15 Phrynosoma cerroense* NE 6 7 3 16 Phrynosoma cornutum LC 1 7 3 11 Phrynosoma coronatum* LC 5 4 3 12 Phrynosoma ditmarsi* DD 5 8 3 16 Phrynosoma goodei NE 4 6 3 13 Phrynosoma hernandesi LC 3 7 3 13 Phrynosoma mcallii NT 4 8 3 15 Phrynosoma modestum LC 4 5 3 12 Phrynosoma orbiculare* LC 5 4 3 12 Phrynosoma platyrhinos LC 3 7 3 13 Phrynosoma solare LC 4 7 3 14 Phrynosoma taurus* LC 5 4 3 12 Phrynosoma wigginsi* NE 5 8 3 16 Sceloporus acanthinus NE 3 7 3 13 Sceloporus adleri* LC 5 7 3 15 Sceloporus aeneus* LC 5 5 3 13 Sceloporus albiventris* NE 5 8 3 16 Sceloporus anahuacus* LC 5 7 3 15 Sceloporus angustus* LC 5 8 3 16 Sceloporus asper* LC 5 6 3 14 Sceloporus bicanthalis* LC 5 5 3 13 Sceloporus bulled* LC 5 7 3 15 Sceloporus carinatus LC 4 5 3 12 Sceloporus cautus* LC 5 7 3 15 Sceloporus chaneyi* EN 5 7 3 15 Sceloporus chrysostictus LC 4 6 3 13 Sceloporus clarkii LC 2 5 3 10 Sceloporus couchii* LC 5 7 3 15 Sceloporus cowlesi NE 4 6 3 13 Sceloporus cozumelae* LC 5 7 3 15 Sceloporus cryptus* LC 5 6 3 14 Sceloporus cupreus* NE 5 8 3 16 Sceloporus cyanogenys* NE 6 7 3 16 Sceloporus cyanostictus* EN 5 8 3 16 Sceloporus druckercolini* NE 5 6 3 14 Sceloporus dugesii* LC 5 5 3 13 Sceloporus edwardtaylori* LC 5 6 3 14 Sceloporus exsul* CR 6 8 3 17 Sceloporus formosus* LC 5 7 3 15 Sceloporus gadoviae* LC 5 3 3 11 Sceloporus goldmani* EN 5 7 3 15 Sceloporus grammicus LC 2 4 3 9 Sceloporus grandaevus* LC 6 7 3 16 Sceloporus halli* DD 6 8 3 17 amphibian-reptile-conservation.org 034 June 2013 I Volume 7 I Number 1 I e61 Wilson et al. Sceloporus heterolepis* LC 5 6 3 14 Sceloporus horrid us* LC 5 3 3 11 Sceloporus hunsakeri* LC 5 6 3 14 Sceloporus ins ignis* LC 5 8 3 16 Sceloporus internasalis LC 4 4 3 11 Sceloporus jalapae* LC 5 5 3 13 Sceloporus jarrovii LC 2 6 3 11 Sceloporus lemosespinali* DD 5 8 3 16 Sceloporus licki* LC 5 5 3 13 Sceloporus lineatulus* LC 6 8 3 17 Sceloporus lineolateralis* NE 5 8 3 16 Sceloporus lundelli LC 4 7 3 14 Sceloporus macdougalli* LC 5 8 3 16 Sceloporus maculosus* VU 5 8 3 16 Sceloporus magister LC 1 5 3 9 Sceloporus marmoratus NE 2 6 3 11 Sceloporus megalepidurus* VU 5 6 3 14 Sceloporus melanorhinus LC 2 4 3 9 Sceloporus merriami LC 4 6 3 13 Sceloporus minor* LC 5 6 3 14 Sceloporus mucronatus* LC 5 5 3 13 Sceloporus nelsoni* LC 5 5 3 13 Sceloporus oberon* VU 5 6 3 14 Sceloporus occidentalis LC 3 6 3 12 Sceloporus ochoterenae* LC 5 4 3 12 Sceloporus olivaceus LC 4 6 3 13 Sceloporus orcutti LC 2 2 3 7 Sceloporus ornatus* NT 5 8 3 16 Sceloporus palaciosi* LC 5 7 3 15 Sceloporus parvus* LC 5 7 3 15 Sceloporus poinsetti LC 4 5 3 12 Sceloporus prezygus NE 4 8 3 15 Sceloporus pyrocephalus* LC 5 4 3 12 Sceloporus salvini* DD 5 7 3 15 Sceloporus samcolemani* LC 5 7 3 15 Sceloporus scalaris* LC 5 4 3 12 Sceloporus serrifer LC 2 1 3 6 Sceloporus shannonorum* NE 5 7 3 15 Sceloporus siniferus LC 2 6 3 11 Sceloporus slevini LC 2 6 3 11 Sceloporus smaragdinus LC 4 5 3 12 Sceloporus smithi* LC 5 7 3 15 Sceloporus spinosus* LC 5 4 3 12 Sceloporus squamosus NE 3 5 3 11 Sceloporus stejnegeri* LC 5 5 3 13 Sceloporus subniger* NE 5 7 3 15 Sceloporus subpictus* DD 6 7 3 16 Sceloporus sugillatus* LC 5 8 3 16 Sceloporus taeniocnemis LC 4 5 3 12 amphibian-reptile-conservation.org 035 June 2013 I Volume 7 I Number 1 I e61 Conservation reassessment of Mexican reptiles Sceloporus tanneri* DD 6 7 3 16 Sceloporus teapensis LC 4 6 3 13 Sceloporus torquatus* LC 5 3 3 11 Sceloporus uniformis NE 3 7 3 13 Sceloporus utiformis* LC 5 7 3 15 Sceloporus vandenburgianus LC 4 7 3 14 Sceloporus variabilis NE 1 1 3 5 Sceloporus virgatus LC 4 8 3 15 Sceloporus zosteromus* LC 5 4 3 12 Uma exsul* EN 5 8 3 16 Uma notata NT 4 8 3 15 Uma paraphygas* NT 6 8 3 17 Uma rufopunctata* NT 5 8 3 16 Urosaurus auriculatus* EN 6 7 3 16 Urosaurus bicarinatus* LC 5 4 3 12 Urosaurus clarionensis* VU 6 8 3 17 Urosaurus gadovi* LC 3 6 3 12 Urosaurus graciosus LC 3 8 3 14 Urosaurus lahtelai* LC 5 8 3 16 Urosaurus nigricaudus LC 3 2 3 8 Urosaurus ornatus LC 2 5 3 10 Uta encantadae* VU 6 8 3 17 Uta lowei* VU 6 8 3 17 Uta nolascensis* LC 6 8 3 17 Uta palmeri* VU 6 8 3 17 Uta squamata* LC 6 8 3 17 Uta stansburiana LC 3 1 3 7 Uta tumidarostra* VU 6 8 3 17 Family Phyllodactylidae (15 species) Phyllodactylus bordai* LC 5 5 3 13 Phyllodactylus bugastrolepis* LC 6 8 3 17 Phyllodactylus davisi* LC 5 8 3 16 Phyllodactylus delcampoi* LC 5 8 3 16 Phyllodactylus duellmani* LC 5 8 3 16 Phyllodactylus homolepidurus* LC 5 7 3 15 Phyllodactylus lanei* LC 5 7 3 15 Phyllodactylus mural is* LC 5 6 3 14 Phyllodactylus nocticolus NE 2 5 3 10 Phyllodactylus partidus* LC 5 8 3 16 Phyllodactylus paucituberculatus * DD 6 7 3 16 Phyllodactylus tuberculosus NE 1 4 3 8 Phyllodactylus u rictus* NT 5 7 3 15 Phyllodactylus xanti* LC 5 7 3 15 Thecadactylus rapicauda NE 3 4 3 10 Family Scincidae (23 species) Mesoscincus altamirani* DD 5 6 3 14 Mesoscincus schwartzei LC 2 6 3 11 Plestiodon bilineatus* NE 5 5 3 13 Plestiodon brevirostris* LC 5 3 3 11 amphibian-reptile-conservation.org 036 June 2013 I Volume 7 I Number 1 I e61 Wilson et al. Plestiodon callicephalus LC 2 7 3 12 Plestiodon colimensis* DD 5 6 3 14 Plestiodon copei* LC 5 6 3 14 Plestiodon dicei* NE 5 4 3 12 Plestiodon dugesi* VU 5 8 3 16 Plestiodon gilberti LC 3 6 3 12 Plestiodon indubitus* NE 5 7 3 15 Plestiodon lagunensis* LC 6 6 3 15 Plestiodon lynxe* LC 5 2 3 10 Plestiodon multilineatus* DD 5 8 3 16 Plestiodon multivirgatus LC 3 8 3 14 Plestiodon nietoi* NE 6 8 3 17 Plestiodon obsoletus LC 3 5 3 11 Plestiodon ochoterenae* LC 5 5 3 13 Plestiodon pan/iauriculatus* DD 5 7 3 15 Plestiodon pan/ulus* DD 5 7 3 15 Plestiodon skiltonianus LC 3 5 3 11 Plestiodon sumichrasti NE 4 5 3 12 Plestiodon tetragrammus LC 4 5 3 12 Family Sphaerodactylidae (4 species) Aristelliger georgeensis NE 3 7 3 13 Gonatodes albogularis NE 3 5 3 11 Sphaerodactylus continentalis NE 4 3 3 10 Sphaerodactylus glaucus NE 4 5 3 12 Family Sphenomorphidae (6 species) Scincella gemmingeri* LC 5 3 3 11 Scincella kikaapoda* NE 3 6 8 3 17 Scincella lateralis LC 3 7 3 13 Scincella silvicola* LC 5 4 3 12 Sphenomorphus assatus NE 2 2 3 7 Sphenomorphus cherriei NE 3 2 3 8 Family Teiidae (46 species) Aspidoscelis angusticeps LC 4 6 3 13 Aspidoscelis bacata* LC 6 8 3 17 Aspidoscelis burti LC 4 8 3 15 Aspidoscelis calidipes* LC 5 6 3 14 Aspidoscelis cana* LC 5 8 3 16 Aspidoscelis carmenensis* LC 6 8 3 17 Aspidoscelis catalinensis* VU 6 8 3 17 Aspidoscelis celeripes* LC 5 7 3 15 Aspidoscelis ceralbensis* LC 6 8 3 17 Aspidoscelis communis* LC 5 6 3 14 Aspidoscelis costata* LC 5 3 3 11 Aspidoscelis cozumela* LC 5 8 3 16 Aspidoscelis danheimae* LC 6 7 3 16 Aspidoscelis deppii LC 1 4 3 8 Aspidoscelis espiritensis* LC 5 8 3 16 Aspidoscelis exanguis LC 4 7 3 14 Aspidoscelis franciscensis* LC 6 8 3 17 amphibian-reptile-conservation.org 037 June 2013 I Volume 7 I Number 1 I e61 Conservation reassessment of Mexican reptiles Aspidoscelis gularis LC 2 4 3 9 Aspidoscelis guttata* LC 5 4 3 12 Aspidoscelis hyperythra LC 2 5 3 10 Aspidoscelis inornata LC 4 7 3 14 Aspidoscelis labial is* VU 5 7 3 15 Aspidoscelis laredoensis LC 4 7 3 14 Aspidoscelis lineattissima* LC 5 6 3 14 Aspidoscelis marmorata NE 4 7 3 14 Aspidoscelis martyris* VU 6 8 3 17 Aspidoscelis maslini LC 4 8 3 15 Aspidoscelis mexicana* LC 5 6 3 14 Aspidoscelis motaguae LC 4 5 3 12 Aspidoscelis neomexicana LC 4 8 3 15 Aspidoscelis opatae* DD 5 8 3 16 Aspidoscelis parvisocia* LC 5 7 3 15 Aspidoscelis picta* LC 6 8 3 17 Aspidoscelis rodecki* NT 5 8 3 16 Aspidoscelis sackii* LC 5 6 3 14 Aspidoscelis semptemvittata LC 3 7 3 13 Aspidoscelis sexlineata LC 3 8 3 14 Aspidoscelis sonorae LC 4 6 3 13 Aspidoscelis stictogramma NE 4 7 3 14 Aspidoscelis tesselata LC 4 7 3 14 Aspidoscelis tigris LC 3 2 3 8 Aspidoscelis uniparens LC 4 8 3 15 Aspidoscelis xanthonota NE 4 7 3 14 Holcosus chaitzami DD 4 7 3 14 Holcosus festiva NE 3 5 3 11 Holcosus undulatus NE 2 2 3 7 Family Xantusiidae (25 species) Lepidophyma chicoasense* DD 6 8 2 16 Lepidophyma cuicateca* NE 3 6 8 2 16 Lepidophyma dontomasi* DD 6 6 2 14 Lepidophyma flavimaculatum NE 1 5 2 8 Lepidophyma gaigeae* VU 5 6 2 13 Lepidophyma lineri* DD 5 8 2 15 Lepidophyma lipetzi* EN 6 8 2 16 Lepidophyma lowei* DD 6 8 2 16 Lepidophyma micropholis* VU 5 8 2 15 Lepidophyma occulor* LC 5 7 2 14 Lepidophyma pajapanense* LC 5 6 2 13 Lepidophyma radula* DD 6 5 2 13 Lepidophyma smithii NE 2 4 2 8 Lepidophyma sylvaticum* LC 5 4 2 11 Lepidophyma tarascae* DD 5 7 2 14 Lepidophyma tuxtlae* DD 5 4 2 11 Lepidophyma zongolica* NE 3 6 8 2 16 Xantusia bolsonae* DD 6 8 3 17 Xantusia extorris* LC 5 7 3 15 amphibian-reptile-conservation.org 038 June 2013 I Volume 7 I Number 1 I e61 Wilson et al. Xantusia gilberti* NE 5 8 3 16 Xantusia henshawi LC 4 5 3 12 Xantusia jaycolei* NE 5 8 3 16 Xantusia sanchezi* LC 5 8 3 16 Xantusia sherbrookei* NE 5 8 3 16 Xantusia wigginsi NE 4 7 3 14 Family Xenosauridae (9 species) Xenosaurus agrenon* NE 5 4 3 12 Xenosaurus grandis* VU 5 1 3 9 Xenosaurus newmanorum* EN 5 7 3 15 Xenosaurus penai* LC 6 7 3 16 Xenosaurus phalaroanthereon* DD 5 8 3 16 Xenosaurus platyceps* EN 5 6 3 14 Xenosaurus rackhami NE 4 4 3 11 Xenosaurus rectocol laris* LC 5 8 3 16 Xenosaurus tzacualtipantecus* NE 6 8 3 17 Family Boidae (2 species) Boa constrictor NE 3 1 6 10 Charina trivirgata LC 4 3 3 10 Family Colubridae (136 species) Arizona elegans LC 1 1 3 5 Arizona pacata* LC 5 5 4 14 Bogertophis rosaliae LC 2 5 3 10 Bogertophis subocularis LC 4 7 3 14 Chilomeniscus savagei* LC 6 7 2 15 Chilomeniscus stramineus LC 4 2 2 8 Chionactus occipitalis LC 4 6 2 12 Chionactus palarostris LC 4 7 2 13 Coluber constrictor LC 1 6 3 10 Conopsis acuta* NE 5 7 2 14 Conopsis amphisticha* NT 5 8 2 15 Conopsis biserial is* LC 5 6 2 13 Conopsis lineata* LC 5 6 2 13 Conopsis megalodon* LC 5 7 2 14 Conopsis nasus* LC 5 4 2 11 Dendrophidion vinitor LC 3 7 3 13 Drymarchon melanurus LC 1 1 4 6 Drymobius chloroticus LC 1 3 4 8 Drymobius margaritiferus NE 1 1 4 6 Ficimia hardyi* EN 5 6 2 13 Ficimia olivacea* NE 5 2 2 9 Ficimia publia NE 4 3 2 9 Ficimia ramirezi* DD 6 8 2 16 Ficimia ruspator* DD 6 8 2 16 Ficimia streckeri LC 3 7 2 12 Ficimia variegata* DD 5 7 2 14 Geagras redimitus* DD 5 7 2 14 Gyalopion canum LC 4 3 2 9 Gyalopion quadrangulare LC 3 6 2 11 amphibian-reptile-conservation.org 039 June 2013 I Volume 7 I Number 1 I e61 Conservation reassessment of Mexican reptiles Lampropeltis alterna LC 4 7 3 14 Lampropeltis californiae NE 2 3 4 3 10 Lampropeltis catalinensis* DD 6 8 3 17 Lampropeltis herrerae* CR 6 8 3 17 Lampropeltis holbrooki NE 2 3 8 3 14 Lampropeltis knoblochi NE 2 2 5 3 10 Lampropeltis mexicana* LC 5 7 3 15 Lampropeltis ruthveni* NT 5 8 3 16 Lampropeltis splendida NE 2 4 5 3 12 Lampropeltis triangulum NE 1 1 5 7 Lampropeltis webbi* DD 5 8 3 16 Lampropeltis zonata LC 3 7 5 15 Leptophis ahaetulla NE 3 3 4 10 Leptophis diplotropis* LC 5 5 4 14 Leptophis mexicanus LC 1 1 4 6 Leptophis modestus VU 3 7 4 14 Liochlorophis vernalis LC 3 8 3 14 Masticophis anthonyi* CR 6 8 3 17 Masticophis au rig ulus* LC 5 4 4 13 Masticophis barbouri* DD 6 8 3 17 Masticophis bilineatus LC 2 5 4 11 Masticophis flagellum LC 1 3 4 8 Masticophis fuiiginosus NE 2 3 4 9 Masticophis lateralis LC 3 3 4 10 Masticophis mentovarius NE 1 1 4 6 Masticophis schotti LC 4 5 4 13 Masticophis slevini* LC 6 8 3 17 Masticophis taeniatus LC 1 5 4 10 Mastigodryas cliftoni* NE 5 6 3 14 Mastigodryas melanolomus LC 1 1 4 6 Opheodrys aestivus LC 3 7 3 13 Oxybelis aeneus NE 1 1 3 5 Oxybelis fulgidus NE 3 2 4 9 Pantherophis bairdi NE 4 7 4 15 Pantherophis emoryi LC 3 6 4 13 Phyllorhynchus browni LC 4 7 2 13 Phyllorhynchus decurtatus LC 4 5 2 11 Pituophis catenifer LC 4 1 4 9 Pituophis deppei* LC 5 5 4 14 Pituophis insulanus* LC 6 6 4 16 Pituophis lineaticollis LC 2 2 4 8 Pituophis vertebral is* LC 5 3 4 12 Pseudelaphe flavirufa LC 2 4 4 10 Pseudelaphe phaescens* NE 5 7 4 16 Pseudoficimia frontalis* LC 5 5 3 13 Pseustes poecilonotus LC 3 4 3 10 Rhinocheilus antonii* NE 5 8 3 16 Rhinocheilus etheridgei* DD 6 7 3 16 Rhinocheilus lecontei LC 1 3 4 8 amphibian-reptile-conservation.org 040 June 2013 I Volume 7 I Number 1 I e61 Wilson et al. Salvadora bairdi* LC 5 6 4 15 Salvadora deserticola NE 4 6 4 14 Salvadora grahamiae LC 4 2 4 10 Salvadora hexalepis LC 4 2 4 10 Salvadora intermedia* LC 5 7 4 16 Salvadora lemniscata* LC 5 6 4 15 Salvadora mexicana* LC 5 6 4 15 Scaphiodontophis annulatus NE 1 5 5 11 Senticolis triaspis NE 2 1 3 6 Sonora aemula* NT 5 6 5 16 Sonora michoacanensis* LC 5 6 3 14 Sonora mutabilis* LC 5 6 3 14 Sonora semiannulata LC 1 1 3 5 Spilotes pullatus NE 1 1 4 6 Stenorrhina degenhardtii NE 3 3 3 9 Stenorrhina freminvillii NE 1 2 4 7 Symphimus leucostomus* LC 5 6 3 14 Symphimus mayae LC 4 7 3 14 Sympholis lippiens* NE 5 6 3 14 Tantilla atriceps LC 2 7 2 11 Tantilla bocourti* LC 5 2 2 9 Tantilla briggsi* DD 6 8 2 16 Tantilla calamarina* LC 5 5 2 12 Tantilla cascadae* DD 6 8 2 16 Tantilla ceboruca* NE 6 8 2 16 Tantilla coronadoi* LC 6 7 2 15 Tantilla cuniculator LC 4 7 2 13 Tantilla deppei* LC 5 6 2 13 Tantilla flavilineata* EN 5 7 2 14 Tantilla gracilis LC 3 8 2 13 Tantilla hobartsmithi LC 3 6 2 11 Tantilla impensa LC 3 5 2 10 Tantilla johnsoni* DD 6 8 2 16 Tantilla moesta LC 4 7 2 13 Tantilla nigriceps LC 3 6 2 11 Tantilla oaxacae* DD 6 7 2 15 Tantilla planiceps LC 4 3 2 9 Tantilla robusta* DD 6 8 2 16 Tantilla rubra LC 2 1 2 5 Tantilla schistosa NE 3 3 2 8 Tantilla sertula* DD 6 8 2 16 Tantilla shawi* EN 5 8 2 15 Tantilla slavensi* DD 5 7 2 14 Tantilla striata* DD 5 7 2 14 Tantilla tayrae* DD 6 7 2 15 Tantilla triseriata* DD 5 6 2 13 Tantilla vulcani NE 4 6 2 12 Tantilla wilcoxi LC 2 6 2 10 Tantilla yaquia LC 2 6 2 10 amphibian-reptile-conservation.org 041 June 2013 I Volume 7 I Number 1 I e61 Conservation reassessment of Mexican reptiles Tantillita brevissima LC 4 3 2 9 Tantillita canula LC 4 6 2 12 Tantillita lintoni LC 4 6 2 12 Trimorphodon biscutatus NE 2 1 4 7 Trimorphodon lambda NE 4 5 4 13 Trimorphodon lyrophanes NE 4 2 4 10 Trimorphodon paucimaculatus* NE 5 6 4 15 Trimorphodon tau* LC 5 4 4 13 Trimorphodon vilkinsonii LC 4 7 4 15 Family Dipsadidae (115 species) Adelphicos latifasciatum* DD 6 7 2 15 Adelphicos newmanorum* NE 5 5 2 12 Adelphicos nigrilatum* LC 5 7 2 14 Adelphicos quadrivirgatum DD 4 4 2 10 Adelphicos sargii LC 4 6 2 12 Amastridium sapperi NE 4 4 2 10 Chersodromus liebmanni* LC 5 5 2 12 Chersodromus rubriventris* EN 5 7 2 14 Coniophanes alvarezi* DD 6 8 3 17 Coniophanes bipunctatus NE 1 5 3 10 Coniophanes fissidens NE 1 3 3 7 Coniophanes imperialis LC 2 3 3 8 Coniophanes lateritius* DD 5 5 3 13 Coniophanes melanocephalus* DD 5 6 3 14 Coniophanes meridanus* LC 5 7 3 15 Coniophanes michoacanensis* NE 3 6 8 3 17 Coniophanes piceivittis LC 1 3 3 7 Coniophanes quinquevittatus LC 4 6 3 13 Coniophanes sarae* DD 5 7 3 16 Coniophanes schmidti LC 4 6 3 13 Coniophanes taylori* NE 5 7 4 16 Cryophis hallbergi* DD 5 7 2 14 Diadophis punctatus LC 1 1 2 4 Dipsas brevifacies LC 4 7 4 15 Dipsas gaigeae* LC 5 8 4 17 Enuiius flavitorques NE 1 1 3 5 Enulius oligostichus* DD 5 7 3 15 Geophis anocu laris* LC 6 8 2 16 Geophis bicolor* DD 5 8 2 15 Geophis blanchardi* DD 5 8 2 15 Geophis cancellatus LC 4 6 2 12 Geophis carinosus LC 2 4 2 8 Geophis chalybeus* DD 6 7 2 15 Geophis dubius* LC 5 6 2 13 Geophis duellmani* LC 5 8 2 15 Geophis dugesi* LC 5 6 2 13 Geophis immaculatus LC 4 8 2 14 Geophis incomptus* DD 6 8 2 16 Geophis isthmicus* DD 6 8 2 16 amphibian-reptile-conservation.org 042 June 2013 I Volume 7 I Number 1 I e61 Wilson et al. Geophis juarezi* DD 6 8 2 16 Geophis juliai* VU 5 6 2 13 Geophis latici rictus* LC 5 4 2 11 Geophis laticol laris* DD 6 8 2 16 Geophis lati frontal is* DD 5 7 2 14 Geophis maculiferus* DD 6 8 2 16 Geophis mutitorques* LC 5 6 2 13 Geophis nasalis LC 4 3 2 9 Geophis nigrocinctus* DD 5 8 2 15 Geophis occabus* NE 3 6 8 2 16 Geophis omiltemanus* LC 5 8 2 15 Geophis petersi* DD 5 8 2 15 Geophis pyburni* DD 6 8 2 16 Geophis rhodogaster LC 3 7 2 12 Geophis russatus* DD 6 8 2 16 Geophis sallei* DD 6 7 2 15 Geophis semidoliatus* LC 5 6 2 13 Geophis sieboldi* DD 5 6 2 13 Geophis tarascae* DD 5 8 2 15 Heterodon kenneriyi NE 3 4 4 11 Hypsigiena affinis* NE 5 7 2 14 Hypsiglena chlorophaea NE 1 5 2 8 Hypsigiena jani NE 1 3 2 6 Hypsiglena ochrorhyncha NE 2 4 2 8 Hypsigiena slevini* NE 5 4 2 11 Hypsiglena tanzeri* DD 5 8 2 15 Hypsiglena torquata* LC 5 1 2 8 Imantodes cenchoa NE 1 3 2 6 Imantodes gemmistratus NE 1 3 2 6 Imantodes tenuissimus NE 4 7 2 13 Leptodeira frenata LC 4 4 4 12 Leptodeira maculata LC 2 1 4 7 Leptodeira nigrofasciata LC 1 3 4 8 Leptodeira punctata* LC 5 8 4 17 Leptodeira septentrionalis NE 2 2 4 8 Leptodeira splendida* LC 5 5 4 14 Leptodeira uribei* LC 5 8 4 17 Ninia diademata LC 4 3 2 9 Ninia sebae NE 1 1 2 5 Pliocercus elapoides LC 4 1 5 10 Pseudoleptodeira latifasciata * LC 5 5 4 14 Rhadinaea bogertorum* DD 6 8 2 16 Rhadinaea cuneata* DD 6 7 2 15 Rhadinaea decorata NE 1 6 2 9 Rhadinaea forbesi* DD 5 8 2 15 Rhadinaea fulvivittis* VU 5 4 2 11 Rhadinaea gaigeae* DD 5 5 2 12 Rhadinaea hesperia* LC 5 3 2 10 Rhadinaea laureata* LC 5 5 2 12 amphibian-reptile-conservation.org 043 June 2013 I Volume 7 I Number 1 I e61 Conservation reassessment of Mexican reptiles Rhadinaea macdougalli* DD 5 5 2 12 Rhadinaea marcel lae* EN 5 5 2 12 Rhadinaea montana* EN 5 7 2 14 Rhadinaea myersi* DD 5 5 2 12 Rhadinaea omiltemana* DD 5 8 2 15 Rhadinaea quinquelineata* DD 5 8 2 15 Rhadinaea taeniata* LC 5 6 2 13 Rhadinella godmani NE 3 5 2 10 Rhadinella hannsteini DD 4 5 2 11 Rhadinella kanalchutchan* DD 6 8 2 16 Rhadinella kinkelini LC 4 6 2 12 Rhadinella lachrymans LC 4 2 2 8 Rhadinella posadasi NE 4 8 2 14 Rhadinella schistosa* LC 5 6 2 13 Rhadinophanes monticola* DD 6 7 2 15 Sibon dimidiatus LC 1 5 4 10 Si bon linearis* DD 6 8 2 16 Sibon nebulatus NE 1 2 2 5 Sibon sanniolus LC 4 6 2 12 Tantalophis discolor* VU 5 6 3 14 Tropidodipsas annulifera* LC 5 4 4 13 Tropidodipsas fasciata* NE 5 4 4 13 Tropidodipsas fischeri NE 4 3 4 11 Tropidodipsas philippi* LC 5 5 4 14 Tropidodipsas repleta* DD 5 8 4 17 Tropidodipsas sartorii NE 2 2 5 9 Tropidodipsas zweifeli* NE 5 7 4 16 Family Elapidae (19 species) Laticauda colubrina LC — — — — Micruroides euryxanthus LC 4 6 5 15 Micrurus bernadi* LC 5 5 5 15 Micrurus bogerti* DD 5 7 5 17 Micrurus browni LC 2 1 5 8 Micrurus diastema LC 2 1 5 8 Micrurus distans* LC 5 4 5 14 Micrurus elegans LC 4 4 5 13 Micrurus ephippifer* VU 5 5 5 15 Micrurus laticol laris* LC 5 4 5 14 Micrurus latifasciatus LC 4 4 5 13 Micrurus limbatus* LC 5 7 5 17 Micrurus nebu laris* DD 5 8 5 18 Micrurus nigrocinctus NE 3 3 5 11 Micrurus pachecogili* DD 6 7 5 18 Micrurus proximans* LC 5 8 5 18 Micrurus tamaulipensis* DD 6 8 5 19 Micrurus tener LC 1 5 5 11 Pelamis platura LC — — — — Family Leptotyphlopidae (8 species) Epictia goudotii NE 1 1 1 3 amphibian-reptile-conservation.org 044 June 2013 I Volume 7 I Number 1 I e61 Wilson et al. Rena boettgeri* NE 5 8 1 14 Rena bressoni* DD 5 8 1 14 Rena dissecta LC 4 6 1 11 Rena dulcis LC 4 8 1 13 Rena humilis LC 4 3 1 8 Rena maxima* LC 5 5 1 11 Rena myopica* LC 5 7 1 13 Family Loxocemidae (1 species) Loxocemus bicolor NE 1 5 4 10 Family Natricidae (33 species) Adelophis copei* VU 5 8 2 15 Adelophis foxi* DD 6 8 2 16 Nerodia erythrogaster LC 3 4 4 11 Nerodia rhombifer LC 1 5 4 10 Storeria dekayi LC 1 4 2 7 Storeria hidalgoensis* VU 5 6 2 13 Storeria storerioides* LC 5 4 2 11 Thamnophis bogerti* NE 5 7 4 16 Thamnophis chrysocephalus* LC 5 5 4 14 Thamnophis conanti* NE 5 8 4 17 Thamnophis cyrtopsis LC 2 1 4 7 Thamnophis elegans LC 3 7 4 14 Thamnophis eques LC 2 2 4 8 Thamnophis errans* LC 5 7 4 16 Thamnophis exsul* LC 5 7 4 16 Thamnophis fulvus LC 4 5 4 13 Thamnophis godmani* LC 5 5 4 14 Thamnophis hammondii LC 4 5 4 13 Thamnophis lineri* NE 5 8 4 17 Thamnophis marcianus NE 1 5 4 10 Thamnophis melanogaster* EN 5 6 4 15 Thamnophis mendax* EN 5 5 4 14 Thamnophis nigronuchalis* DD 5 3 4 12 Thamnophis postremus* LC 5 6 4 15 Thamnophis proximus NE 1 2 4 7 Thamnophis pulch hiatus* LC 5 6 4 15 Thamnophis rossmani* DD 6 8 4 18 Thamnophis rufipunctatus LC 4 7 4 15 Thamnophis scalaris* LC 5 5 4 14 Thamnophis scaliger* VU 5 6 4 15 Thamnophis sirtalis LC 3 7 4 14 Thamnophis sumichrasti* LC 5 6 4 15 Thamnophis validus* LC 5 3 4 12 Family Typhlopidae (2 species) Typhlops microstomus LC 4 7 1 12 Typhlops tenuis LC 4 6 1 11 Family Ungaliophiidae (2 species) Exiliboa placata* VU 5 8 2 15 Ungaliophis continentalis NE 3 5 2 10 amphibian-reptile-conservation.org 045 June 2013 I Volume 7 I Number 1 I e61 Conservation reassessment of Mexican reptiles Family Viperidae (59 species) Agkistrodon bilineatus NT 1 5 5 11 Agkistrodon contortrix LC 3 6 5 14 Agkistrodon taylori* LC 5 7 5 17 Atropoides mexicanus NE 3 4 5 12 Atropoides nummifer* LC 5 3 5 13 Atropoides occiduus NE 4 6 5 15 Atropoides olmec LC 4 6 5 15 Bothriechis aurifer VU 3 6 5 14 Bothriechis bicolor LC 4 5 5 14 Bothriechis rowleyi* VU 5 6 5 16 Bothriechis schlegelii NE 3 4 5 12 Bothrops asper NE 3 4 5 12 Cerrophidion godmani NE 3 3 5 11 Cerrophidion petlalcalensis* DD 5 8 5 18 Cerrophidion tzotzilorum* LC 6 8 5 19 Crotalus angelensis* LC 6 7 5 18 Crotalus aquilus* LC 5 6 5 16 Crotalus atrox LC 1 3 5 9 Crotalus basiliscus* LC 5 6 5 16 Crotalus catalinensis* CR 6 8 5 19 Crotalus cerastes LC 4 7 5 16 Crotalus culminatus* NE 5 5 5 15 Crotalus enyo* LC 5 3 5 13 Crotalus ericsmithi* NE 5 8 5 18 Crotalus estebanensis* LC 6 8 5 19 Crotalus helleri NE 4 3 5 12 Crotalus intermedius* LC 5 5 5 15 Crotalus lannomi* DD 6 8 5 19 Crotalus lepidus LC 2 5 5 12 Crotalus lorenzoensis* LC 6 8 5 19 Crotalus mitchellii LC 4 3 5 12 Crotalus molossus LC 2 1 5 8 Crotalus muertensis* LC 6 8 5 19 Crotalus ornatus NE 4 4 5 13 Crotalus polystictus* LC 5 6 5 16 Crotalus price i LC 2 7 5 14 Crotalus pusillus* EN 5 8 5 18 Crotalus ravus* LC 5 4 5 14 Crotalus ruber LC 2 2 5 9 Crotalus scutulatus LC 2 4 5 11 Crotalus simus NE 3 2 5 10 Crotalus stejnegeri* VU 5 7 5 17 Crotalus tancitarensis* DD 6 8 5 19 Crotalus tigris LC 4 7 5 16 Crotalus totonacus* NE 5 7 5 17 Crotalus transversus* LC 5 7 5 17 Crotalus triseriatus* LC 5 6 5 16 Crotalus tzabcan NE 4 7 5 16 amphibian-reptile-conservation.org 046 June 2013 I Volume 7 I Number 1 I e61 Wilson et al. Crotalus viridis LC 1 6 5 12 Crotalus willardi LC 2 6 5 13 Mixcoatlus barbouri* EN 5 5 5 15 Mixcoatlus browni* NE 5 7 5 17 Mixcoatlus melanurus* EN 5 7 5 17 Ophryacus undulatus* VU 5 5 5 15 Porthidium dunni* LC 5 6 5 16 Porthidium hespere* DD 5 8 5 18 Porthidium nasutum LC 3 6 5 14 Porthidium yucatanicum* LC 5 7 5 17 Sistrurus catenatus LC 3 5 5 13 Family Xenodontidae (8 species) Clelia scytalina NE 4 5 4 13 Conophis lineatus LC 2 3 4 9 Conophis morai* DD 6 7 4 17 Conophis vittatus LC 2 5 4 11 Manolepis putnami* LC 5 5 3 13 Oxyrhopus petolarius NE 3 6 5 14 Tretanorhinus nigroluteus NE 3 5 2 10 Xenodon rabdocephalus NE 3 5 5 13 amphibian-reptile-conservation.org 047 June 2013 I Volume 7 I Number 1 I e61 Cantils (genus Agkistrodon ) are some of the most feared snakes in Mesoamerica, as their bite and powerful venom have caused numerous human fatalities. Equipped with a large and strikingly-marked head, a stout body, and a nervous attitude that often is mis- taken for aggression, these snakes usually are killed on sight. Cantils primarily are found in tropical forests that undergo a prolonged dry season, but occasionally inhabit savannas and areas that flood seasonally after heavy rains. Pictured here is a cantil from Parque Nacional Santa Rosa, in northwestern Costa Rica. Photo by Louis W. Porras. June 2013 I Volume 7 I Number 1 I e63 Amphib. Reptile Conserv. | http://amphibian-reptile-conservation.org 048 Copyright: © 2013 Porras et al. This is an open-access article distributed under the terms of the Creative Commons Attribution-NonCommercial-NoDerivs 3.0 Unported License, which permits unrestricted use for non-commer- cial and education purposes only provided the original author and source are credited. Amphibian & Reptile Conservation 7(1): 48-73. A taxonomic reevaluation and conservation assessment of the common cantil, Agkistrodon bilineatus (Squamata: Viperidae): a race against time 1 Louis W. Porras, 2 Larry David Wilson, 34 Gordon W. Schuett, and 4 Randall S. Reiserer 1 7705 Wyatt EarpAvenue, Eagle Mountain, Utah, 84005, USA 2 CentroZamoranodeBiodiversidad, EscuelaAgncolaPanamericanaZamorano, Francisco Morazdn, HONDURAS; 16010 S. W. 207th Avenue, Miami, Florida, 33187, USA 3 Department of Biology and Center for Behavioral Neuroscience, Georgia State University, Atlanta, Georgia, 30303, USA 4 The Copperhead Institute, P.O. Box 6755, Spartanburg, South Carolina 29304, USA Abstract. — Several lines of evidence suggest that numerous populations of cantils (Agkistrodon bi- lineatus, A. taylori), New World pitvipers with a distribution in Mesoamerica, are in rapid decline. We examined the IUCN conservation status for A. bilineatus, assessed for the entire range of the spe- cies, as well as the Environmental Vulnerability Scores (EVS) provided for certain countries along its distribution. Because of pronounced disparities in these conservation assessments and notable phenotypic differences that coincide with the geographic distribution of certain cantil populations, we conduct a taxonomic reassessment of the common cantil, Agkistrodon bilineatus (Gunther 1863), to determine if the recognized subspecies of A. bilineatus merit specific status. Based on our morphological assessment, biogeographical evidence, and the results of previous DNA-based studies, we elevate the three previously recognized subspecies of A. bilineatus to full species (A. bilineatus, A. russeolus, and A. howardgloydi). Given this taxonomic reassessment, we examine the conservation status of the newly elevated taxa, suggest avenues for future studies within this com- plex of pitvipers, and provide conservation recommendations. Key words. Character evolution, evolutionary species, Mesoamerica, subspecies concept Resumen. — Varias lineas de evidencia sugieren que numerosas poblaciones de cantiles ( Agkistrodon bilineatus, A. taylori), viboras de foseta del Nuevo Mundo con una distribucion en Mesoamerica, es- tan en rapido declive. Examinamos los resultados sobre el estado de conservacion propuestos por la UICN para A. bilineatus, que fueron evaluados para la distribucion total de la especie, asi como los resultados de los Indices de Vulnerabilidad Ambiental (en ingles, Environmental Vulnerability Scores [EVS]) que fueron determinados para esta especie en algunos paises a lo largo de su distri- bucion. Por haber disparidades pronunciadas en estas evaluaciones de conservacion y diferencias fenotipicas notables que coinciden con la distribucion geografica de ciertas poblaciones de can- tiles, en este trabajo realizamos una reevaluacion taxonomica del cantil comun, Agkistrodon biline- atus (Gunther 1863), para determinar si las subespecies reconocidas de A. bilineatus merecen el estatus de especie. Basado en nuestro analisis morfologico, evidencia biogeografica y los resulta- dos de anteriores estudios basados en ADN, elevamos las tres subespecies de A. bilineatus previa- mente reconocidas al nivel de especie (A. bilineatus, A. russeolus y A. howardgloydi). Tomando en cuenta esta nueva evaluacion taxonomica, examinamos el estado de conservacion de los taxones aqui elevados, hacemos sugerencias para estudios futuros dentro de este complejo de viboras de foseta y ofrecemos recomendaciones para su conservacion. Palabras claves. Evolucion de caracteres, especies evolutivas, Mesoamerica, concepto de subespecies Citation: Porras LW, Wilson LD, Schuett GW, Reiserer RS. 2013. A taxonomic reevaluation and conservation assessment of the common cantil, Agkistrodon bilineatus (Squamata: Viperidae): a race against time. Amphibian & Reptile Conservation 7(1): 48-73 (e63). Correspondence. Emails: 1 empub@msn.com (Corresponding author) 2 bufodoc@ aol.com gwschuett@yahoo.com 4 rreiserer@ gmail. com June 2013 I Volume 7 I Number 1 I e63 Amphib. Reptile Conserv. | http://amphibian-reptile-conservation.org 049 Porras et al. Although the restoration of tropical dry forest is still pos- sible, humanity will not give the globe back to its wild- land denizens, and old-growth tropical dry forest will never again cover large areas. Janzen 2004: 80. Introduction The common cantil ( Agkistrodon bilineatus) is a poly- typic species of North American pitviper with a variably fragmented distribution extending from extreme south- western Chihuahua and southern Sonora, Mexico, to northwestern Costa Rica, on the Pacific versant, and parts of the Yucatan Peninsula, northern Belize, Guatemala, and extreme western Honduras on the Atlantic versant; it also occurs in Las Islas Marias, an archipelago of four islands located about 100 km west of the state of Nayarit, Mexico (Gloyd and Conant 1990; Campbell and Lamar 2004; Lemos-Espinal and Smith 2007; Babb and Dugan 2008; Garcfa-Grajales and Buenorostro-Silva 2011; McCranie 2011). With few exceptions, the dominant vegetation zones occupied by A. bilineatus are dry for- est, deciduous forest, thorn scrub, and savanna, primarily areas of low relief that have been exploited heavily for irrigated agriculture and where this species mostly has become a rare snake; the elevational range of A. bilinea- tus extends from near sea level to about 1,500 m (Gloyd and Conant 1990; Conant 1992). Along the Pacific coast of Mesoamerica, tropical dry forests were reported as the most endangered of the major tropical ecosystems, with only 0.09% of that region afforded official conservation status (Janzen 1988). A quarter of a century after Janzen’s elucidative paper, aside from protected areas, dry forests throughout this region have continued to deteriorate. In a monographic study of the Agkistrodon complex, Gloyd and Conant (1990) provided an extensive review of the cantils, including information on their taxonomy, morphology, distribution, and aspects of their natural history. Based on multiple lines of evidence, Parkinson et al. (2002) conducted a phylogeographic analysis of the cantils and elevated A. b. taylori to the rank of full species, emphasizing that the loss of forested areas in the habitat of this species underscored the need for its conservation. More recently, Wilson et al. (2010) com- piled an extensive conservation assessment for the en- tire Mesoamerican herpetofauna, in which numerous authorities provided information on the status of can- tils. Although the methodological approaches of these authors varied, it was clear from the outcome that the conservation status of A. bilineatus showed dramatic differences when analyzed on a country by country or regional basis, since the reported or estimated IUCN rankings for this species extended the gamut from Least Concern to Critically Endangered (Lavin-Murcio and Lazcano 2010; Sasa et al. 2010). Some authors also Amphib. Reptile Conserv. | http://amphibian-reptile-conservation.org 050 provided Environmental Vulnerability Scores (EVS; a conservation measure developed and used by Wilson and McCranie 1992, 2004, and McCranie and Wilson 2002) for certain countries, and their results were more infor- mative. This measure provides a rough gauge of the theo- retical degree that herptofaunal species are vulnerable to environmental degradation; the scores at the upper end of the scale (ranging from 14 to 20) indicate a greater de- gree of concern (Wilson et al. 2013), and the EVS for A. bilineatus was reported as 15 for Honduras, Nicaragua, and Costa Rica, and as 16 for Belize (Sasa et al. 2010; Stafford et al. 2010; Sunyer and Kohler 2010; Townsend and Wilson 2010). Based on our field experiences, recent discussions with several colleagues working in regions where cantils occur, and information from the published literature, we echo the statements of several of the aforementioned au- thorities that in many regions A. bilineatus has declined significantly, largely as a result of human activities. Our principal goal in this paper is to reexamine the conservation status of A. bilineatus , inasmuch as the available information suggests that certain populations are declining or imperiled. In conservation biology the threat status of an organism typically is evaluated at the species level, so first we reevaluate the taxonomic status of the three subspecies of A. bilineatus ( bilineatus , rus- seolus, and howardgloydi ) to determine if any (or all) of them shows sufficient lineage divergence to warrant full species recognition. Accordingly, our conservation as- sessment develops from our taxonomic conclusions. Morphological Assessment Gloyd and Conant (1990) and Campbell and Lamar (2004) provided an extensive amount of biological in- formation on cantils, including excellent drawings of the scalation and pattern of the relevant taxa discussed in this paper, so we relied largely on these sources for our morphological assessment. Unlike previous views (see Gloyd and Conant 1990), the genus Agkistrodon now is restricted to the New World (see Molecular Assessment). As in other pitviper genera, Agkistrodon (sensu stricto) is characterized by the presence of a deep fa- cial pit, a vertically elliptical pupil, a large venom gland in the temporal region, and a canaliculated fang on the maxilla followed by a series of smaller replace- ment fangs. In Agkistrodon , however, the scales on the crown generally are large and plate-like, although often they are fragmented or contain partial sutures, and the skull is relatively broad and equipped with short fangs. Other characters include a pronounced canthus rostra- lis, the presence of a loreal scale in all members except A. piscivorus, a robust (or relatively robust) body, and a moderate to long tail. Scale characters such as supralabi- als, infralabials, and dorsal scale rows at midbody show little variation among the species, although the last of these characters is slightly higher in A. piscivorus. The June 2013 I Volume 7 I Number 1 I e63 Taxonomy and conservation of the common cantil number of ventral scales is lower in A. bilineatus and A. taylori than in A. contortrix and A. piscivorus, and the number of subcaudals is slightly lower in the latter two species. In Agkistrodon, some or most of the subcaudal scales are divided, and the terminal spine on the tail tip is turned downward in all the taxa except A. piscivorus. Moderate hemipenial differences have been reported among the taxa, but the similarities are more pronounced when comparing A. contortrix and A. piscivorus to A. bi- lineatus and A. taylori (Gloyd and Conant 1990; Malnate 1990). The tail tip of neonates and juveniles of all spe- cies of Agkistrodon is brightly colored and typically is yellow, white, or pink (Gloyd and Conant 1990). The coloration of the tail tip changes as animals mature, to a faded yellow, green, gray, black, or sometimes to match the color of the dorsum. Young individuals often use their tail to lure prey (e.g., anurans, lizards) by way of vertical undulations and waving, a behavior termed “caudal lur- ing” (reviewed by Strimple 1988, 1992; Carpenter and Gillingham 1990). 1. The cantils Commonly known as cantils, A. bilineatus and A. taylori are thick-bodied pitvipers (Serpentes: Viperidae) with a large head and a moderately long and slender tail, and their maximum total lengths are similar. As in the other species of Agkistrodon , the scale characters of cantils only show a moderately low range of variation (Table 1). A wide range of color pattern variation is evident in Agkistrodon , and these characters were used to diag- nose the three subspecies of A. bilineatus (Burger and Robertson 1951; Gloyd 1972; Conant 1984), as well to elevate A. taylori to the rank of full species (Parkinson et al. 2000). The coloration of the head is distinctive, as cantils are adorned with five conspicuous pale stripes, one vertically on the front of the snout and two laterally on each side of the head. The dorsal color pattern con- sists of crossbands, at least in juveniles, and this char- acter shows a notable degree of geographic and ontoge- netic variation. The chin color and ventral coloration also demonstrate considerable geographic variation. 2. Color and pattern characteristics of the ornate cantil Among the cantils, the color pattern of A. taylori is the most vivid (Fig.l). The lower facial stripe is broad and extends to cover the lower edge of the supralabials, the dorsal pattern is composed of pronounced black cross- bands separated by gray or pale brown areas that often contain yellowish brown or orange, the chin is patterned with bold markings with wide white or yellow elements, and the venter contains dark gray or black markings Fig. 1 . Adult female Agkistrodon taylori from Aldamas, Tamaulipas, Mexico. The ornate cantil often is vividly marked. Photo by Tim Burkhardt. June 2013 I Volume 7 I Number 1 I e63 Amphib. Reptile Conserv. | http://amphibian-reptile-conservation.org 051 Porras et al. Table 1. Maximum total length and selected scale characters in the three subspecies of Agkistrodon bilineatus and in A. taylori. Min-max values are followed by the mean (in parentheses). Data derived from Gloyd and Conant (1990). Character A. b. bilineatus A. b. russeolus A. b. howardgloydi A. taylori Total length 1,090 mm 1,050 mm* 960 mm 960 mm Ventrals 127-143 (134.5) 131-141 (136.1) 128-135 (131.1) 127-138 (133.7) Subcaudals 52-71 (61.6) 46-62 (55.4) 54-61 (58.8) 40-56 (48.3) Supralabials 5-9 (8.1) 8-9 (8.0) 7-9 (8.0) 7-9 (8.0) Infralabials 9-13(10.7) 8-12(10.8) 9-12(10.9) 9-12 (10.4) Dorsal scale rows (midbody) 21-25 (22.9) 23-25 (23.1) 23-25 (23.4) 21-23 (22.9) * Specimen with an incomplete tail. arranged in a somewhat checkerboard pattern. In contrast to juveniles, adults exhibit a subdued pattern that con- tains brighter colors, but older individuals of both sexes tend to become melanistic, and sexual color dimorphism is present in all age classes (Burchfield 1982). The tail tip of young individuals has been reported as sulphur yellow, ivory white, or salmon pink (Burchfield 1982; Gloyd and Conant 1990); the tail tip of most young individuals, however, is sulphur yellow (LWP, GWS, pers. observ.; Fig. 2). Fig. 2. Neonate female Agkistrodon taylori born in captivity from adults collected in the state of Tamaulipas, Mexico. Sexual color pattern dimorphism is evident in all age classes, except in very old individuals that sometimes darken with age. In young males, the rhombs on the dorsum tend to form bands and the interstitial pattern is reduced. Photo by Breck Bartholomew. 3. Color and pattern characteristics of the common cantil In A. b. bilineatus, both the upper and lower facial stripes are relatively broad, and the lower stripe is continuous and bordered below by dark pigment along the mouth line. From a frontal view, the vertical stripe along the rostral and mental and the lateral head stripes often meet on the tip of the snout. In adults, the dorsal ground color ranges from very dark brown to black, and if crossbands are present often they are difficult to distinguish. The dorsal pattern consists of small white spots or streaks. The chin and throat are dark brown or black with a pat- tern of narrow white lines or markings, and the venter is dark brown or black with pale markings. The coloration of neonates and juveniles is some shade of brown, and consists of brown or chestnut crossbands separated by a paler ground color, with the lateral edges of the cross- bands flecked with white. The crossbands gradually fade with maturity, however, as the overall dorsal coloration darkens (Fig. 3). The tail tip of neonates and juveniles has been reported in numerous publications as bright yel- low (e.g., Allen 1949; Gloyd and Conant 1990). Sexual color dimorphism has not been reported in any age class. In A. b. russeolus, the upper facial stripe is narrow and sometimes is intermittent posterior to the eye, and the lower stripe is broader and continuous and separated from the commissure by a band of dark pigment. From a frontal view, the vertical stripe along the rostral and men- tal and the two upper lateral head stripes typically meet on the tip of the snout. The dorsal ground color of adults generally is pale reddish brown, and the pattern consists of broad, deep reddish brown to brown crossbands that are separated dorsally by areas of paler coloration, and often are edged irregularly with white. The crossbands remain apparent, even in older adults. Laterally, the cen- ters of the crossbands are paler and usually contain one or two dark spots. The pattern on the chin and throat of- ten is reduced, with small whitish spots or lines present on a darker background. Approximately the median third of the venter lacks a pattern or contains a few markings. The coloration of a neonate (150 to 175 mm TL) col- lected near Merida, Yucatan, was described from life by Howard K. Gloyd (Gloyd and Conant 1990: 83) as showing a velvety appearance, and its pattern consisted of rich chestnut-brown crossbands with rufous brown interspaces, which were edged with blackish brown and interrupted lines of white, “and the tip of the tail gray.” A neonate from Dzibilchaltun, Yucatan, showed a similar coloration except that the banding was edged intermit- tently only with white, and the tail tip was pale gray with faint white banding (Fig. 4). This individual was main- tained in captivity and by the time it had grown to a total June 2013 I Volume 7 I Number 1 I e63 Amphib. Reptile Conserv. | http://amphibian-reptile-conservation.org 052 Fig. 3. Young adult Agkistrodon b. bilineatus from Apatzingan, Michoacan, Mexico, at an elevation of 330 m. Adult individuals from much of the west coast of Mexico often lose the dorsal banding (see cover of this issue). Photo by Javier Alvarado. Fig. 4. Neonate Agkistrodon bilineatus russeolus from Dzibilchaltun, Yucatan, Mexico. Note the pale gray tail tip with faint white banding, and the overall dorsal color pattern. Photo by Javier Ortiz. Fig. 5. Juvenile (ca. 400 mm TL) Agkistrodon bilineatus russeo- lus from Dzibilchaltun, Yucatan, Mexico (same individual as in Fig. 4). With growth, the inner portion of the crossbands turned the same color as the interspaces, and the snake’s pattern de- veloped a more fragmented appearance. Photo by Javier Ortiz. length of ca. 400 mm, a marked transformation in color pattern had taken place (Fig. 5). With growth, the inner portion of the crossbands gradually turned the same pale color as the interspaces and the individual’s pattern de- veloped a more fragmented appearance; the color of the tail tip also shifted to include darker gray tones (Fig. 5). Henderson (1978) reported the dorsal pattern of a pre- served young individual (ca. 380 mm) from Orange Walk Town, Orange Walk District, Belize, as faintly banded, and the tail as grayish-yellow with faint narrow bands. Although Gloyd and Conant (1990: 83) reported the tail tip of an individual from the same locality as “bright green,” they did not indicate the total length of the snake and an ontogenetic color shift might have occurred. The fragmentation of the banding in A. b. russeolus is appar- ent in the photograph of an adult collected in the outskirts of Consejo, Corozal, Belize (Fig. 6). Sexual color dimor- phism has not been reported in juveniles or adults of A. b. russeolus. In A. b. howardgloydi, the upper facial stripe is narrow and the posterior part often is absent in adults, and the lower facial stripe is broader and usually divided into June 2013 I Volume 7 I Number 1 I e63 Amphib. Reptile Conserv. | http://amphibian-reptile-conservation.org 053 Taxonomy and conservation of the common cantil Fig. 6. Adult Agkistrodon bilineatus russeolus from the outskirts of Consejo, Corozal, Belize. Note the fragmented color pattern. Photo by Kevin Zansler, courtesy of Robert A. Thomas. Fig. 7. Adult Agkistrodon bilineatus howardgloydi from Volcan Telica, Leon, Nicaragua. The color pattern of individuals from this volcanic region often contains black pigment. Photo by Nony Sonati, courtesy of Javier Sunyer. components that sometimes meet at the suture between the second and third suprlabials, and below is bordered by a dark line; the lower edges of the supralabials also are pale in color. From a frontal view, of the five facial stripes only the top two generally meet on the tip of the snout, but in some individuals all five stripes are con- nected. The dorsal ground color of adults generally is reddish brown or brown. Adults with black pigment, however, are known from Reserva Natural Volcan Telica in northwest- ern Nicaragua, with a pattern con- sisting of darker crossbands that contrast moderately with the dorsal ground color, and along this volcanic area adults sometimes show a dark coloration (J. Sunyer, pers. comm.; Figs. 7, 8). A cantil also was sighted on the eastern shore of Laguna de Xiloa, north of Managua (R. Earley, pers. comm.). The chin and throat are orange yellow, bright orange, or brownish orange with a pattern of a few small white spots, but this col- oration terminates abruptly after the first few ventrals. The venter usually is dark reddish brown. The dorsal coloration of juveniles is tan to reddish orange, or reddish, with distinguishable reddish brown crossbands that are edged intermittently with white and/or black, especially as they approach June 2013 I Volume 7 I Number 1 I e63 Amphib. Reptile Conserv. | http://amphibian-reptile-conservation.org 054 Porras et al. Fig. 8. Young Agkistrodon bilineatus howardgloydi from Volcan Masaya, Masaya, Nicaragua. The color pattern of adults from this area sometimes darkens with age. Photo by Javier Sunyer. Fig 9. Juvenile (311 mm TL) Agkistrodon bilineatus howardgloydi from Parque Nacional Santa Rosa, Guanacaste, Costa Rica. Note the color pattern of the tail tip, which anteriorly to posteriorly turns from very dark to pale gray with corresponding pale gray to white interspaces. Photo by Alejandro Solorzano. the venter. The tail tip of juveniles is banded with a sequential pattern that ranges from very dark gray an- teriorly to paler gray toward the tip, with the interspaces alternating from pale gray to white (Fig. 9). Although Villa (1984: 19) indicated that in Nicaragua “the bright sulphur- yellow tail of the young becomes dark in the adult,” and a photograph of a “juvenile individual” of A. b. howardgloydi with what is indicated as a “yellowish tail” appears on the frontispiece, the robust body features of the snake clearly show that it is not a juvenile and its tail is not yel- low. We question, therefore, whether Villa might not have assumed that the tail color of A. b. howardgloydi would be yellow, as this information long was entrenched in literature re- garding A. b. bilineatus. With regard to sexual color dimorphism, unlike the other subspecies of A. bilineatus, sub-adults and adults of A. b. how- ardgloydi show a moderate degree of sexual color dimorphism; in indi- viduals from Costa Rica, females are distinctly banded and paler in overall coloration, whereas males tend to be darker, with their banding obscured (Figs. 10, 11). Metachrosis, the abil- ity to change color at will or under external stimuli (such as light), was observed in the holotype of A. b. howardgloydi (Conant 1984). The coloration of this individual was paler at night (LWP, pers. observ.). 055 June 2013 I Volume 7 I Number 1 I e63 Amphib. Reptile Conserv. | http://amphibian-reptile-conservation.org Taxonomy and conservation of the common cantil Fig. 10. Adult female Agkistrodon bilineatus howardgloydi from Colonia Jobo de la Cruz, Guanacaste, Costa Rica. The color pattern of subadults and adults is paler in females. Photo by Louis W. Porras. Fig. 11. Adult male A. b. howardgloydi (holotype) from 0.8 kilimeters north of Mirador Canon del Tigre, Parque Nacional Santa Rosa, Guanacaste, Costa Rica. The color pattern of subadults and adults is darker in males. Photo by Louis W. Porras. Amphib. Reptile Conserv. | http://amphibian-reptile-conservation.org 056 June 2013 | Volume 7 | Number 1 | e63 Porras et al. Molecular Assessment Gloyd and Conant (1990) recognized 33 taxa (spe- cies and subspecies) in Agkistrodon (sensu lato), with a distribution in the Old World and the New World, but subsequent studies using molecular (mtDNA) methods partitioned Agkistrodon and demonstrated that the name applies to a monophyletic group of species restricted to the New World (Knight et al. 1992; Kraus et al. 1996; Parkinson et al. 1997, 2002; Parkinson 1999; Castoe and Parkinson 2006; Malhotra et al. 2010). Agkistrodon cur- rently is viewed as containing four species, A. bilineatus, A. contortrix, A. piscivorus, and A. taylori (Parkinson et al. 2000; Campbell and Lamar 2004), although one sub- species of A. piscivorus and two of A. contortrix appear to constitute distinct species (Guiher and Burbrink 2008; Douglas et al. 2009). 1. Molecular studies of cantils Parkinson et al. (2000) provided the first phylogeo- graphic (mtDNA) analysis of cantils, and tested all of the recognized subspecies ( bilineatus , howardgloydi, rus- seolus, and taylori). Using maximum parsimony (MP) and maximum likelihood (ML) methods, these authors recovered the clades ( taylori + ( bilineatus ( howardgloydi -t- russeolus ))). Furthermore, based on additional lines of evidence (e.g., biogeography, morphology) they rec- ommended the elevation of taylori to full species status, whereas the remaining subspecies were thought to be more recently diverged (i.e., having shallower relation- ships). Using other mtDNA regions (ATPase 8 and 6), and both ML and Bayesian methods of analyses, Douglas et al. (2009) corroborated the results of Knight et al. (1992) and Parkinson et al. (2000) with respect to New World Agkistrodon, including the relationships of cantils, although in their study they lacked DNA samples of A. b. russeolus. 2. Current views of cantil systematics and taxonomy Despite efforts by the various aforementioned authori- ties, a considerable gap in our understanding of the tax- onomy and phylogeography of cantils remains. We at- tribute this outcome largely to insufficient sampling, based on the number of specimens used in their analy- ses and the number of localities sampled. For example, Knight et al. (1992) included only two samples of can- tils ( bilineatus and taylori ) and both lacked locality in- formation, although their samples of taylori presumably were collected in Tamaulipas, Mexico (A. Knight, pers. comm.). Similarly, Parkinson et al. (2000) reported on only seven samples of cantils, of which two lacked lo- cality data, and their respective samples of taylori (n = 2) and howardgloydi (n = 2) each came from the same locality (see Parkinson et al. 2000: table 2). In testing Amphib. Reptile Conserv. | http://amphibian-reptile-conservation.org 057 phylogeographic hypotheses in Agkistrodon, Guiher and Burbrink (2008) and Douglas et al. (2009) used extensive sampling of A. contortrix and A. piscivorus, and both studies used cantils as an outgroup. No new localities for cantils, however, were sampled. Presently, only limited mtDNA-based sequence data (no nuclear genes have been tested) are available for a handful of specimens of cantils. No definitive molecular information exists for the no min ate form, A. b. bilinea- tus (i.e., no study has provided precise locality informa- tion) and only one specimen of A. b. russeolus (Yucatan, Mexico) has been subjected to a DNA-based inquiry (Parkinson et al. 2000). Given the extensive range of can- tils, the limited number of specimens sampled and tested thus far (Mexico: Tamaulipas [no specific locality], Yucatan, [no specific locality]; Costa Rica: Guanacaste Province, Santa Rosa) is inadequate to provide a robust view of their phylogeography. Nonetheless, despite these deficiencies, the available molecular (mtDNA) evidence suggests that the three subspecies of cantils (A. b. bilin- eatus, A. b. howardgloydi, and A. b. russeolus ) can be diagnosed as separate evolutionary entities (per Wiley 1978, 1981). Character Mapping Character mapping is a powerful analytical procedure for producing information and gaining insights into character evolution, particularly with respect to origin, direction, and frequency (Brooks and McLennan 1991; Harvey and Pagel 1991; Martins 1996; Fenwick et al. 2011; Maddison and Maddison 2011). Ideally, characters (traits) should be traced onto trees constructed from an explicitly independent data set (Harvey and Pagel 1991; Maddison and Maddison 2011), such as morphological characters mapped onto trees constructed using mol- ecules (e.g., proteins, DNA). 1. Methods We conducted a character mapping analysis (CMA) of the cantils by using morphological data derived from the literature (Gloyd and Conant 1990; Campbell and Lamar 2004), new information presented in this paper, and un- published personal data on all species of Agkistrodon (sensu stricto) (see Appendix 1). All characters were coded as binary (i.e., 0, 1) or multi-state (e.g., 0, 1, 2). Non-discrete multi-state characters (e.g., color pattern) were ordered from lowest to highest values. Character polarity was established by using two congeners (A. con- tortrix and A. piscivorus ) as outgroups. The cottonmouth (A. piscivorus) is confirmed as the sister group to cantils (Douglas et al. 2009). Ten characters were selected as potential apomorphies (shared-derived traits) and were traced onto a fully resolved tree (six taxa) based on the mtDNA-markers used in Parkinson et al. (2000) and Douglas et al. (2009). Character tracing was performed June 2013 I Volume 7 I Number 1 | e63 Taxonomy and conservation of the common cantil separately for each of the 10 traits using outgroup analy- sis and parsimony procedures in Mesquite (Madison and Madison 2011), and then combining the individual re- sults onto a global tree. 2. Results and discussion We found 10 morphological characters (scutellation, color pattern traits) selected for the CMA useful in pro- viding broad support for the topology of the molecular tree, as well as robust evidence for the distinctiveness of the taxa, in particular the three subspecies of A. bilinea- tus (Table 2). We thus assign these characters as putative synapomorphies and autapomorphies for Agkistrodon (Fig. 12). Although we had a priori knowledge of spe- cific and unique traits used to originally diagnose each of the subspecies, the CMA presents them in a phylogenetic and temporal framework. Accordingly, we show trait evolution with respect to origin, direction, and frequency. For example, we recovered dark dorsal coloration (dark brown or black) as the putative ancestral condition of Agkistrodon (Outgroup 1), which is retained in the basal- most cantils (A. taylori and A. b. bilineatus ), but evolved to reddish-brown in the sister clade A. b. howardgloydi + A. b. russeolus. These types of data can be used in CMA to test explicit hypotheses concerning adaptation, such as seeking correlations of body color to climate, habitat types, and a range of other variables (e.g., Martins 1996). Allopatry in A. bilineatus In prioritizing a list of vipers for future conservation mea- sures, Greene and Campbell (1992: 423) considered A. bilineatus (sensu lato) a taxon of special interest because of its “highly fragmented and biogeographically interest- ing distribution.” Parkinson et al. (2002) also commented on the relictual nature of the distribution of cantils, and used allopatry as one of their criteria for elevating A. b. taylori to species level. As presently understood, the distribution of A. b. bilineatus extends along the Pacific coast of Mexico (including the offshore Las Islas Marias) and northern Central America, from extreme southwestern Chihuahua and southern Sonora to central El Salvador; inland in Mexico, this species has been recorded in northwest- ern and extreme southeastern Morelos, as well as in the Rio Grijalva Valley (Central Depression; Johnson et al. 2010) of Chiapas (Gloyd and Conant 1990; Campbell and Lamar 2004; Castro-Franco and Bustos Zagal 2004; Herrera et al. 2006; Lemos-Espinal and Smith 2007; Garcfa-Grajales and Buenorostro-Silva 2011). McCranie (2011) included a photograph of a cantil from extreme western Honduras (Copan, Copan). Based on that pho- tograph, and others provided to us by the collector (R. Garrado, pers. comm.) taken after the animal had reached maturity, the color pattern characteristics of this individ- ual are most similar to those of A. b. bilineatus (Fig. 13). Amphib. Reptile Conserv. | http://amphibian-reptile-conservation.org 058 Table 2. Morphological characters used in the character map- ping analysis (Fig. 12). See text for details. Character State Designation Facial striping absent A0 present Al Upper facial stripe absent B0 variable B1 broad B2 narrow B3 Adult coloration tan CO black/dark brown Cl reddish-brown C2 Adult dorsal band no DO (same as ground color) yes D1 Adult dorsal band brown E0 color (when present) black/dark brown El multi-colored E2 reddish-brown E3 Throat color ground-color F0 cream/white FI multi-colored F2 dark F3 brown F4 yellow-orange F5 Juvenile to adult slight GO color ontogeny pronounced G1 moderate G2 Neonate tail-tip color yellow HO gray HI Neonate tail pattern slight 10 moderate 11 pronounced 12 Sexual color absent JO dimorphism present J1 A photograph of what appears to be A. b. bilineatus , with a locality of Honduras, also appears in Kohler (2001: fig. 264). The distribution of A. b. russeolus primarily extends along the outer part of the Yucatan Peninsula, from west- central Campeche and the northern portion of Yucatan and Quintana Roo on the Gulf side, and in northern Belize on the Caribbean side, although isolated records are avail- able from extreme southeastern Campeche and central Peten, Guatemala (Gloyd and Conant 1990; Campbell 1998; Campbell and Lamar 2004; Kohler 2008). The southernmost population of cantil (A. b. howardgloydi ) June 2013 I Volume 7 I Number 1 I e63 Porras et al. AO- JO Fig. 12. Character mapping analysis of morphological traits in cantils (A. b. bilineatus, A. b. howardgloydi, A. b. russeolus, and A. taylori ). Outgroup 1 = A. piscivorus; Outgroup 2 = A. contortrix. See Table 2 and Appendix 1 . Fig. 13. Young adult Agkistrodon b. bilineatus from La Chorcha Lodge, Copan, Honduras at an elevation of 610 m (2,000 feet). Two sighting of this species have occurred at the lodge, in 2003 and 2008. Photo by Robert Gallardo. occurs along the Pacific coast of Central America from Isla Zacate Grande, in the Golfo de Fonseca, and the adjacent mainland of south- ern Honduras to the southern limit of Parque Nacional Santa Rosa Park in northwestern Costa Rica (Sasa and Solorzano 1995). The taxonomic assignment of certain populations of A. bilineatus, however, remains problematical. A single individual of cantil was re- ported from north of Palma Sola, in central coastal Veracruz, an area disjunct from that of all other popu- lations (Blair et al. 1997). Smith and Chiszar (2001) described the specimen as a new subspecies (A. b. lemosespinali ), but Campbell and Lamar (2004: 266) indicated that this taxon “was diagnosed by sev- eral characteristics, all of which are within the normal range of variation for A. taylori or might be artifacts in a specimen preserved for more than 30 years.” After examining additional specimens of A. taylori from Hidalgo and Veracruz, how- ever, Bryson and Mendoza-Quijano (2007) concluded that the speci- men was most closely related to, if not conspecific with, A. b. bilinea- tus, but that it also differed from all of the subspecies of A. bilineatus in its tail length to total length ra- tio. Bryson and Mendoza-Quijano (2007) further commented that the presence of A. bilineatus in coastal Veracruz lends corroboration to the transcontinental dispersal hypoth- esis presented by Parkinson et al. ( 2002 ). Another isolated population is known from the Atlantic versant of central Guatemala, from the Rfo Chixoy (Negro) Valley (Campbell and Lamar 1989). Gloyd and Conant (1990) commented that two speci- mens from this area show similari- ties in color pattern to each of the three populations of A. bilineatus occurring in Central America. Until additional specimens and/or molec- ular data are available, however, the taxonomic status of this allopatric population is uncertain and remains for future investigation. Similarly, June 2013 I Volume 7 I Number 1 I e63 Amphib. Reptile Conserv. | http://amphibian-reptile-conservation.org 059 Taxonomy and conservation of the common cantil the population in the Central Depression of Chiapas, Mexico, and adjacent western Guatemala merits further examination. In summary, the distribution of A. bilineatus is disjunct or fragmented throughout its extensive range, and thus we contend that three identifiable areas of its distribution are biogeographically distinct. Except for certain issues that remain unresolved (see Discussion), these regions of allopatry constitute the ranges of A. b. bilineatus , A. b. russeolus, and A. b. howardgloydi (see Distribution Map [Fig. 14] below). Our Taxonomic Position Six decades ago, Wilson and Brown (1953) discussed the recognition of subspecies in biology and were among the first to advocate, with compelling academic vigor, to halt the use of trinomials in taxonomy. Since their provoca- tive paper was published, a flurry of literally hundreds of papers on the utility of infraspecific categories has ap- peared, of which many applauded the insights of Wilson and Brown (1953) and supported abandoning the recogni- tion of subspecies (e.g., Edwards 1954; Donoghue 1985; Ball and Avise 1992; Douglas et al. 2002; Zink 2004), whereas others criticized their views as biologically short sighted (e.g., Sibley 1954; Durrant 1955; Crusz 1986; Mallet 1995). Even with the application of an integrative taxonomic approach (reviewed by Padial and de la Riva 2010), a unified concept of species and consequences for solving the problems of species delimitation (see de Queiroz 2007), or a general species concept approach as presented by Hausdorf (2011), no perfect solutions are available to resolve all of the conflicting viewpoints. Nevertheless, Padial and de la Riva (2010: 748) argued that on the basis of the evolutionary species concept, “the point of separation from [a] sister lineage is what marks the origin of a species... and neither subspecies nor ‘sub- speciation’ are logically needed.” Importantly, this state- ment implies that there are no “stages of speciation,” i.e., subspecies are not “on their way” to becoming species. We also share the opinion of Johnson et al. (2010: 327), who asserted that the species level is “the lowest evolu- tionary lineage segment that should be used in a formal phylogenetically based taxonomy... In this modern taxo- nomic hierarchy, all taxa except for subspecies are hy- pothesized to consist of separate evolutionary lineages, and thus subspecies should not be recognized as a formal taxonomic unit.” Moreover, today new subspecies rarely are described in most major zoological journals, although many authors retain already-recognized subspecies as a provisional measure (e.g., Oatley et al. 2011). Here, we adopt the position on subspecies outlined by Wilson and Brown (1953) and subsequently supported by hundreds of biologists (reviewed by Burbrink et al. 2000; Douglas et al. 2002; Johnson et al. 2010). Amphib. Reptile Conserv. | http://amphibian-reptile-conservation.org 060 Taxonomic Conclusions The taxonomic overview and analysis we provide for the three putative subspecies of the common cantil (A. b. bilineatus, A. b. russeolus, and A. b. howardgloydi ) substantiates that sufficient morphological (color and pattern), molecular (mtDNA), and ecological (biogeo- graphical) data are available to consider these taxa as separate and diagnosable entities with their own evolu- tionary trajectories (see Wiley 1981; Wiley and May den 2000; Douglas et al. 2002). As we view it necessary to adopt and identify a species concept (Padial and de la Riva 2010), we used the evolutionary species concept (ESC) introduced by Wiley (1978, 1981). We agree with others that the ESC is preferred among the species hy- potheses, since it best accommodates both morphologi- cal and molecular information (Wiley and May den 2000; Schwentner et al. 2011). Accordingly, we elevate the three subspecies of A. bilineatus to full species and suggest the following com- mon names: Agkistrodon bilineatus (common cantil), A. russeolus (Yucatecan cantil), and A. howardgloydi (southern cantil). We indicate the reported localities for all the cantils, including A. taylori, in a distribution map (Fig. 14). Conservation Assessment Up to 2006, the conservation status of Agkistrodon bili- neatus (sensu lato) was judged by the IUCN as Least Concern, but in 2007, presumably as a result of the rep- tile assessment undertaken in September 2005, in Jalisco, Mexico, the status was changed to Near Threatened (IUCN Red List website; accessed 20 February 2013). Given that we elevated each of the three subspecies of A. bilineatus to full species, we will assess their conserva- tion status individually. 1. Application of the IUCN rankings The IUCN categories for assigning conservation status are the most widely used scheme for attempting to as- sess the degree of extinction risk for taxa at the species level (www.iucnredlist.org). The criteria used for this assessment are stipulated in the Guidelines for Using the IUCN Red List Categories and Criteria (Version 8.1; August 2010). Those with the greatest application to Mesoamerican reptile populations involve the extent of occurrence (i.e., geographic range), and at least two criteria regarding the degree of range fragmentation, the degree of decline in one of a number of distributional or populational characteristics, or the degree of fluctuations in any of these characteristics. The extent of occurrence is related to the threat categories as follows: Critically Endangered (< 100 km 2 ); Endangered (< 5,000 km 2 ); and Vulnerable (< 20,000 km 2 ). June 2013 I Volume 7 I Number 1 I e63 Porras et al. Q Agkistrodon t&ylori £ Agkistrodon bilineatus O Agkistrodon russeolus © Agkistrodon howardgloydi O Undetermined San Juanita Maria Madre Q' . I M^rra Magdalena ? <] Man’s Cleofas Las [si as Manas = IMI \ IH 2in» mi Fig. 14. Distribution map of the reported localities for cantils, including some indicated in this paper. Green is used to designate localities from where we regard the systematic status of cantils as undetermined. Under our new taxonomic arrangement, the distribu- tion of A. bilineatus (sensu stricto) is extended to include extreme western Honduras, in the vicinity of the city of Copan on the Caribbean versant (McCranie 2011). Thus, its extent of distribution well exceeds the 20,000 km 2 that forms the upper cutoff for a Vulnerable species; it also is greater than the 250,000 km 2 indicated by Garcia (2006) as the combined extent of the six dry forest ecore- gions in Pacific coastal Mexico, in addition to its range in Central America. Given its approximate geographic distribution, it clearly lies outside of the upper size limits for any of the IUCN threat categories. In addition, this species does not appear to qualify as Near Threatened, given that “the taxon should be close to qualifying for the Vulnerable category. The estimates of population size or habitat should be close to the Vulnerable thresholds, especially when there is a high degree of uncertainty” (IUCN 2010: 63). If, however, A. bilineatus cannot be judged as Near Threatened, only three other categories are available, viz., Extinct, Least Concern, and Data Deficient. The species is not Extinct, or as we maintain in this paper not of Least Concern, and also does not clas- sify as Data Deficient because enough information was Amphib. Reptile Conserv. | http://amphibian-reptile-conservation.org 061 available for it to be judged as Near Threatened (Lee and Hammerson 2007). In light of this information, we con- tend that A. bilineatus (sensu stricto) should be judged as Near Threatened. A broad-scale assessment of this snake’s conservation status throughout its distribution is extremely critical, however, since much of its area of occurrence has been subjected to considerable human population growth. In Mexico, A. bilineatus primarily occurs in the coastal portion of nine states from Sonora to Chiapas, as well as in Morelos. According to information obtained from Wikipedia (www.wikipedia.org), here and else- where in this section, these 10 states have a combined human population of 33,432,935 (29.0% of the 2012 population of Mexico). With a growth rate of 1.4% for the country (Population Reference Bureau 2010) and an estimated doubling time of 50 years, if these growth rates remain comparable the population of these states will reach 66,865,870 by the year 2063. Although these figures and projections apply to an area greater than the total range of A. bilineatus in Mexico, they signal grave concern for the survival of these populations. June 2013 I Volume 7 I Number 1 I e63 Taxonomy and conservation of the common cantil The prospects for the future of A. bilineatus in Guatemala and El Salvador are equally as disturbing. Guatemala is the most rapidly growing country in Central America, with a human population 13,824,463 in 2011, a growth rate of 2.8%, and an estimated doubling time of 25 years, and El Salvador already has become the most densely populated region in Mesoamerica. These statis- tics, therefore, portend a gloomy picture for the flora and fauna of these countries. Consequently, in light of these data, we consider A. bilineatus as Near Threatened, while conceding that fu- ture population analyses might demonstrate a threatened status. The distribution of A. russeolus is much greater than 100 km 2 (the upper cutoff point for a Critically Endangered species), but significantly less than 5,000 km 2 (the upper cutoff point for an Endangered species). Thus, based on the extent of occurrence, A. russeolus should be judged as an Endangered species. According to the maps in Gloyd and Conant (1990), Lee (1996), Campbell and Lamar (2004), and Kohler (2008), A. rus- seolus is known from up to twelve localities, depend- ing on the level of discrimination. Most of these locali- ties are from the state of Yucatan, from the vicinity of Merida, Motul, and Piste. Given this number of locations ( n = 12), A. russeolus should be assessed as Vulnerable, since the criterion for this category is < 10, as opposed to Endangered, which is < 5. These records are historical, however, with some dating prior to 1895 (sensu Gloyd and Conant 1990), and to our knowledge no modem sur- vey has been undertaken to ascertain the viability of can- til populations in these regions. The human population of the three Mexican states occupying the Yucatan Peninsula, Campeche, Yucatan, and Quintana Roo, is over 4,000,000 (Population Refer- ence Bureau 2010). Most of the historical records for A. russeolus are from the state of Yucatan, the most populous of the three with a current population of about 2,000,000. Specimens assigned to A. russeolus have been reported from seasonally dry forest in northern Belize, from Corozal and northern Belize Districts (Stafford and Meyer 2000), and the savanna area of central Peten, Guatemala (Campbell 1998). Lee and Hammerson (2007) indicated that the major factor affecting the long-term viability of populations of A. bilineatus (sensu lato) is “the extreme pressure from persecution leading to population reductions of close to 30% over the last 15 to 30 years...” According to J. Lee (pers. comm.), this evaluation cannot be applied precisely to A. russeolus, but would point to a Critically Endangered status based on criterion Cl, i.e., an estimate of continuing decline of at least 25% in 3 years or one generation (IUCN 2010). Lee (1996: 399) commented that, u Agkistrodon bilineatus [sensu lato] is a danger- ously venomous snake that is widely feared by the na- tive people of Yucatan. It is believed to be capable of Amphib. Reptile Conserv. | http://amphibian-reptile-conservation.org 062 prodigious jumps and to deliver venom both through its bite and with its tail, which is thought to act as a stinger...” Lee (1996: 416) also discussed the historical and the modern attitude toward snakes in general and A. russeolus (as A. bilineatus ) in particular, in his chapter on ethnoherpetology in the Yucatan Peninsula, indicating that the cantil or uolpoch (the Mayan name) “is consid- ered by many contemporary Maya to be the most danger- ous of all Yucatecan snakes.” This attitude translates into this snake being killed on sight (J. Lee, pers. comm.). Consequently, based on the available information on the conservation status of A. russeolus, we consider this spe- cies as Endangered. A conservation assessment needs to be undertaken, however, to determine if this categoriza- tion is appropriate, or whether the category of Critically Endangered would be more applicable. Agkistrodon howardgloydi is distributed in appar- ently fragmented populations that extend from Isla Zacate Grande in the Golfo de Fonseca and the adja- cent mainland of southern Honduras (McCranie 2011), western Nicaragua in the area west of Rio Tipitapa and the northwestern shore of Lago de Nicaragua (Kohler 1999, 2001), and in extreme northwestern Costa Rica from Bahia Salinas, near the Nicaraguan border, to the s sectors of Santa Rosa and Guanacaste, both in Area de Conservacion Guanacaste (Conant 1984; Solorzano 2004). Gloyd and Conant (1990: 92) discussed additional Nicaraguan localities that would extend the distribution northeastward into the southwestern tip of Departamento Jinotega, but this record is one of several supplied to the authors by Jaime Villa. Unfortunately, these specimens were in Villa’s “personal collection that was destroyed during the earthquake and fire that devastated Managua beginning on December 23, 1972.” Like Kohler (1999, 2001) , we discounted these records until museum speci- mens are available from those areas to provide verifica- tion. The extent of this species’ range, therefore, appar- ently is greater than 100 km 2 but less than 5,000 km 2 , so on the basis of its extent of occurrence it would be assessed as Endangered. With respect to the number of localities, three have been reported for Honduras, includ- ing one based on a photograph in Kohler et al. (2006), five from Nicaragua (Kohler 2001; a sight record in this paper), and five from Costa Rica (Conant 1984; Savage 2002) ; most of these localities in Costa Rica, however, fall within Parque Nacional Santa Rosa, so their total number could be considered as few as two. Thus the total number of localities would range from 10 to 13, which techni- cally would place this species in the Near Threatened cat- egory, but again historical records (Nicaragua) date back to 1871 (Gloyd and Conant 1990). As a consequence, this species would appear to fall in the Vulnerable category. Furthermore, given the localized distribution of A. how- ardgloydi in Costa Rica, it is noteworthy that this species was not reported from the country until 1970 (Bolanos and Montero 1970). June 2013 I Volume 7 I Number 1 I e63 Porras et al. Agkistrodon howardgloydi occurs in disjunct popula- tions in Honduras, Nicaragua, and Costa Rica, in low- land dry forest — the most endangered of the major for- est types in Mesoamerica (Janzen 2004). In Honduras, nearly all of this forest has been removed from the Pacific coastal plain. A telling feature in McCranie (2011: table 22) is that of the protected areas in Honduras currently supporting “some good forest,” not one contains lowland dry forest. Based on figures from 2001, the departments of Choluteca and Valle each rank among the top five in human population density in the country. As noted by Solorzano et al. (1999), M. Sasa was unsuccessful in finding this species at several localities in the Golfo de Fonseca and indicated that most of the locals were un- aware of its existence. These disturbing reports and ob- servations suggest that low population densities (or lo- cal extirpation) might be the trend. Similarly, McCranie (2011) noted that professional collectors in Choluteca failed to identify this species from photographs. Also, three of us (LWP, LDW, GWS) have been unsuccessful in finding this species on Isla Zacate Grande, in the Golfo de Fonseca, and on the adjacent mainland. According to Sunyer and Kohler (2010: 494), similar population trends prevail in Nicaragua, since A. how- ardgloydi (as A. bilineatus ) is restricted to lowland dry forest in the western part of the country, and “this forma- tion has undergone severe human alteration.” Although A. howardgloydi apparently occurs in at least three pro- tected areas, 75% of the protected areas in Nicaragua “contain less than 50% of their original forest cover...” (Sunyer and Kohler 2010: 505). The five known locali- ties for this species in Nicaragua (Kohler 2001; this pa- per) all are from the most heavily populated region in the country, an area that likely harbored more extensive populations of this species in the past. In Costa Rica, the conservation of A. howardgloydi is more promising, as most of the restricted range of ✓ this species is located within the Area de Conservacion Guanacaste. In this region, populations have been re- ported as “relatively stable and protected” (Solorzano 2004: 622). At Parque Nacional Santa Rosa, for exam- ple, 21 individuals were obtained for study from 1993 to 1996 (Solorzano et al. 1999). Nonetheless, Sasa et al. (2010: table 8) indicated that although the distribution of this species has been reduced by slightly more than 20% from a potential distribution of 6,883 km 2 , only a little more than 13% of that reduced distribution (5,465 km 2 ) is located within reserves. Like other venomous snakes, we can assume that this species is killed on sight in the 87% of the reduced range outside of protected areas. An important factor in this species’ favor is that the human population growth rate of Costa Rica (1.2%) is the lowest in Central America, and that Guanacaste Province, which encompasses the snake’s entire range in Costa Rica, is the most sparsely populated of all the provinces. Although the population of A. howardgloydi in pro- tected areas of Costa Rica apparently remains stable, Amphib. Reptile Conserv. | http://amphibian-reptile-conservation.org 063 throughout most of the range populations have been ex- tirpated (or are nearing extirpation). Thus, in light of the conservation prospects for A. howardgloydi, we consider this species as Endangered with the understanding that a range-wide conservation assessment is required, espe- cially in Honduras and Nicaragua. 2. Application of the EVS The conservation status algorithm known as the Environmental Vulnerability Score (EVS) was developed by Wilson and McCranie (1992) for use with amphibians in Honduras and subsequently applied to both amphib- ians and reptiles in this country (Wilson and McCranie 2004). The EVS was utilized in a broader fashion in most of the chapters dealing with Central American countries in Wilson et al. (2010), and in all cases used at the country level. As noted in the Introduction of this paper, the EVS for A. bilineatus (sensu lato) in four Central American countries fell within the upper end of the vulnerability scale (Wilson and McCranie 2004). Originally, the EVS algorithm was constructed for use strictly within Honduras, and thus had limited utility outside of that country. For example, the scale used for Honduras was as follows: 1 = widespread in and outside of Honduras 2 = distribution peripheral to Honduras, but wide- spread elsewhere 3 = distribution restricted to Nuclear Middle America (exclusive of Honduran endemics) 4 = distribution restricted to Honduras 5 = known only from the vicinity of the type locality In its original form, four of the five levels of this scale could not be used outside of Honduras. For the EVS to have a broader application, therefore, it required recon- struction and this recently was accomplished for Belize (Stafford et al. 2010), Nicaragua (Sunyer and Kohler 2010), and Costa Rica (Sasa et al. 2010). In order to use the EVS measure independent of coun- try divisions, it requires additional reconstruction, as follows: 1 = distribution extending from North America (United States and Canada) to South America 2 = distribution extending from North America to Mesoamerica or from Mesoamerica to South America 3 = distribution restricted to Mesoamerica 4 = distribution restricted to a single physiographic region within Mesoamerica 5 = known only from the vicinity of the type locality The other components of the gauge require only mini- mal reconstruction. The ecological distribution compo- nent can be revised as follows: 1 = occurs in eight or more formations 2 = occurs in seven formations 3 = occurs in six formations 4 = occurs in five formations June 2013 I Volume 7 I Number 1 I e63 Taxonomy and conservation of the common cantil 5 = occurs in four formations 6 = occurs in three formations 7 = occurs in two formations 8 = occurs in one formation The only modification of this component is that the first level was changed from “occurs in eight formations” to “occurs in eight or more formations” (see Wilson and McCranie 2004). This change appears acceptable, since very few species in Mesoamerica occupy more than eight formations (see Wilson and Johnson 2010: table 16). The component for the degree of human persecution in reptiles (a different measure was used for amphibians) is the same as used by Wilson and McCranie (2004), as follows: 1 = fossorial, usually escape human notice 2 = semifossorial, or nocturnal arboreal or aquatic, non-venomous and usually non-mimicking, sometimes escape human notice 3 = terrestrial and/or arboreal or aquatic, generally ig- nored by humans 4 = terrestrial and/or arboreal or aquatic, thought to be harmful, might be killed on sight 5 = venomous species or mimics thereof, killed on sight 6 = commercially or non-commercially exploited for hides and/or meat and/or eggs Based on these changes to the EVS, the calculated scores for the three species of cantils are as follows: A. bilineatus : 3 + 5 + 5 = 13 A. russeolus: 4 + 6 + 5 = 15 A. howardgloydi : 4 + 8 + 5 = 17 Consequently, the value for A. bilineatus falls at the up- per end of the medium vulnerability category, and the values for A. russeolus and A. howardgloydi fall into the high vulnerability category. In summary, the IUCN categorizations and EVS val- ues for these three taxa are as follows: A. bilineatus (Near Threatened and 13); A. russeolus (Endangered and 15); and A. howardgloydi (Endangered and 17). Interestingly, the IUCN has assessed A. taylori as a species of Least Concern (Lavin et al. 2007), whereas the EVS for this taxon is reported as 17 (Wilson et al. 2013). Discussion We provided a substantive review of the taxonomy and conservation status of the common cantil (A. bilineatus, sensu lato). Our taxonomic assessment led us to elevate the three subspecies of A. bilineatus to full species (A. bilineatus, A. howardgloydi, and A. russeolus), based on multiple lines of evidence. Nonetheless, we are not con- fident that this arrangement necessarily captures the full diversity of this widely distributed group of pitvipers. Amphib. Reptile Conserv. | http://amphibian-reptile-conservation.org 064 Accordingly, we identified several regions where ad- ditional sampling must be accomplished, but overall we recommend a thorough phylogeographic analysis employing morphological analyses and the use of both mtDNA and nuclear (e.g., introns, microsatellites) mark- ers. Owing largely to the isolation of certain populations, we suspect that additional species will be discovered within this complex. The population of A. bilineatus in southern Sonora and adjacent southwestern Chihuahua, Mexico, for ex- ample, occurs in a distinctive habitat (“Sonoran-Sinaloan transition subtropical dry forest” according to the WWF [see Garcia 2006]), the color pattern of adults differs somewhat from that of typical A. bilineatus (Fig. 15), and a moderate hiatus exists from the closest-known popu- lation to the south (49 miles [78.8 kilometers] south of Culiacan, Sinaloa, Mexico; Hardy and McDiarmid 1969; Campbell and Lamar 2004). Another example is the insular population on Las Islas Marias. On this offshore group of islands, two speci- mens collected in 1881 were reported from the “Tres Marias” (without naming a specific island), and one specimen from Isla Maria Grande was collected in 1 897 (Boulenger 1896; Stejneger 1899; see Zweifel 1960). Interestingly, Gloyd and Conant (1990) indicated that the cantil with the greatest total length is among these speci- mens, as well as the A. b. bilineatus (sensu lato) with the lowest number of subcaudals. Gloyd and Conant (1990), however, considered this latter specimen as aber- rant, but commented (p. 69) that “Whether other aberrant specimens occurred on the islands probably will never be known, inasmuch as the species may now have been extirpated from the archipelago.” Casas-Andreu (1992) indicated the presence of A. bilineatus on other islands of the Las Islas Marias chain (on Isla San Juanito and Isla Maria Magdalena). According to G. Casas-Andreu (pers. comm.), however, these records were not based on new material, as no cantils were encountered dur- ing his survey in 1986, but rather they were obtained from the literature. Inasmuch as no literature citations or museum numbers for these specimens appear in Casas- Andreu (1992), our knowledge of the distribution of A. bilineatus on Las Islas Marias remains sketchy. Although some areas of “good habitat” were present in the archi- pelago in 1986 (G. Casas-Andreu, pers. comm.), habitat destruction, a growing human population (including a large penal colony), the presence of agricultural camps and domestic animals, the outright killing of fauna, and the introduction of rats and feral cats all had become a significant problem (Casas-Andreu 1992). In 2000, the archipelago and its surrounding waters were declared an international protected area (Reserva de la Biosfera Islas Marfas). In spite of the lack of information on A. bilinea- tus from these islands, the only reptiles protected under the Secretarfa del Medio Ambiente y Recursos Naturales (SEMARNAT) are Crocodylus acutus (special protec- tion), Iguana iguana (special protection), Ctenosaura June 2013 I Volume 7 I Number 1 I e63 Porras et al. Fig. 15. Adult Agkistrodon bilineatus found by Larry Jones and Thomas Skinner in August of 2005, ca. 12 km NW of Alamos, Sonora, Mexico. This individual later was released. Photo by James C. Rorabaugh. Fig. 16. Young cantil from Aldea La Laguna, Nenton, Huehuetenango, Guatemala. The specific allocation of this population remains uncertain (see Fig. 14). Photo by Manuel Acevedo. pectinata (threatened), and Eretmochelys imbricata (in danger of extinction) (Anonymous 2007). A determina- tion of the actual distribution and population status of A. bilineatus on Las Islas Marias, therefore, is a conserva- tion priority. The taxonomic status of A. b. lemosespinali , which tentatively was assigned to A. b. bilineatus by Bryson and Mendoza-Quijano (2007), remains unresolved. Known from a single specimen from Palma Sola, in coastal central Veracruz, Mexico, this area was noted by Smith and Chizar (2001: 133) as highly agricultural and located next to a nuclear power plant regarded by “many local residents and environ- mentalists in general as having con- taminated the surrounding area with radioactivity.” These authors further indicated that if “A. b. lemosespinali ever occurred in that area, it is likely now to be extinct, or it likely would have been found [again] long ago.” Other disjunct populations of cantils merit a closer examination at both morphological and molecu- lar levels, such as those from the Central Depression of Chiapas and the headwaters of the Rio Grijalva that extend into northwestern Guatemala (Fig. 16), the Rio Chixoy and Motagua valleys of Guatemala, as well as isolated populations of A. russeolus (Gloyd and Conant 1990; Campbell and Lamar 2004; McCranie 2011). Assigning protected areas for the conservation of cantil populations is not simply a matter of determining regions that exist within the range of the three species, as these have been shown to vary in their level of protection. Jaramillo et al. (2010: 650) presented a model that could be used to analyze systems of pro- tected areas in Mesoamerica, and based on six requisites concluded that the system of protected areas in Panama is impressive due to the number of areas included and their collective territory; a detailed ex- amination of the features, however, demonstrated that all but one of the 97 areas failed, to some degree, “in meeting the necessary requirements for the long-term protection of its biotic resources.” In Honduras, McCranie (2011) indicated that, “at first glance, Honduras appears to have in place a robust system of protected areas, especially when compared to nearby countries. However, most of those areas exist on paper only.” Similarly, Acevedo et al. (2010) stated that, “the existing system of protected areas in Guatemala is insufficient to protect the country’s herpetofauna, be- cause most of the legally designated areas must be con- sidered as ‘paper parks’.” Essentially the same story can be told about systems of protected areas in other coun- tries where cantils occur (see various chapters in Wilson Amphib. Reptile Conserv. | http://amphibian-reptile-conservation.org 065 June 2013 | Volume 7 | Number 1 | e63 Taxonomy and conservation of the common cantil et at. 2010), an unfortunate aspect of reality in ongoing efforts to conserve biodiversity. Unfortunately, because of the continuing destruction of natural habitats and the potential for the extirpation of cantil populations, the answers to some of the aforemen- tioned questions are on the brink of being lost forever, if not lost already. This problem is critical, and we view it as a race against time to generate the necessary informa- tion that could help set aside protected areas to conserve disjunct and relictual populations of cantils for posterity. Conservation Recommendations Our recommendations for the long-term conservation of A. bilineatus, A. howardgloydi, A. russeolus , and A. tay- lori are as follows: 1. In light of the paucity of information regarding the relative health of populations of these species, it will be essential to undertake population assessments for all the cantils at or near localities where they have been recorded, most critically for A. howardgloydi and A. russeolus because of their relatively limited geographic ranges. 2. Once these surveys are completed, a conservation management plan should be developed to ascertain if populations of all four species are located within established protected areas, or if new areas should be considered. Such a plan is critical to the survival of cantils, especially since outside of protected areas these snakes generally are killed on sight or other- wise threatened by persistent habitat destruction or degradation. 3. Inasmuch as not all protected areas can be expected to provide adequate levels of protection to support viable populations of cantils, long-term population monitor- ing will be essential. 4. Given the elevation of these taxa to full species, conservation agencies can now use these vipers as “flagship species” in efforts to publicize conserva- tion efforts in their respective countries at all lev- els of interest and concern, including education and ecotourism. 5. We recommend the establishment of zoo conserva- tion (e.g., AZA) and outreach programs, such as those currently in progress for the venomous Guatemalan beaded lizard (e.g., www.ircf.org; see Domfnguez- Vega et al. 2012) and a wide variety of highly en- dangered anuran species (e.g., www.zooatlanta.org). Captive assurance colonies might help maximize fu- ture options for the recovery of wild populations. Amphib. Reptile Conserv. | http://amphibian-reptile-conservation.org 066 6. One major conclusion of this paper is that our knowl- edge of the taxonomy and phylogeography of cantils remains at an elementary level. Thus, as conservation assessments proceed, it will be important to obtain tis- sue samples from a sufficiently broad array of popu- lations to allow for more robust molecular analyses. Similarly, we need more detailed morphological as- sessments and more sophisticated levels of analyses, such as geometric morphometric approaches (Davis 2012 ). Acknowledgments. — We thank the following peo- ple for submitting or helping us obtain photographs for this paper: Manuel Acevedo, Javier Alvarado Dias, Breck Bartholomew, Tim Burkhardt, Eric Dugan, Robert Gallardo (La Chorcha Lodge), Javier Ortiz, James C. Rorabaugh, Alejandro Solorzano, Ireri Suazo- Ortuno, Javier Sunyer, Robert A. Thomas, R. Wayne Van Devender, and Kevin Zansler. Additionally, Chris Mattison graciously provided a photo of Agkistrodon bilinatus for the cover of this issue. We also are grate- ful to the following individuals for providing regional biological information on cantils: Gustavo Casas-Andreu (Las Islas Marias), Alec Knight (Tamaulipas), Javier Ortiz (Yucatan), Julian C. Lee (Yucatan Peninsula), Manuel Acevedo (Guatemala), Ryan Earley and Javier Sunyer (Nicaragua), and Mahmood Sasa and Alejandro Solorzano (Costa Rica). Lor other courtesies, we appre- ciate the efforts and cooperation provided by Vicente Mata-Silva, Robert A. Thomas, and Josiah H. Townsend. Lran Platt assisted with image cleanup and the layout of this paper. The molecular work we discussed was made possible through the courtesy of Michael Douglas and Marlis Douglas. Several anonymous reviewers provided valuable insights that helped to improve this paper. Over the years, numerous people have accompanied one or more of the authors into the field in search of cantils, and we reminisce about the good times spent with Ed Cassano, the late Roger Conant, W. W. Lamar, James R. McCranie, John Rindfleish, Alejandro Solorzano, and Mahmood Sasa. Of these, we are especially indebted to Roger Conant, for without his encouragement and in- spiration this paper might never have come to fruition. Beyond this, we wish to dedicate this paper to this re- markable man, whose influence has been so broadly felt in our own lives and among herpetologists far and wide.* *This paper is part of a special issue of Amphibian & Reptile Conservation that deals with the herpetofauna of Mexico. In ad- dition to Dr. Conant ’s seminal work on Agkistrodon (with Dr. Howard K. Gloyd), readers should be reminded that he also pro- duced important works on this country’s Nerodia (then Natrix ) and Thamnophis. June 2013 I Volume 7 I Number 1 I e63 Porras et al. 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Eagle Mountain Publishing, LC, Eagle Mountain, Utah, USA. Zink RM. 2004. The role of subspecies in obscuring avian biological diversity and misleading conservation policy. Proceedings of the Royal Society of London B 271: 561-564. Zweifel RG. 1960. Results of the Puritan-American Museum of Natural History expedition to western Mexico. 9. Herpetology of the Tres Marias Islands. Bulletin of the American Museum of Natural History 119 (article 2): 77-128. Received: 18 March 2013 Accepted: 24 May 2013 Published: 20 June 2013 June 2013 I Volume 7 I Number 1 I e63 Amphib. Reptile Conserv. | http://amphibian-reptile-conservation.org 070 Taxonomy and conservation of the common cantil Louis W. Porras is President of Eagle Mountain Publishing, LC, a company that has published Biology of the Vipers (2002), Biology of the Boas and Pythons (2007), Amphibians, Reptiles, and Turtles in Kansas (2010), Conservation of Mesoamerican Amphibians and Reptiles (2010), and Amphibians and Reptiles of San Luis Potosi (2013). For many years Louis served as Vice-President and President of the International Herpetological Symposium, and during his tenure was instrumental (along with Gordon W. Schuett) in launching the journal Herpetological Natural History. A native of Costa Rica, Porras has authored or co-authored over 50 papers in herpetology. During the course of his studies he has traveled extensively throughout the Bahamas and Latin America. Two taxa, Sphaerodactylus nigropunctatus porrasi, from the Ragged Islands, and Porthidium porrasi , from Costa Rica, have been named in his honor. Larry David Wilson is a herpetologist with lengthy experience in Mesoamerica, totaling six collective years (combined over the past 47). Larry is the senior editor of the recently published Conservation of Mesoamerican Amphibians and Reptiles and a co-author of seven of its chapters. He retired after 35 years of service as Professor of Biology at Miami-Dade College in Miami, Florida. Larry is the author or co- author of more than 290 peer-reviewed papers and books on herpetology, including the 2004 Amphibian & Reptile Conservation paper entitled “The conservation status of the herpetofauna of Honduras.” His other books include The Snakes of Honduras, Middle American Herpetology, The Amphibians of Honduras, Amphibians & Reptiles of the Bay Islands and Cay os Cochinos, Honduras, The Amphibians and Reptiles of the Honduran Mosquitia, and Guide to the Amphibians & Reptiles ofCusuco National Park, Honduras. He also served as the Snake Section Editor for the Catalogue of American Amphibians and Reptiles for 33 years. Over his career, Larry has authored or co-authored the descriptions of 69 currently recognized herpetofaunal species and six species have been named in his honor, including the anuran Craugastor lauraster and the snakes Cerrophidion wilsoni, Myriopholis wilsoni, and Oxybelis wilsoni. Gordon W. Schuett is an evolutionary biologist and herpetologist who has conducted extensive research on reptiles. His work has focused primarily on venomous snakes, but he has also published on turtles, lizards, and amphibians. His most significant contributions to date have been studies of winner-loser ef- fects in agonistic encounters, mate competition, mating system theory, hormone cycles and reproduction, caudal luring and mimicry, long-term sperm storage, and as co-discoverer of facultative parthenogenesis in non-avian reptiles. He served as chief editor of the peer-reviewed book Biology of the Vipers and is presently serving as chief editor of an upcoming peer-reviewed book The Rattlesnakes of Arizona (rattlesnakesofarizona.org). Gordon is a Director and scientific board member of the newly founded non- profit The Copperhead Institute (copperheadinstitute.org). He was the founding Editor of the journal Herpetological Natural History. Dr. Schuett is an adjunct professor in the Department of Biology at Georgia State University. Randall S. Reiserer is an integrative biologist whose research focuses on understanding the interrelation- ships among ecology, morphology, and behavior. Within the broad framework of evolutionary biology, he studies cognition, neuroscience, mimicry, life-history evolution, and the influence of niche dynamics on patterns of evolutionary change. His primary research centers on reptiles and amphibians, but his academic interests span all major vertebrate groups. His studies of behavior are varied and range from caudal luring and thermal behavior in rattlesnakes to learning and memory in transgenic mice. His studies of caudal luring in snakes established methods for studying visual perception and stimulus control. He commonly employs phylogenetic comparative methods and statistics to investigate and test evolutionary patterns and adaptive hypotheses. Dr. Reiserer is an editor of the upcoming peer-reviewed book, The Rattlesnakes of Arizona. Amphib. Reptile Conserv. | http://amphibian-reptile-conservation.org 071 June 2013 I Volume 7 I Number 1 I e63 Porras et al. Appendix 1 . Morphological characters of the subspecies of Agkistrodon bilineatus (ingroup) and two outgroups (A. contortrix and A.piscivorus) used for character mapping analysis in this study. Unless otherwise indicated, characters are based on adult stages. *Not used in analysis. Ingroup (cantils) Agkistrodon bilineatus bilineatus Upper facial stripe (lateral view): relatively broad and white. Lower facial stripe (lateral view): relatively broad and continuous with dark pigment below; white.* Dorsal coloration of adults: very dark brown to black; crossbands usually absent; if present, difficult to distinguish; pattern composed of small white spots or streaks. Chin and throat: dark brown or black, with narrow white lines or markings. Venter: dark brown or black with pale markings.* Coloration of neonates/juveniles: some shade of brown with crossbands separated by a paler ground color; lateral edges of crossbands flecked with white. Tail tip of neonates: bright yellow. Sexual color dimorphism: absent. Agkistrodon bilineatus howardgloydi Upper facial stripe (lateral view): narrow and white; posterior portion often absent in adults. Lower facial stripe (lateral view): broader than upper stripe, and divided into two components; stripe bordered below by dark line, followed by pale pigment to lower edge of supralabials; white.* Dorsal coloration of adults: reddish brown or brown; pattern of dark crossbands contrasts moderately with dorsal ground color. Chin and throat: orange yellow, bright orange, or brownish orange with few white spots. Venter: dark reddish brown.* Coloration of neonates/juveniles: tan to reddish orange, or reddish, with reddish brown crossbands edged intermit- tently with white and/or black, especially as they approach venter. Tail tip of neonates/juveniles: banded with sequential pattern ranging from very dark gray anteriorly to paler gray toward the tip, with interspaces alternating from pale gray to white. Sexual color dimorphism: moderate sexual color dimorphism present in sub-adults and adults. Agkistrodon bilineatus russeolus Upper facial stripe (lateral view): narrow and white; sometimes intermittent posterior to eye. Lower facial stripe (lateral view): broader than upper stripe and continuous, with narrow band of dark pigment below; white.* Dorsal coloration of adults: pale reddish brown; broad deep reddish brown to brown crossbands separated by paler areas, and strongly edged irregularly with white; crossbands remain apparent, even in older adults; laterally, centers of crossbands paler and usually contain one or two dark spots. Chin and throat: pattern often reduced; small whitish spots or lines evident on a darker background. Venter: approximately the median third is not patterned.* Coloration of neonates/juveniles: pattern of brown crossbands with paler brown interspaces; banding intermittently edged with white; with growth, inner portion of crossbands turns same color as interspaces, thereby developing a highly fragmented pattern. Tail tip of neonates/juveniles: pale gray with faint white banding; darker gray tones evident with growth. Sexual color dimorphism: absent. Agkistrodon taylori Upper facial stripe (lateral view): relatively broad and white. Lower facial stripe (lateral view): broad and continuous, and extends to lower edge of supralabials. Dorsal coloration of adults: pronounced black crossbands separated by gray, pale brown, or lavender areas that often contain yellow-brown or orange.* Chin and throat: bold markings, with white, yellow and or orange elements. Venter: dark gray or black markings arranged in a somewhat checkerboard pattern. Amphib. Reptile Conserv. | http://amphibian-reptile-conservation.org 072 June 2013 | Volume 7 | Number 1 | e63 Taxonomy and conservation of the common cantil Coloration of neonates/juveniles: strongly patterned, but with markings like those of adults but less intense. Tail tip of neonates/juveniles: yellow (rarely, white). Sexual color dimorphism: present in all age classes; sometimes difficult to detect in older adults that darken. Outgroups Agkistrodon piscivorus (outgroup 1) Upper facial stripe (lateral view): variable in size and appearance; pale but not white. Lower facial stripe (lateral view): relatively broad and continuous with dark pigment below.* Dorsal coloration of adults: very dark brown to black; crossbands present in some populations, difficult to distin- guish; pattern composed of small white spots or streaks. Chin and throat: pale, cream to white. Venter: dark brown or black with pale markings.* Coloration of neonates/juveniles: pale ground color with pronounced bands; strong ontogenetic change Tail tip of neonates: bright yellow. Sexual color dimorphism: absent. Agkistrodon contortrix (outgroup 2) Upper facial stripe (lateral view): absent. Lower facial stripe (lateral view): absent.* Dorsal coloration of adults: light tan ground color; brown crossbands of varying size present. Chin and throat: tan; typically same as ground color of face and dorsum. Venter: pale tan with dark tan markings.* Coloration of neonates/juveniles: ground color pale tan; similar to adults but subdued. Tail tip of neonates: bright yellow. Sexual color dimorphism: absent. Amphib. Reptile Conserv. | http://amphibian-reptile-conservation.org 073 June 2013 I Volume 7 I Number 1 I e63 Dr. Daniel D. Beck (right) with Martin Villa at the Centro Ecologia de Sonora, in Hermosillo, Mexico. Dr. Beck is holding a near- record length Rio Fuerte beaded lizard ( Heloderma horridum exasperation). Photo by Thomas Wiewandt. July 2013 | Volume 7 | Number 1 | e67 Amphib. Reptile Conserv. | http://amphibian-reptile-conservation.org 074 Copyright: © 2013 Reiserer et al. This is an open-access article distributed under the terms of the Creative Com- mons Attribution-NonCommercial-NoDerivs 3.0 Unported License, which permits unrestricted use for non-com- mercial and education purposes only provided the original author and source are credited. Amphibian & Reptile Conservation 7(1): 74-96. Taxonomic reassessment and conservation status of the beaded lizard, Heloderma horridum (Squamata: Helodermatidae) Randall S. Reiserer, 1 2 Gordon W. Schuett, and 3 Daniel D. Beck 'The Copperhead Institute, P. O. Box 6755, Spartanburg, South Carolina 29304, USA department of Biology and Center for Behavioral Neuro- science, Georgia State University, 33 Gilmer Street, SE, Unit 8, Atlanta, Georgia, 30303-3088, USA 3 Department of Biological Sciences, Central Washington University, Ellensburg, Washington 98926, USA Abstract. — The beaded lizard ( Heloderma horridum) and Gila monster ( H . suspectum) are large, highly venomous, anguimorph lizards threatened by human persecution, habitat loss and degrada- tion, and climate change. A recent DNA-based phylogenetic analysis of helodermatids (Douglas et al. 2010. Molecular Phylogenetics and Evolution 55: 153-167) suggests that the current infraspecific taxonomy (subspecies) of beaded lizards underestimates their biodiversity, and that species status for the various subspecies is warranted. Those authors discussed “conservation phylogenetics,” which incorporates historical genetics in conservation decisions. Here, we reassess the taxonomy of beaded lizards utilizing the abovementioned molecular analysis, and incorporate morphology by performing a character mapping analysis. Furthermore, utilizing fossil-calibrated sequence diver- gence results, we explore beaded lizard diversification against a backdrop of the origin, diversifica- tion, and expansion of seasonally dry tropical forests (SDTFs) in Mexico and Guatemala. These for- ests are the primary biomes occupied by beaded lizards, and in Mesoamerica most are considered threatened, endangered, or extirpated. Pair-wise net sequence divergence (%) values were greatest between H. h. charlesbogerti and H. h. exasperatum (9.8%), and least between H. h. alvarezi and H. h. charlesbogerti (1%). The former clade represents populations that are widely separated in distribu- tion (eastern Guatemala vs. southern Sonora, Mexico), whereas in the latter clade the populations are much closer (eastern Guatemala vs. Chiapas, Mexico). The nominate subspecies ( Heloderma h. horridum) differed from the other subspecies of H. horridum at 5.4% to 7.1%. After diverging from a most-recent common ancestor ~35 mya in the Late Eocene, subsequent diversification (cladogen- esis) of beaded lizards occurred during the late Miocene (9.71 mya), followed by a lengthy stasis of up to 5 my, and further cladogenesis extended into the Pliocene and Pleistocene. In both beaded lizards and SDTFs, the tempo of evolution and diversification was uneven, and their current distribu- tions are fragmented. Based on multiple lines of evidence, including a review of the use of trinomi- als in taxonomy, we elevate the four subspecies of beaded lizards to full species: Heloderma alvarezi (Chiapan beaded lizard), H. charlesbogerti (Guatemalan beaded lizard), H. exasperatum Rio Fuerte beaded lizard), and H. horridum (Mexican beaded lizard), with no changes in their vernacular names. Finally, we propose a series of research programs and conservation recommendations. Key words. mtDNA, ATPase, nuclear genes, character mapping, genomics, seasonally dry tropical forests, reptiles Resumen. — El escorpion ( Heloderma horridum) y el monstruo de Gila (H. suspectum) son lagartijas grandes, anguimorfas, y muy venenosas que estan sufriendo diversas amenazas como resultado de la persecucion humana, degradacion y perdida del habitat y el cambio climatico global. Un analisis filogenetico reciente basado en ADN de este grupo (Douglas et al. 2010. Molecular Phylogenetics and Evolution 55: 153-167) sugiere que la actual taxonomia intraespecifica (subespecies) del es- corpion esta subestimando la diversidad biologica, y el reconocimiento de especies es justificable. Estos autores discuten la utilidad del enfoque denominado “conservacion filogenetica”, que hace hincapie en la incorporacion de la genetica historica en las decisiones de conservacion. En este estudio, reevaluamos la taxonomia del escorpion utilizando el analisis molecular antes mencionado e incorporamos la morfologia en un analisis de mapeo de caracteres. Asi mismo, con los resultados de la secuencia de divergencia calibrada con fosiles, se explora la diversificacion del escorpion en forma yuxtapuesta al origen, la diversificacion y la expansion de los bosques tropicales estacional- mente secos (SDTFs) en Mexico y Guatemala. Estos bosques son los principales biomas ocupados por los escorpiones, y en Mesoamerica la mayoria son considerados amenazados, en peligro o July 2013 | Volume 7 | Number 1 | e67 Amphib. Reptile Conserv. | http://amphibian-reptile-conservation.org 075 Reiserer et al. extirpados. Los valores de la secuencia de divergencia neta por pares (%) fueron mayores entre H. h. charlesbogerti y H. h. exasperatum (9,8%) y menores entre H. h. alvarezi y H. h. charlesbogerti (1%). El primer grupo representa a poblaciones que estan muy distantes una de la otra en su distri- bucion (este de Guatemala vs. sur de Sonora, Mexico), mientras que las poblaciones en el segundo grupo estan mucho mas relacionadas (este de Guatemala vs. Chiapas, Mexico). La subespecie de- nominada (Heloderma h. horridum) difirio de las otras subespecies de H. horridum entre un 5,4% a 7,1%. Despues de la separacion de un ancestro comun mas reciente, ~35 mda a finales del Eoceno, ocurrio una diversificacion (cladogenesis) posterior de Heloderma a finales del Mioceno tardfo (9,71 mda), seguida de un estancamiento prolongado de hasta 5 mda, con una cladogenesis posterior que se extendio hasta el Plioceno y Pleistoceno. En ambos grupos, escorpiones y bosques tropi- cales estacionalmente secos, los procesos de evolucion y diversificacion fueron desiguales, y su distribucion fue fragmentada. Hoy en dia, el escorpion esta distribuido de manera irregular a lo largo de su amplio rango geografico. Basandonos en varias lineas de evidencia, incluyendo una re- vision del uso de trinomios taxonomicos, elevamos las cuatro subespecies del escorpion al nivel de especie: Heloderma alvarezi (escorpion de Chiapas), H. charlesbogerti (escorpion Guatemalteco), H. exasperatum (escorpion del Rio Fuerte), y H. horridum (escorpion Mexicano), sin cambios en los nombres vernaculos. Por ultimo, proponemos una serie de programas de investigacion y recomen- daciones para su conservacion. Palabras claves. ADNmt, ATPasas, genes nucleares, mapeo de caracteres, genomica, bosque tropical estacionalmente seco, reptiles Citation: Reiserer RS, Schuett GW, Beck DD. 2013. Taxonomic reassessment and conservation status of the beaded lizard, Heloderma horridum (Squamata: Helodermatidae). Amphibian & Reptile Conservation 7(1): 74-96 (e67). The century-long debate over the meaning and utility of the subspecies concept has produced spirited print but only superficial consensus. I suggest that genuine con- sensus about subspecies is an impossible goal ... the sub- species concept itself is simply too heterogeneous to be classified as strict science. Fitzpatrick 2010: 54. Introduction The beaded lizard ( Heloderma horridum) is a large, high- ly venomous, anguimorph (Helodermatidae) squamate with a fragmented distribution in Mesoamerica that ex- tends from northwestern Mexico (Sonora, Chihuahua) to eastern Guatemala (Bogert and Martin del Campo 1956; Campbell and Vannini 1988; Campbell and Lamar 2004; Beck 2005; Beaman et al. 2006; Anzueto and Campbell 2010; Wilson et al. 2010, 2013; Dominguez- Vega et al. 2012). Among the reptilian fauna of this region, the beaded lizard (in Spanish, known as the “escorpion”) is well known to local inhabitants, yet its natural history is surrounded by mystery, notoriety and misconception. Consequently, it is frequently slaughtered when encoun- tered (Beck 2005). Adding to this anthropogenic pressure, beaded lizard populations, with rare exceptions (Lemos-Espinal et al. 2003; Monroy-Vilchis et al. 2005), occur primarily in seasonally dry tropical forests, SDTFs (Campbell and Lamar 2004; Beck 2005; Campbell and Vannini 1988; Dominguez- Vega et al. 2012), the most endangered biome in Mesoamerica owing to persistent deforesta- tion for agriculture, cattle ranching, and a burgeoning human population (Janzen 1988; Myers et al. 2000; Trejo and Dirzo 2000; Hoekstra et al. 2005; Miles et al. 2006; Stoner and Sanchez- Azofeifa, 2009; Williams-Linera and Lorea 2009; Beck 2005; Pennington et al. 2006; Wilson et al. 2010, 2013; Dirzo et al. 2011; De-Nova et al. 2012; Dominguez- Vega et al. 2012; Golicher et al. 2012). Furthermore, drought and fires escalate the above threats (Beck 2005; Miles et al. 2006), and recent predic- tive models of climate change show that the persistence of SDTFs in this region is highly dubious (Trejo and Dirzo 2000; Miles et al. 2006; Golicher et al. 2012). Despite its large size and charismatic nature, our knowledge of the ecology, geographical distribution, and status of populations of H. horridum remains lim- ited (Beck and Lowe 1991; Beck 2005; Ariano- Sanchez 2006; Douglas et al. 2010; Domiguez-Vega et al. 2012). Furthermore, based on multiple lines of evidence, a taxo- nomic reevaluation of this group of lizards is long over- due (Beck 2005; Douglas et al. 2010). Here, we continue the dialogue concerning the infra- specifc (subspecific) taxonomy and conservation status of beaded lizards. We reviewed recent publications by Beck (2005) and Dominguez-Vega et al. (2012), and aug- ment their conclusions based on personal (DDB) field re- search in Mexico. We reassess the taxonomic status of the populations of H. horridum using morphology, bio- geography, and a recent molecular-based (mtDNA, nDNA) analysis conducted by Douglas et al. (2010). Although Douglas et al. (2010) commented on the mo- July 2013 | Volume 7 | Number 1 | e67 Amphib. Reptile Conserv. | http://amphibian-reptile-conservation.org 076 Taxonomy and conservation of beaded lizards lecular diversity of Heloderma, especially in H. horri- dum, they did not provide explicit taxonomic changes. In this paper, therefore, we reevaluate and expand upon their conclusions. To gain insights into phenotypic (mor- phological) evolution of extant Heloderma , with em- phasis on H. horridum, we conduct a character mapping analysis (Brooks and McLennan 1991; Harvey and Pagel 1991; Martins 1996; Maddison and Maddison 2011), uti- lizing the phylogenetic information (trees) recovered by Douglas et al. (2010). Overview of Morphology and Molecules in the genus Heloderma 1. Morphological assessment Published over half a century ago, Bogert and Martin del Campo’s (1956) detailed and expansive monograph of extant and fossil helodermatid lizards remains the defini- tive morphological reference (reviewed in Campbell and Lamar, 2004; Beck, 2005), and it contains the diagno- ses and descriptions of two new subspecies (. Heloderma horridum alvarezi and H. h. exasperatum). Thirty-two years later, Campbell and Vannini (1988) described a new subspecies (H. h. charlesbogerti ), from the Rio Mo- tagua Valley in eastern Guatemala, in honor of Charles Bogert’ s pioneering work on these lizards. With few ex- ceptions, such as Conrad et al. (2010) and Gauthier et al. (2012), who examined higher-level relationships of the Helodermatidae and other anguimorphs, a modern phy- logeographic analysis of morphological diversity for ex- tant helodermatids is lacking. However, as we illustrate in our character mapping analysis, the morphological characters used by Bogert and Martin del Campo (1956) in diagnosing and describing the subspecies of beaded lizards, though somewhat incomplete, remains useful in analyzing phenotypic variation. 2. Diagnosis, description, and distribution of Heloderma horridum Diagnosis and description . — Bogert and Martin del Campo (1956) and Campbell and Vannini (1988) pro- vided diagnoses and descriptions of the subspecies of Heloderma horridum. Recent information on the biol- ogy, systematics, and taxonomy of H. horridum and H. suspectum is summarized and critiqued by Campbell and Lamar (2004) and Beck (2005), and Beaman et al. (2006) provided a literature reference summary of the Heloder- matidae. Presently, four subspecies of H. horridum are recognized (Figs. 1-5). Mexican beaded lizard: H. h. horridum (Wiegmann 1829) Rio Fuerte beaded lizard: H. h. exasperatum Bogert and Martin del Campo 1956 Amphib. Reptile Conserv. | http://amphibian-reptile-conservation.org 077 Chiapan beaded lizard: H. h. alvarezi Bogert and Mar- tin del Campo 1956 Guatemalan beaded lizard: H. h. charlesbogerti Campbell and Vannini 1988 The four subspecies of H. horridum were diagnosed and described on the basis of scutellation, color pattern, and geographical distribution, and we refer the reader to the aforementioned works for detailed descriptions and taxonomic keys. The characters used by Bogert and Mar- tin del Campo (1956) and Campbell and Vannini (1988) to diagnose the subspecies have been reevaluated as to their stability, albeit informally (Campbell and Lamar 2004; Beck 2005). Poe and Wiens (2000) and Douglas et al. (2007) discussed the problem of character stabil- ity in phylogenetic analyses. Kraus (1988), for example, commented that reasonable evidence for character stabil- ity, and thus its usefulness as a shared-derived character (apomorphy), was the occurrence of a discrete trait in adults at a frequency of 80% or greater. In our character mapping analysis using published morphological char- acters (discussed below), character stability was a major assumption. Consequently, further research is warranted for substantiation. Geographic distribution . — The geographic distribu- tion of Heloderma horridum extends from southern So- nora and adjacent western Chihuahua, in Mexico, south- ward to eastern and southern Guatemala (Campbell and Lamar 2004; Beck 2005; Anzueto and Campbell 2010; Domiguez-Vega et al. 2012). The Rio Fuerte Beaded Lizard (H. h. exasperatum) in- habits the foothills of the Sierra Madre Occidental, with- in the drainage basins of the Rio Mayo and Rio Fuerte of the Sonoran-Sinaloan transition subtropical dry forest in southern Sonora, extreme western Chihuahua, and north- ern Sinaloa (Campbell and Lamar 2004; Beck 2005). Its distribution closely matches the fingers of SDTFs within this region, but it has also been encountered in pine-oak forest at 1,400 m near Alamos, Sonora (Schwalbe and Lowe 2000). Bogert and Martin del Campo (1956) com- mented that as far as their records indicated, a consider- able hiatus existed between the distribution of H. h. exas- peratum (to the north) and H. h. horridum (to the south), but owing to the narrow contact between the supranasal and postnasal in H. h. horridum from Sinaloa, intergra- dation might be found in populations north of Mazatlan. Based on this information, Beck (2005: 24) stated, “...in tropical dry forest habitats north of Mazatlan, Sinaloa, H. h. exasperatum likely intergrades with H. h. horridum .” Definitive data on intergradation remains unreported, however, and published distribution maps have incorpo- rated that assumption (e.g., Campbell and Lamar 2004; Beck 2005). Campbell and Lamar (2004, p. 104) show a single example of H. suspectum from El Dorado in west-central Sinaloa, Mexico (deposited in the American Museum of Natural History [90786]), a locality 280 km south from northern records in Rio del Fuerte, Sinaloa. July 2013 | Volume 7 | Number 1 | e67 Reiserer et al. Fig. 1. A. Adult Rio Fuerte bended lizaid ( Heloderma h o f' fid it m exasperatum ) in a defensive display (Alamos, Sonoia). B. Adult Rio Fuerte beaded lizard raiding a bird nest (Alamos, Sonora). Photos by Thomas Wiewandt. Fig. 2. Adult Mexican beaded lizard ( H . h. horridum ) observed on 11 July 2011 at Emiliano Zapata, municipality of La Huerta, coastal Jalisco, Mexico. Photo by Javier Alvarado. July 2013 | Volume 7 | Number 1 | e67 Amphib. Reptile Conserv. | http://amphibian-reptile-conservation.org 078 Taxonomy and conservation of beaded lizards Fig. 3. Adult Chiapan beaded lizard ( Heloclerma horridum alvarezi ) from Sumidero Canyon in the Rio Grijalva Valley, east of Tuxtla Gutierrez, Chiapas, Mexico. Photo by Thomas Wiewandt. Fig. 4. Adult Guatemalan beaded lizard ( Heloderma horridum charlesbogerti ) from the Motagua Valley, Guatemala. Photo by Daniel Ariano-Sdnchez. Amphib. Reptile Conserv. | http://amphibian-reptile-conservation.org 079 July 2013 | Volume 7 | Number 1 | e67 Reiserer et al. Owing to this unusual location, we suggest a re-examina- tion of this museum specimen to verify its identity. Neo- nates and juveniles of H. h. exasperatum resemble adults in color pattern (Fig. 5a), but they show greater contrast (i.e., a pale yellow to nearly white pattern on a ground color of brownish-black). Also, their color pattern can be distinguished from that of adults (e.g., no yellow speck- ling between the tail bands), and an ontogenetic increase in yellow pigment occurs (Bogert and Martin del Campo 1956; Beck 2005). The Mexican beaded lizard ( H . h. horridum ), the subspecies with the most extensive distribution, occurs primarily in dry forest habitats from southern Sinaloa southward to Oaxaca, including the states of Jalisco, Nayarit, Colima, Michoacan, and Guerrero, and inland into the states of Mexico and Morelos (Campbell and Lamar 2004; Beck 2005). Monroy-Vilchis et al. (2005) Fig. 5. A. Juvenile Heloderma horridum exasperatum (in situ, / Alamos, Sonora, Mexico). Photo by Stephanie Meyer. B. Neonate Heloderma h. horridum (wild-collected July 2011, Chamela, Jalisco). Photo by Kerry Holcomb. C. Neonate Heloderma horridum alvarezi (Rio Lagartero Depression, extreme western Guatemala). Photo by Quetzal Dwyer. D. Neonate Heloderma horridum charlesbogerti (hatched at Zoo Atlanta in late 2012). Photo by David Brothers, courtesy of Zoo Atlanta. recorded an observation of this taxon at mid eleva- tions (e.g., 1861 m) in pine-oak woodlands in the state s of Mexico. Campbell and Vannini (1988), citing Alva- rez del Toro (1983), indicated the probability of areas of intergradation between H. h. horridum and H. h. al- varezi , in the area between the Isthmus of Tehuantepec s and Cintalapa, Chiapas. Nonetheless, Alvarez del Toro (1983) stated that individuals of beaded lizards with yel- low markings (a coloration character present in H. h. horridum ) are found in the region from Cintalapa to the Isthmus of Tehuantepec, as well as in dry areas along the coast from Arriaga (near the Isthmus of Tehuantepec) to Huixtla (near the Guatemalan border). Literature infor- mation on intergradation between these two subspecies is inconclusive and, therefore, will require further inves- tigation. Neonates and juveniles of H. h. horridum , like those of H. h. exasperatum , resemble adults in color pat- tern (Fig. 5b), but their color contrast is greater (Bogert and Martin del Campo 1956; Beck 2005). The Chiapan beaded lizard ( H . h. alvarezi ) inhab- its dry forests in the Central Depression (Rio Grijalva July 2013 | Volume 7 | Number 1 | e67 Amphib. Reptile Conserv. | http://amphibian-reptile-conservation.org 080 Taxonomy and conservation of beaded lizards Depression) of central Chiapas and the Rio Lagartero Depression in extreme western Guatemala (Campbell and Lamar 2004; Beck 2005; Johnson et al. 2010; Wil- son et al. 2010: p. 435). This taxon is unique among the subspecies in that it undergoes an ontogenetic increase in melanism, whereby it tends to lose the juvenile color pattern (Bogert and Martin del Campo 1956; Beck 2005). Neonates and juveniles often are distinctly marked with yellow spots and bands, including on the tail (Fig. 5c), whereas the color pattern of adults gradually transforms to an almost uniform dark brown or gray. Black individu- als, however, are uncommon. Yellow banding on the tail, a characteristic typical of the other subspecies of beaded lizards, (Fig. 2), is essentially absent in adults (Bogert and Martin del Campo 1956; Beck 2005). The Guatemalan beaded lizard ( H . h. charlesbogerti ) inhabits the Rio Motagua Valley, in the Atlantic versant of eastern Guatemala (Campbell and Vannini 1988). Re- cently, however, Anzueto and Campbell (2010) reported three specimens from two disjunct populations on the Pacific versant of Guatemala, to the southwest of the Motagua Valley. Neonates resemble adults in color pat- tern, though they tend to be paler (Fig. 5d). In summary, the distribution of H. horridum is frag- mented throughout its extensive range and corresponds closely with the patchy distribution of SDTFs in Mexico and Guatemala (Beck 2005; Miles et al. 2006; Domin- guez- Vega et al. 2012). The distribution of the Guate- malan beaded lizard ( H . h. charlesbogerti ) is distinctly allopatric (Campbell and Vannini 1988; Beck 2005; Ariano-Sanchez 2006; Anzueto and Campbell 2010). 3. Molecular assessment Douglas et al. (2010) provided the first detailed molec- ular-based (mtDNA, nDNA) analysis of the phylogeo- graphic diversity of helodermatid lizards, which is avail- able at www.cnah.org/cnah_pdf.asp. Two authors (GWS, DDB) of this paper were co-authors. Specifically, Doug- las et al. (2010) used a “conservation phylogenetics” approach (Avise 2005, 2008; Avise et al. 2008), which combines and emphasizes the principles and approaches of genetics and phylogeography and how they can be ap- plied to describe and interpret biodiversity. Methods . — Douglas et al. (2010) sampled 135 locality- specific individuals of Heloderma (48 H. horridum , 87 H. suspectum) from throughout their range (their ingroup). The outgroup taxa included multiple lineages of lizards and snakes, with an emphasis on anguimorphs. Based on both morphological and DNA-based analyses, all author- ities have recognized the extant helodermatid lizards as monotypic (a single genus, Heloderma ), and as members of a larger monophyletic assemblage of lizards termed the Anguimorpha (Pregill et al. 1986; Estes et al. 1988; Townsend et al. 2004; Wiens et al. 2010, 2012; Gauthier et al. 2012). This lineage includes the well-known va- ranids ( Varanus ), alligator lizards and their relatives Amphib. Reptile Conserv. | http://amphibian-reptile-conservation.org 081 (Anguidae), as well as such relatively obscure taxa as the Old World Lanthanotidae ( Lanthanotus ) and Shinisauri- dae ( Shinisaurus ), and the New World Xenosauridae (Xe- nosaurus). The mtDNA analyses in Douglas et al. (2010) were rooted with the tuatara ( Sphenodon punctatus), and Bayesian and maximum parsimony (MP) analyses were conducted using Mr. Bayes (Hulsenbeck and Rohnquist 2001 ). Douglas et al. (2010) used sequence data from both mi- tochondrial (mt) DNA and nuclear (n) DNA as molecular markers in their phylogenetic analyses. Specifically, they discussed reasons for selecting mtDNA regions ATPase 8 and 6, and the nDNA introns alpha-enolase (ENOL) and ornithine decarboxylase (OD). The utility of com- bining mt- and nDNAs (supertree) in recovering phylo- genetic signals has been discussed (Douglas et al. 2007, 2010), yet each of these markers and the procedure of combining sequence data have both benefits and pitfalls (Wiens 2008; Castoe et al. 2009). Long-branch attraction and convergence, for example, can result in misleading relationships (Bergsten 2005; Wiens 2008; Castoe et al. 2009). The tools for detecting and potentially correcting these problems have been discussed (e.g., Castoe et al. 2009; Assis and Rieppel 2011). Results and discussion. — Douglas et al. (2010) recov- ered the genus Heloderma as monophyletic (Heloder- matidae), with H. horridum and H. suspectum as sister taxa. In a partitioned Bayesian analysis of mtDNA, He- lodermatidae was recovered as sister to the anguimorph clade {Shinisaurus {Abronia + Elgaria )), which in turn was sister to the clade Lanthanotus + Varanus. Recent molecular studies of squamates by Wiens et al. (2012, see references therein) recovered a similar topology to that of Douglas et al. (2010). However, an extensive morpho- logical analysis by Gauthier et al. (2012) supported a tra- ditional topology of Heloderma as sister to varanids and Lanthanotus borneensis (see Estes et al. 1986; Pregill et al. 1988). In Douglas et al. (2010), a partitioned Bayes- ian analysis of the nuclear marker alpha-enolase (intron 8 and exon 8 and 9), however, recovered Heloderma as sis- ter to a monophyletic Varanus. Using a combined analy- sis of morphology (extant and fossil data), mitochondrial, and nuclear markers, Lee (2009) recovered Varanidae as sister to the clade Helodermatidae + Anguidae. In a com- bined approach, Wiens et al. (2010) recovered results that were similar to those of Lee (2009). A recent DNA-based analysis of Squamata by Pyron et al. (2013) examined 4151 species (lizards and snakes), and they recovered Helodermatidae as sister to the clade Anniellidae + An- guidae. Moreover, they recovered the clade Varanidae + Lanthanotidae as sister to Shinisauridae. How do systematists deal with this type of incon- gruity (discordance) in studies that use different types (e.g., morphology vs. molecular) of phylogenetic mark- ers? Recently, Assis and Rieppel (2011) and Losos et al. (2012) discussed the common occurrence of discordance between molecular and morphological phylogenetic July 2013 | Volume 7 | Number 1 | e67 Reiserer et al. analyses. Specifically, with respect to discordance, As- sis and Rieppel (2011) stated that, “...the issue is not to simply let the molecular signal override the morphologi- cal one. The issue instead is to make empirical evidence scientific by trying to find out why such contrastive signals are obtained in the first place.” We concur with their opinions, and thus further research is warranted to resolve such conflicts in the phylogeny of anguimorph squamates. Relationships among the four subspecies of H. hor- ridum recovered in the analysis by Douglas et al. (2010, p. 158-159, fig. 3a, b) are depicted in Fig. 6. This topol- ogy was derived from a partitioned Bayesian analysis of the mtDNA regions ATPase 8 and 6. The Gila monster (H. suspectum ) was the immediate outgroup. Two sets of sister pairs of beaded lizards were recovered: H. h. exas- peratum (HHE) + H. h. horridum (HHH), and H. h. al- varezi (HHA) + H. h. charlesbogerti (HHC). The current subspecific designations for H. horridum were robustly supported (concordant) by these genetic analyses. Un- like results obtained for Gila monsters ( H . suspectum ), haplotype and genotype data for H. horridum were both diverse and highly concordant with the designated sub- species and their respective geographic distributions. Douglas et al. (2010) generated pair-wise net sequence divergence (%) values based on their recovered relation- ships (Table 1, Fig. 6). The greatest divergence was be- tween HHE and HHC (9.8%), and the least between HHA and HHC (1%). The former pair represents populations widely separated in distribution (southern Sonora, Mex- ico vs. eastern Guatemala), whereas the latter are much more closely distributed (Chiapas, Mexico vs. eastern Guatemala). The nominate subspecies ( Heloderma h. horridum) differed from the other three subspecies of beaded lizards, from 5.4% to 7.1%. Table 1. Pair-wise net sequence divergence (%) values between the four subspecies of the beaded lizard ( Heloderma horridum ) derived from a partitioned Bayesian analysis of the mtDNA re- gions ATPase 8 and 6 (modified from Douglas et al. 2010, pp. 157-159, 163; fig. 3a, b, tables 1 and 3). Values in parentheses denote evolutionary divergence times, which represent mean age. Mean age is the time in millions of years (mya) since the most-recent common ancestor (tree node) and is provided for the sister clades HHE-HHH and HHA-HHC (Fig. 6). Beaded lizards and Gila monsters (H. suspectum ) are hypothesized to have diverged from a most-recent common ancestor in the late Eocene ~35 mya (Douglas et al. 2010, p. 163). Percent se- quence divergence was greatest for HHC-HHE, and was lowest for HHA-HHC. See text for further details. HHA HHC HHE HHH HHA HHC HHE 1% (3.02) 9.3% 9.8% HHH 5.4% 6.2% 7.1% (4.42) — HHA = H. h. alvarezi', HHC = H. h. charlesbogerti ; HHE = H. h. exas- peratum; HHH = H. h. horridum. Fig. 6. Character mapping analysis. Tree topology and node dates based on Douglas et al. (2010). Morphological characters (Table 2) were mapped via parsimony and outgroup methods using the software program Mesquite (Maddison and Maddison 2011). Node 1 = Late Eocene (—35 million years ago, mya); Node 2 = 9.71 mya; Node 3 = 4.42 mya; and Node 4 = 3.02 mya (see Table 1). See text for details of the analysis. July 2013 | Volume 7 | Number 1 | e67 Amphib. Reptile Conserv. | http://amphibian-reptile-conservation.org 082 Taxonomy and conservation of beaded lizards Table 2. Morphological characters used for the character mapping analysis (see Table 1, Fig. 6). See text for details. Character State Designation Tail length 41-55% of snout-to-vent length A0 > 65% of snout-to-vent length A1 Number of caudal vertebrae 25-28 B0 40 B1 Number of transverse rows of ventromedial absent CO caudal scales (vent to tail tip) greater than 62 present Cl Usually one pair of enlarged preanal scales present DO absent D1 First pair of infralabials usually in contact with present E0 chin shields absent El Number of maxillary teeth 8-9 F0 6-7 FI Upper posterior process of splenial bone overlaps inner surface of coronoid GO does not overlap coronoid G1 Number of black tail bands (including black 4-5 HO terminus on tail of juveniles) 6-7 HI Adult total length < 570 mm 10 > 600 mm 11 Tongue color black or nearly so JO pink J1 Supranasal-postnasal association in contact KO separated by first canthal K1 Association of second supralabial and in contact LO prenasal/nasal plates separated by lorilabial LI Shape of mental scute shield-shaped (elongate and triangular) MO wedge-shaped (twice as long as wide) Ml Dominant adult dorsal coloration orange, pink NO black or dark brown N1 yellow N2 Adult dorsal yellow spotting absent OO extremely low 01 low 02 med 03 high 04 Mental scute scalloped edges absent PO moderately scalloped edges PI Enlarged preanal scutes in some females absent QO present Qi Ontogenetic melanism absent RO present R1 Spots on tail in adults absent SO present SI Bands on tail black TO yellow T1 absent T2 Amphib. Reptile Conserv. | http://amphibian-reptile-conservation.org 083 July 2013 | Volume 7 | Number 1 | e67 Reiserer et al. 4. Character mapping analysis A character mapping analysis (CMA) is one of several ro- bust tools used in comparative biology to comprehend the distribution of traits (e.g., morphology), often by explic- itly utilizing molecular phylogenetic information (Brooks and McLennan 1991; Harvey and Pagel 1991; Martins 1996; Freeman and Herron 2004; Maddison and Mad- dison 2011; for a critique, see Assis and Rieppel 2011). Specifically, the CMA aims to provide insights to the ori- gin, frequency, and distribution of selected traits formally expressed onto a tree (e.g., Schuett et al. 2001, 2009; Fen- wick et al. 2011). These procedures also are potentially useful in disentangling homology from homoplasy (Free- man and Herron 2004). Furthermore, the CMA provides a framework for testing hypotheses of adaptive evolution and the identification of species (Harvey and Pagel 1991; Futuyma 1998; Freeman and Herron 2004; Schuett et al. 2001, 2009; Maddison and Maddison 2011). However, CMA does not replace a strict phylogenetic analysis of morphological traits (Assis and Rieppel 2011). Here, we used character mapping to investigate the morphological traits of the four subspecies of H. horri- dum, to gain insights on the distribution, divergence, and homology (e.g., shared-derived traits, such as possible autapomorphies) of these traits. Methods . — We used published morphological data on Heloderma (Bogert and Martin del Campo 1956; Camp- bell and Vannini 1988; Campbell and Lamar 2004; Beck 2005) and selected 20 morphological characters for the CMA (Table 2). All characters were coded as binary (i.e., 0, 1) or multi-state (e.g., 0, 1, 2). Non-discrete multi-state characters (e.g., color pattern) were ordered from low- est to highest values. Character polarity was established by using H. suspectum as the outgroup. The CMA traced each character independently by using the outgroup anal- ysis and parsimony procedures in Mesquite (Maddison and Maddison 2011), and we combined the individual results onto a global tree. Results and discussion . — The CMA results (Fig. 6) show that multiple morphological traits are putative apo- morphies or autapomorphies (traits unique to a single taxon) for the various H. horridum clades (subspecies) delimited in the molecular tree recovered by Douglas et al. (2010). Although we had a priori knowledge of spe- cific and unique traits (presumptive autapomorphies) used to diagnose each of the subspecies, the CMA pres- ents them in a phylogenetic and temporal framework. Our results show trends in scutellation (e.g., presence- absence, relative positions), relative tail length, and body color pattern, including ontogenetic melanism. Are the characters we used in the CMA stable in the subspecies? That question remains for future investigation; however, we have no evidence to the contrary. Indeed, we antici- pate that these characters, and others likely to be revealed through detailed studies, will exhibit stability. Importantly, each of these traits is amenable to further investigation and formal tests. For examples, what is the evolutionary and ecological significance of tongue color differences in beaded lizards (always pink) and Gila monsters (always black), the extreme differences in adult dorsal color pattern in H. h. exasperatum (yellow is predominant) vs. H. h. alvarezi (dark brown and pat- ternless predominate), and ontogenetic melanism in H. h. alvarezi ? As we discussed, beaded lizards occupy similar seasonally dry tropical forests, yet each of the subspe- cies exhibits pronounced molecular and morphological differentiation. Similar types of questions concerning adaptation have used a CMA to explore social systems and sexual dimor- phisms in lizards (Carothers 1984), male fighting and prey subjugation in snakes (Schuett et al. 2001), types of bipedalism in varanoids (Schuett et al. 2009), and di- rection of mode of parity (oviparous vs. viviparous) in viperids (Fenwick et al. 2011). Subspecies and the Taxonomy of Beaded Lizards Introduced in the late 19 th century by ornithologists to de- scribe geographic variation in avian species, the concept of subspecies and trinomial taxonomy exploded onto the scene in the early 20 th century (Bogert et al. 1943), but not without controversy. The use of subspecies has been both exalted and condemned by biologists (see perspec- tives by Mallet 1995; Douglas et al. 2002; Zink 2004; Fitzpatrick 2010). Thousands of papers have been pub- lished in an attempt to either bolster the utility and prom- ulgation of subspecies, or to denounce the concept as meaningless and misleading in evolutionary theory (Wil- son and Brown 1953; Zink 2004). What is the problem? One common critical response is that the subspecies con- cept lacks coherence in meaning, and hence is difficult to comprehend (Futuyma 1998; Zink 2004). Moreover, the use of subspecies often masks real diversity (cryptic species, convergence) or depicts diversity that is non-ex- istent or only trivial (e.g., lack of support in DNA-based analyses; Zink 2004). Indeed, as John Fitzpatrick attests (2010, p. 54), “The trinomial system cannot accurately represent the kind of information now available about ge- netic and character variation across space. Instead, even more accurate tools are being perfected for quantitative, standardized descriptions of variation. These analyses — not subspecies classifications — will keep providing new scientific insights into geographic variation.” Even with the identification of a variety of problems, many authors recommend that complete abandonment of the trinomial category in taxonomy is not necessary nor advised (e.g., Mallett 1995, Hawlitschek et al. 2012). Unfortunately, a consensus among biologists concerning the use of subspecies is not likely to emerge (Fitzpatrick 2010). In step with Fitzpatrick’s (2010) comments, we July 2013 | Volume 7 | Number 1 | e67 Amphib. Reptile Conserv. | http://amphibian-reptile-conservation.org 084 Taxonomy and conservation of beaded lizards contend that the plethora of variation detected in organ- isms must be approached in a modern sense that does not rely upon a cumbersome and outdated taxonomic system. Indeed, we anticipate that the description of geographic variation in organisms, once emancipated from infraspecific taxonomy, will actually accelerate our understanding of variation and its complexities. In our view, the confusion in recognizing subspecies can also mislead conservation planning, and it has on more than one occasion (e.g., the dusky seaside sparrow, see Avise and Nelson 1989). We thus agree with Wilson and Brown (1953), Douglas et al. (2002), Zink (2004), Fitzpatrick (2010) and others in their insightful criticisms leveled at the subspecies concept and the use of trinomials in taxon- omy. Other authors have echoed similar views (Burbrink et al., 2000; Burbrink 2001; Douglas et al. 2007; Tobias et al. 2010; Braby et al. 2011; Hoisington-Lopez 2012; Porras et al. 2013). Given our reassessment of molecular (mt- and nDNAs), phylogeographic, morphological, and biogeo- graphic evidence, we elevate the subspecies of Heloder- ma horridum to the rank of full species (Wiley, 1978; Zink 2004; Tobias et al. 2010; Braby et al. 2011; Porras et al. 2013). Indeed, Douglas et al. (2010, p. 164) stated that, “... unlike H. suspectum, our analyses support the subspecific designations within H. horridum. However, these particular lineages almost certainly circumscribe more than a single species . . . Thus, one benefit of a con- servation phylogenetic perspective is that it can properly identify biodiversity to its correct (and thus manageable) taxonomic level.” Accordingly, based on multiples lines of concordant evidence, we recognize four species of beaded lizards. They are: Mexican beaded lizard: Heloderma horridum (Wieg- mann 1829) Rio Fuerte beaded lizard: Heloderma exasperatum (Bogert and Martin del Campo 1956) Chiapan beaded lizard: Heloderma alvarezi (Bogert and Martin del Campo 1956) Guatemalan beaded lizard: Heloderma charlesbogerti (Campbell and Vannini, 1988) In the above arrangement, we do not recognize subspe- cies and vernacular names remain unchanged. The geo- graphic distribution of the four species of beaded lizards is presented in Fig. 7. Locality data for the map were derived from Bogert and Martin del Campo (1956), Campbell and Vannini (1988), Schwalbe and Lowe (2000), Lemos-Espinal et al. (2003), Campbell and La- mar (2004), Beck (2005), Monroy-Vilchis et al. (2005), Ariano-Sanchez and Salazar (2007), Anzueto and Camp- bell (2010), Domiguez-Vega et al. (2012), and Sanchez- De La Vega et al. (2012). The “?” on the map (coastal Oaxaca, municipality: San Pedro Tututepec) denotes a jet-black adult specimen photographed by Vicente Mata- Silva (pers. comm.) in December 2010. The validity of Amphib. Reptile Conserv. | http://amphibian-reptile-conservation.org 085 this record is questionable owing to its striking coloration resemblance to H. alvarezi from the Central Depression (Rio Grijalva Depression) of Chiapas and extreme west- ern Guatemala, rather than to H. horridum. Although the individual might represent an isolated population of H. alvarezi , further study in this area of Oaxaca is required to rule out human activity as an agent (e.g., displace- ment). Beaded Lizards and Seasonally Dry Tropical Forests The key to understanding the evolution and biogeogra- phy of beaded lizards and the prospects for implementing meaningful conservation measures is through a recogni- tion of the biomes they occupy, which we emphasize are the widely but patchily distributed low elevation season- ally dry tropical forests (SDTFs; see Trejo and Dirzo 2000; Campbell and Lamar 2004; Beck 2005; Ariano- Sanchez 2006; Miles et al. 2006; Pennington et al. 2006; Dirzo et al. 2011; Domiguez-Vega et al. 2012). The evolution of SDTFs in Mesoamerica is a complex evolutionary scenario (Stuart 1954, 1966), and our un- derstanding of their origin and temporal diversification is in its infancy (Janzen, 1988; Becerra 2005; Pennington et al. 2006; Dirzo et al. 2011; De-Nova et al. 2012). One approach to grapple with complex issues such as the ori- gin and historical construction of SDTFs in Mesoamerica has been to examine a single but highly diverse plant tax- on within a phylogenetic (phylogenomic) backdrop. This approach, accomplished by Becerra (2005) and more re- cently by De-Nova et al. (2012), uses the woody plant (tree) Bursera (Burseracae, Sapindales), a highly diverse genus (> 100 species) with a distribution in the New World and emblematic of most dry forest landscapes (De-Nova et al. 2012). Owing to this diversity, coupled with extensive endemism, this taxon has yielded valuable information that serves as a reasonable proxy for diver- sification and expansion of the SDTF biomes (Dick and Pennington 2012). Hence, plant (angio sperm) species richness and expansion of SDTF biomes in Mesoamerica is hypothesized to parallel the diversification of Bursera (Dick and Wright 2005). Based on both plastid and nuclear genomic markers that were analyzed using fossil-calibrated techniques and ancestral habitat reconstruction, the origin of Bursera in Mesoamerica is hypothesized to be in northwestern Mex- ico in the earliest Eocene (-50 mya), with subsequent ex- tensive diversification and southern expansion along the Mexican Transvolcanic Belt in the Miocene, especially -7-10 mya (De-Nova et al. 2012). Accelerated clade di- versification of Bursera and its sister genus Commiphora occurred during the Miocene, a period of increased arid- ity likely derived from seasonal cooling and rain shadow effects (Dick and Wright 2005). Although causal con- nections are complex, they include global tectonic pro- July 2013 | Volume 7 | Number 1 | e67 Reiserer et al. Fig. 7. The distribution of beaded lizards in Mexico and Guatemala. Colored dots represent verified sightings (populations) and museum records. Note the fragmented populations of all four species, which closely approximates the patchy distribution of sea- sonal dry tropical forests (see map in Brown and Lowe [1980]). See text for explanation of question marks (“?”) and other details. cesses, orogenic activities (uplifting of the Sierra Madre Occidental and Sierra Made Oriental) and local volca- nism (Dick and Wright 2005; De-Nova et al. 2012). De- Nova et al. (2012) concluded by emphasizing that their phylogenomic analysis of Bursera points to high species diversity of SDTFs in Mesoamerica that derives from within-habitat speciation rates that occurred in the enve- lope of increasing aridity from the early Miocene to the present. Furthermore, they stated (p. 285), “This scenario agrees with previous suggestions that [angiosperm] lin- eages mostly restricted to dry environments in Mexico resulted from long periods of isolated evolution rather than rapid species generation....” Beaded Lizard Evolution and Diversification The phylogenetic analyses of Heloderma horridum (sensu lato) by Douglas et al (2010) provided fossil- calibrated estimates of divergence times, which allow us to draw connections to the origin and diversification of SDTFs in Mesoamerica (Table 1, Fig. 6). Based on those analyses, H. horridum (sensu lato) and H. suspectum are hypothesized to have diverged from a most-recent com- mon ancestor in the late Eocene (~35 mya), which cor- responds to the establishment of Bursera in northwestern Mexico. Subsequent diversification (cladogenesis) of the beaded lizards occurred during the late Miocene (9.71 mya), followed by a lengthy period of stasis of up to 5 my, with subsequent cladogenesis extending into the Pliocene and Pleistocene. Of particular interest is that this scenario approximately parallels the diversification and southern expansion of SDTFs (Dick and Wright 2005; De-Nova et al. 2012). Accordingly, based on the above discussion of SDTFs and phylogenetic analyses, we suggest that beaded lizard lineage diversification resulted from long periods of isolated (allopatric) evo- lution in SDTFs. Douglas et al. (2010) referred to the fragmented tropical dry forests of western Mexico as “engines” for diversification. The extralimital distribu- tion of H. exasperatum and H. horridum into adjacent pine-oak woodland and thorn scrub biomes appears to be relatively uncommon (Schwalbe and Lowe 2000; Beck 2005; Monroy-Vilchis et al. 2005). July 2013 | Volume 7 | Number 1 | e67 Amphib. Reptile Conserv. | http://amphibian-reptile-conservation.org 086 Taxonomy and conservation of beaded lizards Conservation of Beaded Lizards A primary aim of this paper is to provide a useful and accurate synthesis of information on the taxonomy of beaded lizards that will lead to informed decisions re- garding their conservation (see Douglas et al., 2010). Until recently, H. horridum (sensu lato) was designated as Vulnerable on the World Conservation Union (IUCN) Red List. In 2007, that designation was changed to Least Concern based on more stringent criteria (Canseco- Marquez and Munoz 2007; categories and criteria ver- sion 3.1). The 2007 IUCN Red List also determined that, “Additional research is needed into the taxonomic status, distribution and threats to this species” (Canseco-Mar- quez and Munoz 2007). The critically endangered status of H. h. charlesbogerti (sensu lato) in Guatemala (Ari- ano-Sanchez 2006; Ariano-Sanchez and Salazar 2007) has not altered the current IUCN Red List designation of this taxon, because population trends of other beaded lizards in Mexico remain “unknown” (www.iucnredlist. org/search; see International Reptile Conservation Foun- dation, IRCF; www.ircf.org). As more information on the population status of the newly elevated beaded lizards becomes available, in view of their fragmented distribu- tions and threats to their habitats, the IUCN likely will designate these taxa as Vulnerable or a higher threat cat- egory (see our EVS analysis below). For example, H. ex- asperatum, H. alvarezi, and H. charlesbogerti all occupy limited areas of SDTF (Beck 2005). In Mexico, helodermatid lizards are listed as “threat- ened” (amenazadas) under the Mexican law (NOM- 059-SEMARNAT-2010), legislation comparable to that in the United States Endangered Species Act. The threat- ened category from Mexican law coincides, in part, with the “Vulnerable” category of the IUCN Red List. This document defines “threatened” as species or populations that could become at risk of extinction in a short to me- dium period if negative factors continue to operate that reduce population sizes or alter habitats. Heloderma h. charlesbogerti (sensu lato) is listed on the Guatemalan Lista Roja (Red List) as “endangered,” with approxi- mately 200-250 adult individuals remaining in under 26,000 ha of its natural habitat of SDTF and thorn scrub, (Ariano-Sanchez 2006). Furthermore, H. h. charlesbogerti (sensu lato) is listed on CITES Appendix I, a designation that includes spe- cies threatened with extinction (see CITES document appended to Ariano-Sanchez and Salazar 2007). Trade in CITES Appendix I species is prohibited except under exceptional circumstances, such as for scientific research (CITES 2007). The remaining taxa of Heloderma hor- ridum (sensu lato) ( H . h. alvarezi, H. h. exasperatum, and H. h. horridum) are listed on Appendix II of CITES (CITES 2007). International trade in Appendix II species might be authorized under an export permit, issued by the originating country only if conditions are met that show trade will not be detrimental to the survival of the Amphib. Reptile Conserv. | http://amphibian-reptile-conservation.org 087 species in the wild. The United States Fish & Wildlife Service issues permits only if documentation is provided proving legal origin, including a complete paper trail back to legal founder animals. This procedure allows the importation of beaded lizards into the United States to be tightly regulated (in theory), and also subjects such imports to provisions of the Lacey Act that control com- merce in illegally obtained fish and wildlife (Beck 2005). Beaded Lizards: Denizens of Endangered SDTFs Although occasional sightings of beaded lizards have been reported from mid elevation pine-oak woodlands, all four species primarily inhabit lowland SDTFs and rarely in associated thorn scrub, in both Mexico and Guatemala (Schwalbe and Lowe 2000; Lemos-Espinal et al. 2003; Campbell and Lamar 2004; Beck 2005; Monroy-Vilchis et al. 2005; Ariano and Salazar 2007; Domiguez-Vega et al. 2012). Thus, the optimal measure to reduce threats to beaded lizards is to maintain the integrity of their tropi- cal dry forest habitats. Current threats to beaded lizards throughout their range include habitat loss, road mor- tality, poaching, and illegal trade (Beck 2005; Miles et al. 2006; Golicher et al. 2012). Habitat loss takes many forms, from the conversion of SDTFs to areas of agricul- ture and cattle ranching, to forest fragmentation owing to roads and other forms of development (Pennington et al. 2006). Degradation from human-introduced invasive (exotic) organisms and fire also are contributing factors (Beck 2005). When the Spaniards arrived in the Western Hemi- sphere, Mesoamerican SDTFs covered a region stretch- ing from Sonora (Mexico) to Panama, an area roughly the size of France (-550,000 km 2 ). Today, only 0.1% of that region (under 500 km 2 ) has official conservation status, and less than 2% remains sufficiently intact to attract the attention of conservationists (Janzen 1988; Hoekstra et al. 2005). Of all 13 terrestrial biomes analyzed by Hoek- stra et al. (2005), the SDTF biome has the third highest conservation risk index (ratio of % land area converted per % land area protected), far above tropical wet forest and temperate forest biomes (Miles et al. 2006). Mexico ranks among the most species rich countries in the world (Garcia 2006; Urbina-Cardona and Flores- Villela 2010; Wilson and Johnson 2010; Wilson et al. 2010, 2013). Nearly one-third of all the Mexican herpe- tofaunal species are found in SDTFs (Garcia 2006; De- Nova et al. 2012). Neotropical dry forests span over 16 degrees of latitude in Mexico, giving way to variation in climatic and topography that results in a diversity of tropical dry forest types, and a concurrent high propor- tion of endemism of flora and fauna (Garcia 2006; De- Nova et al. 2012; Wilson et al. 2010; 2013). Mexican seasonally tropical dry forest, classified into seven ecore- gions that encompass about 250,000 km 2 , has enormous conservation value and has been identified as a hotspot July 2013 | Volume 7 | Number 1 | e67 Reiserer et al. for conservation priorities (Myers et al. 2000; Sanchez- Azofeifa et al. 2005; Garcia 2006; Urbina-Cardona and Flores-Villela, 2010; Wilson et al. 2010, Mittermeier et al. 2011). The vast majority (98%) of this region, how- ever, lies outside of federally protected areas (De-Nova et al. 2012). With few exceptions, most of the protected areas in Mexico occur in the states of Chiapas and Jalis- co, leaving much of the region (e.g., Nayarit and Sinaloa) without government (federal) protection (Garcia 2006). In Guatemala, less than 10% of an estimated 200,000 ha of original suitable habitat have been established as protected critical habitat in the Motagua Valley for the endangered H. charlesbogerti (Najera Acevedo 2006). A strong effort led by local citizens, conservation workers, biologists, government officials, NGOs, and conserva- tion organizations (e.g., The Nature Conservancy, Inter- national Reptile Conservation Association, Zoo Atlanta, and Zootropic) negotiated to have H. h. charlesbogerti (sensu lato) placed on CITES Appendix I, to purchase habitat, conduct research, employ local villagers in mon- itoring the lizards, and promote environmental education (Lock 2009). Similar efforts for beaded lizards have been underway for many years in Chiapas (Mexico), spear- headed at ZooMAT (Ramirez-Velazquez 2009), and in Chamela, Jalisco (www.ibiologia.unam.mx/ebchamela/ www/reserva.html). Such efforts will need to expand in the years ahead and will doubtless play a crucial role if we hope to retain the integrity of existing SDTFs inhab- ited by beaded lizards throughout their range. Discussion In this paper, we reassessed the taxonomy of Heloderma horridum (sensu lato) using both published information and new analyses (e.g., CMA). We concluded that diver- sity in beaded lizards is greater than explained by infra- specific differences and that the recognition of subspecies is not warranted, as it obscures diversity. Our decision to elevate the four subspecies of H. horridum to full species status is not entirely novel (Beck 2005; Douglas et al. 2010). Furthermore, our taxonomic changes are based on integrative information (i.e., morphology, mt- and nDNA sequence information, biogeography) and changing per- spectives on the utility of formally recognizing infraspe- cific diversity using a trinomial taxonomy (Wilson and Brown 1953; Douglas et al. 2002; Zink 2004; Porras et al. 2013). This decision not only adds to a better under- standing of the evolution of helodermatids, but also pro- vides an important evolutionary framework from which to judge conservation decisions with prudence (Douglas et al. 2002). Below, we delineate and discuss prospective research and conservation recommendations for beaded lizards based on our present review. Borrowing some of the guidelines and recommendations for future research and conservation for cantils, also inhabitants of SDTFs, by Porras et al. (2013), we outline similar ones for the four Amphib. Reptile Conserv. | http://amphibian-reptile-conservation.org 088 species of beaded lizards ( H . alvarezi, H. charlesbogerti , H. exasperatum, and H. horridum ). Future Research and Conservation Recommendations I . Throughout this paper we emphasized the importance of SDTFs in the distribution of beaded lizards, yet most SDTFs within their distribution are not Protected Natural Areas (PNAs; Beck 2005; Urbina-Cardona and Flores- Villela 2009; Domiguez-Vega et al. 2012). Accordingly, emphasis should be placed on those areas of SDTFs for prospective research, new conservation projects, and for establishing new PNAs. The protection of beaded liz- ards must be placed into a larger context of conservation planning. Proper stewardship of SDTFs and other biomes must include meaningful (scientific) protective measures for all of the flora and fauna, rather than piecemeal (e.g., taxon-by-taxon) approaches that lack a cohesive conser- vation plan (Douglas et al. 2010). We applaud the efforts of Domiguez-Vega et al. (2012) in identifying conservation areas for beaded liz- ards; however, we do not agree with all of their conclu- sions. In particular, based on field experiences by one of us (DDB), we contend that the potential (predicted) range of H. exasperatum in Sonora (Mexico) based on the results of their habitat suitability modeling, appears exaggerated and thus may be misleading. In our opin- ion, their distribution maps (figs. 2 and 3) overestimate the extent of true SDTFs in Sonora, showing their occur- rence in a type of biome that is more accurately classi- fied as Sinaloan Thorn Scrub (see the excellent maps in Brown and Lowe 1980; Robichaux and Yetman 2000). In Sonora, beaded lizards ( H . exasperatum) are rarely found in association with pure thorn scrub, while Gila monsters, in contrast, are frequently encountered in that type of habitat (Schwalbe and Lowe 2000; Beck 2005). 2. With few exceptions, the population viability of beaded lizards is largely unknown (Beck 2005; Ariano- Sanchez 2006; Ariano- Sanchez et al. 2007; Domiguez-Vega et al. 2012). We highly recommend that modem assessments of the four species occur at or near localities where they have been recorded (e.g., Jimenez- Valverde and Lobo 2007). Whereas H. charlesbogerti , and to a lesser degree H. alvarezi (Ramirez-Velazquez 2009), are receiving in- ternational conservation attention, we feel that similar consideration is necessary for H. exasperatum owing to its relatively limited geographic range (Sonora, Chi- huahua, Sinaloa), the large extent of habitat destruction and fragmentation (Fig. 8), and limited areas receiving protection (Trejo and Dirzo 2000; Domiguez-Vega et al. 2012; see http://www.conanp.gob.mx/regionales/). In 1996, about 92, 000 hectares in the Sierra de Alamos and the upper drainage of the Rio Cuchujaqui were declared a biosphere reserve by the Secretary of the Environment and Natural Resources (SEMARNAT 2010), called the July 2013 | Volume 7 | Number 1 | e67 Taxonomy and conservation of beaded lizards / ✓ Area de Protection de Fauna y Flora Sierra de Alamos y Rio Cuchujaqui (Martin and Yetman 2000; S. Meyer, pers. comm.). Efforts continue in Sonora to set aside ad- ditional habitat for conservation, but, other than Alamos, no other areas with true SDTFs presently exist (Robich- aux and Yetman 2000; S. Meyer, pers. comm.). 3. Conservation management plans for each of the spe- cies of beaded lizards should be developed from an integrative perspective based on modern population assessments, genetic information, and ecological (e.g., soil, precipitation, temperature) and behavioral data (e.g., social structure, mating systems, home range size). Such a conservation plan is in place for the Guatemalan beaded lizard ( H . charlesbogerti ) by CONAP-Zootropic (www.ircf.org/downloads/PCHELODERMA-2Web. pdf). Also, aspects of burgeoning human population growth must be considered, since outside of PNAs these large slow -moving lizards generally are slaughtered on sight, killed on roads by vehicles (Fig. 9), and threatened by persistent habitat destruction primarily for agriculture and cattle ranching (Fig. 10). For discussions on conser- vation measures in helodermatid lizards, see Sullivan et al. (2004), Beck (2005), Kwiatkowski et al. (2008), Doug- las et al. (2010), Domfguez-Vega et al. (2012), and Ariano- Sanchez and Salazar (2013). In Mexico, the IUCN lists Heloderma horridum (sensu lato) under the category of Least Concern. Recently, Wilson et al. (2013) reported the Environmen- tal Vulnerability Score (EVS) for H. horridum (sensu lato) as 11. Briefly, an EVS analysis as- sesses the potential threat sta- tus of a given species based on multiple criteria and provides a single score or index value (Wil- son and McCranie 2004; Porras et al. 2013; Wilson et al. 2013). High EVS scores (e.g., 17), for example, signify vulnerability. With the taxonomic changes we proposed for beaded lizards, an EVS assessment is thus required for each species. Using the new criteria developed by Wilson et al. (2013; see Porras et al. 2013), we recalculated the EVS for the species of beaded lizards, which are presented below: Fig. 8. Destruction of seasonally dry tropical forest near Alamos, Sonora, Mexico. Photo by Daniel D. Beck. H. horridum'. 5 + 4 + 5 = 14 H. exasperatum : 5 + 7 + 5 = 17 H. alvarezi'. 4 + 6 + 5 = 15 H. charlesbogerti'. 4 + 8 + 5 = 17 Fig. 9. A dead-on-the-road (DOR) H. exasperatum (sensu stricto) near Alamos, Sonora, Mexico. Vehicles on paved roads are an increasing threat to beaded lizards, Gila monsters, and other wildlife. Photo by Thomas Wiewandt. These recalculated values fall into the high vulnerability category (Wilson et al. 2013; Porras et al. 2013), underscoring the urgency for the development of conserva- tion management plans and long- term population monitoring of all species of beaded lizards. These values thus need to be reported to the appropriate IUCN commit- tees, so immediate changes in sta- tus can be made and conservation actions implemented. July 2013 | Volume 7 | Number 1 | e67 Amphib. Reptile Conserv. | http://amphibian-reptile-conservation.org 090 Reiserer et al. Fig. 10. Agave cultivation in Mexico results in the destruction of seasonally dry tropical forests. Photo by Thomas Wiewandt. natural habitat, small population size (200-250 adults) and endan- gered status, H. charlesbogerti is currently listed as CITES Appen- dix I (Ariano- Sanchez and Sala- zar 2007). Given the taxonomic elevation of these taxa, conserva- tion agencies can use these char- ismatic lizards as flagship species in efforts to publicize conserva- tion efforts in their respective countries at all levels of interest and concern, including educa- tion and ecotourism (Beck 2005). Eli Lilly Co., Disney Worldwide Conservation Fund and The Na- ture Conservancy support the conservation of H. charlesboger- ti (Ariano-Sanchez and Salazar 2012). Such corporate involve- ment provides funds and positive public exposure (e.g., social net- work advertising) that otherwise would not be possible. Fig. 11. Antonio Ramirez Ramirez- Velazquez, a herpetologist, discusses the beauty and importance of beaded lizards ( H . alvarezi, sensu stricto) to a group of enthusiastic children and their teacher at Zoo Miguel Alvarez del Toro (ZooMAT) in Tuxtla Gutierrez, Chiapas, Mexico. The zoo was named in honor of its founding director, Senor Miguel Alvarez del Toro, who had a keen academic and conservation interest in beaded lizards. He collected the type specimen of H. alvarezi (described in Bogert and Martin del Campo, 1956), which was named in his honor. ZooMAT offers hands-on environmental education programs to schoolchildren and other citizens of southern Mexico. Photo by Thomas Wiewandt. 4. We recommend the establishment of zoo conservation (AZA) educational outreach programs, both ex situ and in situ, such as those currently in progress for H. charles- bogerti (www.IRCF.org;www.zooatlanta.org) and for H. alvarezi in Chiapas (Ramirez-Velazquez, 2009, see Fig. 11). Because of its limited range, destruction of its 5. One of the major conclusions of this paper is that our knowl- edge of the taxonomy and phy- logeography of beaded lizards remains at an elementary level. As discussed, a robust phylogeo- graphic analysis using morpho- logical characters is not avail- able. Our character mapping exercise, for various reasons, is not a substitute procedure for detailed phylogenetic analyses using morphology (Assis 2009; Assis and Rieppel 2011). Other authors have made similar pleas concerning the importance of morphology, including fossils, in phylogenetic reconstruction (Poe and Wiens 2000; Wiens 2004, 2008; Gauthier et al. 2012). Moreover, further studies on the historical biogeography of he- lodermatids (e.g., ancestral area reconstruction) are needed (e.g., Ronquist 1997, 2001; Ree and Smith 2008). Detailed morpho- logical analyses can be conducted with new tools such as computed tomography (CT) scans of osteological char- acters of both extant and fossil specimens (Gauthier et al. 2012), and geometric morphometric approaches to exter- nal characters (Davis 2012). Furthermore, in the expand- ing field of “venomics” new venom characters in beaded July 2013 | Volume 7 | Number 1 | e67 Amphib. Reptile Conserv. | http://amphibian-reptile-conservation.org 090 Taxonomy and conservation of beaded lizards lizards will likely be discovered, which might prove use- ful in phylogenetic analyses (Fry et al. 2009, 2010). As we progress into the “Age of Genomics” with ever-growing computational advancements (e.g., bio- informatics; Homer et al. 2009), new and exciting meth- ods to explore organismal diversity are opening, includ- ing such next-generation approaches as pyrosequencing (microsatellite isolation), establishing transcriptome databases, and whole-genome sequencing (Wiens 2008; Castoe et al. 2011; Culver et al. 2011). Currently, plans are underway to apply pyrosequencing methods to helo- dermatids to generate a nearly inexhaustible supply of microsatellite markers for a variety of proposed analy- ses (W. Booth and T. Castoe, pers. comm.). Standing on the shoulders of The Human Genome Project (Culver et al. 2011), and reaping the success of genome projects in other reptilian taxa (Castoe et al. 2011), it is now possible to establish a “Helodermatid Genome Project.” Beaded lizards and the Gila monster are especially good candi- dates for such an investment, especially given the impor- tance of their venom components in medical research and recent pharmaceutical applications (Beck 2005; Douglas et al. 2010; Fry et al. 2009, 2010). 6. An important take-home message from Douglas et al. (2010) is that future conservation efforts will require a robust understanding of phylogenetic diversity (e.g., conservation phylogenetics) to make sensible (logical) and comprehensive conservation plans. For example, the range of H. horridum (sensu stricto) is the most expansive of the species of beaded lizards and has not been fully explored with respect to genetic diversity. Accordingly, sampling throughout its range may yield cryptic genetic diversity, perhaps even new species. We emphasize that viable conservation planning must incorporate all intel- lectual tools available, including those that incorporate old methods (e.g., paleoecological data) but viewed through a new lens (Douglas et al. 2007, 2009; Willis et al. 2010). Wisely, Greene (2005) reminds us that we are still grappling with understanding basic and essential issues concerning the natural history of most organisms. To that end, we must continue in our efforts to educate students and the public of the need for and importance of this branch of science. 7. The new taxonomic arrangement of beaded lizards we proposed will affect other fields of science, such as conservation biology and human medicine (Beck, 2005; Douglas et al., 2010). In Fry et al. (2010, p. 396, table 1), toxins are matched to the subspecies of beaded lizards and Gila monsters. Yet as noted by Beck (2005) and Douglas et al. (2010), the banded Gila monster (H. s. cinctum) is not a valid subspecies, which is based on several levels of analysis (i.e., morphology, geographic distribution, and haplotype data). Individuals assigned to H. s. cinctum based on color and pattern, for example, have been found in southwestern Arizona near the Mexi- can border and in west-central New Mexico (Beck 2005). Furthermore, most venom researchers, including those who study helodermatids, often obtain samples from cap- tive subjects in private collections and zoological institu- tions. Many of these animals have been bred in captivity and result from crossing individuals of unknown origin or from different populations (D. Boyer, pers. comm). Among other negative outcomes, such “mutts” will con- found results of the true variation of venoms. Geographic and ontogenetic variation in venom constituents is well established in other squamates (Minton and Weinstein 1986; Alape-Giron et al. 2008; Gibbs et al. 2009), which is apparently the case in helodermatids (Fry et al. 2010). Thus, we strongly encourage researchers investigating helodermatid venoms for molecular analysis and phar- maceutical development to use subjects with detailed lo- cality information, as well as age, gender, and size, and to provide those data in their publications. 8. Owing to problems that many scientists, their stu- dents, and other interested parties from Mesoamerica have in gaining access to primary scientific literature, we highly recommend that authors seek Open Access peer-reviewed journals as venues for their publications on beaded lizards, an important factor in our choice for selecting the present journal (www.redlist-ARC.org) as a venue for our data and conservation message. Acknowledgments. — We thank Larry David Wilson for inviting us to participate in the Special Mexico Is- sue. A Heritage Grant from the Arizona Game and Fish Department and a Research Incentive Award/Scholarly Research and Creative Activities Award (Arizona State University) awarded to GWS funded parts of this re- search. Zoo Atlanta (Dwight Lawson, Joe Mendelson III) and Georgia State University (Department of Biology) provided various levels of support. Warren Booth, Donal Boyer, Dale DeNardo, Andres Garcia, Stephanie Mey- er, and Tom Wiewandt were always willing to discuss beaded lizard and tropical dry forest biology with us. We thank Brad Lock, Louis Porras, and Larry David Wilson for their suggestions and valuable insights in improving an earlier version of this manuscript. Also, three review- ers, including Daniel Ariano-Sanchez, provided key information and sharpened our focus, though we bear the burden of any blunders. We thank Javier Alvarado, Daniel Ariano-Sanchez, David Brothers, Quetzal Dwyer, Kerry Holcomb, Vicente Mata-Silva, Stephanie Meyer, Adam Thompson, and Tom Wiewandt for graciously supplying us with images. Vicente Mata-Silva kindly as- sisted us in preparing the resumen and locating literature on Heloderma. July 2013 | Volume 7 | Number 1 | e67 Amphib. Reptile Conserv. | http://amphibian-reptile-conservation.org 091 Reiserer et al. Literature Cited Alape-Giron A, Sanz L, Escolano J, Flores-Dfaz M, Madrigal M, Sasa M, et al. 2008. Snake venomics of the lancehead pitviper Bothrops asper : geographic, individual, and ontogenetic variations. Journal of Proteome Research 7: 3556-3571. Alvarez del Toro M. 1983 (1982). Los Reptiles de Chi- apas (3 rd edition). Publicacion del Instituto de Historia Natural, Tuxtla Gutierrez, Chiapas, Mexico. Anzueto VR, Campbell JA. 2010. Guatemalan beaded lizard ( Heloderma horridum charlesbogerti ) on the Pacific versant of Guatemala. 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Tree species diversity driven by environmental and anthropogenic factors in tropical dry forest fragments in central Veracruz, Mexico. Biodiversity and Conservation 18: 3269- 3293. Zink RM. 2004. The role of subspecies in obscuring avi- an biological diversity and misleading conservation policy. Proceedings of the Royal Society of London B 271: 561-564. Received: 23 May 2013 Accepted: 12 June 2013 Published: 29 July 2013 Randall S. Reiserer is an integrative biologist whose research focuses on understanding the interrelationships among ecology, morphology, and behavior. Within the broad framework of evolutionary biology, he studies cognition, neuroscience, mimicry, life-history evolution, and the influence of niche dynamics on patterns of evolutionary change. His primary research centers on reptiles and amphibians, but his academic interests span all major vertebrate groups. His studies of behavior are varied and range from caudal luring and themial behavior in rattlesnakes to learning and memory in transgenic mice. Randall established methods for study- ing visual perception and stimulus control in is studies of caudal luring in snakes. He commonly employs phylogenetic comparative methods and statistics to investigate and test evolutionary patterns and adaptive hypotheses. Dr. Reiserer is an editor of the upcoming peer-reviewed book, The Rattlesnakes of Arizona. Gordon W. Schuett is an evolutionary biologist and herpetologist who has conducted extensive research on rep- tiles. His work has focused primarily on venomous snakes, but he has also published on turtles, lizards, and amphibians. Among his most significant contributions are studies of winner-loser effects in agonistic encoun- ters, mate competition, mating system theory, hormone cycles and reproduction, caudal luring and mimicry, long-term sperm storage, phylogeographic analyses of North American pitvipers, and as a co-discoverer of facultative parthenogenesis in non-avian reptiles. He served as chief editor of the peer-reviewed book Biology of the Vipers and is presently serving as chief editor of an upcoming peer-reviewed book The Rattlesnakes of Arizona (rattlesnakesofarizona.org). Gordon is a Director and scientific board member of the newly founded non-profit The Copperhead Institute (copperheadinstitute.org). He was the founding Editor of the journal Herpetological Natural History. Dr. Schuett resides in Arizona and is an adjunct professor in the Department of Biology at Georgia State University. Daniel D. Beck is an ecologist and herpetologist who has conducted research on the ecology, physiology, and be- havior of rattlesnakes and helodermatid lizards. He has pioneered many of the field studies on helodermatid lizards in the past 30 years, including topics ranging from energy metabolism and habitat use to combat and foraging behaviors in locations ranging from the deserts of Utah, Arizona, and New Mexico, to the tropical dry forests of Sonora and Jalisco, Mexico. His book, Biology of Gila Monsters and Beaded Lizards (2005), presents a synthesis of much of our knowledge of these charismatic reptiles. Dr. Beck is Professor of Biology at Central Washington University, in Ellensburg, Washington, where he lives in a straw bale house with his wife, biologist Kris Ernest, and their two teenage children. July 2013 | Volume 7 | Number 1 | e67 Amphib. Reptile Conserv. | http://amphibian-reptile-conservation.org 096 Pseudoeurycea naucampatepetl. The Cofre de Perote salamander is endemic to the Sierra Madre Oriental of eastern Mexico. This relatively large salamander (reported to attain a total length of 150 mm) is recorded only from, “a narrow ridge extending east from Cofre de Perote and terminating [on] a small peak (Cerro Volcancillo) at the type locality,” in central Veracruz, at elevations from 2,500 to 3,000 m (Amphibian Species of the World website). Pseudoeurycea naucampatepetl has been assigned to the P. bellii complex of the P. bellii group (Raffaelli 2007) and is considered most closely related to P gigantea, a species endemic to the La Joya-Jalapa region of Veracruz and adjacent northeastern Hidalgo (Parra-Olea et al. 2001). This salamander is known from only five specimens and has not been seen for 20 years, despite thorough surveys in 2003 and 2004 (EDGE; www.edgeofexistence.org), and thus it might be extinct. The habitat at the type locality (pine-oak forest with abundant bunch grass) lies within Lower Montane Wet Forest (Wilson and Johnson 2010; IUCN Red List website [accessed 21 April 2013]). The known specimens were “found beneath the surface of roadside banks” (www.edgeofexistence.org) along the road to Las Lajas Microwave Station, 15 kilometers (by road) south of Highway 140 from Las Vigas, Veracruz (Amphibian Species of the World website). This species is terrestrial and presumed to reproduce by direct development. Pseudoeurycea naucampatepetl is placed as number 89 in the top 100 Evolutionarily Distinct and Globally Endangered amphib- ians (EDGE; www.edgeofexistence.org). We calculated this animal’s EVS as 17, which is in the middle of the high vulnerability category (see text for explanation), and its IUCN status has been assessed as Critically Endangered. Of the 52 species in the genus Pseudoeurycea, all but four are endemic to Mexico (see Appendix of this paper and Acevedo et al. 2010). Photo by James Hanken. August 2013 | Volume 7 | Number 1 Amphib. Reptile Conserv. http://amphibian-reptile-conservation.org 97 e69 Copyright: © 2013 Wilson et al. This is an open-access article distributed under the terms of the Creative Com- mons Attribution-NonCommercial-NoDerivs 3.0 Unported License, which permits unrestricted use for non-com- mercial and education purposes only provided the original author and source are credited. Amphibian & Reptile Conservation 7(1): 97-127. A conservation reassessment of the amphibians of Mexico based on the EVS measure ^arry David Wilson, 2 Jerry D. Johnson, and 3 Vicente Mata-Silva 1 Centro Zamorano de Biodiversidad, Escuela Agricola Panamericana Zamorano, Departamento de Francisco Morazdn, HONDURAS ^Depart- ment of Biological Sciences, The University of Texas at El Paso, El Paso, Texas 79968-0500, USA Abstract . — Global amphibian population decline is one of the better documented symptoms of bio- diversity loss on our planet, and one of the environmental super-problems humans have created. Most people believe that we should manage nature for our benefit, instead of understanding that we are part of the natural world and depend on it for our survival. As a consequence, humans keep unraveling Earth’s life-support systems, and to reverse this trend must begin to develop a sustain- able existence. Given this reality, we examine the conservation status of the 378 species of amphib- ians in Mexico, by using the Environmental Vulnerability Score (EVS) algorithm. We summarize and critique the IUCN Red List Assessments for these creatures, calculate their EVS, and compare the results of both conservation assessments. We also compare the EVS for Mexican amphibians with those recently reported for Mexican reptiles, and conclude that both groups are highly imperiled, especially the salamanders, lizards, and turtles. The response of humans to these global impera- tives has been lackluster, even though biological scientists worldwide have called attention to the grave prospects for the survival of life on our planet. As part of the global community, Mexico must realize the effects of these developments and the rapid, comprehensive need to conserve the coun- try’s hugely significant herpetofauna. Based on this objective, we provide five broad-based recom- mendations. Key words. EVS, anurans, salamanders, caecilians, IUCN categorizations, survival prospects Resumen . — La disminucion global de las poblaciones de anfibios es uno de los sintornas mas docu- mentados sobre la perdida de biodiversidad en nuestro planeta, que a su vez es uno de los super- problemas ambientales creados por los seres humanos. La mayoria de los seres humanos creemos que podemos y debemos manejar la naturaleza para nuestro propio beneficio, en lugar de compren- der que somos parte y dependemos de ella misma. Como consecuencia de ello, estamos desarticu- lando los sistemas biologicos del planeta, y para revertir esta tendencia debemos desarrollar una existencia sostenible. Ante esta realidad, examinamos el estado de conservacion de las 378 espe- cies de anfibios mexicanos utilizando el algoritmo de Medida de Vulnerabilidad Ambiental (EVS). Resumimos y criticamos las evaluaciones de la Lista Roja para estos organismos, calculamos su EVS, y comparamos los resultados con los resultados de la categorizacion de la UICN. Tambien comparamos el EVS de los anfibios mexicanos con los publicados recientemente para los reptiles de Mexico, concluyendo que ambos grupos estan en un peligro altamente significativo, principal- mente las salamandras, las lagartijas y las tortugas. La respuesta humana a esta crisis global ha sido mediocre, a pesar de que la comunidad mundial de biologos se une al llamado de atencion sobre las perspectivas graves que amenazan la supervivencia de la vida en nuestro planeta. Como parte de la comunidad mundial, el pais de Mexico debe de considerar los efectos de estos cambios, y la rapida necesidad de conservar de manera integral la herpetofauna altamente significativa de este pais. Basandonos en este objetivo, proporcionamos cinco recomendaciones generalizadas. Palabras claves. EVS, anuros, salamandras, cecilios, categorizacion de UICN, perspectivas de supervivencia Citation: Wilson LD, Johnson JD, Mata-Silva V. 2013. A conservation reassessment of the amphibians of Mexico based on the EVS measure. Amphibian & Reptile Conservation 7(1): 97-127(e69). Correspondence. Emails: 1 bufodoc@aol.com (Corresponding author) 2 jjohnson@utep.edu 3 vmata@ utep.edu August 2013 | Volume 7 | Number 1 | e69 Amphib. Reptile Conserv. | http://amphibian-reptile-conservation.org gs Wilson et al. How will humans react to an increased awareness that Earth’s biodiversity is diminishing? What are these loss- es telling us about our place on the planet, our role in the biosphere? What is our role in conserving biodiversity as we become custodians of a planet that has clear limita- tions ? And how can we pass to future generations the wisdom needed to make sound environmental decisions? The answers to these questions will tell us much about ourselves, and science will take us only part of the way along that journey. Collins and Crump 2009: 205. Introduction Global amphibian population decline is a well-known environmental issue to conservation biologists and her- petologists (Collins and Crump 2009; Stuart et al. 2010). This issue, however, often does not make it onto lists of the world’s most significant problems. A survey of European Union citizens conducted in the fall of 2011 identified the following problems of greatest concern: (1) poverty, hunger and lack of drinking water (28% of those surveyed); (2) climate change (20%); (3) the eco- nomic situation (16%); (4) international terrorism (11%); (5) the availability of energy (7%); (6) the increasing global population (5%); (7) the spread of infectious dis- ease (4%); (8) armed conflict (4%); the proliferation of nuclear weapons (3%); and (10) don’t know (2%). Such surveys expose several underlying concerns. One is that amphibian population decline is not on the list, but neither is the larger issue of biodiversity decline. Another concern is that this “pick the biggest problem” approach does not acknowledge that all of these issues are intertwined and capable of creating “environmental super-problems,” as explained by Bright (2000). Further, with respect to the natural world Bright (2000: 37) indi- cated that “we will never understand it completely, it will not do our bidding for free, and we cannot put it back the way it was.” These features are characteristic of biodiver- sity and biodiversity decline, and indicative of how little we know about the current status of biodiversity. Mora et al. (2011) provided an estimate of the total amount of biodiversity, which they indicated at approximately 8.7 million (±1.3 mill ion SE), with about 86% of the existing land species and 91% of the oceanic species still await- ing description. The description of new taxa is only the initial step toward understanding how the natural world works. The world will not do our bidding for free, since we cannot obtain an appreciable quantity of anything from nature without sacrificing something in the process. In transforming our planet to fill the needs of our species, we have destroyed the habitats of countless creatures (in- cluding amphibians) that also have evolved over time. We cannot reverse this damage, as evidenced by the fact that we have been unable to provide permanent solutions to any of the significant environmental problems. Such is Amphib. Reptile Conserv. | http://amphibian-reptile-conservation.org 99 the case with biodiversity decline, since no retreat from species extinction is possible. Biodiversity decline is an environmental super-prob- lem, as contributing factors include habitat modification, fragmentation, and loss, pollution and disease, over-har- vesting, exotic species, and extinction (Vitt and Caldwell 2009). These problems interact to enmesh species into an extinction vortex, defined as “a downward population spiral in which inbreeding and genetic drift combine to cause a small population to shrink and, unless the spiral is reversed, to become extinct” (Campbell et al. 2008: 1251). Theoretically, this effect should significantly im- pact species with narrower distributions. The extent of biodiversity decline is unknown, al- though most estimates indicate that we know very little about this topic. With respect to animals, we know sub- stantially more about the diversity of vertebrates than in- vertebrates. Among the vertebrates subjected to a global analysis, a greater proportion of amphibians have been documented as threatened than birds or mammals (Stuart et al. 2010). Reptiles and fishes, however, remain unas- sessed. The data presented in Stuart et al. (2010) essentially were the same as in Stuart (2004). The number of am- phibians known globally now exceeds 7,000 (7,139; www.amphibiaweb.org [accessed 8 June 2013]), which is 24.3% greater than the one cited by Stuart et al. (2010). The description of new species of amphibians obviously is a “growth industry,” and the rate of discovery does not appear to be slowing. Thus, we expect that the number of new amphibian taxa from Mexico will continue to in- crease. Another major fault with assessing the “world’s great- est problems” is that their causes are not identified. As noted by Wilson et al. (2013: 23), “no permanent solution to the problem of biodiversity decline (including herpe- tofaunal decline) will be found in Mexico (or elsewhere in the world) until humans recognize overpopulation as the major cause of degradation and loss of humankind’s fellow organisms.” Further, they stated (Pp. 23-24) that, “solutions will not be available until humanity begins to realize the origin, nature, and consequences of the mis- match between human worldviews and how our planet functions.” Miller and Spoolman (2012: 20) defined this “planetary management worldview” as maintaining that “we are separate from and in charge of nature, that nature exists mainly to meet our needs and increasing wants, and that we can use our ingenuity and technology to manage the earth’s life-support systems, mostly for our benefit, into the distant future.” Unfortunately, over the span of about 10,000 years, humans have dismantled the planet’s life-support sys- tems, and today we are living unsustainably (Miller and Spoolman 2012). So, until and unless we develop an en- vironmentally sustainable society, no lasting, workable solutions to environmental problems will be found, in- cluding that of biodiversity decline. August 2013 | Volume 7 | Number 1 | e69 Conservation reassessment of Mexican amphibians Incilius pisinnus. The Michoacan toad, a state endemic, is known only from the Tepalcatepec Depression. This toad’s EVS has been assessed as 15, placing it in the lower portion of the high vulnerability category, and its IUCN status as Data Deficient. This individual came from Apatzingan. Photo by Ivan Trinidad Ahumada- Carrillo. Craugastor hobartsmithi. The distribution of the endemic Smith’s pygmy robber frog is along the southwestern portion of the Mexi- can Plateau, from Nayarit and Jalisco to Michoacan and the state of Mexico. Its EVS has been determined as 15, placing it in the lower portion of the high vulnerability category, and its IUCN status as Endangered. This individual is from the Sierra de Manantlan in Jalisco. Photo by Ivan Trinidad Ahumada-Carrillo. August 2013 | Volume 7 | Number 1 | e69 Amphib. Reptile Conserv. | http://amphibian-reptile-conservation.org 100 Wilson et al. Nonetheless, building a sustainable society requires steps that only a few people appear willing to take. Thus, efforts by conservation biologists to reverse biodiversity decline, including amphibian population decline, must proceed with the realization that we will only be design- ing short-term solutions that deal with the symptoms of the problems rather than their causes. Within this real- ization, we undertake the following reassessment of the conservation status of the amphibians of Mexico. A Revised Environmental Vulnerability Measure In conducting a conservation reassessment of Mexican reptiles, Wilson et al. (2013) revised the Environmen- tal Vulnerability Score (EVS) from that used in various chapters of Wilson et al. (2010). Similarly, we modified the EVS measure for use with Mexican amphibians, es- pecially by substituting the human persecution scale used for reptiles with a reproductive mode scale, as did Wilson and McCranie (2004) and other authors who used this measure with Central American amphibians (see Wilson et al. 2010). Wilson et al. (2013) indicated that the EVS measure originally was designed for use in cases where the details of the population status of a species, upon which many of the criteria for IUCN status categorization depend, were not available, as well as to provide an estimate of the susceptibility of amphibians and reptiles to future en- vironmental threats. The advantages for using the EVS measure are indicated below (see EVS for Mexican am- phibians). The EVS algorithm we developed for use with Mexi- can amphibians consists of three scales, for which the values are added to produce the Environmental Vulner- ability Score. The first scale deals with geographic distri- bution, as follows: 1 = distribution broadly represented both inside and outside Mexico (large portions of range are both inside and outside Mexico) 2 = distribution prevalent inside Mexico, but limited outside Mexico (most of range is inside Mexico) 3 = distribution limited inside Mexico, but prevalent outside Mexico (most of range is outside Mex- ico) 4 = distribution limited both inside and outside Mexi- co (most of range is marginal to areas near bor- der of Mexico and the United States or Central America) 5 = distribution within Mexico only, but not restricted to vicinity of type locality 6 = distribution limited to Mexico in the vicinity of type locality Amphib. Reptile Conserv. | http://amphibian-reptile-conservation.org 101 The second scale deals with ecological distribution, as follows: 1 = occurs in eight or more formations 2 = occurs in seven formations 3 = occurs in six formations 4 = occurs in five formations 5 = occurs in four formations 6 = occurs in three formations 7 = occurs in two formations 8 = occurs in one formation The third scale is concerned with the type of reproductive mode, as follows: 1 = both eggs and tadpoles in large to small bodies of lentic or lotic water 2 = eggs in foam nests, tadpoles in small bodies of lentic or lotic water 3 = tadpoles occur in small bodies of lentic or lotic water, eggs outside of water 4 = eggs laid in moist situation on land or moist ar- boreal situations, direct development, or vivipa- rous 5 = eggs and tadpoles in water-retaining arboreal bro- meliads or water-filled tree cavities Once these three components are added, their EVS can range from 3 to 19. Wilson and McCranie (2004) allo- cated the range of scores for Honduran amphibians into three categories of vulnerability to environmental degra- dation, as follows: low (3-9); medium (10-13); and high (14-19). We use the same categorization. Recent Changes to the Mexican Amphibian Fauna Our knowledge of the composition of the Mexican am- phibian fauna keeps changing due to discovery of new species and the systematic adjustment of certain known species, which adds or subtracts from the list of taxa that appeared in Wilson et al. (2010). Since that time, the fol- lowing seven species have been described or resurrected: Incilius aurarius: Mendelson et al. 2012. Journal of Herpetology 46: 473-479. New species. Incilius mccoyi : Santos-Barrera and Flores Villela. 2011. Journal of Herpetology 45: 211-215. New spe- cies. Craugastor saltator: Hedges et al. 2008. Zootaxa 1737: 1-182. Resurrected from synonymy of C. mexi- canus. August 2013 | Volume 7 | Number 1 | e69 Conservation reassessment of Mexican amphibians Eleutherodactylus modestus. The endemic blunt-toed chirping frog is known from Colima and southwestern Jalisco. Its EVS has been calculated at 16, placing it in the middle portion of the high vulnerability category, and its IUCN status as Vulnerable. This individual is from the Sierra de Manantlan in Jalisco. Photo by Ivan Trinidad Ahumada- Carrillo. Dendropsophus sartori. The endemic Taylor’s yellow treefrog is distributed along the Pacific slopes from Jalisco to Oaxaca. Its EVS has been determined as 14, at the lower end of the high vulnerability category, and its IUCN status as of Least Concern. This individual came from the Municipality of Minatitlan, Colima. Photo by Jacobo Reyes-Velasco. Amphib. Reptile Conserv. | http://amphibian-reptile-conservation.org 102 August 2013 | Volume 7 | Number 1 | e69 Wilson et al. Charadrahyla tecuani: Campbell et al. 2009. Copeia 2009: 287-295. New species. Gastrophryne mazatlanensis : Streicher et al. 2012. Molecular Phylogenetics and Evolution 64: 645-653. Resurrected from synonymy of G. olivacea. Bolitoglossa chinanteca : Rovito et al. 2012. ZooKeys 185: 55-71. New species. Pseudoeurycea cafetalera : Parra-Olea et al. 2010. Zootaxa 2725: 57-68. New species. This represents an increase of 2.0% over the 373 species listed by Wilson and Johnson (2010). The following species have undergone status changes, and include some taxa discussed in the addendum to Wil- son and Johnson (2010): Diaglena spatulata: Smith et al. 2007. Evolution 61: 2075-2085. Transfer from genus Triprion. Hypopachus ustus : Streicher et al. 2012. Molecular Phylogenetics and Evolution 64: 645-653. Transfer from genus Gastrophryne. Spelling of specific epithet corrected by Frost (2013). Trachycephalus typhonius : Lavilla et al. 2010. Zoo- taxa 2671: 17-30. New name for T. venulosus. Ixalotriton niger : Wake. 2012. Zootaxa 3484: 75-82. Resurrection of genus. Ixalotriton parva: Wake. 2012. Zootaxa 3484: 75-82. Resurrection of genus. IUCN Red List Assessment of Mexican Amphibians The IUCN assessment of Mexican amphibians was con- ducted as part of a Mesoamerican Workshop held in 2002 at the La Selva Biological Station in Costa Rica (see fore- word in Kohler 2011). The results of this workshop were incorporated into a general worldwide overview called the Global Amphibian Assessment (Stuart et al. 2004; Stuart et al. 2008; Stuart et al. 2010). This overview un- covered startling conclusions, of which the most impor- tant was that nearly one-third (32.3%) of the world’s am- phibian species are threatened with extinction, i.e., were assessed as Critically Endangered, Endangered, or Vul- nerable. This proportion did not include 35 species con- sidered as Extinct or Extinct in the Wild, and by adding them 1,891 of 5,743 species (32.9%) were considered as Table 1 . IUCN Red List categorizations for Mexican amphibian families. Number of species IUCN Red List categorizations Families Critically Endangered Endangered Vulnerable Near Threatened Least Concern Data Deficient Not Evaluated Bufonidae 35 1 7 2 3 19 1 2 Centrolenidae 1 — — — — 1 — — Craugastoridae 39 7 8 7 3 6 6 2 Eleutherodactylidae 23 2 4 7 — 4 5 1 Hylidae 97 29 18 10 4 25 8 3 Leiuperidae 1 — — — — 1 — — Leptodactylidae 2 — — — — 2 — — Microhylidae 6 — — 1 — 4 — 1 Ranidae 28 4 2 5 2 12 2 1 Rhinophrynidae 1 — — — — 1 — — Scaphiopodidae 4 — — — — 2 — 2 Subtotals 237 43 39 31 12 77 22 12 Ambystomatidae 18 9 2 — — 2 3 2 Plethodontidae 118 36 37 11 9 10 12 3 Salamandridae 1 — 1 — — — — — Sirenidae 2 — — — — 2 — — Subtotals 139 45 40 11 9 14 15 5 Dermophiidae 2 — — 1 — — 1 — Subtotals 2 — — 1 — — 1 — Totals 378 88 79 44 21 91 38 17 August 2013 | Volume 7 | Number 1 | e69 Amphib. Reptile Conserv. | http://amphibian-reptile-conservation.org 103 Conservation reassessment of Mexican amphibians Smilisca dentata. The endemic upland burrowing treefrog occurs only in southwestern Aguascalientes and adjacent northern Jalisco. Its EVS has been assessed as 14, placing it at the lower end of the high vulnerability category, and its IUCN status as Endangered. This individual was found in the Municipality of Ixtlahuacan del Rio, Jalisco. Photo by Jacobo Reyes-Velasco. Lithobates johni. Moore’s frog is an endemic anuran whose distribution is limited to southeastern San Luis Potosf, eastern Hidalgo, and northern Puebla. Its EVS has been assessed as 14, placing it at the lower end of the high vulnerability category, and its IUCN status as Endangered. This individual came from Rio Claro, Municipality of Molango, Hidalgo. Photo by Uriel Hernandez- Salinas. August 2013 | Volume 7 | Number 1 | e69 Amphib. Reptile Conserv. | http://amphibian-reptile-conservation.org 104 Wilson et al. threatened, near extinction, or extinct. Notably, another 1,290 species (22.5%) were evaluated as Data Deficient, i.e., too poorly known to allocate to any of the other IUCN categories. By adding these species to the previous figure of 1,891, an astonishing amount of amphibian spe- cies (3,181 [55.4%]) known at that time were considered threatened, near extinction, extinct, or too poorly known to assess. These horrific pronouncements gave rise to a worldwide cottage industry that continues to evaluate the state of amphibian population decline, as registered in a number of websites, most prominently AmphibiaWeb and the Global Amphibian Assessment. The IUCN Red List website lists the current catego- rizations for the world’s amphibians using the standard IUCN system. We accessed this website in order to sum- marize the current situation for Mexican amphibians (Table 1). The data in this table are more complete than those for reptiles, as reported by Wilson et al. (2013). All but 17 of the current 378 known Mexican amphibian spe- cies have been assigned to an IUCN category, and as for the reptiles (see Wilson et al. 2013) we placed these 17 amphibian taxa (4.5%) in a Not Evaluated (NE) category. The remaining categorizations are: Critically Endangered (CR; 88; 23.2%); Endangered (EN; 79; 20.8%); Vulner- able (VU; 44; 11.6%); Near Threatened (NT; 21; 5.5%); Least Concern (LC; 92; 24.2%); and Data Deficient (DD; 38; 10.0%). Thus, 211 species (55.7%) are placed in one of the three threat categories (CR, EN, or VU), a propor- tion significantly higher from that reported for these cat- egories on a global scale (CR+EN+VU = 1,856 species, 32.3%; Stuart et al., 2010). If the DD species are added to those in the threat categories, then 249 (65.7%) are either threatened with extinction or too poorly known to allow for assessment, a proportion significantly beyond that for the global situation (CR+EN+VU+DD = 3,146 species; 54.8%; Stuart et al. 2010). The largest proportion of threatened species are in the anuran families Craugastoridae (22 of 39 species; 56.4%), Eleutherodactylidae (13 of 24 species; 54.2%), and Hylidae (57 of 97 species; 58.8%), and the salaman- der families Ambystomatidae (11 of 19 species; 57.9%) and Plethodontidae (84 of 118 species; 71.2%). Collec- tively, the 297 species in these five families make up 78.4% of the amphibian taxa in Mexico, and the 187 threatened species in these families comprise 88.6% of the 211 total. Table 2. Environmental Vulnerability Scores for Mexican amphibian species, arranged by family. Shaded area to left encompasses low vul- nerability scores, and to the right high vulnerability scores. Families , Environmental Vulnerability Scores of species J 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 Bufonidae 35 1 — 1 2 2 2 3 2 6 4 5 5 2 — — — — Centrolenidae 1 1 Craugastoridae 39 — — — — — 1 1 1 1 1 4 4 10 3 5 8 — Eleutherodactylidae 23 2 3 — — 3 4 8 3 — Hylidae 97 1 2 — — 4 4 7 5 9 11 16 22 12 1 1 1 1 Leiuperidae 1 1 Leptodactylidae 2 — — 1 1 — — — — — — — — — — — — — Microhylidae 6 — 1 — — 1 2 1 1 Ranidae 28 1 — 1 — 1 2 2 2 2 5 4 5 3 — — — — Rhinophrynidae 1 1 Scaphiopodidae 4 1 — — 1 — — — 1 — 1 — — — — — — — Subtotals 237 4 3 3 4 9 12 14 13 20 25 29 36 30 8 14 12 1 Subtotals % — 1.7 1.3 1.3 1.7 3.8 5.1 5.9 5.4 8.4 10.5 12.2 15.2 12.7 3.4 5.9 5.1 0.4 Ambystomatidae 18 — — — — — — — 2 — — 4 5 7 — — — — Plethodontidae 118 — — — — — — 1 — 2 3 3 8 16 13 36 36 — Salamandridae 1 1 Sirenidae 2 — — — — — — — — — 2 — — — — — — — Subtotals 139 — — — — — — 1 2 2 6 7 13 23 13 36 36 — Subtotals % — — — — — — — 0.7 1.4 1.4 4.3 5.0 9.4 16.6 9.4 25.9 25.9 — Dermophiidae 2 1 1 — — — — — — — Subtotals 2 1 1 — — — — — — — Subtotals % — 50.0 50.0 — — — — — — — Totals 378 4 3 3 4 9 12 15 15 23 32 36 49 53 21 50 48 1 Totals % — 1.1 0.8 0.8 1.1 2.3 3.2 4.0 4.0 6.1 8.4 9.5 12.9 14.0 5.6 13.2 12.7 0.3 August 2013 | Volume 7 | Number 1 | e69 Amphib. Reptile Conserv. | http://amphibian-reptile-conservation.org 105 Conservation reassessment of Mexican amphibians Triprion petasatus. The Yucatecan casque-headed treefrog is restricted primarily to the Yucatan Peninsula, occurring in the Mexican states of Yucatan, Campeche, and Quintana Roo, as well as in northern Guatemala and northern Belize. A disjunct population also has been recorded from Santa Elena, Departamento de Cortes, Honduras. Its EVS has been calculated as 10, placing it at the lower end of the medium vulnerability category, and its IUCN status is of Least Concern. Although this treefrog is broadly distributed in the Yucatan Peninsula, it usually is found only during the rainy season when males and females congregate around restricted bodies of water (solution pits, cenotes, and ephemeral ponds) on this flat limestone platform. During the dry season, these frogs retreat into tree holes and rock crevices, and sometimes use their head to plug the opening. This individual is from the state of Yucatan. Photo by Ed Cassano. August 2013 | Volume 7 | Number 1 | e69 Amphib. Reptile Conserv. | http://amphibian-reptile-conservation.org 106 Wilson et al. These data from the IUCN Red List show a frighten- ing picture for the amphibian fauna of Mexico, acknowl- edged as a major herpetodiversity hotspot in the world on the basis of its diversity and endemism (Wilson and Johnson 2010). Mexico’s level of amphibian endemism (66.8%) also has been reported as greater than that for the country’s reptiles (57.2%; Wilson and Johnson 2010). Even more frightening is the fact that Mexican salaman- ders are more threatened than anurans (Table 1). Of the 139 recognized species of salamanders, 96 (69.1%) were assessed into one of the threat categories, as compared to anurans (114 of 236 [48.3%]). In addition, a much smaller proportion of salamander species were judged as Least Concern (14 [10.1%]), as compared to anurans (78 [33.1%]). Critique of the IUCN Assessment Although the conservation status of amphibians in Mex- ico is better understood than that for reptiles (see Wil- son et al. 2013), a need for reassessment still is required for several reasons. About 10% of Mexico’s amphib- ians have been judged as Data Deficient, and thus their conservation status remains undetermined. In addition, because certain species have been described recently (see above), 4.5% have not been evaluated (see www. iucnredlist.org; accessed 08 May 2013). Also, by adding the DD and NE species, 55 (14.5%) of Mexico’s amphib- ians presently are not assigned to any of the other IUCN categories. Thus, we consider it worthwhile to subject the Mexican amphibians to the same assessment measure ap- plied by Wilson et al. (2013) for reptiles, to allow for a comparison between these two groups. For these reasons, we will reassess the Mexican amphibian fauna using the Environmental Vulnerability Score (EVS). EVS for Mexican Amphibians The EVS provides several advantages for assessing the conservation status of amphibians and reptiles. First, this measure can be applied as soon as a species is named, because the information necessary for its application generally is known at that point. Second, calculating the EVS is economical because it does not require expen- sive, grant- supported workshops, such as those undertak- en for the Global Amphibian Assessment (sponsored by the IUCN). Third, the EVS is predictive, as it measures susceptibility to anthropogenic pressure and can pinpoint taxa with the greatest need of immediate attention and continued scrutiny. Finally, it is simple to calculate and does not “penalize” poorly known species. Thus, given the geometric pace at which environmental threats wors- en, since they are commensurate with the rate of human population growth, it is important to use a conservation assessment measure that can be applied simply, quickly, and economically. Amphib. Reptile Conserv. | http://amphibian-reptile-conservation.org 107 We calculated the EVS using the above-mentioned methodology. This step allowed us to determine the con- servation status of all the currently recognized Mexican amphibian species (378), including the 55 species placed in the DD category or not evaluated by the IUCN (www. iucnredlist.org; see Appendix 1, Table 2). Theoretically, the EVS can range from 3 to 20 (in Mexico, from 3 to 19). A score of 3 is indicative of a spe- cies that ranges widely both within and outside of Mexi- co, occupies eight or more forest formations, and lays its eggs in small to large lentic or lotic bodies of water. Four such species (one each in the families Bufonidae, Hyli- dae, Ranidae, and Scaphiopodidae) are found in Mexico. At the other extreme, a score of 20 relates to a species that is known only from the vicinity of the type locality, occupies a single forest formation, and its eggs and tad- poles are found in water-retaining arboreal bromeliads or water-filled tree cavities (no such species occur in Mex- ico). Thus, all the scores fall within the range of 4-19. In the Introduction, we expressed an interest in at- tempting to determine the impact of small populations on amphibian species survival in Mexico. The data in Appendix 1 allow us to approximate an answer to this question, inasmuch as one of the components of the EVS assesses the extent of geographic distribution on a sliding scale (1-6), on which higher numbers signify increas- ingly smaller geographic ranges. Using this range, the distribution of the 378 Mexican species is as follows: 1 = 13 species (3.4%); 2 = 20 (5.3%); 3 = 28 (7.4%); 4 = 64 (16.9%); 5 = 126 (33.3%); and 6 = 127 (33.6%). Obvi- ously, the higher the value of the geographic range, the higher the number and percentage of the taxa involved. These figures indicate that about one-third of the amphib- ian species in Mexico are known only from the vicinity of their respective type localities. The range of another one-third is somewhat broader, but still limited to the confines of Mexico. As a consequence, the survival pros- pects of about two-thirds of Mexico’s amphibians are tied to changes in their natural environment, as well as to the conservation atmosphere in this nation. We summarized the EVS for Mexican amphibians by family in Table 2. The EVS range falls into the follow- ing three portions: low (3-9), medium (10-13), and high (14-19). The range and average EVS for the major amphib- ian groups are as follows: anurans = 3-19 (12.4); sala- manders = 9-18 (15.9); and caecilians = 11-12 (11.5). Salamanders generally are significantly more susceptible than anurans to environmental degradation and caeci- lians somewhat less susceptible than anurans (although only two caecilian species are involved). The average scores either fall in the medium category, in the case of anurans and caecilians, or in the middle portion of the high category, in the case of salamanders. The average EVS for all amphibian species is 13.7, a value near the lower end of the high range of vulnerability. August 2013 | Volume 7 | Number 1 | e69 Conservation reassessment of Mexican amphibians Ambystoma velasci. The endemic Plateau tiger salamander, as currently recognized, is distributed widely from northwestern Chi- huahua southward along the eastern slopes of the Sierra Madre Oriental, and from southern Nuevo Leon in the Sierra Madre Ori- ental, westward to Zacatecas and southward onto the Transverse Volcanic Axis of central Mexico. Its EVS has been determined as 10, placing it at the lower end of the medium vulnerability category, and its IUCN status is of Least Concern. Even though this species does not appear threatened, this is likely an artifact of the composite nature of this taxon. This individual was found at Santa Cantarina, Hidalgo. Photo by Raciel Cruz-Elizalde. Bolitoglossa frcinklini. Franklin’s salamander is distributed along Pacific slopes from southern Chiapas, Mexico, southeastward to south-central Guatemala. Its EVS has been determined as 14, placing it at the lower end of the high vulnerability category, and its IUCN status as Endangered. This individual came from Cerro Mototal, in the Municipality of Motozintla, Chiapas. Photo by Sean M. Rovito. August 2013 | Volume 7 | Number 1 | e69 Amphib. Reptile Conserv. | http://amphibian-reptile-conservation.org 108 Wilson et al. An EVS of 14, at the lower end of the high vulnerabil- ity category, was found in the highest percentage (15.2) of anuran species. For salamanders, the respective values are 25.9% for an EVS of both 17 and 18, near the upper end of the range for the high vulnerability category, and for caecilians 50.0% for an EVS of both 11 and 12. The total EVS scores generally increased from the low end of the scale (3) through most of the high end (14-18), with a single exception (a decrease from 53 to 21 species at scores 15 and 16). An EVS of 15 was found in the peak number of taxa (53), a score that falls within the high range of vulnerability. Of the 378 total taxa, 50 (13.2%) fall into the low vulnerability category, 106 (28.0%) into the medium category, and 222 (58.7%) into the high category. Thus, six of every 10 Mexican amphibian species were judged as having the highest degree of vulnerability to environ- mental degradation, and slightly more than one-seventh the lowest degree. This considerable increase in the absolute and rela- tive numbers from the low portion, through the medium portion, to the high portion differs somewhat from the results published for amphibians and reptiles for sev- eral Central American countries in Wilson et al. (2010). Acevedo et al. (2010) reported 89 species (23.2%) with low scores, 179 (46.7%) with medium scores, and 115 (30.0%) with high scores for Guatemala. The same trend was reported for Honduras, where Townsend and Wilson (2010) indicated the corresponding values for amphib- ians and reptiles as 71 (19.7%), 169 (46.8%), and 121 (33.5%). The comparable data for the Panamanian her- petofauna in Jaramillo et al. (2010) are 143 (33.3%), 165 (38.4%), and 122 (28.4%). The principal reason that EVS scores are relatively high in Mexico is because of the high level of endemism and the concomitantly narrow range of geographical and ecological occurrence (Appendix 1). Of the 253 endemic amphibian species (139 anurans, 113 salamanders, and one caecilian), 125 (49.4%) were allocated a geographic distribution score of 6, signifying that these creatures are known only from the vicinity of their respective type localities; the remainder of the endemic species (128 [50.6%]) are more broadly distributed within the country (Appendix 1). Of the 378 Mexican amphibian species, 128 (33.9%) are limited in ecological distribution to one formation (Appendix 1). Therefore, we emphasize that close to one- half of the country’s endemic amphibian species are not known to occur outside of the vicinity of their type local- ities. In addition, essentially one-third are not known to occur outside of a single forest formation. This situation imposes serious challenges in our attempt to conserve the endemic component of the strikingly important Mexican amphibian fauna. Comparison of IUCN Categorizations and EVS Values Table 3. Comparison of Environmental Vulnerability Scores (EVS) and IUCN categorizations for Mexican amphibians. Shaded area at the top encompasses low vulnerability category scores, and that at the bottom high vulnerability category scores. IUCN categories EVS Critically Endangered Endangered Vulnerable Near Threatened Least Concern Data Deficient Not Evaluated Totals 3 4 5 6 7 8 1 — 2 2 4 3 3 3 8 6 — 1 2 4 3 3 4 9 12 9 1 1 1 1 10 — 1 15 10 1 2 1 — 9 — 2 15 11 1 2 7 — 13 — — 23 12 5 4 3 4 13 2 1 32 13 4 12 5 5 6 3 1 36 14 12 11 7 2 8 6 3 49 15 22 8 5 2 3 10 3 53 16 4 9 4 2 1 1 — 21 17 15 17 6 2 1 7 2 50 18 19 21 1 13 3 1 — 9 1 48 1 Totals 88 79 44 21 91 38 17 378 August 2013 | Volume 7 | Number 1 | e69 Amphib. Reptile Conserv. | http://amphibian-reptile-conservation.org 109 Conservation reassessment of Mexican amphibians Ixalotriton niger. The black jumping salamander is known only from the immediate vicinity of the type locality in northwestern Chiapas. Its EVS has been calculated as 18, placing it in the upper portion of the high vulnerability category, and its IUCN status as Critically Endangered. This individual came from the type locality and was used as part of the type series in the description of the species by Wake and Johnson (1989). The genus Ixalotriton is endemic to Mexico, and contains one other species (I. parvus). Photo by David B. Wake. Pseudoeurycea longicauda. The endemic long-tailed false brook salamander is distributed in the Transverse Volcanic Axis of east- ern Michoacan and adjacent areas in the state of Mexico. Its EVS has been determined as 17, placing it in the middle of the high vulnerability category, and its IUCN status as Endangered. This individual came from Zitacuaro, Michoacan, near the border with the state of Mexico. Photo by Ivan Trinidad Ahumada- Carrillo. August 2013 | Volume 7 | Number 1 | e69 Amphib. Reptile Conserv. | http://amphibian-reptile-conservation.org 110 Wilson et al. We noted in Wilson et al. (2013: 18) that, “Since the IUCN categorizations and EVS values both measure the degree of environmental threat impinging on a given spe- cies, a certain degree of correlation between the results, using the two measures, is expected.” They further in- dicated that Townsend and Wilson (2010) demonstrated this to be the case with the Honduran herpetofauna. Wil- son et al. (2013: 22) concluded, however, that, “the re- sults of the EVS analysis are nearly the reverse of those obtained from the IUCN categorizations.” We compared the results of these two conservation measures in Table 3, expecting that our results for the Mexican amphibians would be more consistent with those obtained for the Honduran herpetofauna (Townsend and Wilson 2010) than those garnered for the Mexican rep- tiles (Wilson et al. 2013). 1. Nature of the IUCN categorizations in Table 3 Like Wilson et al. (2013), we used the “Not Evaluated” category (IUCN 2010), since 17 species (4.5%) have not been evaluated at the IUCN Red List website, and 38 (10.1%) were evaluated as “Data Deficient” (www. iucnredlist.org; accessed 08 May 2013). Thus, the IUCN conservation status of 55 (14.6%) of the total amphibian species remained undetermined. A greater proportion of the Mexican amphibians, however, were assessed based on the IUCN categorizations (323 species [85.4%]) than the Mexican reptiles (Wilson et al. 2013). 2. Pattern of mean EVS vs. IUCN categorizations In order to more precisely determine the relationship be- tween the IUCN categorizations and the EVS, we cal- culated the mean EVS for each of the IUCN columns in Table 3, including for the NE species and the total species. The results are as follows: CR (88 spp.) = 15.5 (range 7-19); EN (79 spp.) = 15.1 (9-18); VU (44 spp.) = 13.8 (8-18); NT (21 spp.) = 13.3 (8-18); LC (91 spp.) = 10.0 (3-17); DD (38 spp.) = 15.6 (12-18); NE (17 spp.) = 12.6 (6-18); and total (378 spp.) = 13.7 (3-19). The results of these data show that the mean EVS decreases steadily from the CR category (15.5) through the EN (15.1), VU (13.8), and NT (13.3) categories to the LC category (10.0). This pattern of decreasing values was expected. In addition, the mean value for the DD species (15.6) is closest to that for the CR species. As we stated with regard to Mexican reptiles (Wilson et al. 2013: 22), “this indicates what we generally have suspected about the DD category, i.e., that the species placed in this cat- egory likely will fall into the EN or CR categories when (and if) their conservation status is better understood. Placing species in this category is of little benefit to de- termining their conservation status, however, since once sequestered with this designation their significance tends to be downplayed.” Wilson et al. (2013) demonstrated that this problem was more significant with Mexican reptiles, given that 118 species were evaluated as DD, which provided the impetus to work on the 38 amphibian Table 4. Comparison of Environmental Vulnerability Scores for Mexican amphibian and reptile species, arranged by major groups. Shaded area to the left encompasses low vulnerability scores, and to the right high vulnerability scores. Number of Environmental Vulnerability Scores major groups species 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 Anurans 237 4 3 3 4 9 12 14 13 20 25 29 36 30 8 14 12 1 — Percentages — 1.7 1.3 1.3 1.7 3.8 5.1 5.9 5.4 8.4 10.5 12.2 15.2 12.7 3.4 5.9 5.1 0.4 — Salamanders 139 — — — — — — 1 2 2 6 7 13 23 13 36 36 — — Percentages — — — — — — — 0.7 1.4 1.4 4.3 5.0 9.4 16.6 9.4 25.9 25.9 — — Caecilians 2 1 1 Percentages 50.0 50.0 Amphibian Totals 378 4 3 3 4 9 12 15 15 23 32 36 49 53 21 50 48 1 — Percentages — 1.0 0.8 0.8 1.0 2.4 3.2 4.0 4.0 6.1 8.5 9.5 13.0 14.0 5.5 13.2 12.7 0.3 — Crocodilians 3 1 1 — 1 — — — — Percentages — 33.3 33.3 — 33.3 — — — — Turtles 42 — — — — — 1 — 3 1 1 3 8 6 4 3 5 6 1 Percentages — — — — — — 2.4 — 7.1 2.4 2.4 7.1 19.0 14.3 9.5 7.1 11.9 14.3 2.4 Lizards 409 — — 1 3 6 11 12 15 26 39 49 54 67 77 37 10 2 — Percentages — — — 0.2 0.7 1.5 2.7 2.9 3.7 7.1 9.5 12.0 13.2 16.4 18.8 9.0 2.4 0.5 — Snakes 382 1 1 7 10 9 19 17 30 25 31 46 52 50 44 24 9 7 — Percentages — 0.3 0.3 1.8 2.6 2.4 5.0 4.5 7.9 6.5 8.1 12.0 13.6 13.1 11.5 6.3 2.4 1.8 — Reptile Totals 836 1 1 8 13 15 31 30 46 53 71 99 115 123 126 64 24 15 1 Percentages — 0.1 0.1 1.0 1.6 1.8 3.7 3.6 5.5 6.3 8.5 11.8 13.8 14.7 15.1 7.8 2.9 1.8 0.1 August 2013 | Volume 7 | Number 1 | e69 Amphib. Reptile Conserv. | http://amphibian-reptile-conservation.org 111 Conservation reassessment of Mexican amphibians Dermophis oaxacae. The endemic Oaxacan caecilian is distributed in Colima, Jalisco, Michoacan, Guerrero, Oaxaca, and Chiapas. Its EVS has been calculated as 12, placing it in the middle portion of the medium vulnerability category, and its IUCN status as Data Deficient. This individual was found on the road at Ixtlahuacan, Colima. Photo by Jacobo Reyes-Velasco. species assessed as DD with those occupying the threat categories (CR, EN, and VU) to arrive at a total of 249 species (65.9% of the total amphibian fauna). The EVS range for these DD species (12-18) falls within that for the threat species as a whole (7-19) and the mean for all the four categories becomes 15.1, the same as that for the EN species alone. So, if the DD species can be consid- ered “threat species in disguise,” then close to two-thirds of the Mexican amphibian species would be considered under the threat of extinction. The EVS for the 17 Mexican amphibian species that have not been evaluated by the IUCN range from 6 to 1 8 (mean = 12.6). These species are of significant conserva- tion interest, inasmuch as the EVS of nine of them falls into the range of high vulnerability. Based on the pattern of relationships between the LC species and their corresponding EVS, this IUCN cate- gory apparently has become a “dumping ground” for a sizable number of Mexican amphibians (9 1 ; 24. 1 % of the amphibian fauna) and like Wilson et al. (2013: 22) con- cluded for Mexican reptiles, we concur that “A more dis- cerning look at both the LC and NE species might dem- onstrate that many should be partitioned into other IUCN categories, rather than the LC.” The range of EVS values for this category (3-17) is almost as broad as the range of EVS (3-19) for amphibians as a whole; 37 (40.7%) of these 91 species are relegated to the low vulnerabil- ity range (3-9), 41 (45.0%) to the medium vulnerability range, and 13 (14.3%) to the high vulnerability range. Again, these results indicate that the LC category likely has been used rather indiscriminately and that the EVS algorithm provides a more useful conservation measure than the IUCN system of categories. Comparison of EVS Values for Mexican Amphibians and Reptiles One of our major reasons for writing this paper was to determine the EVS values for Mexican amphibians, so they could be compared to those calculated for Mexican reptiles in Wilson et al. (2013). Thus, we summarized the data in Table 2, and reduced them to the major group level in Table 4. We also reduced the data in Wilson et al. (2013: table 2) and placed them in our Table 4. The data in this table indicate that the range of EVS values are comparable for amphibians (3-19) and rep- tiles (3-20). The EVS for the number of amphibian spe- cies essentially increases until a score of 15 is reached (53 species), and at 16 drops considerably (21 species) only to spike back up at 17 and 18 (50 and 48 species, respectively). The highest EVS value (19) was assigned to a single species (the fringe-limbed hylid Ecnomiohyla echinata ). For the reptiles, the numbers and percentages also increase, with the peak (126 [15.1%]) reached at an August 2013 | Volume 7 | Number 1 | e69 Amphib. Reptile Conserv. | http://amphibian-reptile-conservation.org 112 Wilson et al. EVS of 16, and decreasing rapidly thereafter. As with amphibians, only a single species (the soft-shelled turtle Apalone atra ) was assigned the highest EVS (20). When the EVS values are arranged into low, medium, and high categories, the numbers and percentages of spe- cies are as follows (amphibians, followed by reptiles): low = 50 (13.2%), 99 (11.8%); medium = 106 (28.0%), 269, (32.2%); and high = 222 (58.8%), 468, (56.0%). The percentages for these two groups are comparable and arranged in the same order. The greatest concern is that in both amphibians and reptiles more than one-half of the species fall into the upper portion of the high vulnerabil- ity category, indicating that the Mexican herpetofauna is seriously imperiled. Of the major groups of amphibians and reptiles, Mex- ican salamanders were judged the most imperiled. Of the 139 species known from the country, 121 (87.1%) were assessed in the high vulnerability category. The compa- rable figure for anurans is 101 (42.6%), less than one- half of that for salamanders. Among the reptiles, lizards were judged more threatened than snakes. Of the lizards, 247 (60.4%) fall within the high vulnerability category; the comparable figures for snakes are 186 and 48.7%. Turtles, although fewer in numbers, are more threatened than other reptiles, with 33 species (78.6%) in the high vulnerability category. In the final analysis, although amphibians are ac- knowledged widely as threatened on a global basis, a fair accounting of the worldwide conservation status of most reptiles remains unavailable. Our use of the EVS mea- sure for Mexican amphibians and reptiles demonstrates that both groups are in grave peril, and we expect that this situation will worsen exponentially in the coming decades. Discussion Global amphibian population decline has occupied the attention of herpetologists since the late 1980s (Gas- con et al. 2007). In the years that followed, the Global Amphibian Assessment (GAA) was undertaken (Stuart et al. 2004), which uncovered the startling conclusions discussed in the Introduction. As noted in the foreword to Gascon et al. (2007: 2), “the first GAA documented the breadth of amphibian losses worldwide and made it clear that business as usual — the customary conserva- tion approaches and practices — were not working.” As a result, an Amphibian Conservation Summit was con- vened in September 2005, which resulted in a putatively comprehensive Amphibian Conservation Action Plan (ACAP; Gascon et al. 2007). The ACAP declaration pro- posed (p. 59) that, “Four kinds of intervention are needed to conserve amphibians, all of which need to be started immediately: 1 . Expanded understanding of the causes of declines and extinctions Amphib. Reptile Conserv. | http://amphibian-reptile-conservation.org 113 2. Ongoing documentation of amphibian diversity, and how it is changing 3. Development and implementation of long-term conservation programmes 4. Emergency responses to immediate crises.” We maintain that the ACAP does an admirable job of ex- amining many of the issues directly related to amphibian decline, but this examination essentially stops after con- sidering the proximate symptoms of the problem. None- theless, as noted by Wilson and Townsend (2010: 774), “problems created by humans, i.e., overpopulation and its sequelae, are not solved by treating only their symp- toms, e.g., organismic endangerment.” Consequently, trying to deal with a symptom of overpopulation and resource overuse and abuse, such as amphibian decline, will create only limited short-term responses, instead of lasting solutions to the fundamental problems tied to the impact of humans. Thus, ultimately, amphibian decline will not be successfully addressed. The fundamental problem is that humans have not created a sustainable existence for themselves. Under- standing why not is simple through examination of the principles of sustainability elaborated by Miller and Spoolman (2012: 6), as follows: • “Nature has sustained itself for billions of years by relying on solar energy, biodiversity, and nutrient cy- cling. • Our lives and economies depend on energy from the sun and on natural resources and natural services 0 natural capital ) provided by the earth. • As our ecological footprints grow, we are depleting and degrading more of the earth’s natural capital. • Major causes of environmental problems are popula- tion growth, wasteful and unsustainable resource use, poverty, and not including the harmful environmental costs of resource use in the market prices of goods and services. • Our environmental worldview plays a key role in determining whether we live unsustainably or more sustainably. • Living sustainably means living off the earth’s natu- ral income without depleting or degrading the natural capital that supplies it.” Living unsustainably is a consequence of unregulated human population growth that generates the overuse and abuse of renewable and non-renewable resources, and dependence on a cost-accounting system that ignores factoring in clean up expenses in determining how goods and services are priced. Life-sustaining resources are not distributed equitably among people, but along a scale ranging from very high to very low. Poverty is the conse- quence of existing at the low end of the scale, where peo- ple are unable to meet their basic needs for adequate food and water, clothing, or shelter (Raven and Berg 2004). August 2013 | Volume 7 | Number 1 | e69 Conservation reassessment of Mexican amphibians Environmental scientists use the concept of ecological footprint to express “the average amount of land and ocean needed to supply an individual with food, energy, water, housing, transportation, and waste disposal” (Ra- ven and Berg 2004: G-5). The global ecological footprint has increased over the years to the point that the Global Footprint Network calculated it would take “1.5 years to generate the renewable resources used in 2008” (WWF Fiving Planet Report 2012: 40). “Humanity’s annual de- mand on the natural world has exceeded what the Earth can renew in a year since the 1970s,” which has created a so-called “ecological overshoot” (WWF Fiving Planet Report 2012: 40). Thus, Earth’s capital (its biocapacity) is being depleted on a continually growing basis, and the planet is becoming less capable of supporting life in gen- eral, and human life in particular. Estimates indicate that by the year 2050, under a “business as usual” scenario, it would require an equivalent of 2.9 planets to support the amount of humanity expected to exist at that time (WWF Fiving Planet Report 2012: 101). The World Wildlife Fund promulgated its “One Planet perspective,” which “explicitly proposes to manage, gov- ern and share natural capital within the Earth’s ecologi- cal boundaries. In addition to safeguarding and restoring this natural capital, WWF seeks better choices along the entire system of production and consumption, supported by redirected financial flows and more equitable resource governance. All of this, and more, is required to decou- ple human development from unsustainable consump- tion (moving away from material and energy-intensive commodities), to avoid greenhouse gas emissions, to maintain ecosystem integrity, and to promote pro-poor growth and development” (WWF Fiving Planet Report 2012: 106). Only within this context will the provisions of ACAP have the desired effects, i.e., to preserve the portion of natural capital represented by amphibians. Thus, in writ- ing about the conservation status of the amphibians of Mexico, we are constructing our conclusions and recom- mendations in light of these global imperatives. Conclusions and Recommendations We structured our conclusions and recommendations af- ter those of Wilson and Townsend (2010) for the entire Mesoamerican herpetofauna, refining them specifically for the Mexican amphibian fauna, as follows: 1. Given that Mexico contains the highest level of amphibian diversity and endemicity in the Me- soamerican biodiversity hotspot, our most funda- mental recommendation is that protection of this aspect of the Mexican patrimony should be made a major component of the management strategy of the Secretarra de Medio Ambiente y Recursos Na- turales (SEMARNAT). In turn, that strategy needs to be incorporated into an overall plan for a sus- Amphib. Reptile Conserv. | http://amphibian-reptile-conservation.org 114 tainable future for Mexico, of which the most criti- cal component is to “explicitly integrate population dynamics (size, growth rate, composition, location and migration) and per capita consumption trends into national planning policies to support a better balance between population and available resourc- es” (WWF Fiving Planet Report 2012: 121). 2. All organisms have intrinsic and extrinsic value, especially as components of healthily functioning ecosystems, but we believe that although conserva- tion efforts should extend to all species in a given area, most interest should be focused on species with a limited distribution (i.e., endemic species). The rationale for this position is that funds to sup- port conservation initiatives have remained scarce, although this situation will have to change in the near future. The principal regions of Mexican am- phibian endemism are the Sierra Madre Oriental, the Sierra Madre del Sur, and the Mesa Central, in the order listed. Unfortunately, about 39% of Mexico’s population occupies the Mesa Central (Flores- Villela et al. 2010). Inasmuch as this con- centrated population will continue to grow into the foreseeable future, not only as a consequence of the rate of natural increase (1.4% in Mexico), but also because of the increase in the percentage of the population attracted to the large cities of the Mesa Central (Guadalajara, Feon, Mexico, Mo- relia, Salamanca, and others; Flores- Villela et al. 2010), it is critically important to make the am- phibian fauna of the Mesa Central a fundamental component of the national plan for biodiversity protection by SEMARNAT. 3. Oscar Flores- Villela and his colleagues produced highly significant conservation analyses (Flores- Villela 1993; Flores- Villela and Gerez 1994; Ochoa-Ochoa and Flores- Villela 2006; Flores- Vil- lela et al. 2010) that have documented the centers of diversity and endemism of the Mexican herpe- tofauna. Given the large disparity between these centers and the placement of protected areas in the country, we can only echo the conclusions of Flores- Villela et al. (2010: 313) that, “Given the great importance of the herpetofauna of the Central Highlands of Mexico, both in terms of its diversity and endemicity, appropriate steps need to be taken quickly to establish protected areas around the cen- ter of herpetofaunal endemism in the Sierra Madre del Sur, and to reassess the ability of the protected areas already established in the Mesa Central to encompass their centers of endemism.” A simi- lar recommendation can be made with respect to the other centers, e.g., the Sierra Madre Oriental, which has been even more ignored than areas in the Central Highlands (Favrn et al. 2010). August 2013 | Volume 7 | Number 1 | e69 Wilson et al. 4. Finding ways to use biodiversity sustainably must become a fundamental goal for all humanity. The steps necessary to achieve this end are not difficult to envision; the problem lies in marshaling the par- adigm shift necessary to make the transition. The major steps involve: (a) creating a reality-based educational system that will prepare people for the world as it is and will come to be, instead of the way people wish it were; (b) integrating education- al reform into a broad-based plan for governmen- tal and economic reform founded on principles of equality, shared responsibility, and commitment to a sustainable future for humanity and the natural world; (c) using governmental and economic re- form to design a global society structured to exist within the limits of nature; and (d) basing a soci- ety on the notion that everyone must work toward this end. Within such overarching goals, the task of learning the best way to catalogue, protect, and make sustainable use of the world’s organisms is a huge undertaking. New molecular-based technol- ogy, however, is allowing for a better understand- ing of biological diversity, which is much greater than we previously envisioned. Because of the ac- celerating rate at which we are losing biological di- versity, biologists are faced with helping humanity adopt a worldview in which all species matter, and that the sustainability of humans will depend on reforming our society based on the framework for survival tested by the process of natural selection over the last 3.5 billion years life has occupied our planet (Beattie and Ehrlich 2004). 5. In 2012, the United Nations Secretary-General’s High-level Panel on Global Sustainability pro- duced a seminal report entitled “Resilient People, Resilient Planet: A Future Worth Choosing.” In a vision statement (p. 13), the panel introduced the concept of “tipping points,” as follows: “The cur- rent global development model is unsustainable. We can no longer assume that our collective ac- tions will not trigger tipping points as environmen- tal thresholds are breached, risking irreversible damage to both ecosystems and human communi- ties. At the same time, such thresholds should not be used to impose arbitrary growth ceilings on de- veloping countries seeking to lift their people out of poverty. Indeed, if we fail to resolve the sus- tainable development dilemma, we run the risk of condemning up to 3 billion members of our human family to a life of endemic poverty. Neither of these outcomes is acceptable, and we must find a new way forward.” The panel also pointed out (p. 14) that “it is time for bold global efforts, includ- ing launching a major global scientific initiative, to strengthen the interface between science and policy. We must define, through science, what sci- entists refer to as ‘planetary boundaries,’ ‘environ- mental thresholds,’ and ‘tipping points.” On p. 23, they emphasize that, “awareness is growing of the potential for passing ‘tipping points’ beyond which environmental change accelerates, has the poten- tial to become self-perpetuating, and may be dif- ficult or even impossible to reverse.” Environmen- tal scientists have warned of this eventuality for decades; most of the world’s people just have not listened. The Stockholm Resilience Centre (www. stockholmresilience.org), however, has exposed a number of “planetary boundaries,” defined as certain thresholds or tipping points beyond which there is the “risk of irreversible and abrupt environ- mental change” (Box 2 on p. 24 of the UN panel re- port). The Stockholm Resilience Centre sponsored a group of scientists (Rockstrom et al. 2009) that identified nine planetary boundaries, including: “climate change, rate of biodiversity loss, biogeo- chemical flows (both nitrogen and phosphorus), stratospheric ozone depletion, ocean acidification, global freshwater use, change in land use, atmo- spheric aerosol loading and chemical pollution.” The scientists estimated that “human activity ap- pears to have already transgressed the [planetary] boundaries associated with climate change, rate of biodiversity loss and changes to the global nitrogen cycle.” Furthermore, “humanity may soon be ap- proaching the boundaries for interference with the global phosphorous cycle, global freshwater use, ocean acidification and global change in land use.” Finally, they concluded that, “the boundaries are strongly interlinked, so that crossing one may shift others and even cause them to be overstepped.” As a consequence of these realities, governments across the globe are faced with the choice of con- tinuing to do “business as usual,” ultimately spill- ing over all the planetary boundaries and ending up in a world in which all of our options have been ex- hausted except for the last one... the option to fail, or to pull together to develop a human existence lying within planetary boundaries in order to define a “safe operating space for humanity.” Our chances to avoid the one and succeed with the other will depend on how well humanity is able to embrace new ways of thinking about our problems and en- list the help of groups of people who traditionally have been marginalized — especially women and the young. These words apply to Mexico, as they do to all other countries in the world. The three authors of this work are herpetologists who specialize in research on amphibians and reptiles in Me- soamerica. This paper focuses on the conservation status of the amphibians of Mexico, and follows a si mil ar effort on the reptiles (Wilson et al. 2013). We demonstrated by using both the IUCN categorizations and EVS measure August 2013 | Volume 7 | Number 1 | e69 Amphib. Reptile Conserv. | http://amphibian-reptile-conservation.org 115 Conservation reassessment of Mexican amphibians that the Mexican amphibian fauna is one of the most se- riously threatened of any existing in the world. All indi- cations suggest that humans have transgressed the plan- etary boundaries associated with biodiversity loss, and there is no time to lose to reverse this dismantling trend or our descendants will be left to conclude that our gen- eration condemned them to an environmentally impover- ished world by our inaction. In the final analysis, life on Earth has survived five prior mass extinction events; hu- manity’s job now is to survive the one of its own making. Acknowledgments. — We are thankful to the follow- ing individuals for improving the quality of this contribu- tion: Javier Alvarado-Dfaz, Irene Goyenechea, and Louis W. Porras. We are most grateful to Louis, who applied his amazing editing skills to the job of making our work better than what we initially produced. Literature Cited Acevedo M, Wilson LD, Cano EB, Vasquez-Almazan C. 2010. Diversity and conservation status of the Gua- temalan herpetofauna. Pp. 406-435 In: Conservation of Mesoamerican Amphibians and Reptiles. 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Streicher JW, Cox CF, Campbell JA, Smith EN, de Sa RO. 2012. Rapid range expansion in the Great Plains narrow-mouthed toad ( Gastrophryne olivacea) and a revised taxonomy for North American microhylids. Molecular Phylogenetics and Evolution 64: 645-653. Stuart SN, Chanson JS, Cox NA, Young BE, Rodrigues ASF, Fischman DF, Waller RW. 2004. Status and trends of amphibian declines and extinctions world- wide. Science 306(5702): 1783-1786. Stuart SN, Hoffman M, Chanson JS, Cox NA, Berridge RJ, Ramani P, Young BE. 2008. Threatened Amphib- ians of the World. Fynx Ediciones, Barcelona, Spain. Stuart SN, Chanson JS, Cox NA, Young BE. 2010. The global decline of amphibians: current trends and fu- ture prospects. Pp. 2-15 In: Conservation of Meso- american Amphibians and Reptiles. Editors, Wilson ED, Townsend JH, Johnson JD. Eagle Mountain Pub- lishing, EC, Eagle Mountain, Utah, USA. Townsend JH, Wilson ED. 2010. Conservation of the Honduran herpetofauna: Issues and imperatives. Pp. 460-435. In: Conservation of Mesoamerican Amphib- ians and Reptiles. Editors, Wilson ED, Townsend JH, Johnson JD. Eagle Mountain Publishing, LC, Eagle Mountain, Utah, USA. United Nations Secretary-General’s High-level Panel on Global Sustainability. 2012. Resilient People, Resil- ient Planet: A Future Worth Choosing. United Na- tions, New York, New York, USA. Vitt LJ, Caldwell JP. 2009. Herpetology (3 rd edition). Academic Press, Burlington, Maine, USA. Wake DB. 2012. Taxonomy of salamanders of the family Plethodontidae (Amphibia: Caudata). Zootaxa 3484: 75-82. Wake DB, Johnson JD. 1989. Anew genus and species of plethodontid salamander from Chiapas, Mexico. Con- tributions in Science Natural History Museum of Los Angeles County 411: 1-10. Wilson LD, Johnson JD. 2010. Distributional patterns of the herpetofauna of Mesoamerica, a biodiversity hotspot. Pp. 30-235 In: Conservation of Mesoameri- can Amphibians and Reptiles. Editors, Wilson LD, Townsend JH, Johnson JD. Eagle Mountain Publish- ing, LC, Eagle Mountain, Utah. Wilson LD, Mata-Silva V, Johnson JD. 2013. A conser- vation reassessment of the reptiles of Mexico based on the EVS measure. Amphibian & Reptile Conserva- tion 7: 1-47. Wilson LD, McCranie JR. 2004. The conservation status of the herpetofauna of Honduras. Amphibian & Rep- tile Conservation 3: 6-33. Wilson LD, Townsend JH. 2010. The herpetofauna of Mesoamerica: Biodiversity significance, conservation status, and future challenges. Pp. 76-812 In: Conser- vation of Mesoamerican Amphibians and Reptiles. Editors, Wilson LD, Townsend JH, Johnson JD. Eagle Mountain Publishing, LC, Eagle Mountain, Utah, USA. Wilson LD, Townsend JH, Johnson JD. 2010. Conserva- tion of Mesoamerican Amphibians and Reptiles. Ea- gle Mountain Publishing, LC, Eagle Mountain, Utah, USA. World Wildlife Fund. 2012. Living Planet Report 2012. WWF International, Gland, Switzerland. Received: 05 March 2013 Accepted: 26 April 2013 Published: 02 August 2013 August 2013 | Volume 7 | Number 1 | e69 Amphib. Reptile Conserv. | http://amphibian-reptile-conservation.org 117 Conservation reassessment of Mexican amphibians Larry David Wilson is a herpetologist with lengthy experience in Mesoamerica, totaling six collective years (combined over the past 47). Larry is the senior editor of the recently published Conservation of Mesoameri- can Amphibians and Reptiles and a co-author of seven of its chapters. He retired after 35 years of service as Professor of Biology at Miami-Dade College in Miami, Florida. Larry is the author or co-author of more than 290 peer-reviewed papers and books on herpetology, including the 2004 Amphibian & Reptile Conserva- tion paper entitled “The conservation status of the herpetofauna of Honduras.” His other books include The Snakes of Honduras, Middle American Herpetology, The Amphibians of Honduras, Amphibians & Reptiles of the Bay Islands and Cayos Cochinos, Honduras, The Amphibians and Reptiles of the Honduran Mosquitia, and Guide to the Amphibians & Reptiles of Cusuco National Park, Honduras. He also served as the Snake Section Editor for the Catalogue of American Amphibians and Reptiles for 33 years. Over his career, Larry has authored or co-authored the descriptions of 69 currently recognized herpetofaunal species and six spe- cies have been named in his honor, including the anuran Craugastor lauraster and the snakes Cerrophidion wilsoni, Myriopholis wilsoni, and Oxybelis wilsoni. Jerry D. Johnson is Professor of Biological Sciences at The University of Texas at El Paso, and has exten- sive experience studying the herpetofauna of Mesoamerica. He is the Director of the 40,000 acre “Indio Mountains Research Station,” was a co-editor on the recently published Conservation of Mesoamerican Amphibians and Reptiles, and is Mesoamerica/Caribbean editor for the Geographic Distribution section of Herpetological Review. Johnson has authored or co-authored over 80 peer-reviewed papers, including two 2010 articles, “Geographic distribution and conservation of the herpetofauna of southeastern Mexico” and “Distributional patterns of the herpetofauna of Mesoamerica, a biodiversity hotspot.” Vicente Mata-Silva is a herpetologist interested in ecology, conservation, and the monitoring of amphibians and reptiles in Mexico and the southwestern United States. His bachelor’s thesis compared herpetofaunal richness in Puebla, Mexico, in habitats with different degrees of human related disturbance. Vicente’s master’s thesis focused primarily on the diet of two syntopic whiptail species of lizards, one unisexual and the other bisexual, in the Trans-Pecos region of the Chihuahuan Desert. Currently, he is a postdoctoral research fellow at the University of Texas at El Paso, where his work focuses on rattlesnake populations in their natural habitat. His dissertation was on the ecology of the rock rattlesnake, Crotalus lepidus, in the northern Chihuahuan Desert. To date, Vicente has authored or co-authored 34 peer-reviewed scientific publications. August 2013 | Volume 7 | Number 1 | e69 Amphib. Reptile Conserv. http://amphibian-reptile-conservation.org 118 Wilson et al. Appendix 1 . Comparison of the IUCN Ratings from the Red List Website (updated to 08 May 2013) and Environmental Vulnerability Scores for 378 Mexican Amphibians. See text for explanations of the IUCN and EVS rating systems. * = species endemic to Mexico. Species IUCN rating Environmental Vulnerability Score Geographic Distribution Ecological Distribution Reproductive Mode Total Score Order Anura (237 species) Family Bufonidae (35 species) Anaxyrus boreus NT 3 4 1 8 Anaxyrus californicus EN 4 7 1 12 Anaxyrus cognatus LC 3 5 1 9 Anaxyrus compactilis* LC 5 8 1 14 Anaxyrus debilis LC 1 5 1 7 Anaxyrus kelloggi* LC 5 8 1 14 Anaxyrus mexicanus* NT 5 7 1 13 Anaxyrus punctatus LC 1 3 1 5 Anaxyrus retiformis LC 4 7 1 12 Anaxyrus speciosus LC 4 7 1 12 Anaxyrus woodhousii LC 3 6 1 10 Incilius alvarius LC 4 6 1 11 Incilius aurarius NE 4 8 1 13 Incilius bocourti LC 4 6 1 11 Incilius campbelli NT 4 8 1 13 Incilius canaliferus LC 4 3 1 8 Incilius cavifrons* EN 5 7 1 13 Incilius coccifer LC 3 5 1 9 Incilius cristatus* CR 5 8 1 14 Incilius cycladen* VU 5 8 1 14 Incilius gemmifer* EN 6 8 1 15 Incilius luetkenii LC 3 3 1 7 Incilius macrocristatus VU 4 6 1 11 Incilius marmoreus* LC 5 5 1 11 Incilius mazatlanensis* LC 5 6 1 12 Incilius mccoyi* NE 5 8 1 14 Incilius nebulifer LC 1 4 1 6 Incilius occidental is* LC 5 5 1 11 Incilius perplexus* EN 5 5 1 11 Incilius pisinnus* DD 6 8 1 15 Incilius spiculatus* EN 5 7 1 13 Incilius tacanensis EN 4 4 1 9 Incilius tutelarius EN 4 5 1 10 Incilius valliceps LC 3 2 1 6 Rhinella marina LC 1 1 1 3 Family Centrolenidae (1 species) Hyalinobatrachium fleischmanni LC 3 4 3 10 Family Craugastoridae (39 species) Craugastor alfredi VU 2 5 4 11 Craugastor amniscola DD 4 6 4 14 Craugastor augusti LC 2 2 4 8 Craugastor batrachylus* DD 6 8 4 18 August 2013 | Volume 7 | Number 1 | e69 Amphib. Reptile Conserv. | http://amphibian-reptile-conservation.org 1 1 9 Conservation reassessment of Mexican amphibians Craugastor berkenbuschii* NT 5 5 4 14 Craugastor brocchi VU 4 6 4 14 Craugastor decoratus* VU 5 6 4 15 Craugastor galacticorhinis* NE 6 8 4 15 Craugastor glaucus * CR 6 8 4 18 Craugastor greggi CR 4 7 4 15 Craugastor guerreroensis* CR 6 8 4 18 Craugastor hobartsmithi* EN 5 6 4 15 Craugastor laticeps NT 4 4 4 12 Craugastor lineatus CR 4 7 4 15 Craugastor loki LC 2 4 4 10 Craugastor matudai VU 4 7 4 15 Craugastor megalotympanum* CR 6 8 4 18 Craugastor mexicanus* LC 5 7 4 16 Craugastor montan us* EN 6 8 4 18 Craugastor occidental is* DD 5 4 4 13 Craugastor omiltemanus* EN 5 7 4 16 Craugastor palenque DD 4 7 4 15 Craugastor pelorus * DD 5 6 4 15 Craugastor polymniae * CR 6 8 4 18 Craugastor pozo* CR 6 7 4 17 Craugastor pygmaeus VU 2 3 4 9 Craugastor rhodopis* VU 5 5 4 14 Craugastor rugulosus* LC 5 4 4 13 Craugastor rupinius LC 4 5 4 13 Craugastor saltator* NE 5 6 4 15 Craugastor silvicola * EN 6 8 4 18 Craugastor spatulatus* EN 5 7 4 16 Craugastor stuarti EN 4 7 4 15 Craugastor tarahumaraensis* VU 5 8 4 17 Craugastor taylori* DD 6 8 4 18 Craugastor uno* EN 5 8 4 17 Craugastor vocal is* LC 5 4 4 13 Craugastor vulcani* EN 6 7 4 17 Craugastor yucatanensis * NT 5 8 4 17 Family Eleutherodactylidae (23 species) Eleutherodactylus albolabris* NE 6 7 4 17 Eleutherodactylus angustidigitorum * VU 5 8 4 17 Eleutherodactylus cystignathoides LC 2 6 4 12 Eleutherodactylus dennisi* EN 6 8 4 18 Eleutherodactylus dilatus* EN 5 8 4 17 Eleutherodactylus grand is* CR 6 8 4 18 Eleutherodactylus guttilatus LC 2 5 4 11 Eleutherodactylus interorbital is * DD 5 6 4 15 Eleutherodactylus leprus VU 2 6 4 12 Eleutherodactylus longipes* VU 5 6 4 15 Eleutherodactylus maurus* DD 5 8 4 17 Eleutherodactylus modestus* VU 5 7 4 16 Eleutherodactylus nitidus* LC 5 3 4 12 August 2013 | Volume 7 | Number 1 | e69 Amphib. Reptile Conserv. | http://amphibian-reptile-conservation.org 1 20 Wilson et al. Eleutherodactylus nivicolimae * VU 6 7 4 17 Eleutherodactylus pallidus* DD 5 8 4 17 Eleutherodactylus pipilans LC 2 5 4 11 Eleutherodactylus rubrimaculatus VU 4 7 4 15 Eleutherodactylus rufescens* CR 6 7 4 17 Eleutherodactylus saxatilis* EN 5 8 4 17 Eleutherodactylus syristes* EN 5 7 4 16 Eleutherodactylus teretistes* DD 5 7 4 16 Eleutherodactylus verrucipes* VU 5 7 4 16 Eleutherodactylus verruculatus* DD 6 8 4 18 Family Hylidae (97 species) Acris blanchardi NE 3 8 1 12 Agalychnis callidryas LC 3 5 3 11 Agalychnis dacnicolor* LC 5 5 3 13 Agalychnis moreletii CR 1 3 3 7 Anotheca spinosa LC 3 6 5 14 Bromeliohyla bromeliacia EN 4 7 5 16 Bromeliohyla dendroscarta* CR 5 7 5 17 Charadrahyla altipotens* CR 5 6 1 12 Charadrahyla chaneque* EN 5 7 1 13 Charadrahyla nephila* VU 5 7 1 13 Charadrahyla taeniopus* VU 5 7 1 13 Charadrahyla tecuani* NE 6 8 1 15 Charadrahyla trux* CR 6 7 1 14 Dendropsophus ebraccatus LC 3 6 3 10 Dendropsophus microcephalus LC 3 3 1 7 Dendropsophus robertmertensi LC 4 4 1 9 Dendropsophus sartori* LC 5 8 1 14 Diaglena spatulata* LC 5 7 1 13 Duellmanohyla chamulae* EN 6 7 1 13 Duellmanohyla ignicolor* EN 6 7 1 14 Duellmanohyla schmidtorum VU 4 3 1 8 Ecnomiohyla echinata* CR 6 8 5 19 Ecnomiohyla miotympanum* NT 5 3 1 9 Ecnomiohyla valancifer* CR 6 7 5 18 Exerodonta abdivita* DD 6 8 1 15 Exerodonta bivocata* DD 6 8 1 15 Exerodonta chimalapa* EN 6 5 1 12 Exerodonta juanitae* VU 5 8 1 14 Exerodonta melanomma* VU 5 5 1 11 Exerodonta pinorum* VU 5 7 1 13 Exerodonta smaragdina* LC 5 6 1 12 Exerodonta sumichrasti* LC 5 3 1 9 Exerodonta xera* VU 5 8 1 14 Hyla arboricola* DD 5 6 1 12 Hyla arenicolor LC 2 4 1 7 Hyla euphorbiacea* NT 5 7 1 13 Hyla eximia* LC 5 4 1 10 Hyla plicata* LC 5 5 1 11 August 2013 | Volume 7 | Number 1 | e69 Amphib. Reptile Conserv. | http://amphibian-reptile-conservation.org 121 Conservation reassessment of Mexican amphibians Hyla walkeri VU 4 6 1 11 Hyla wrightorum LC 2 6 1 9 Megastomatohyla mixe* CR 6 8 1 15 Megastomatohyla mixomaculata* EN 5 8 1 14 Megastomatohyla nubicola* EN 5 8 1 14 Megastomatohyla pellita* CR 6 7 1 14 Plectrohyla acanthodes CR 4 7 1 12 Plectrohyla ameibothalame* DD 6 8 1 15 Plectrohyla arborescandens* EN 5 5 1 11 Plectrohyla avia CR 4 8 1 13 Plectrohyla bistincta* LC 5 3 1 9 Plectrohyla calthula* CR 5 8 1 14 Plectrohyla calvicollina* CR 6 7 1 14 Plectrohyla celata* CR 6 7 1 14 Plectrohyla cembra* CR 5 8 1 14 Plectrohyla charadricola* EN 5 8 1 14 Plectrohyla chryses* CR 6 7 1 14 Plectrohyla crassa* CR 5 8 1 14 Plectrohyla cyanomma* CR 5 8 1 14 Plectrohyla cyclada* EN 5 8 1 14 Plectrohyla ephemera* CR 6 8 1 15 Plectrohyla guatemalensis CR 4 4 1 9 Plectrohyla hartwegi CR 4 5 1 10 Plectrohyla hazelae* CR 5 6 1 12 Plectrohyla ixil CR 4 7 1 12 Plectrohyla labedactyla* DD 6 8 1 15 Plectrohyla lacertosa* EN 5 8 1 14 Plectrohyla matudai VU 4 6 1 11 Plectrohyla miahuatlanensis* DD 6 8 1 15 Plectrohyla mykter* EN 5 7 1 13 Plectrohyla pachyderma* CR 6 8 1 15 Plectrohyla pentheter* EN 5 7 1 13 Plectrohyla psarosema* CR 6 8 1 15 Plectrohyla pych nochi la* CR 6 8 1 15 Plectrohyla robertsorum* EN 5 7 1 13 Plectrohyla sabrina* CR 5 8 1 14 Plectrohyla sagorum EN 4 5 1 10 Plectrohyla siopela* CR 6 8 1 15 Plectrohyla thorectes* CR 5 7 1 13 Pseudacris cadaverina LC 4 6 1 11 Pseudacris dark i LC 3 8 1 12 Pseudacris hypochondriaca NE 4 4 1 9 Ptychohyla acrochorda* DD 6 7 1 14 Ptychohyla erythromma* EN 5 7 1 13 Ptychohyla euthysanota NT 4 3 1 8 Ptychohyla leonhardschultzei* EN 5 6 1 12 Ptychohyla macrotympanum CR 4 6 1 11 Ptychohyla zophodes* DD 5 7 1 13 Scinax staufferi LC 2 1 1 4 August 2013 | Volume 7 | Number 1 | e69 Amphib. Reptile Conserv. | http://amphibian-reptile-conservation.org 1 22 Wilson et al. Smilisca baudinii LC 1 1 1 3 Smilisca cyanosticta NT 4 7 1 12 Smilisca dentata* EN 5 8 1 14 Smilisca fodiens LC 2 5 1 8 Tlalocohyla godmani* VU 5 7 1 13 Tlalocohyla loquax LC 3 3 1 7 Tlalocohyla picta LC 2 5 1 8 Tlalocohyla smithii* LC 5 5 1 11 Trachycephalus typhonius LC 1 2 1 4 Triprion petasatus LC 4 5 1 10 Family Leiuperidae (1 species) Engystomops pustulosus LC 3 2 2 7 Family Leptodactylidae (2 species) Leptodactylus fragilis LC 1 2 2 5 Leptodactylus melanonotus LC 1 3 2 6 Family Microhylidae (6 species) Gastrophryne elegans LC 2 5 1 8 Gastrophryne mazatlanensis NE 2 5 1 8 Gastrophryne olivacea LC 3 5 1 9 Hypopachus barberi VU 4 5 1 10 Hypopachus ustus LC 2 4 1 7 Hypopachus variolosus LC 2 1 1 4 Family Ranidae (28 species) Lithobates berlandieri LC 4 2 1 7 Lithobates brownorum NE 4 3 1 8 Lithobates catesbeianus LC 3 6 1 10 Lithobates chichicuahutla* CR 6 8 1 15 Lithobates chiricahuensis VU 4 6 1 11 Lithobates dunni* EN 5 8 1 14 Lithobates forreri LC 1 1 1 3 Lithobates johni* EN 5 8 1 14 Lithobates lemosespinali* DD 5 8 1 14 Lithobates macroglossa VU 4 7 1 12 Lithobates maculatus LC 3 1 1 5 Lithobates magnaocularis* LC 5 6 1 12 Lithobates megapoda* VU 5 8 1 14 Lithobates montezumae* LC 5 7 1 13 Lithobates neovolcanicus* NT 5 7 1 13 Lithobates omiltemanus* CR 5 7 1 13 Lithobates psilonota* DD 5 8 1 14 Lithobates pueblae* CR 6 8 1 15 Lithobates pustulosus* LC 5 3 1 9 Lithobates sierramadrensis* VU 5 7 1 13 Lithobates spectabilis* LC 5 6 1 12 Lithobates tarahumarae VU 2 5 1 8 Lithobates tlaloci* CR 6 8 1 15 Lithobates vaillanti LC 3 5 1 9 Lithobates yavapaiensis LC 4 7 1 12 Lithobates zweifeli* LC 5 5 1 11 August 2013 | Volume 7 | Number 1 | e69 Amphib. Reptile Conserv. | http://amphibian-reptile-conservation.org 1 23 Conservation reassessment of Mexican amphibians Rana boylii NT 3 8 1 12 Rana draytonii LC 3 6 1 10 Family Rhinophrynidae (1 species) Rhinophrynus dorsalis LC 2 5 1 8 Family Scaphiopodidae (4 species) Scaphiopus couchii LC 1 1 1 3 Spea bombifrons NE 3 6 1 10 Spea hammondii LC 3 8 1 12 Spea multiplicata NE 1 4 1 6 Order Caudata (139 species) Family Ambystomatidae (18 species) Ambystoma altamirani* EN 5 7 1 13 Ambystoma amblycephalum* CR 6 6 1 13 Ambystoma andersoni* CR 6 8 1 15 Ambystoma bombypellum* CR 6 8 1 15 Ambystoma dumerilii* CR 6 8 1 15 Ambystoma flavipiperatum* DD 6 7 1 14 Ambystoma granulosum* CR 6 7 1 14 Ambystoma leorae* CR 6 8 1 15 Ambystoma lermaense* CR 6 8 1 15 Ambystoma mavortium NE 3 6 1 10 Ambystoma mexicanum* CR 6 8 1 15 Ambystoma ordinarium* EN 5 7 1 13 Ambystoma rivulare* DD 5 7 1 13 Ambystoma rosaceum* LC 5 8 1 14 Ambystoma silvense* DD 5 8 1 14 Ambystoma subsalsum* NE 5 8 1 14 Ambystoma taylori* CR 6 8 1 15 Ambystoma velasci* LC 5 4 1 10 Family Plethodontidae (118 species) Aneides lugubris LC 3 7 4 14 Batrachoseps major LC 4 6 4 14 Bolitoglossa alberchi* LC 6 5 4 15 Bolitoglossa chinanteca NE 6 8 4 18 Bolitoglossa engelhardti EN 4 7 4 15 Bolitoglossa flavimembris EN 4 7 4 15 Bolitoglossa flaviventris EN 4 5 4 13 Bolitoglossa franklini EN 4 6 4 14 Bolitoglossa hartwegi NT 4 4 4 12 Bolitoglossa hermosa* NT 5 7 4 16 Bolitoglossa lincolni NT 4 5 4 13 Bolitoglossa macrinii* NT 5 6 4 15 Bolitoglossa mexicana LC 4 3 4 11 Bolitoglossa mulleri VU 4 7 4 15 Bolitoglossa oaxacensis* DD 5 8 4 17 Bolitoglossa occidentalis LC 4 3 4 11 Bolitoglossa platydactyla* NT 5 6 4 15 Bolitoglossa riletti* EN 6 6 4 16 Bolitoglossa rostrata VU 4 6 4 14 August 2013 | Volume 7 | Number 1 | e69 Amphib. Reptile Conserv. | http://amphibian-reptile-conservation.org 1 24 Wilson et al. Bolitoglossa rufescens LC 1 4 4 9 Bolitoglossa stuarti DD 4 7 4 15 Bolitoglossa veracrucis* EN 6 7 4 17 Bolitoglossa yucatana LC 4 7 4 15 Bolitoglossa zapoteca* DD 6 8 4 18 Chiropterotriton arboreus* CR 6 8 4 18 Chiropterotriton chiropterus* CR 6 6 4 16 Chiropterotriton chondrostega* EN 5 8 4 17 Chiropterotriton cracens* EN 6 7 4 17 Chiropterotriton dimidiatus * EN 6 7 4 17 Chiropterotriton lavae* CR 6 8 4 18 Chiropterotriton magnipes* CR 6 6 4 16 Chiropterotriton mosaueri* DD 6 8 4 18 Chiropterotriton multidentatus * EN 5 6 4 15 Chiropterotriton orculus* VU 6 8 4 18 Chiropterotriton priscus* NT 6 6 4 16 Chiropterotriton terrestris* CR 6 8 4 18 Cryptotriton alvarezdeltoroi* EN 6 8 4 18 Dendrotriton megarhinus* VU 6 7 4 17 Dendrotriton xolocalcae* VU 6 8 4 18 Ensatina eschscholtzii LC 3 7 4 14 Ensatina klauberi NE 4 6 4 14 Ixalotriton niger* CR 6 8 4 18 Ixalotriton parvus* CR 6 8 4 18 Nyctanolis pernix EN 4 7 4 15 Oedipina elongata LC 4 7 4 15 Parvimoige townsendi* CR 5 7 4 16 Pseudoeurycea ahuitzotl* CR 6 8 4 18 Pseudoeurycea altamontana* EN 5 8 4 17 Pseudoeurycea amuzga* DD 6 8 4 18 Pseudoeurycea anitae* CR 6 8 4 18 Pseudoeurycea aquatica* CR 6 8 4 18 Pseudoeurycea aurantia* VU 6 8 4 18 Pseudoeurycea bellii* VU 5 3 4 12 Pseudoeurycea boneti* VU 6 7 4 17 Pseudoeurycea brunnata CR 4 7 4 15 Pseudoeurycea cafetalera NE 6 7 4 17 Pseudoeurycea cephalica* NT 5 5 4 14 Pseudoeurycea cochranae* EN 6 7 4 17 Pseudoeurycea conanti* EN 5 7 4 16 Pseudoeurycea firscheini* EN 6 8 4 18 Pseudoeurycea gadovii* EN 5 4 4 13 Pseudoeurycea galaenae* NT 6 8 4 18 Pseudoeurycea gigantea* CR 5 7 4 16 Pseudoeurycea goebeli CR 4 7 4 15 Pseudoeurycea juarezi* CR 6 7 4 17 Pseudoeurycea leprosa* VU 5 7 4 16 Pseudoeurycea lineola* EN 5 5 4 14 Pseudoeurycea longicauda* EN 5 8 4 17 August 2013 | Volume 7 | Number 1 | e69 Amphib. Reptile Conserv. | http://amphibian-reptile-conservation.org 1 25 Conservation reassessment of Mexican amphibians Pseudoeurycea lynchi* CR 5 8 4 17 Pseudoeurycea maxima* DD 5 8 4 17 Pseudoeurycea melanomolga* EN 6 6 4 16 Pseudoeurycea mixcoatl* DD 6 8 4 17 Pseudoeurycea mixteca* LC 5 8 4 17 Pseudoeurycea mystax* EN 6 8 4 18 Pseudoeurycea naucampatepetl* CR 6 7 4 17 Pseudoeurycea nigromaculata* CR 5 8 4 17 Pseudoeurycea obesa* DD 6 8 4 18 Pseudoeurycea orchileucos* EN 6 8 4 18 Pseudoeurycea orchimelas* EN 6 7 4 17 Pseudoeurycea papenfussi* NT 6 7 4 17 Pseudoeurycea praecellens* CR 6 8 4 18 Pseudoeurycea quetzalanensis* DD 6 7 4 17 Pseudoeurycea rex CR 4 4 4 12 Pseudoeurycea robertsi* CR 6 8 4 18 Pseudoeurycea ruficauda* DD 6 8 4 18 Pseudoeurycea saltator* CR 6 8 4 18 Pseudoeurycea scandens* VU 6 7 4 17 Pseudoeurycea smithi* CR 5 6 4 15 Pseudoeurycea tenchalli* EN 6 7 4 17 Pseudoeurycea teotepec* EN 6 8 4 18 Pseudoeurycea tlahcuiloh* CR 6 7 4 17 Pseudoeurycea tlilicxitl* DD 5 8 4 17 Pseudoeurycea unguidentis* CR 6 7 4 17 Pseudoeurycea werleri* EN 6 7 4 17 Thorius adelos* EN 6 8 4 18 Thorius arbor eus* EN 6 8 4 18 Thorius aureus* CR 6 7 4 17 Thorius boreas* EN 6 8 4 18 Thorius dubitus* EN 5 7 4 16 Thorius grandis* EN 6 5 4 15 Thorius infernal is* CR 6 8 4 18 Thorius insperatus* DD 6 8 4 18 Thorius lunaris* EN 6 8 4 18 Thorius macdougalli* VU 6 6 4 16 Thorius magnipes* CR 6 7 4 17 Thorius minutissimus* CR 6 7 4 17 Thorius minydemus* CR 6 8 4 18 Thorius munificus* CR 6 8 4 18 Thorius narismagnus* CR 6 8 4 18 Thorius narisovalis* CR 6 7 4 17 Thorius omiltemi* EN 6 8 4 18 Thorius papaloae* EN 6 7 4 17 Thorius pennatulus* CR 5 6 4 15 Thorius pulmonaris* EN 6 7 4 17 Thorius schmidti* EN 6 7 4 17 Thorius smithi* CR 6 7 4 17 Thorius spilogaster* CR 6 7 4 17 August 2013 | Volume 7 | Number 1 | e69 Amphib. Reptile Conserv. | http://amphibian-reptile-conservation.org 1 26 Wilson et al. Thorius troglodytes* EN 6 6 4 16 Family Salamandridae (1 species) Notophthalmus meridionalis EN 2 8 1 12 Family Sirenidae (2 species) Siren intermedia LC 3 8 1 12 Siren lacertina LC 3 8 1 12 Order Gymnophiona (2 species) Family Dermophiidae (2 species) Dermophis mexicanus VU 4 3 4 11 Dermophis oaxacae* DD 5 3 4 12 Amphib. Reptile Conserv. | http://amphibian-reptile-conservation.org 1 27 August 2013 | Volume 7 | Number 1 | e69 Crotalus tancitarensis. The Tancftaro cross-banded mountain rattlesnake is a small species (maximum recorded total length = 434 mm) known only from the upper elevations (3,220-3,225 m) of Cerro Tancftaro, the highest mountain in Michoacan, Mexico, where it inhabits pine-fir forest (Alvarado and Campbell 2004; Alvarado et al. 2007). Cerro Tancftaro lies in the western portion of the Transverse Volcanic Axis, which extends across Mexico from Jalisco to central Veracruz near the 20°N latitude. Its entire range is located within Parque Nacional Pico de Tancftaro (Campbell 2007), an area under threat from manmade fires, logging, avocado culture, and cattle raising. This attractive rattlesnake was described in 2004 by the senior author and Jonathan A. Campbell, and placed in the Crotalus intermedins group of Mexican montane rattlesnakes by Bryson et al. (2011). We calculated its EVS as 19, which is near the upper end of the high vulnerability category (see text for explanation), its IUCN status has been reported as Data Deficient (Campbell 2007), and this species is not listed by SEMARNAT. More information on the natural history and distribution of this species is available, however, which affects its conservation status (especially its IUCN status; Alvarado-Dfaz et al. 2007). We consider C. tancitarensis one of the pre-eminent flagship reptile species for the state of Michoacan, and for Mexico in general. Photo by Javier Alvarado-Diaz. September 2013 | Volume 7 | Number 1 | e71 Amphib. Reptile Conserv. | http://amphibian-reptile-conservation.org 128 Copyright: © 2013 Alvarado-Dfaz et al. This is an open-access article distributed under the terms of the Creative Commons Attribution-NonCommercial-NoDerivs 3.0 Unported License, which permits unrestricted use for non-commercial and education purposes only provided the original author and source are credited. Amphibian & Reptile Conservation 7(1): 128-170. Patterns of physiographic distribution and conservation status of the herpetofauna of Michoacan, Mexico Wavier Alvarado-Diaz, 2 lreri Suazo-Ortuno, 3 Larry David Wilson, and 4 Oscar Medina-Aguilar 12A Instituto de Investigaciones sobre los Recursos Naturales, Universidad Michoacana de San Nicolas de Hidalgo, Av. San Juanito Itzicuaro s/n Col. Nva. Esperanza, Morelia, Michoacan, MEXICO, 58337 ''Centro Zamorano de Biodiversidad, Escuela Agricola Panamericana Zamorano, Departamento de Francisco Morazdn, HONDURAS Abstract . — At their respective levels, the country of Mexico and the state of Michoacan are major cen- ters of herpetofaunal diversity and endemicity. Three of us (JAD, ISO, OMA) conducted extensive fieldwork in Michoacan from 1998 to 2011, and recorded 169 herpetofaunal species. With additional species reported in the literature and specimens available in scientific collections, the number of species in Michoacan has grown to 215. We examined the distribution of these species within the framework of the five physiographic provinces within the state, i.e., the Coastal Plain, the Sierra Madre del Sur, the Balsas-Tepalcatepec Depression, the Transverse Volcanic Axis, and the Cen- tral Plateau, which briefly are characterized geomorphologically and climatically. The herpetofauna consists of 54 amphibians and 161 reptiles (17.5% of the total for Mexico), classified in 38 families and 96 genera. Almost one-half of Michoacan’s herpetofaunal species occur in a single physio- graphic province, and the percentage of species decreases with an increase in the number of prov- inces. The province with the most species is the Sierra Madre del Sur, with slightly fewer numbers in the Balsas-Tepalcatepec Depression and the Transverse Volcanic Axis. An intermediate number is found in the Coastal Plain, and the lowest in the Central Plateau province. We constructed a Co- efficient of Biogeographic Resemblance matrix and found the greatest degree of herpetofaunal re- semblance between the Balsas-Tepalcatepec Depression and the Sierra Madre del Sur. The greatest resemblance of the Coastal Plain herpetofauna is to that of Balsas-Tepalcatepec Depression, that of the Transverse Volcanic Axis to that of the Central Plateau, and vice versa. Of the species limit- ed to one physiographic province, 47 occur only in the Transverse Volcanic Axis, 23 in the Coastal Plain, 15 in the Balsas-Tepalcatepec, 14 in the Sierra Madre del Sur, and one in the Central Plateau. We employed three systems for determining the conservation status of the herpetofauna of Micho- acan: SEMARNAT, IUCN, and EVS. Almost one-half of the species in the state are not assessed by the SEMARNAT system, with the remainder allocated to the Endangered (four species), Threatened (31), and Special Protection (79) categories. The IUCN system provides an assessment for 184 of the 212 native species, allocating them to the Critically Endangered (five species), Endangered (10), Vulnerable (12), Near Threatened (four), Least Concern (127), and Data Deficient (26) categories. The EVS system provides a numerical assessment for all of the native non-marine species (four ma- rine species occur in the state), with the values ranging from three to 19. The resulting 208 species were placed in low, medium, and high categories of vulnerability, as follows: low (17 amphibians, 39 reptiles); medium (23 amphibians, 45 reptiles); and high (13 amphibians, 71 reptiles). The EVS system is the only one that provides an assessment for all the species (except for the four marine taxa), as well as the only one that considers the distributional status of Michoacan’s herpetofauna (state-level endemic, country-level endemic, and non-endemic). Furthermore, the values indicate that ca. 40% of the state’s herpetofauna is categorized at the highest level of environmental vul- nerability. Based on these conclusions, we provide recommendations for protecting Michoacan’s herpetofauna in perpetuity. Key words. Amphibians, reptiles, physiographic provinces, conservation status, recommendations Correspondence. Emails: 1 jvr.alvarado@gmail.com (Corresponding author) 2 ireri. suazo@gmail.com 3 bufodoc@aol. com 4 mineo_osc@hotmail. com September 2013 | Volume 7 | Number 1 | e71 Armphib. Reptile Conserv. | http://amphibian-reptile-conservation.org 129 Physiographic distribution and conservation of Michoacan herpetofauna Resumen . — Mexico es un importante centro de diversidad y endemismo herpetofaunistico y el es- tado de Michoacan tambien presenta estas caracteristicas. Durante el periodo de 1998-2011, tres de nosotros (JAD, ISO, OMA) conducimos un extenso trabajo de campo en Michoacan, registrando 169 especies de anfibios y reptiles. Con la adicion de especies reportadas en la literatura y los registros disponibles en colecciones cientificas, el numero total de especies de la herpetofauna michoacana es de 215. Examinamos la distribution de estas especies en Michoacan, considerando las cinco provincias fisiograficas representadas en el Estado: la Llanura Costera, la Sierra Madre del Sur, la Depresion del Balsas-Tepalcatepec, el Eje Volcanico Transversal, y la Meseta Central, las que de manera resumida son caracterizadas en base a su geomorfologia y clima. La herpeto- fauna consiste de 54 anfibios y 161 reptiles (17.5% del total de Mexico), clasificadas en 38 familias y 96 generos. Casi la mitad de las especies de la herpetofauna de Michoacan ocurre en una sola provincia fisiografica, con un cada vez menor porcentaje de especies a medida que el numero de provincias se incrementa. El mayor numero de especies se encuentra en la Sierra Madre del Sur, con cifras ligeramente menores en la Depresion del Balsas-Tepalcatepec y el Eje Volcanico Trans- versal. Un numero intermedio de especies se encuentra en la provincia Planicie Costera y el menor numero se encuentra en la provincia Meseta Central. Implementamos una matriz del Coeficiente de Semejanza Biogeografica, la que muestra que el mayor grado de semejanza herpetofaunistica se encuentra entre la Depresion del Balsas-Tepalcatepec y la Sierra Madre del Sur. La mayor similitud de la herpetofauna de la Planicie Costera es con la herpetofauna de la Depresion Balsas-Tepal- catepec, la del Eje Volcanico Transversal con la de la Meseta Central y viceversa. De las especies restringidas a una sola provincia fisiografica, 47 ocurren solamente en el Eje Volcanico Transversal, 23 en la Planicie Costera, 15 en la Depresion del Balsas-Tepalcatepec, 14 en la Sierra Madre del Sury y una en la Meseta Central. Usamos tres sistemas para determinar el estado de conservacion: SEMARNAT, UICN, y EVS. Casi la mitad de las especies de Michoacan no han sido evaluadas por el sistema de SEMARNAT, y las evaluadas han sido asignadas a las categorias de Peligro (cuatro especies), Amenazadas (31), y Proteccion Especial (79). El sistema de la UICN ha evaluado 184 de las 212 especies nativas de Michoacan, asignadas a las siguientes categorias: Peligro Critico (cinco especies), En Peligro (10), Vulnerable (12), Casi Amenazado (cuatro), Preocupacion Menor (127), y Datos Insuficientes (26). El sistema EVS proporciona una evaluacion numerica para todas las espe- cies nativas que no son marinas (cuatro especies marinas ocurren en el estado), con valores de tres a 18. Las 209 especies evaluadas mediante el EVS fueron asignadas a las categorias de baja, media y alta vulnerabilidad de la siguiente manera: baja (17 anfibios, 39 reptiles); media (23 anfibios, 45 reptiles); y alta (13 anfibios, 71 reptiles). El sistema EVS es el unico de los tres que proporciona una evaluacion de todas las especies (excepto para los cuatro taxa marinos) y el unico que considera el estado distribucional de los componentes de la herpetofauna de Michoacan (endemico a nivel estatal, endemico a nivel de pais, y no endemico). Ademas, los valores muestran que cerca del 40% de la herpetofauna del estado se encuentra en la categoria mas alta de vulnerabilidad ambiental. En base a estas conclusiones, proponemos recomendaciones para la proteccion a perpetuidad de la herpetofauna de Michoacan. Palabras claves. Anfibios, reptiles, provincias fisiograficas, estatus de conservacion, recomendaciones Citation: Alvarado-Dfaz J, Suazo-Ortuno I, Wilson LD, Medina-Aguilar O. 2013. Patterns of physiographic distribution and conservation status of the herpetofauna of Michoacan, Mexico. Amphibian & Reptile Conservation 7(1): 128-170(e71). The publication of On the Origin of Species in 1859 is a recognized watershed in biological science. Perhaps the greatest threat to Western ideology was not the com- mon origin of all beings, as is assumed, but rather the possibility of a common ending: that all beings, humans among them were subjected to the same forces and vul- nerabilities. Chernela 2012: 22. Amphib. Reptile Conserv. | http://amphibian-reptile-conservation.org Introduction Mesoamerica is one of the principal biodiversity hotspots in the world (Wilson and Johnson 2010), and the coun- try of Mexico comprises about 79% of the land surface of Mesoamerica (CIA World Factbook). The document- ed amphibian fauna of Mexico currently consists of 379 species, including 237 anurans, 140 salamanders, and two caecilians (Wilson et al. 2013b). Based on this 130 September 2013 | Volume 7 | Number 1 | e71 Alvarado-Diaz et al. Incilius pisinnus. The Michoacan toad is a state endemic, with a distribution in the Balsas-Tepalcatepec Depression and the Sierra Madre del Sur. Its EVS was estimated as 15, which is unusually high for a bufonid anuran, its IUCN ranking has been judged as Data Deficient, and a SEMARNAT status has not been provided. This individual is from Apatzingan, Michoacan. Photo by Oscar Medina- Aguilar. Eleutherodactylus rufescens. The blunt-toed chirping frog is endemic to the Sierra de Coalcoman region of the Sierra Madre del Sur. Its EVS has been assessed as 17, placing this species in the middle of the high vulnerability category, this frog is considered as Critically Endangered by IUCN, and as a Special Protection species by SEMARNAT. This individual was found at Dos Aguas in the Sierra de Coalcoman (Sierra Madre del Sur) in Michoacan. Photo by Oscar Medina- Aguilar. September 2013 | Volume 7 | Number 1 | e71 Amphib. Reptile Conserv. | http://amphibian-reptile-conservation.org 131 Physiographic distribution and conservation of Michoacan herpetofauna figure, Mexico is the country with the 5 th largest number of amphibian species in the world (Llorente-Bousquets and Ocegueda 2008; Stuart et al. 2010a), after Brazil, Colombia, Ecuador, and Peru. The country also is in- habited by 849 species of reptiles, including 798 squa- mates, 48 turtles, and three crocodylians (Wilson et al. 2013a), which globally is the second largest reptile fauna (Llorente-Bousquets and Ocegueda 2008), after Australia. The total number of 1,227 species makes the Mexican herpetofauna the second largest in the world (Llorente-Bousquets and Ocegueda 2008), comprising 7.3% of the global herpetofauna (7,044 amphibian spe- cies, according to the Amphibian Species of the World website, accessed 21 Lebruary 2013, and 9,766 reptile species, according to the Reptile Database website, also accessed 21 Lebruary 2013, for a total of 16,810). Beyond its highly significant herpetofaunal diversity, Mexico also contains an amazing amount of endemicity. Currently, 254 of 379 (67.0%) of the known amphibi- an species and 480 of 849 (56.5%) of the known reptile species are endemic (Wilson et al. 2013a,b). The com- bined figure for both groups is 734 species (59.8%), a percentage 2.4 times as high as the next highest rate of endemicity for the Central American countries (24.8% for Honduras; Townsend and Wilson 2010). Michoacan (the formal name is Michoacan de Ocam- po) is the 16 th largest state in Mexico, with an area of 58,599 km 2 (www.en.wikipedia.org/wiki/List_of_Mexi- can_states_by_area), which comprises about 3.0% of the country’s land surface. The state is located in southwest- ern Mexico between latitudes 20°23'44" and 18°09'49" N and longitudes 100°04'48” and 103°44'20" W, and is bounded to the northwest by Colima and Jalisco, to the north by Guanajuato and Queretaro, to the east by Mexico, and to the southeast by Guerrero. Michoacan is physiographically and vegetationally diverse, inasmuch as elevations range from sea level to 3,840 m (at the top of Volcan Tancftaro). The state encompasses a portion of the Pacific coastal plain, a long stretch of the Balsas-Te- palcatepec Depression, a segment of the Sierra Madre del Sur called the Sierra de Coalcoman, and a significant portion of the Transverse Volcanic Axis. Mexico is known for its high level of herpetofaunal endemism, but compared with the country the herpeto- fauna of Michoacan is several percentage points higher, with a number of the country endemics limited in distri- bution to the state (see below). Any attempt to assess the conservation status of a herpetofaunal group depends on an accurate accounting of the distribution and composi- tion of the species involved. Thus, our objectives with this study are to update the list of amphibians and rep- tiles in Michoacan, to discuss their distribution among the physiographic provinces, and to use these data to gauge the conservation status of the entire herpetofau- na using various measures. Linally, based on our con- servation assessment, we provide recommendations to enhance current efforts to protect the state’s amphibians and reptiles. Diaglena spatulata. The shovel-headed treefrog is distributed along the Pacific coastal lowlands from Sinaloa to Oaxaca, and thus is a Mexican endemic hylid anuran. In Michoacan, it occurs in the Balsas-Tepalcatepec Depression and along the Coastal Plain. Its EVS was gauged as 13, placing it at the upper end of the medium vulnerability category, IUCN has assessed this anuran as Least Concern, and it is not listed by SEMARNAT. This individual was photographed at the Reserva de la Biosfera Chamela-Cuixmala on the coast of Jalisco. Photo by Oscar Medina- Aguilar. September 2013 | Volume 7 | Number 1 | e71 Amphib. Reptile Conserv. | http://amphibian-reptile-conservation.org 132 Alvarado-Dfaz et al. Materials and Methods 1. Sampling procedures From 1998 to 2011, three of us (JAD, ISO, OMA) con- ducted fieldwork in 280 localities (58 municipalities) of Michoacan, representing all of the state’s physiographic provinces, with significant attention paid to poorly sam- pled areas, as part of the “Diversidad Herpetofaunfstica del Estado de Michoacan” project undertaken by person- nel from the Laboratorio de Herpetologla of the Instituto de Investigaciones sobre los Recursos Naturales (INI- RENA) of the Universidad Michoacana de San Nicolas de Hidalgo (UMSNH). Importantly, due to unsafe conditions in certain parts of the state in recent years, large areas have not been explored. During each visit to the sampling sites, we used visual encounter surveys (Crump and Scott 1994) to locate amphibians and rep- tiles during the day and at night. This work was conduct- ed under scientific collecting permits (DGVS/FAUT- 0113), and used the collection techniques described by Casas et al. (1991). In cases where we could not identify individuals in the field, they were sacrificed and subse- quently deposited in the herpetological collections of INIRENA-UMSNH. We identified specimens by using taxonomic keys and other information in Smith and Tay- lor (1945, 1948, 1950), Duellman (1961, 1965, 2001), Casas- Andreu and McCoy (1979), Ramfrez-Bautista (1994), Flores- Villela et al. (1995), and Huacuz (1995), and updated scientific names by using Flores-Villela and Canseco-Marquez (2004), Faivovich et al. (2005), Wil- son and Johnson (2010), and Wilson et al. (2013a,b). 2. Updating the herpetofaunal list In addition to the specimens recorded during the field- work, the list of species was augmented using material donated by others. We also used records from the Colec- cion Nacional de Anfibios y Reptiles-UNAM (CNAR), the California Academy of Sciences (CAS), the Universi- ty of Colorado Museum of Natural History, Herpetology Collection (CUMNH), the Museum of Natural Sciences, Louisiana State University (LSUMZ), the Field Muse- um of Natural History (FMNH), and the Royal Ontario Museum (ROM). Additionally, we included records for Michoacan from the Catalogo de la Biodiversidad en Michoacan (SEDUE [Secretarfa de Desarrollo Urbano y Ecologfa], UMSNH 2000), la Biodiversidad en Micho- acan Estudio de Estado (Villasenor 2005), various dis- tribution notes published in Herpetological Review and otherwise posted at the IUCN Red List website, as well as data presented by Flores-Villela and Canseco-Marquez (2004), Vargas-Santamarfa and Flores-Villela (2006), Gonzalez-Hemandez and Garza-Castro (2006), Medi- na- Aguilar et al. (20 1 1 ) , and Torres (20 11). We follow the taxonomy used in Wilson (2013 a, b), with the exception Amphib. Reptile Conserv. | http://amphibian-reptile-conservation.org 133 of the deletion of the nominal species Anolis schmidti, which recently was synonymized by Nieto et al. (2013). 3. Systems for determining conservation status We used the following three systems to determine the conservation status of the 212 native species of amphibi- ans and reptiles in Michoacan: SEMARNAT, IUCN, and EVS. The SEMARNAT system, established by the Sec- retarfa de Medio Ambiente y Recursos Naturales, em- ploys three categories — Endangered (P), Threatened (A), and Subject to Special Protection (Pr). The results of the application of this system are reported in the NORMA Oficial Mexicana NOM-059-SEMARNAT-2010 (www. semarnat.gob.mx). For species not assessed by this sys- tem, we use the designation “No Status.” The IUCN system is utilized widely to assess the con- servation status of species on a global basis. The catego- ries used are explained in the document IUCN Red List of Categories and Criteria (2010), and include Extinct (EX), Extinct in the Wild (EW), Critically Endangered (CR), Endangered (EN), Vulnerable (VU), Near Threat- ened (NT), Least Concern (LC), Data Deficient (DD), and Not Evaluated (NE). The categories Critically En- dangered, Endangered, and Vulnerable collectively are termed “threat categories,” to distinguish them from the other six. The EVS system was developed initially for use in Honduras by Wilson and McCranie (2004), and subse- quently was used in several chapters on Central American countries in Wilson et al. (2010). Wilson et al. (2013a, b) modified this system and explained its use for the am- phibians and reptiles of Mexico, and we follow their pre- scriptions. The EVS measure is not designed for use with marine species (e.g., marine turtles and sea snakes), and generally is not applied to non-native species. Physiography and Climate 1. Physiographic provinces Based on geological history, morphology, structure, hy- drography, and soils, five physiographic provinces can be recognized within the state of Michoacan, including the Pacific Coastal Plain, the Sierra Madre del Sur, the Balsas-Tepalcatepec Depression, the Transverse Volcanic Axis, and the Central Plateau (Fig. 1). The Coastal Plain province comprises a narrow strip of land between the Pa- cific Ocean and the Sierra Madre del Sur, and consists of small alluvial plains extending from the mouth of the Rio Balsas to the east and the Rio Coahuayana to the west. The Sierra Madre del Sur (Sierra de Coalcoman) lies between the Coastal Plain and the Balsas-Tepalcatepec Depression, extends for over 100 km in a north- west-southeast direction, and contains elevations reach- ing about 2,200 m. The Balsas-Tepalcatepec Depression September 2013 | Volume 7 | Number 1 | e71 Physiographic distribution and conservation of Michoacan herpetofauna is located between the Sierra Madre del Sur to the south- west and the Transverse Volcanic Axis to the northeast. This intermontane area is a broad structural basin that lies at elevations ranging from 200 to 700 m. As noted by Duellman (1961 : 10), “the western part of this basin. . . is the valley of the Rio Tepalcatepec, a major tributary of the Rio Balsas. The eastern part of the basin is the valley of the Rio Balsas.” The Transverse Volcanic Axis is located to the south of the Central Plateau and crosses Mexico at about the 20 th parallel. The region is composed of volcanic ejecta and is volcanically active. This area is home to Mexico’s highest mountains, such as Pico de Orizaba (5,636 m) and Popocatepetl (5,426 m), which in Michoacan is represented by Pico de Tancftaro, with an elevation of 3,850 m. In addition, several endorheic lakes are located in this province, including Patzcuaro, Zirahuen, and Cuitzeo. The Central Plateau is a vast ta- bleland bordered on the south by the Transverse Volca- nic Axis, on the west by the Sierra Madre Occidental, on the east by the Sierra Madre Oriental, and on the north by the Rio Bravo (Rio Grande). Elevations in this prov- ince range from 1,100 m in the northern portion of the country to 2,000 m. In Michoacan, this province is rep- resented by a relatively small area (3,905 km 2 ) along the northern border of the state; the Rio Lerma flows from it, and empties into the Pacific Ocean (Duellman 1961). Amphib. Reptile Conserv. | http://amphibian-reptile-conservation.org 134 2. Climate Given its location in the tropical region of Mexico, south of the Tropic of Cancer, temperatures in Michoacan vary as a consequence of differences in elevation and the ef- fects of prevailing winds. To illustrate variation in ambi- ent temperatures in the state, we extracted data for one locality from each of the five physiographic provinces from the Servicio Meteorologico Nacional, Michoacan and placed them in Table 1 . These data are organized in the table from top to bottom based on the elevation of the localities (from low to high). As expected, a decrease in the mean annual temperature occurs from lower to high- er elevations. The same pattern is seen for annual mini- mum and maximum temperatures, except for the Coastal Plain compared to the Balsas-Tepalcatepec Depression (33.0 vs. 34.4 °C). As expected in the tropics, relatively little tempera- ture variation occurs throughout the year. The differenc- es between the low and high mean monthly temperatures (in °C) for the localities in the five physiographic prov- inces are as follows: Coastal Plain (Lazaro Cardenas, 50 m) = 1.9; Balsas-Tepalcatepec Depression (Apatzingan, 320 m) = 5.5; Sierra Madre del Sur (Coalcoman, 1,100 m) = 5.2; Central Plateau (Morelia, 1,915 m) = 5.9; and Transverse Volcanic Axis (Patzcuaro, 2,035 m) = 6.6. September 2013 | Volume 7 | Number 1 | e71 Alvarado-Diaz et al. The lowest mean monthly temperatures are for January, and the highest for May or June. Essentially the same pattern occurs with minimum and maximum monthly temperatures, except for minor departures in a few areas (Table 1). The highest mean monthly temperature (34.4 °C) is at Apatzingan in the Balsas-Tepalcatepec Depression. Duellman (1961) stated that the highest mean annu- al temperatures (29.3 °C) in this depression have been recorded at Churumuco (251 m), as reported by Con- treras (1942). More recent data at the Servicio Meteo- rologico Nacional website for Michoacan indicates that the highest daily temperature of 46 °C was recorded at this locality on 9 April 1982. At the other extreme are temperatures on the peak of Volcan Tancftaro, where the mean annual temperature is less than 10 °C and it snows during the winter. In tropical locales, heavy or light precipitation typ- ically occurs during the rainy and dry seasons, respec- tively. In Michoacan, the rainy season extends from June to October, when 80% or more of the annual precipita- tion is deposited. As with temperature data, we extract- ed information on mean annual precipitation and vari- ation in monthly precipitation recorded at one locality for each of the five physiographic provinces, and placed the data in Table 2. The results demonstrate that at each locality the highest amount of precipitation occurs from June to October. The percentage of annual precipitation Table 1. Monthly minimum, mean (in parentheses), maximum, and annual temperature data (in °C) for the physiographic provinces of Michoacan, Mexico. Localities and their elevation for each of the provinces are as follows: Coastal Plain (Lazaro Cardenas, 50 m); Balsas-Tepalcatepec Depression (Apatzingan, 320 m); Sierra Madre del Sur (Coalcoman de Vazquez Pallares, 1,100 m); Central Plateau (Morelia, 1,915 m); Transverse Volcanic Axis (Patzcuaro, 2,035 m). Data (1971-2000) from the Sistema Meteorologico Nacional, Michoacan (smn.cna.gob.mx/index). Physiographic Province Jan. Feb. March April May June July Aug. Sept. Oct. Nov. Dec. Annual Coastal Plain 20.6 (26.6) 32.6 20.6 (26.8) 33.1 20.8 (27.0) 33.2 21.2 (27.3) 33.5 22.8 (28.3) 33.8 23.9 (28.5) 33.1 23.4 (28.0) 32.7 23.7 (28.1) 32.6 23.3 (27.7) 32.0 23.5 (28.1) 32.6 22.7 (27.9) 33.2 21.1 (27.1) 33.2 22.3 (27.6) 33.0 Balsas- Tepalcatepec Depression 16.7 (24.6) 32.5 17.6 (25.9) 34.1 19.1 (27.7) 36.3 20.7 (29.2) 37.6 22.3 (30.3) 38.3 22.7 (29.1) 35.6 21.6 (27.3) 33.1 21.6 (27.3) 33.1 21.7 (27.3) 33.0 21.5 (27.7) 33.8 19.5 (26.4) 33.3 17.7 (25.1) 32.5 20.2 (27.3) 34.4 Sierra Madre del Sur 10.2 (19.9) 29.7 10.7 (20.8) 30.9 11.6 (22.1) 32.7 12.3 (23.5) 34.6 14.3 (24.8) 35.3 17.9 (25.1) 32.4 18.2 (24.1) 30.1 17.4 (23.8) 30.2 17.7 (23.8) 30.0 16.7 (23.7) 30.8 13.9 (22.2) 30.4 11.9 (21.0) 30.0 14.4 (22.9) 31.4 Central Plateau 6.8 (15.8) 24.7 7.6 (17.0) 26.4 9.6 (19.0) 28.4 11.1 (20.4) 29.7 12.6 (21.7) 30.9 13.3 (21.2) 29.1 12.8 (19.6) 26.5 13.1 (19.8) 26.4 12.9 (19.4) 26.0 11.3 (18.7) 26.1 9.3 (17.7) 26.2 7.3 (16.4) 25.5 10.6 (18.9) 27.2 Transverse Volcanic Axis 3.3 (12.9) 22.5 4.0 (14.1) 24.1 5.4 (16.0) 26.6 7.3 (17.8) 28.2 9.4 (19.1) 28.7 12.5 (19.5) 26.4 12.0 (18.0) 23.9 11.9 (18.0) 24.1 11.5 (17.7) 23.9 9.2 (16.7) 24.1 5.9 (14.8) 23.7 4.3 (13.4) 22.6 8.1 (16.5) 24.9 Table 2. Monthly and annual precipitation data (in mm.) for the physiographic provinces of Michoacan, Mexico. Localities and their elevation for each of the provinces are as follows: Coastal Plain (Lazaro Cardenas, 50 m); Sierra Madre del Sur (Coalcoman de Vazquez Pallares, 1,100 m); Balsas-Tepalcatepec Depression (Apatzingan, 320 m); Transverse Volcanic Axis (Patzcuaro, 2,035 m); Central Plateau (Morelia, 1,915 m). The shaded area indicates the months of the rainy season. Data taken from Servicio Meteorologico Nacional, Michoacan (smn.cna.gob.mx/index). Physiographic Province Jan. Feb. March Apr. May June July Aug. Sept. Oct. Nov. Dec. Annual Coastal Plain 7.5 0.4 1.0 0.0 17.0 240.4 269.0 257.0 374.2 150.1 23.7 34.0 1,374.3 Balsas- Tepalcatepec Depression 19.8 22.0 9.0 2.5 24.1 138.0 167.9 160.8 133.6 78.8 36.9 15.3 808.7 Sierra Madre del Sur 33.7 42.8 24.8 7.8 37.2 272.2 284.1 258.0 225.7 166.8 93.0 42.1 1,488.2 Central Plateau 15.8 5.6 7.5 9.9 37.9 146.5 166.1 167.8 131.6 51.6 10.4 4.2 754.9 Transverse Volcanic Axis 27.1 5.0 5.1 9.7 37.8 150.3 219.6 204.1 157.9 71.2 17.6 13.4 918.8 September 2013 | Volume 7 | Number 1 | e71 Amphib. Reptile Conserv. | http://amphibian-reptile-conservation.org 135 Physiographic distribution and conservation of Michoacan herpetofauna during this period ranges from 81.1% at Coalcoman in the Sierra Madre del Sur to 93.9% at Lazaro Cardenas on the Coastal Plain (mean 86.9%). Generally, the driest month is April (except on the Central Plateau, where it is December) and the wettest month is July (except on the Central Plateau, where it is August). Annual precip- itation is lowest on the Central Plateau, with 754.9 mm for the capital city of Morelia, and highest at Coalcoman in the Sierra Madre del Sur, with 1,488.2 mm (Table 2). Composition of the Herpetofauna Field surveys and a review of the published literature and databases yielded a total of 215 species of amphibians and reptiles for the state of Michoacan (54 amphibians, 161 reptiles). Of the amphibians, 44 are anurans (81.1%, including the non-native Lithobates catesbeianus ), nine are salamanders (17.0%), and one is a caecilian (1.9%). Of the 161 reptiles, 153 are squamates (95.0%, including the non-native Hemidactylus frenatus and Ramphoty- phlops braminus ), seven are turtles (4.4%), and one is a crocodylian (0.6%). The number of species occurring in Michoacan is 17.5% of the total for the Mexican herpe- tofauna (1,227 species; Wilson et al. 2013a,b; Table 3). Table 3. Composition of the amphibians and reptiles of Mexico and the state of Michoacan. In each column, the number to the left is that indicated in Wilson et al. (2013a,b) for the country of Mexico; the number to the right is that recorded in this study for the state of Michoacan. These numbers include the marine and non-native taxa. Taxa Families Genera Species Anura 11/9 35/19 237/44 Caudata 4/2 15/2 139/9 Gymnophiona 1/1 1/1 2/1 Subtotals 16/12 51/22 378/54 Squamata 31/21 139/68 798/153 Testudines 9/4 18/5 48/7 Crocodylia 2/1 2/1 3/1 Subtotals 42/26 159/74 849/161 Totals 58/38 210/96 1,227/215 1. Families The herpetofauna of Michoacan (215 species) is clas- sified in 38 families (65.5% of the number in Mexico), with the 54 species of amphibians in 12 of the 16 fami- lies known from the country (75.0%; Wilson et al. 2013a, b; Table 3). About one-half of the amphibian species are classified in one of three families (Hylidae, Ranidae, and Amphib. Reptile Conserv. | http://amphibian-reptile-conservation.org 136 Ambystomatidae). The 161 species of reptiles are clas- sified in 26 fa mili es (including the family Gekkonidae, occupied by a single non-native species, H. frenatus, and the family Typhlopidae, occupied by a single non-native species, R. braminus ), 61.9% of the 42 families found in Mexico (Wilson et al. 2013a; Table 3). One-half of the species of reptiles in the state are classified in one of three families (Phrynosomatidae, Colubridae, and Dipsadidae). 2. Genera The herpetofauna of Michoacan is represented by 96 genera (45.7% of the 210 known from Mexico; Wilson et al. 2013a,b), with the amphibians composed of 22 genera (43.1% of the 51 known from the country). The reptiles consist of 74 genera (46.5% of the country total of 159). The largest amphibian genera are Incilius (four species), Craugastor (five), Eleutherodactylus (five), Lithobates (11), and Ambystoma (seven). Together, these 32 species comprise 59.3% of the amphibians known from the state (Table 3). The most sizable reptilian genera are Scelopo- rus (16), Geophis (nine), Thamnophis (nine), Crotalus (eight), Aspidoscelis (seven), Phyllodactylus (five), Plestiodon (five), Coniophanes (five), and Leptodeira (five). These 69 species constitute 42.9% of the reptiles known from the state (Table 3). 3. Species Mexico is home to 378 amphibian species, of which 54 (14.3%) occur in Michoacan (Table 3). Anurans are better represented in the state (18.6% of 237 Mexican species) than salamanders (6.5% of 139). Only two cae- cilian species are known from Mexico, and one occurs in Michoacan (50.0%). Mexico also is inhabited by 849 reptile species, of which 161 (19.0%) are found in Mi- choacan. Squamates are somewhat better represented in the state (19.2% of 798) than turtles (14.6% of 48). Only three crocodylian species occur in Mexico, and one is found in Michoacan (Table 3). Patterns of Physiographic Distribution We recognize five physiographic provinces in Micho- acan (Fig. 1), and their herpetofaunal distribution is in- dicated in Table 4 and summarized by family in Table 5. Of the 215 species recorded from the state, 100 (46.5%, 24 amphibians, 76 reptiles) are limited in dis- tribution to a single physiographic province. In addi- tion, 64 (29.8%, 15 amphibians, 49 reptiles) are known from two provinces, 37 (17.2%, eight amphibians, 29 reptiles) from three, 11 (5.1%, seven amphibians, four reptiles) from four, and only three (1.4%, 0 amphibians, three reptiles) from all five provinces (Table 4). In both amphibians and reptiles, the number of species steadily drops from the lowest to the highest occupancy figures. This distributional feature is significant to conservation September 2013 | Volume 7 | Number 1 | e71 Alvarado-Diaz et al. efforts, inasmuch as the more restricted their distribu- tion the more difficult it will be to provide species with effective protective measures. This feature is obvious when examining the mean occupancy figure, which is 2.0 for amphibians and 1.8 for reptiles, indicating that on average both groups occupy two or slightly fewer physiographic provinces. The three most broadly dis- tributed species (i.e., occurring in all five provinces) all are reptiles and include the anole Anolis nebulosus , the whipsnake Masticophis mentovarius, and the mud tur- tle Kinosternon integrum (Table 4). The most broadly distributed amphibians all are anurans and include the following seven species: the toad Rhinella marina , the chirping frog Eleutherodactylus nitidus, the treefrogs Exerodonta smaragdina and Hyla arenicolor, the white- lipped frog Leptodactylus fra gilis, the sheep frog Hypo- pachus variolosus, and the leopard frog Lithobates neo- volcanicus (Table 4). Similar numbers of species have been recorded from the Balsas-Tepalcatepec Depression, the Sierra Madre del Sur, and the Transverse Volcanic Axis. A small- er number occupies the Coastal Plain and the smallest number is found on the Central Plateau. The distinction between the species numbers in the higher-species areas (Balsas-Tepalcatepec Depression, Sierra Madre del Sur, and the Transvese Volcanic Axis) and the lower-species areas (Coastal Plain and Central Plateau) is more marked for amphibians than for reptiles (Table 5). Table 4. Distribution of the native and non-native amphibian and reptiles of Michoacan, Mexico, by physiographic province. Taxa Physiographic Provinces Coastal Plain (COP) Balsas- Tepalcatepec Depression (BTD) SierraMadre del Sur (SMS) Transverse Volcanic Axis (TVA) Central Plateau (CEP) Amphibia (54 species) Anura (44 species) Bufonidae (6 species) Anaxyrus compactilis + + Incilius marmoreus + + + Incilius occidentalis + + Incilius perplexus + + Incilius pisinnus + + Rhinella marina + + + + Craugastoridae (5 species) Craugastor augusti + + Craugastor hobartsmithi + Craugastor occidentalis + Craugastor pygmaeus + + + Craugastor vocalis + + + Eleutherodactylidae (5 species) Eleutherodactylus angustidigitorum + Eleutherodactylus maurus + Eleutherodactylus modestus + Eleutherodactylus nitidus + + + + Eleutherodactylus rufescens + Hylidae (11 species) Agalychnis dacnicolor + + + Diaglena spatulata + + Exerodonta smaragdina + + + + Hyla arenicolor + + + + Hyla eximia + + September 2013 | Volume 7 | Number 1 | e71 Amphib. Reptile Conserv. | http://amphibian-reptile-conservation.org 137 Physiographic distribution and conservation of Michoacan herpetofauna Hyla plicata + Plectrohyla bistincta + + Smilisca baudinii + + + Smilisca fodiens + + Tlalocohyla smithii + + + Trachycephalus typhonius + Leptodactylidae (2 species) Leptodactylus fragilis + + + + Leptodactylus melanonotus + + + Microhylidae (2 species) Hypopachus ustus + Hypopachus variolosus + + + + Ranidae (11 species) Lithobates berlandieri + Lit ho bates catesbeianus + Lithobates dunni + Lithobates forreri + + Lithobates magnaocularis + Lithobates megapoda + + Lithobates montezumae + + Lithobates neovolcanicus + + + + Lithobates pustulosus + + + Lithobates spectabilis + Lithobates zweifeli + + Rhinophrynidae (1 species) Rhinophrynus dorsalis + Scaphiopodidae (1 species) Spea multiplicata + + Caudata (9 species) Ambystomatidae (6 species) Ambystoma amblycephalum + Ambystoma andersoni + Ambystoma dumerilii + Ambystoma ordinarium + Ambystoma rivulare + Ambystoma velasci + Plethodontidae (3 species) Pseudoeurycea bellii + Pseudoeurycea leprosa + Pseudoeurycea longicauda + Gymnophiona (1 species) Caeciliidae (1 species) Dermophis oaxacae + + Reptilia (161 species) Crocodylia (1 species) Crocodylidae (1 species) Crocodylus acutus + September 2013 | Volume 7 | Number 1 | e71 Amphib. Reptile Conserv. | http://amphibian-reptile-conservation.org 138 Alvarado-Dfaz et al. Squamata (153 species) Bipedidae (1 species) Bipes canaliculatus + Anguidae (6 species) Abronia deppii + Barisia imbricata + Barisia jonesi + Barisia rudicollis + Elgaria kingii + Gerrhonotus liocephalus + Corytophanidae (1 species) Basiliscus vittatus + + + Dactyloidae (2 species) Anolis dunni + + Anolis nebulosus + + + + + Eublepharidae (1 species) Coieonyx eiegans + + Gekkonidae (1 species) Hemidactylus frenatus + + + Helodermatidae (1 species) Heloderma horridum + + + Iguanidae (3 species) Ctenosaura clarki + Ctenosaura pectinata + + + Iguana iguana + + + Mabuyidae (1 species) Marisora brachypoda + + Phrynosomatidae (20 species) Phrynosoma asio + + Phrynosoma orbiculare + Sceloporus aeneus + Sceloporus asper + + + Sceloporus bulled + Sceloporus dugesii + + Sceloporus gadoviae + + Sceloporus grammicus + Sceloporus heterolepis + + Sceloporus horridus + + + + Sceloporus insignis + Sceloporus melanorhinus + + + Sceloporus pyrocephalus + + + Sceloporus scalaris + + Sceloporus siniferus + + Sceloporus spinosus + + Sceloporus torquatus + + Sceloporus utiformis + + + + Urosaurus bicarinatus + + + + September 2013 | Volume 7 | Number 1 | e71 Amphib. Reptile Conserv. | http://amphibian-reptile-conservation.org 139 Physiographic distribution and conservation of Michoacan herpetofauna Urosaurus gadovi + + Phyllodactylidae (5 species) Phyllodactylus davisi + Phyllodactylus duellmani + + Phyllodactylus homolepidurus + Phyllodactylus lane i + + + + Phyllodactylus paucituberculatus + Scincidae (6 species) Mesoscincus altamirani + + Plestiodon colimensis + + Plestiodon copei + Plestiodon dugesii + Plestiodon indubitus + + Plestiodon parvulus + Sphenomorphidae (1 species) Scincella assata + + + Teiidae (8 species) Aspidoscelis calidipes + + Aspidoscelis communis + + + Aspidoscelis costata + + Aspidoscelis deppei + + + Aspidoscelis gularis + + Aspidoscelis lineatissima + + + Aspidoscelis sacki + Holcosus undulatus + + + Xantusiidae (1 species) Lepidophyma tarascae + + Boidae (1 species) Boa constrictor + + + Colubridae (28 species) Conopsis biserialis + Conopsis lineatus + + Conopsis nasus + Drymarchon melanurus + + + Drymobius margaritiferus + + + Geagras redimitus + Gyalopion canum + Lampropeltis ruthveni + Lampropeltis triangulum + Leptophis diplotropis + + + Masticophis flagellum + + Masticophis mentovarius + + + + + Masticophis taeniatus + + Mastigodryas melanolomus + + Oxybelis aeneus + + + Pituophis deppei + + September 2013 | Volume 7 | Number 1 | e71 Amphib. Reptile Conserv. | http://amphibian-reptile-conservation.org 140 Alvarado-Dfaz et al. Pituophis lineaticollis + + Pseudoficimia frontalis + + + Salvadora bairdi + + Salvadora mexicana + + Senticolis triaspis + + Sonora michoacanensis + + Symphimus leucostomus + Tantilla bocourti + Tantilla calamarina + + + Tantilla cascadae + Trimorphodon biscutatus + + + Trimorphodon tau + + + Dipsadidae (33 species) Coniophanes fissidens + + Coniophanes lateritius + + Coniophanes michoacanensis + Coniophanes piceivittis + Coniophanes sarae + Diadophis pu net at us + Dipsas gaigeae + Enuiius flavitorques + + Enuiius oligostichus + Geophis bicolor + Geophis dugesii + Geophis incomptus + Geophis maculiferus + Geophis nigrocinctus + Geophis petersii + + Geophis pyburni + Geophis sieboldi + Geophis tarascae + Hypsiglena torquata + + Imantodes gemmistratus + Leptodeira maculata + + + Leptodeira nigrofasciata + Leptodeira septentrionalis + Leptodeira splendida + + + Leptodeira uribei + Pseudoleptodeira latifasciata + + Rhadinaea hesperia + + Rhadinaea laureata + Rhadinaea taeniata + Si bon nebulata + + Tropidodipsas annulifera + Tropidodipsas fasciata + Tropidodipsas philippii + + September 2013 | Volume 7 | Number 1 | e71 Amphib. Reptile Conserv. | http://amphibian-reptile-conservation.org 141 Physiographic distribution and conservation of Michoacan herpetofauna Elapidae (4 species) Micrurus distans + + Micrurus laticollaris + Micrurus tener + Pelamis platura + Leptotyphlopidae (4 species) Epictia goudotii + + Rena bressoni + Rena humilis + Rena maxima + Loxocemidae (1 species) Loxocemus bicolor + + Natricidae (11 species) Adelophis copei + Storeria storerioides + + Thamnophis cyrtopsis + + Thamnophis eques + + Thamnophis melanogaster + Thamnophis postremus + Thamnophis proximus + Thamnophis pulchrilatus + Thamnophis scalaris + Thamnophis scaliger + + Thamnophis validus + Typhlopidae (1 species) Ramphotyphlops braminus + + + Viperidae (10 species) Agkistrodon bilineatus + + + Crotalus aquilus + Crotalus basiliscus + + + Crotalus culminatus + Crotalus molossus + Crotalus polystictus + Crotalus pusillus + + Crotalus tancitarensis + Crotalus triseriatus + Porthidium hespere + Xenodontidae (2 species) Conophis vittatus + + + Manolepis putnami + + Testudines (7 species) Cheloniidae (2 species) Chelonia mydas + Lepidochelys olivacea + Dermochelyidae (1 species) Dermochelys coriacea + September 2013 | Volume 7 | Number 1 | e71 Amphib. Reptile Conserv. | http://amphibian-reptile-conservation.org 142 Alvarado-Dfaz et al. Geoemydidae (2 species) Rhinoclemmys pulcherrima + Rhinoclemmys rubida + + + Kinosternidae (2 species) Kinosternon hirtipes + + Kinosternon integrum + + + + + Table 5. Summary of the distributional occurrence of families of amphibians and reptiles in Michoacan by physiographic province. Families Number of Species Distributional Occurrence Coastal Plain (COP) Balsas- Tepalcatepec Depression (BTD) Sierra Madre del Sur (SMS) Transverse Volcanic Axis (TVA) Central Plateau (CEP) Bufonidae 6 2 4 5 2 2 Craugastoridae 5 — 2 3 5 — Eleutherodactylidae 5 — 2 3 2 1 Hylidae 11 5 7 6 5 4 Leptodactylidae 2 2 2 2 1 — Microhylidae 2 1 1 1 1 1 Ranidae 11 — 6 4 7 3 Rhinophrynidae 1 — 1 — — — Scaphiopodidae 1 — — — 1 1 Subtotals 44 10 25 24 24 12 Ambystomatidae 6 — — — 6 — Plethodontidae 3 — — — 3 — Subtotals 9 — — — 9 — Caeciliidae 1 1 — — 1 — Subtotals 1 1 — — 1 — Totals 54 11 25 24 34 12 Crocodylidae 1 1 — — — — Subtotals 1 1 — — — — Cheloniidae 2 2 — — — — Dermochelyidae 1 1 — — — — Geoemydidae 2 2 1 1 — — Kinosternidae 2 1 1 1 2 2 Subtotals 7 6 2 2 2 2 Bipedidae 1 — 1 — — — Anguidae 6 — — 2 4 — Corytophanidae 1 1 1 1 — — Dactyloidae 2 1 2 2 1 1 Eublepharidae 1 1 1 — — — Gekkonidae 1 1 1 1 — — Helodermatidae 1 1 1 1 — — Iguanidae 3 2 3 2 — — Mabuyidae 1 1 1 — — — Phrynosomatidae 20 6 9 13 12 4 September 2013 | Volume 7 | Number 1 | e71 Amphib. Reptile Conserv. | http://amphibian-reptile-conservation.org 143 Physiographic distribution and conservation of Michoacan herpetofauna Phyllodactylidae 5 3 3 2 1 — Scincidae 6 2 1 3 3 — Sphenomorphidae 1 1 1 1 — — Teiidae 8 4 7 6 1 1 Xantusiidae 1 1 1 — — — Subtotals 58 25 33 34 22 6 Boidae 1 1 1 1 — — Colubridae 28 11 13 13 15 6 Dipsadidae 33 8 10 19 9 — Elapidae 4 1 2 1 1 — Leptotyphlopidae 4 — 4 1 — — Loxocemidae 1 — 1 1 — — Natricidae 11 2 1 2 7 3 Typhlopidae 1 — 1 1 1 — Viperidae 10 3 3 3 6 — Xenodontidae 2 2 2 1 — — Subtotals 95 28 38 43 39 9 Totals 161 60 73 79 63 17 Sum Totals 215 71 98 103 97 29 Anurans are more broadly represented in the Balsas-Tepalcatepec Depression, where 25 species classified in all but one of the nine families occurring in the state are found. These anurans are represented most narrowly on the Coastal Plectrohyla bistincta. The Mexican fringe-limbed treefrog is distributed from Durango and Veracruz southward to Mexico and Oaxaca. Its EVS has been assessed as 9, placing at the upper end of the low vulnerability category, this species is considered as Least Concern by IUCN, and as a Special Protection species by SEMARNAT. This individual came from San Jose de las Tomes, near Morelia, in Michoacan. Photo by Javier Alvarado -Diaz. Plain, where only 10 species assigned to four families occur. One or more species in the families Bufonidae, Hylidae, and Microhylidae are distributed in each of the five provinces (Table 5). As expected, the family Hylidae is best represented in each of the provinces except for the Transverse Volcanic Axis, where more ranids (sev- en species) than hylids (five) occur. All nine species of salamanders are limited in occurrence to the Transverse Volcanic Axis and the single caecilian to the Transverse Volcanic Axis and the Coastal Plain (Table 5). Lizards are best represented in the Sierra Madre del Sur, with 34 species, but the Balsas-Tepalcatepec De- pression falls only one behind, with 33 (Table 5). Both of these figures comprise more than one-half of the 58 species of lizards known from the state. Fewer than one- half of this number occurs on the Coastal Plain (25) and the Transverse Volcanic Axis (22). Only a few species (six) occur on the Central Plateau. In the families Dacty- loidae, Phrynosomatidae, and Teiidae, one or more spe- cies is distributed in each of the five provinces (Table 5). Due to the size of the Phrynosomatidae in Michoacan (20 species), this family is the best represented in each of the provinces. Several lizard families are represented by a single species in each of the provinces, but only one with a single species (the Bipedidae) is limited to a single province (Table 5). The largest number of snake species is known from the Sierra Madre del Sur, with 43 species. Fewer num- bers are found in the Transverse Volcanic Axis (39), Bal- sas-Tepalcatepec Depression (38), Coastal Plain (28), and the Central Plateau (nine). One or more represen- tatives of only two snake families, the Colubridae and Natricidae, are found in each of the five provinces (Table 5). Interestingly, although the Colubridae in Michoacan is represented by five fewer species than the Dipsadidae, it is the best-represented family in all of the provinces except for the Sierra Madre del Sur, in which the Dip- September 2013 | Volume 7 | Number 1 | e71 Amphib. Reptile Conserv. | http://amphibian-reptile-conservation.org 144 Alvarado-Dfaz et al. Amby stoma velasci. The plateau tiger salamander is found along the Transverse Volcanic Axis in Michoacan and elsewhere, thence northward into both the Sierra Madre Occidental to northwestern Chihuahua and the Sierra Madre Oriental to southern Nuevo Leon. Its EVS has been assigned a value of 10, placing it at the lower end of the medium vulnerability category, its status has been judged as Least Concern by IUCN, and it is considered a Special Protection species by SEMARNAT. This individual came from Los Azufres, in the Tranverse Volcanic Axis. Photo by Javier Alvarado-Dfaz. sadidae is the best represented. Only three snake fam- ilies are represented by a single species (including the Typhlopidae, containing the non-native blindsnake Ram- photyphlops braminus ), but in all three cases they occur in two or three provinces (Table 5). Relatively few species of turtles have been recorded in Michoacan, and given that three of the seven are sea turtles, most of them (six) are known from the Coastal Plain (obviously, sea turtles come on land for egg depo- sition). Only two species of the families Geoemydidae and/or Kinosternidae are found in the