Wilson & Reeder’s Mammal Species of the World – 3rd edition
“A checklist of species is an invaluable tool for both researchers and the interested public.” Thus began the first edition of this work, published by the Association of Systematics Collections (ASC) and Allen Press in 1982. That first edition was prepared by 189 professional mammalogists from 23 countries. It was coordinated by a special Checklist Committee of the American Society of Mammalogists. During the ensuing decade, it became the industry standard for mammalian taxonomy, providing an authoritative reference for nonspecialists and establishing an overall taxonomic hypothesis for testing by systematic mammalogists.
The American Society of Mammalogists anticipated the need for revision and established a Standing Checklist Committee under the chairmanship of Karl F. Koopman in June 1982, concurrent with the publication of the first edition. Duane A. Schlitter joined Koopman as co-chair in 1985, and they coordinated the committee’s efforts until 1990. At that time, Don E. Wilson assumed the chairmanship of the committee, with a mandate to expand the committee and produce a second edition of the checklist. With support from the Office of Biodiversity Programs at the Smithsonian Institution’s National Museum of Natural History, and with additional funding from the Seidell Fund of the Smithsonian Institution, DeeAnn M. Reeder joined the project in August 1991. At this time, we shifted from the egalitarian approach of multiple authors per section and assigned authorship of the various taxonomic groups to specialists in the field. In 1993, the second edition of Mammal Species of the World was published, and the database from which it was derived became available to the public at: http://www.nmnh.si.edu/msw.
In 2002, the various authors, some of whom were new to the project, began in earnest to update the text for the third edition. This third edition is significantly enhanced by the inclusion of common names, recognition of subspecies, and inclusion of authorities for all synonyms. This additional information, coupled with the virtual explosion in taxonomic literature over the past decade, has resulted in the near doubling in size of the text between the second and third edition. Students of mammalian taxonomy have made significant advances in recent years, especially with the advent and refinement of additional molecular techniques. Beyond the additions due to revisions of known mammals that have occurred over the decades, a significant number of new mammalian species have been described, totaling 171 new species between the first and second edition of Mammal Species of the World (1982-1992) and 260 new species between the second and third edition (1993-2003).
Because of the inherent fluidity of mammalian taxonomy, with dramatic changes occurring in relatively short periods of time due to new data and interpretations and new species discoveries, we anticipate continued changes to the arrangement presented here. The checklist will be maintained and updated in a web-based database, where all data will be freely available to the public in a variety of ways and usable formats. We anticipate updating the database at least annually. We welcome your suggestions, comments, and additions, and would particularly appreciate receiving copies of pertinent literature for preparation of future editions.
Don E. Wilson
National Museum of Natural History
DeeAnn M. Reeder
When a work as complex as Mammal Species of the World is finished, it is almost impossible to identify all of the many contributions deserving mention that have made the feat possible. We are ultimately most grateful to the hundreds of professional mammalogists who devoted their careers to the systematic study of Mammalia. Each has made a particular contribution to this synthesis.
The project owes much to Stephen R. Edwards and Robert S. Hoffmann, who initiated the effort that led to the first edition. That first edition was capably edited by James H. Honacki, Kenneth E. Kinman, and James W. Koeppl. The original Checklist Committee, chaired by Alfred L. Gardner and including Robert L. Brownell, Jr., Robert S. Hoffmann, Karl F. Koopman, Guy G. Musser, and Duane A. Schlitter, worked intensively to complete the first edition.
Elaine Hoagland, who succeeded Steve Edwards as executive director of the Association of Systematics Collections, facilitated transfer of the copyright to the American Society of Mammalogists. Duane Schlitter and Karl Koopman served as co-chairmen of the Checklist Committee for many years, keeping the project functioning during the decade between the first and second editions.
The production of the second edition was greatly facilitated by the creation of a database system for housing and searching the data from the text, which has been available on the internet at: http://www.nmnh.si.edu/msw. A number of Smithsonian Institution Information Resource Management staff were involved in this initiative, including Joe Russo and Barbara Weitbrecht. W. Christopher Wozencraft managed the thousands of references for the second edition. Ralph Walker of Information Resource Management and Peter Cannell of Smithsonian Institution Press diligently worked with the editors to facilitate final production of the second edition. The web version of the second edition was created and maintained by Don Gourley and subsequently by Dennis Hasch, both webmasters for the National Museum of Natural History. This has proven to be one of the museum’s most popular websites and has been a valuable resource to the research community and the public at large.
The Division of Mammals at the National Museum of Natural History, Smithsonian Institution, provided an intellectual home for the editors for the second and third edition and the assistance of the curatorial, collections, and library staff is much appreciated. Kristofer M. Helgen reviewed most of the text and contributed significantly to the integrity of the data. A myriad of professional mammalogists and others from around the world made significant contributions by reviewing the text, offering advice, examining and/or providing specimens, providing unpublished data, and otherwise contributing intellectually, including: H. Abe, Alexei Abramov, John Aguilar, Ticul Alvarez-Castaneda, Robert P. Anderson, Ken Aplin, Robert Asher, J.-C.P. Auffray, Stephane Aulagnier, Patrick Barriere, Kurt Bauer, Simon Bearder, Susan K. Bell, Wim Bergmans, Mary Boise, Robert Bradley, Doug Brandon-Jones, W. G. Breed, Norman Bridges, Patricia Brunauer, Leslie Carraway, Francois Catzeflis, Chris Chimimba, Gordon B. Corbet, Woody Cotterill, João Crawford Cabral, Gabor Csorba, Nick Czeplewski, Daryl P. Domning, Judith Eger, Guillermo d'Elia, Louise Emmons, James W. Demastes, Christiane Denys, Jerry Dragoo, Jean-Francois Ducroz, Jakob Fahr, Katherine E. Ferrell, Robert D. Fisher, Tim Flannery, Larry J. Flynn, Fred Ford, Nikolai A. Formozov, Rosa García-Perea, Alfred L. Gardner, Philippe Gaubert, Valerius Geist, F. N. Golenishchev, Enrique Gonzalez, Steve Goodman, Antonia Gorog, Laurent Granjon, David J. Hafner, Mark S. Hafner, Meredith Happold, Rudolf Haslauer, Larry Heaney, Robert Hoffmann, M. Hugueney, Rainer Hutterer, Peter Jackson, Sharon Jansa, Paula Jenkins, Kate Jones, Y. Kaneko, Julian Kerbis Peterhans, Andrew Kitchener, Dieter Kock, Karl Koopman, Boris Kryštufek, Marcia Lara, Leonid A. Lavrenchenko, Emilie Lecompte, Yuri Leite, Georges Lenglet, Enrique P. Lessa, Burton Lim, Alicia Linzey, Darrin Lunde, Milos Macholan, V. G. Malikov, Joe T. Marshall, Jr., Erik Meijaard, Pierre Mein, Jim Meng, J. Michaux, A. Miljutin, Alvaro Mones, Brian Mould, Masaharu Motokawa, Guy Musser, J. Obuch, Jose Ochoa, Link Olson, Ulyses Pardinas, Bruce Patterson, James L. Patton, I. Ya. Pavlinov, Caroline Pollock, E. G. Potapova, Fiona Reid, Dale Rice, Ken Richardson, Eric Rickart, Luis A. Ruedas, Sue Ruff, Anthony Rylands, Eric Sargis, Erik Seiffert, Chad Schennum, Duane Schlitter, G. Shenbrot, Chris Smeenk, Angela Smith, William Stanley, Gerhard Storch, Peter Taylor, Adrian Tejedor, M. Tranier, Victor Van Cakenberg, Harry Van Rompaey, Erik Van der Straeten, Walter N. Verheyen, Geraldine Veron, Vitaly Volobouev, John H. Wahlert, Robert Wayne, Lars Werdelin, Daniel F. Williams, John Wible, Neal Woodman, Jiong Wozencraft, Yan Xie, and Mikhail Zaitsev.
Finally, we would like to express our gratitude to the National Museum of Natural History for making funds available for the completion of this project.
LIST OF CONTRIBUTORS
Gary N. Bronner
Department of Zoology
Robert L. Brownell, Jr.
Southwest Fisheries Center
Michael D. Carleton
Division of Mammals
Staatliches Museum für Naturkunde
Alfred L. Gardner
U.S. Geological Survey
Colin P. Groves
Department of Archaeology and Anthropology
35 Downhills Park Road
Kristofer M. Helgen
School of Earth and Environmental Sciences
Robert S. Hoffmann
Division of Mammals
Mary Ellen Holden
Division of Vertebrate Zoology
Zoologisches Forschunginstitut und Museum Alexander Koenig
Paulina D. Jenkins
Mammal Group, Department of Zoology
C. William Kilpatrick
Department of Biology
James G. Mead
Division of Mammals
Guy G. Musser
Division of Vertebrate Zoology
James L. Patton
Museum of Vertebrate Zoology
DeeAnn M. Reeder
Department of Biology
Duane A. Schlitter
Department of Wildlife and Fisheries Sciences
Department of Biology
Nancy B. Simmons
Division of Vertebrate Zoology
Andrew T. Smith
School of Life Sciences
Brian J. Stafford
Division of Mammals
Richard W. Thorington, Jr.
Division of Mammals
Don E. Wilson
Division of Mammals
Charles A. Woods
Florida Museum of Natural History
W. Christopher Wozencraft
Division of Natural Science
The dynamic, rapidly changing state of mammalian taxonomy, documented by an enormous literature, long hampered the compilation of a detailed, complete world checklist. It was only about a century ago that the first complete appraisal of all mammals of the world was produced by Trouessart (1898-99, 1904-05). A more modern compilation was provided when E. P. Walker and colleagues brought out, in 1964, the first edition of Mammals of the World. This compendium, now in its sixth edition (Nowak, 1999), is arranged systematically (with considerable supplementary natural history data) to the generic level, and later editions list the species in each genus in addition to furnishing an illustration of at least one member of each genus. V. E. Sokolov based his Systematics of Mammals, published in Russian (1973-79), on Walker’s Mammals of the World. He provided a list of species with a brief summary of geographic distribution in each generic account. Corbett and Hill (1980, 1991) listed the species of the world, abbreviated distributions, common names, literature citations to major regional distributional works, and some additional revisionary works where appropriate. McKenna and Bell (1997) provided a complete phylogeny of mammals above the species level, including fossil as well as recent forms. That work provided a starting point for this edition of Mammal Species of the World, and deviations from their arrangement are noted in the comment sections of the accounts that follow. In the short time since the publication of McKenna and Bell (1997), an explosion of literature based on new techniques of molecular systematics has resulted in wholesale changes in our thinking about mammalian phylogeny. Those changes are reflected in the following pages, but this work is primarily a checklist at the species level, and higher-level relationships are used primarily to provide structure rather than to reflect phylogeny.
This volume, like previous editions (1982, 1993), will undoubtedly be used by many readers who are not systematic mammalogists. Do not be alarmed or disheartened by the debate over definition of species limits within many groups of mammals. Differences of opinion are aired in the comments sections in order to emphasize areas needing additional taxonomic study. Mammals are no worse off in this regard than other groups of animals, and in fact are probably better known than most, with the possible exception of birds. One recurring suggestion from users of previous editions spurred us to include common names in this edition. The publication of the first complete list of common names of mammal species of the world (Wilson and Cole, 2000) made this possible. Contributors to this edition used those names as a starting point, but were urged to adopt alternatives if there were compelling reasons to do so. As a result, this volume can be viewed as a second edition of Wilson and Cole (2000).
THE PROCESS OF COMPILATION
Knowledge of the systematics of mammals is distributed over an extensive assortment of works. The process of compiling and editing the information contained in this edition drew heavily on lessons learned from prior editions.
The first edition (Honacki et al., 1982) evolved from three sources: a manuscript written by Kenneth E. Kinman, a preliminary list developed by the U.S. Fish and Wildlife Service, and the International Species Inventory (ISIS) List (Seal and Makey, 1974). A draft checklist developed from those sources was made available to the professional mammalogy community. Members of the American Society of Mammalogists were invited to contribute to the list at whatever level they wished. As a result, appropriate portions of the draft checklist were sent to 255 individuals. These reviewers were asked to provide the following information: author of the scientific name of the species, and citation; type locality; distribution; citations of revisions or reviews; important synonyms; and explanatory comments if necessary. One hundred fifty mammalogists provided reviews, covering 85 percent of the species in the list. Additional reviewers were eventually found for the remaining species. Compiling the various drafts included the necessity of comparing the various contributions and including information on which there was agreement. In cases where there was not agreement, errors may have been retained. To determine valid names of taxa in cases of disagreement, the most recent reviewer was followed. Some of these names have stood the test of time, and others have not.
The penultimate draft was forwarded to the Checklist Committee of the American Society of Mammalogists. This group (Robert L. Brownell, Jr., Alfred L. Gardner [Chair] Robert S. Hoffmann, Karl F. Koopman, Guy G. Musser, and Duane A. Schlitter) furnished the final review of the text and provided much useful additional information. The committee detected many problems resulting from the egalitarian effort, not all of which were satisfactorily resolved in the first edition.
Two categories of information were added after the review process. The protected status of affected taxa was listed, based on information from the Federal Register. The ISIS numbers for each species were listed, based on Seal and Makey (1974). In the second edition the protected status was updated and retained, but the ISIS numbers were not, due to their limited usefulness to most users.
The second edition was compiled and edited in a distinctly different fashion from the first. Although the original decision to consult many professional mammalogists was theoretically sensible, it introduced a practical information management dilemma for the first edition. The wealth of material and the challenge of uniting numerous different, frequently conflicting opinions caused weaknesses in the original volume. Nevertheless, the taxonomic hypotheses outlined in that volume provided the basis for the refinements apparent in the second edition. The second edition was envisioned as the second step in organizing a continuing taxonomic database for mammals.
Because comprehensive survey of the literature is now prohibitive for a single mammalogist (or perhaps because they don’t make us like they used to), the labor of composing the second edition was partitioned among 20 authors, specialists on their respective taxa. Although some authors had been maintaining records for their groups of interest for some time, preparation of the second edition began in earnest in the fall of 1990, when electronic and paper copies of the appropriate portions of the original text were distributed to the members of the committee along with a mandate to produce an up-to-date revision. Most of 1991 was spent in the production of that updated text. The authors examined the literature on mammals, from the initial works to the most recent, and provided references published in many languages. As each portion was received from the authors, it was converted to an electronic database, edited, regenerated in word processing format, and returned to the authors for revisions. After a second round of editing, reviews were sought from other members of the community of systematic mammalogists. Although both editors and reviewers made suggestions to the authors, each section was the product of an individual author’s scholarship, and represented that author’s best hypothesis of the relationships of a particular group at that time.
The information leading to that edition was compiled with computer technology that permitted rapid manuscript revision and organization. This allowed automatic indexing of the many names used and standardization of literature citations, as well as considerably more consistency in citations, punctuation, format, and of spelling of place names. In spite of this, infelicities crept in, and the third printing of the second edition contained corrections of some typographical mistakes, misspellings, incorrect dates, and overlooked synonyms. Some literature citations were further standardized at that time.
Details on the 4,629 species of mammals covered by the second edition (up from 4,170 in the first edition) were the product of myriad former scholars united by a common curiosity about the diversity of mammals (Table 1). A major feature of the second edition, like the original, was that it identified gaps in our knowledge in need of further study, and served as a starting point for the third edition.
The third edition was prepared in much the same manner as the second, although the number of contributors was increased slightly. During the past decade, information on the systematic relationships of mammals has continued to increase. The advent of modern molecular techniques has allowed increasingly detailed comparisons of species limits and evolutionary relationships. The number of species represented in the current edition is 5,416, up from 4,629 in the second edition. Although most of this increase is due to taxonomic revision, a significant proportion is due to newly described species. In addition to currently recognized species, this edition contains 37,378 synonyms and full references to 9,373 scientific publications.
ORGANIZATION OF THE BOOK
Species recognized herein are limited to existing or recently extinct species (possibly alive during the preceding 500 years); in instances where the persistence of a species is doubtful, the comment section so indicates. 500 years is an arbitrary span of time, selected to better allow us to judge the “recent” distribution of mammals on earth. It has nothing to do with Western humans’ arrival into the new world, or with any other historical event. Obviously, determining the exact time of extinction of any species is difficult at best, and undoubtedly we have included some species that push our arbitrary limit.
List of Museum Abbreviations
Abbreviations for specific museums are used in the text to indicate specimen repositories in some accounts. See page xxxv for full museum names.
Various workers have reviewed the higher categories of mammals since Simpson (1945) produced his definitive classification, including Anderson and Jones (1967, 1984), McKenna (1975), Corbet and Hill (1980, 1986, 1991), and McKenna and Bell (1997). We used McKenna and Bell (1997) as the starting point for this edition. Modifications to their arrangement have been duly noted and documented in the text. In addition to a wealth of changes at the species level since the second edition, our understanding of the phylogenetic relationships of mammals at the higher category level is changing rapidly, and currently presents a moving target. The only mandatory categories are order, family, genus, and species. We are recognizing 29 orders in this edition, up from 26 in the second edition. Xenarthra is split into the orders Cingulata and Pilosa, and Insectivora is split into Afrosoricida, Erinaceomorpha, and Soricomorpha. Within each order, individual authors have chosen the higher categories that best represent our knowledge of the group. Comment fields provide additional justification for the use or non-use of additional categories (e.g., suborder, superfamily, subfamily, tribe, etc.) within each order. Generic names are ordered alphabetically within subfamilies (or families, if no subfamilies are recognized), and species names are ordered alphabetically within genera, without exception. If subgenera are thought to be useful, they are listed in the comments section of each species. Most authors have indicated subspecies as well. Readers are urged to review the introductory comments for each section.
Scientific Name and Authority
Each currently used scientific name is followed by the name of the author(s) and the year in which it was described; e.g., Vulpes vulpes (Linnaeus, 1758). A species originally named by its author in a genus other than the one in which it is now placed has parentheses surrounding the author and date. In this example Linnaeus, when he first named the red fox vulpes, placed it in the genus Canis instead of Vulpes. Should it be returned to Canis, the parentheses would be removed, i.e., Canis vulpes Linnaeus, 1758. Following the species name, authority, and date is the citation for the work in which the original description appeared.
Usually, only the first page on which the species name appears follows the title citation. In some cases, for a variety of reasons, more than one page may be cited, as well as references to figures or plates. We have attempted to cite the first page on which the name appears, but if it is not listed unequivocally, the reader is advised to consult the original source.
Family names follow the principle of coordination outlined in Article 36 of the International Code of Zoological Nomenclature (1999). Generic accounts include the type species for which the generic name was proposed. In general, the name appears as it was originally described, including authority. When the type species is no longer a recognized species, the species with which it is currently synonymized is listed in parentheses in its original form with its authority.
Unlike previous editions, we have provided a common name for each recognized species. The starting point for these names is Wilson and Cole (2000), but each author was encouraged to examine those names and to provide a different one if there was good reason to do so. Thus, this list can be viewed as a second edition of Wilson and Cole (2000). There are no rules governing vernacular names, but Wilson and Cole (2000) outlined several reasons for adopting a single such name for each species of mammal.
The type locality is the geographical site where the type material of a species was obtained. Type localities quoted exactly from the original description are enclosed in quotation marks. Information not surrounded by quotation marks has been arranged where possible with the current country name followed by state, province, or district, and specific locality. Elevation above sea level has been included when available, as have global coordinates in some cases. When appropriate, restrictions of the type locality made by revisers have been included as well.
The geographical range of each species is summarized using contemporary political units or, in some cases, geographical names. However, geographical names are usually used only when the entire area is included in the range of the species. We have attempted to standardize usage and spelling, but some inconsistencies may remain, particularly in transliterations from other alphabets. Country names have changed drastically during the time the text was being prepared; we have attempted to give the most current name whenever possible. Current country names have been standardized according to the list of Independent States of the World, produced by the U.S. Department of State as of February 7, 2003. With the exception of the Republic of Congo and the Democratic Republic of Congo (herein abbreviated as Dem. Rep. Congo), the official “short-forms” of country names were used. Geographic names are subject to rapid changes in many parts of the world. Readers are urged to consult geographic thesauruses and local governmental lists when questions arise. Distribution records resulting from human introduction are sometimes noted. Maps of the distributions of many species are provided in the cited literature. If a species is known only from the type locality, that is noted.
Mammal species covered by the regulations for the U.S. Endangered Species Act (U.S. ESA) as of February 12, 2004; those listed in the 2003 International Union for Conservation of Nature and Natural Resources (IUCN) Red List of Threatened Animals, and those listed in the appendices of the Convention on International Trade in Endangered Species of Wild Fauna and Flora (CITES) as of March 2, 2004 are noted in the text in this category. Additionally, some authors have included local governmental regulations in this field as well as general comments regarding the conservation status of
particular species. For U.S. ESA listings, the categories of Endangered and Threatened (and supplemental categories such as proposed listings, emergency listings, and delisted taxa) are included. For the IUCN listings, the categories of Extinct, Extinct in the Wild, Critically Endangered, Endangered, Vulnerable, Lower Risk - near threatened (nt), Lower Risk - least concern (lc), Lower Risk - conservation dependent (cd), and Data Deficient are included. IUCN listings for species either officially "Not Evaluated" or not yet on the IUCN list are not included. For the CITES listings, Appendix I, II, and III are listed, where Appendix I includes species threatened with extinction that are or may be affected by trade, Appendix II includes species that although not necessarily threatened may become so unless trade in them is strictly controlled, as well as nonthreatened species that must be subject to regulation in order to control threatened species, and Appendix III includes species that any Party identifies as being subject to regulation within its jurisdiction for purposes of preventing or restricting exploitation, and for which it needs the cooperation of other Parties in controlling trade. Readers are cautioned against interpreting the lack of information in a status field as an indication that a particular species is not threatened. In many cases, recent taxonomic changes that are not yet reflected in the evaluations of the major conservation agencies will result in a taxon not being listed that is in fact endangered. Readers should consult the following websites for updates to the status fields: U.S. ESA (http://endangered.fws.gov/); IUCN (http://www.redlist.org); and CITES http://www.cites.org).
Authors have attempted to provide taxonomic lists consistent with recent literature, tempered by their own individual judgments. Considerable effort has been expended to compile a complete list of synonyms that have been used in the scientific literature for each taxon. These are usually either names of later origin than that used (junior synonyms) or names that are invalid systematically, for various reasons. Also included here are subspecies names, including any that might be currently recognized. Currently recognized subspecies are listed in boldface type, followed by their junior synonyms. Names listed before a boldfaced subspecies are synonyms of the nominate form. Subsequent emendations, misspellings, incorrect allocations, and partial synonyms are not included, for the most part. Theoretically, any scientific name used for a mammal should be found in this volume, either as a currently recognized species or as a synonym. One addition to this edition is the inclusion of the authority and date for each synonym, which should make it easier to find the original description of the name.
Taxonomic and nomenclatorial alternatives are accompanied by appropriate documentation, including opinions of the International Commission on Zoological Nomenclature (ICZN); revisions and additional literature sources are also cited. In the interest of brevity, secondary reference sources are sometimes cited to document taxonomic evidence, and reference to the primary sources can be found therein. Personal opinions of the author(s) and unpublished information are sometimes included here as well. When appropriate, other data such as references discussing type locality, occurrence of hybridization, and species known only from a single or few specimens are also included in the comments section. Absence of a comments section may indicate either that the species is taxonomically uncontroversial or that it is too poorly known to require comment.
The literature cited section contains the works consulted in the compilation of this text. Works appearing after 2003 are generally not cited, although several works in press at the end of 2003 are included. References cited as authorities for original descriptions are not included in the literature cited but are given after each currently recognized name in the text in sufficient detail to allow the reader to locate them. Author and date citations in the synonym section are not included in the literature cited as they are provided only as authorities for the names.
The index contains all taxonomic names contained in this volume, including synonyms. Page references to all currently recognized generic and species names employed in this volume are in italic boldface type. Species names are individually listed in alphabetical order.
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