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.

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.

Department of Zoology
University of Cape Town
Private Bag, Rondebosch 7701
South Africa

Southwest Fisheries Center
National Marine Fisheries Service
Pacific Grove, CA 93950
USA

Division of Mammals
National Museum of Natural History
Washington, DC 20013-7012
USA

U.S. Geological Survey
National Museum of Natural History
Washington, DC 20013-7012
USA

Department of Archaeology and Anthropology
The Australian National University
GPO Box 4
Canberra ACT 2601
Australia

School of Earth and Environmental Sciences
University of Adelaide
Adelaide, SA 5005
Australia

Division of Mammals
National Museum of Natural History
Washington, DC 20013-7012
USA

Division of Vertebrate Zoology
American Museum of Natural History
Correspondence Address:
1714 Henley Lane
Charleston, SC 29412
USA

Zoologisches Forschunginstitut und Museum Alexander Koenig
Adenauerallee 160
53113 Bonn
Germany

Mammal Group, Department of Zoology
The Natural History Museum
Cromwell Road
London SW7 5BD
UK

Division of Mammals
National Museum of Natural History
Washington, DC 20013-7012
USA

Division of Vertebrate Zoology
American Museum of Natural History
Correspondence Address:
1714 Henley Lane
Charleston, SC 29412
USA

Department of Wildlife and Fisheries Sciences
Texas A&M University
2258 TAMU
College Station, TX 77843-2258
USA

Department of Biology
University of Asmara
P. O. Box 1220 Asmara
Eritrea (Horn of Africa)
Correspondence Address:
Elephant Research Foundation
106 East Hickory Grove Road
Bloomfield Hills, Michigan 48304-1729
USA

Division of Vertebrate Zoology
American Museum of Natural History
New York, NY 10024
USA

School of Life Sciences
Box 874501
Arizona State University
Tempe, AZ 85287-4501
USA

Division of Mammals
National Museum of Natural History
Washington, DC 20013-7012
USA

Florida Museum of Natural History
University of Florida
Gainesville, FL 32611-2035
USA

Division of Natural Science
Bethel College
1001 W. McKinley Ave.
Mishawaka, IN 46545
USA

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).

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.

						Number of Genera	Number of Species	Number of New Species

Class Mammalia						1229	5416	260
	Order Monotremata					3	5	1
					Family Tachyglossidae	2	4	1
					Family Ornithorhynchidae	1	1	–
	Order Didelphimorphia					17	87	2
					Family Didelphidae	17	87	2
	Order Paucituberculata					3	6	1
					Family Caenolestidae	3	6	1
	Order Microbiotheria					1	1	–
					Family Microbiotheriidae	1	1	–
	Order Notoryctemorphia					1	2	–
					Family Notoryctidae	1	2	–
	Order Dasyuromorphia					22	71	5
					Family Thylacinidae	1	1	–
					Family Myrmecobiidae	1	1	–
					Family Dasyuridae	20	69	5
	Order Paramelemorphia					8	21	–
					Family Thylacomyidae	1	2	–
					Family Chaeropodidae	1	1	–
					Family Peramelidae	6	18	–
	Order Diprotodontia					39	143	8
		Suborder Vombatiformes				3	4	–
					Family Phascolarctidae	1	1	–
					Family Vombatidae	2	3	–
		Suborder Phalangeriformes				20	63	3
				Superfamily Phalangeroidea		8	32	3
					Family Burramyidae	2	5	–
					Family Phalangeridae	6	27	3
				Superfamily Petauroidea		12	31	–
					Family Pseudocheiridae	6	17	–
					Family Petauridae	3	11	–
					Family Tarsipedidae	1	1	–
					Family Acrobatidae	2	2	–
		Suborder Macropodiformes				16	76	5
					Family Hypsiprymnodontidae	1	1	–
					Family Potoroidae	4	10	–
					Family Macropodidae	11	65	5
	Order Afrosoricida					19	51	6
		Suborder Tenrecomorpha				10	30	5
					Family Tenrecidae	10	30	5
		Suborder Chrysochloridea				9	21	1
					Family Chrysochloridae	9	21	1
	Order Macroscelidea					4	15	–
					Family Macroscelididae	4	15	–
	Order Tubulidentata					1	1	–
					Family Orycteropodidae	1	1	–
	Order Hyracoidea					3	4	–
					Family Procaviidae	3	4	–
	Order Proboscidea					2	3	–
					Family Elephantidae	2	3	–
	Order Sirenia					3	5	–
					Family Dugongidae	2	2	–
					Family Trichechidae	1	3	–
	Order Cingulata					9	21	1
					Family Dasypodidae	9	21	1
	Order Pilosa					5	10	1
		Suborder Folivora				2	6	1
					Family Bradypodidae	1	4	1
					Family Megalonychidae	1	2	–
		Suborder Vermilingua				3	4	–
					Family Cyclopedidae	1	1	–
					Family Myrmecophagidae	2	3	–
	Order Scandentia					5	20	–
					Family Tupaiidae	4	19	–
					Family Ptilocercidae	1	1	–
	Order Dermoptera					2	2	–
					Family Cynocephalidae	2	2	–
	Order Primates					69	376	24
		Suborder Strepsirrhini				23	88	9
			Infraorder Lemuriformes			14	59	8
				Superfamily Cheirogaleoidea		5	21	6
					Family Cheirogaleidae	5	21	6
				Superfamily Lemuroidea		9	38	1
					Family Lemuridae	5	19	–
					Family Lepilemuridae	1	8	–
					Family Indridae	3	11	1
			Infraorder Chiromyiformes			1	1	–
					Family Daubentoniidae	1	1	–
			Infraorder Lorisiformes			8	28	2
					Family Lorisidae	5	9	1
					Family Galagidae	3	19	1
		Suborder Haplorrhini				46	288	14
			Infraorder Tarsiiformes			1	7	–
					Family Tarsiidae	1	7	–
			Infraorder Simiiformes			45	281	14
				Parvorder Platyrrhini		16	128	11
					Family Cebidae	6	56	8
					Family Aotidae	1	8	–
					Family Pitheciidae	4	40	3
					Family Atelidae	5	24	–
				Parvorder Catarrhini		29	153	3
				Superfamily Cercopithecoidea		21	132	4
					Family Cercopithecidae	21	132	4
				Superfamily Hominoidea		8	21	–
					Family Hylobatidae	4	14	–
					Family Hominidae	4	7	–
	Order Rodentia					481	2277	128
		Suborder Sciuromorpha				61	307	1
					Family Aplondontiidae	1	1	–
					Family Sciuridae	51	278	–
					Family Gliridae	9	28	1
		Suborder Castorimorpha				13	102	2
					Family Castoridae	1	2	–
					Family Heteromyidae	6	60	2
					Family Geomyidae	6	40	–
		Suborder Myomorpha				326	1569	96
				Superfamily Dipodoidea		16	51	–
					Family Dipodidae	16	51	–
				Superfamily Muroidea		310	1518	96
					Family Platacanthomyidae	2	2	–
					Family Spalacidae	6	36	4
					Family Calomyscidae	1	8	–
					Family Nesomyidae	21	61	6
					Family Cricetidae	130	681	53
					Family Muridae	150	730	33
		Suborder Anomaluromorpha				4	9	–
					Family Anomaluridae	3	7	–
					Family Pedetidae	1	2	–
		Suborder Hystricomorpha				77	290	29
			Infraorder Ctenodactylomorphi			4	5	–
					Family Ctenodactylidae	4	5	–
			Infraorder Hystricognathi			73	285	29
					Family Bathyergidae	5	16	2
					Family Hystricidae	3	11	–
					Family Petromuridae	1	1	–
					Family Thryonomyidae	1	2	–
					Family Erethizontidae	5	16	2
					Family Chinchillidae	3	7	–
					Family Dinomyidae	1	1	–
					Family Caviidae	6	18	2
					Family Dasyproctidae	2	13	–
					Family Cuniculidae	1	2	–
					Family Ctenomyidae	1	60	5
					Family Octodontidae	8	13	3
					Family Abrocomidae	2	10	2
					Family Echimyidae	21	90	13
					Family Myocastoridae	1	1	–
					Family Capromyidae	8	20	–
					Family Heptaxodontidae	4	4	–
	Order Lagomorpha					13	92	5
					Family Ochotonidae	1	30	2
					Family Prolagidae	1	1	–
					Family Leporidae	11	61	3
	Order Erinaceomorpha					10	24	1
					Family Erinaceidae	10	24	1
	Order Soricomorpha					45	428	18
					Family Nesophontidae	1	9	–
					Family Solenodontidae	1	4	1
					Family Soricidae	26	376	17
					Family Talpidae	17	39	–
	Order Chiroptera					202	1116	49
					Family Pteropodidae	42	186	6
					Family Rhinolophidae	1	77	4
					Family Hipposideridae	9	81	7
					Family Megadermatidae	4	5	–
					Family Rhinopomatidae	1	4	–
					Family Craseonycteridae