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Novitatesamerican MUSEUM PUBLISHED by the AMERICAN MUSEUM of NATURAL HISTORY CENTRAL PARK WEST at 79TH STREET NEW YORK, N.Y NovitatesAMERICAN MUSEUM PUBLISHED BY THE AMERICAN MUSEUM OF NATURAL HISTORY CENTRAL PARK WEST AT 79TH STREET NEW YORK, N.Y. 10024 U.S.A. NUMBER 2646 APRIL 21, 1978 RICHARD G. VAN GELDER A Review of Canid Classification AMERICAN MUSEUM Novitates PUBLISHED BY THE AMERICAN MUSEUM OF NATURAL HISTORY CENTRAL PARK WEST AT 79TH STREET, NEW YORK, N.Y. 10024 Number 2646, pp. 1-10 April 21, 1978 A Review of Canid Classification RICHARD G. VAN GELDER' ABSTRACT A review of the varied classifications of Recent tereutes, Cuon, Lycaon, and Otocyon), and one members of the family Canidae involving mor- polytypic genus, Canis, with eight subgenera (Canis, phological, numerical, and reproductive analyses Dusicyon, Pseudalopex, Lycalopex, Cerdocyon, suggests that the taxonomy should be arranged with Atelocynus, Vulpes, and Alopex). six monotypic genera (Chrysocyon, Speothos, Nyc- INTRODUCTION In the past few years reclassifications of the the recognition of Cerdocyon and Atelocynus as family Canidae have suggested different com- valid genera. They had been included in Dusi- positions of the genera. The generic allocations cyon by Simpson (1945). Stains (1967, 1975) of the canids had been fairly stable for a num- recognized Dasycyon, a genus named by Krum- ber of years. A dozen genera and 40 or fewer biegel (1949, 1953). species were usually recognized. Lately, the South American canids have Most studies followed Simpson's (1945) ar- been studied by Langguth (1969, 1975), and the rangement of three subfamilies: Caninae, Simo- entire family was subjected to a numerical anal- cyoninae, and Otocyoninae. In the subfamily ysis by Clutton-Brock, Corbet, and Hills Caninae, Simpson had put eight genera: Canis, (1976). The conclusions of these studies are Alopex, Vulpes, Fennecus, Urocyon, Nyc- different, and in attempting to recommend tax- tereutes, Dusicyon, and Chrysocyon. His sub- onomic changes that seemed warranted on the family Simocyoninae contained the genera basis of intergeneric hybridization, I (Van Speothos, Cuon, and Lycaon. Otocyon was the Gelder, 1977) found it necessary to attempt a sole representative of the subfamily Oto- consistent arrangement of the genera that dif- cyoninae. fers from that of Langguth (1975) and Clutton- Until 1969, subsequent compilations and Brock, Corbet, and Hills (1976) or any other classifications had, in general, continued to use current classification. The present paper is an these genera, the most consistent change being analysis of these different classifications. 'Curator, Department of Mammalogy, the American Museum of Natural History. Copyright (©) The American Museum of Natural History 1978 ISSN 0003-0082 / Price $1.00 2 AMERICAN MUSEUM NOVITATES NO. 2646 ACKNOWLEDGMENTS Cerdocyon H. Smith, 1839. One species: C. thous. I am indebted to Drs. Sydney Anderson, Dusicyon H. Smith, 1839. Nine species: D. Guy Musser, and Karl F. Koopman, and es- australis, culpaeus, culpaeolus, griseus, pecially to Dr. Alfredo Langguth for comments fulvipes, gymnocercus, sechurae, vetulus, and criticism. and inca. Fennecus Desmarest, 1804. One species: F. DISCUSSION zerda. Urocyon Baird, 1858. Two species: U. cin- Other than the genera that Simpson (1945) ereoargenteus, and littoralis. included in the Caninae, there has been little Vulpes Bowdich, 1821. Ten species: V. dispute concerning the content of the genera of bengalensis, cana, chama, corsac, ferrilata, canids that Simpson had placed in the Simo- macrotis, pallida, rueppelli, velox, and vul- cyoninae and Otocyoninae. The alliance of pes. Speothos, Cuon, and Lycaon in a single sub- family had been questioned (Pocock, 1941; Additional clarification of these genera and Ellerman and Morrison-Scott, 1951; Todd, species can be made and, in fact, Stains evi- 1970), and Simpson himself (1945) admitted dently made some changes between his 1967 that it was a heterogeneous group. Clutton- and 1975 listings. Canis dingo is now generally Brock, Corbet, and Hills (1976) found that sub- regarded as a distinctive feral domestic dog. familial separations in the Canidae were not Canis familiaris is used for domestic dogs, al- warranted. They concluded that each of these though taxonomically it should probably be genera represented a monotypic, highly spe- synonymous with Canis lupus. In 1975 Stains cialized entity whose relationship to the others had dropped Dusicyon culpaeolus, evidently was distant, although perhaps closer to one an- following Langguth (1967). He did not regard other than to those genera that were usually Dusicyon fulvipes as specifically distinct from placed in the subfamily Caninae. Similarly, D. griseus. Dusicyon inca also was no longer Clutton-Brock, Corbet, and Hills did not recog- recognized by Stains (1975), again following nize the subfamily Otocyoninae for Otocyon. Langguth (1967). Stains listed only six species The distinction of each of these four genera of Dusicyon in 1975: D. australis, culpaeus, that were formerly split off from the Caninae is griseus, gymnocercus, sechurae, and vetulus. unquestioned. There seems also to be no ques- Stains (1975) listed Urocyon littoralis in his ac- tion about the recognition of Nyctereutes as a count of U. cinereoargenteus, pointing out that distinctive canid genus, although Frechkop these insular populations are regarded by some (1959) proposed that it belonged with the pro- as subspecies of U. cinereoargenteus, and by cyonids. Chrysocyon also is universally recog- others as subspecies of a separable species, U. nized as a clearcut genus. It is the other genera littoralis. within the Caninae that have received most of Stains (1967, 1975) persisted in recognizing the current attention and rearrangement that Dasycyon hagenbecki, known only from a sin- have resulted in the unequal classifications. gle skin. Cabrera (1958) and Hershkovitz (1961) For clarity in the following discussion, I have indicated that this genus and species is must define the extent of the genera that I shall most probably a domestic dog. mention. For these purposes I follow the list- For the South American canids, in addition ings of Stains (1967, 1975): to Urocyon, Chrysocyon, and Speothos, Cabrera (1958) recognized three genera: Dusi- Alopex Kaup, 1829. One species: A. lagopus. cyon, Atelocynus, and Cerdocyon. Simpson Atelocynus Cabrera, 1940. One species: A. mi- (1945) had put these in the genus Dusicyon and crotis. followed Osgood (1934) in subdividing it into Canis Linnaeus, 1758. Nine species: C. three subgenera: Dusicyon (including Pseuda- adustus, aureus, mesomelas, dingo, famil- lopex), Cerdocyon (including Atelocynus), and iaris, latrans, lupus, niger, and simensis. Lycalopex. Cabrera (1958) divided his genus 1978 VAN GELDER: CANID CLASSIFICATION 3 Dusicyon into two subgenera: Dusicyon, and At the end of 1975, the species in the genus Lycalopex. The latter contained only Dusicyon Canis included C. adustus, aureus, mesomelas, vetulus. familiaris, latrans, lupus, and niger, in the subgenus Canis; C. simensis in the subgenus Langguth (1969) arranged the South Ameri- Simenia (Ellerman, Morrison-Scott, and Hay- can canids at quite different levels. His classifi- man, 1953); C. australis in the subgenus Dusi- cation, which excluded Urocyon, had cyon; and C. culpaeus, gymnocercus, griseus; Chrysocyon, Dusicyon, and Cerdocyon as gen- and sechurae in the subgenus Pseudalopex. era. The first two were monotypic. Dusicyon In 1976 Clutton-Brock, Corbet, and Hills was divided into two subgenera, Dusicyon for published a classification based on a numerical D. australis alone, and Pseudalopex for D. analysis of the family Canidae. For the South culpaeus, gymnocercus, and griseus. His genus American canids they recognized only Chryso- Cerdocyon was divided into three subgenera: cyon and Speothos as monotypic genera, put Cerdocyon, Atelocynus, and Speothos. Urocyon in Vulpes, and included all the other Subsequently (1975), Langguth changed his South American canids in the genus Dusicyon, classification of the South American canines. more or less returning these species to the ar- He gave generic rank to the differentiated rangement (without subgenera) of Simpson in kinds, Cerdocyon, Speothos, Lycalopex, and 1945. Atelocynus, and he-placed the Patagonian can- Excluding the placement of Urocyon in Vul- ids, formerly called Dusicyon, in the genus pes by Clutton-Brock, Corbet, and Hills (1976), Canis. He recognized the subgenus Dusicyon which was a novel proposal-discussed later in for the Falkland wolf, C. australis only, and the present paper-the alternatives for the put the other species (culpaeus, gymnocercus, classification of South American canids at pres- and sechurae), in a second subgenus of Canis, ent are: Pseudalopex. Clutton-Brock, Corbet, and Hills, Langguth, 1976 1975 culpaeus valid species, placed in the genus valid species, placed in the genus Dusicyon Canis, subgenus Pseudalopex culpaeolus possibly conspecific with culpaeus, skin is culpaeus, skull is gymno- but needs study cercus; mismatch (Langguth, 1967) gymnocercus possibly conspecific with culpaeus, valid species, placed in the genus but needs study Canis, subgenus Pseudalopex inca possibly conspecific with culpaeus, skin is gymnocercus skull is culpaeus; but needs study mismatch (Langguth, 1967) griseus valid species, placed in the genus valid species, placed in the genus Dusicyon Canis, subgenus Pseudalopex fulvipes possibly conspecific with griseus, but conspecific with griseus, valid as a needs more study subspecies sechurae valid species, placed in the genus valid species, placed
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