PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON ll3(l):319-333. 2000.
Status of the name Odocoileus hemionus crooki (Mammalia: Cervidae)
James R. Heffelfinger
Arizona Game and Fish Department, 555 N. Greasewood Road, Tucson, Arizona 85745, U.S.A.
Abstract. —The present name of the desert mule deer {Odocoileus hemionus crooki) is based on a specimen collected in southwestern New Mexico near the Mexican border. This specimen was originally described as a new species (Dorcelaphus crooki) of black-tailed deer, not as a mule deer, because many
of its characteristics are intermediate between mule deer and white-tailed deer. In the same publication, Dorcelaphus hemionus eremicus is described from western Sonora, Mexico, as a new subspecies of desert mule deer. A number of mammalogists believed the type specimen of crooki to be a hybrid between desert mule deer and Coues white-tailed deer (O. virginianus couesi), while
others hypothesized it represented extremes of normal variation in mule deer.
I have reassessed the type specimen of Dorcelaphus crooki and reaffirm that
it is a hybrid, invalidating the use of the crooki subspecies name. Consequently, the oldest available name for the desert mule deer is O. h. eremicus. Resumen. —El nombre actual del venado bura del desierto (Odocoileus hem- ionus crooki) se basa en un ejemplar colectado en el suroeste de Nuevo Mexico cerca de la frontera con Mexico. E. A. Mearns describio este ejemplar como una nueva especie (Dorcelaphus crooki) de venado cola negra, no como un venado bura, debido a que muchos de sus caracteres son intermedios entre los del venado cola blanca y los del venado bura. En la misma publicacion, Mearns describio tambien a Dorcelaphus hemionus eremicus del occidente de Sonora, Mexico, como una nueva subespecie de venado bura del desierto. Un gran numero de mastozoologos ha considerado que el ejemplar tipo de crooki es un hibrido entre el venado bura del desierto y el venado cola blanca de Coues (O. virginianus couesi), mientras que otros han hipotetizado que representaba ex- tremos de la variacion normal del venado bura. Despues de reevaluar el ejem- plar tipo de Dorcelaphus crooki concluyo que es en realidad un hibrido. Por lo tanto, el nombre valido mas antiguo para el venado bura del desierto es O. h. eremicus.
Desert mule deer (presently known as cific name of this taxon has been conten- Odocoileus hemionus crooki Mearns, 1897) tious due to uncertainties regarding the hy- inhabit the southwestern United States from brid status of the type specimen. West Texas through southern New Mexico and southern Arizona, and southward into Historical Review Sonora, Chihuahua, Coahuila, Zacatecas, and Durango, Mexico (Leopold 1959, Cow- Mearns (1897) described Dorcelaphus an 1961, Wallmo 1981). In addition, a small crooki as a new species of black-tailed deer herd has been translocated to Nuevo Leon, because of its similarity to the Columbian
Mexico (Morrison et al. 1992; Fig. 1). black-tailed deer (O. h. columbianus). The Since its original description, the subspe- type specimen (National Museum of Natu- 320 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
Fig. 1 . Current distribution of desert mule deer in the southwestern United States and northern Mexico. Type localities identified for a) Dorcelaphus hemionus eremicus Mearns, 1897; b) Dorcelphus crooki Mearns, 1897; and c) Odocoileus hemionus canus Merriam, 1901.
ral History [USNM] 20572/35752) was col- lers. This description was based on the lected in 1892 by E. A. Mearns on the sum- skin of the male and two sets of antlers, mit of the Dog Mountains, Hidalgo County each from different deer; one of which (formerly part of Grant Co.), New Mexico came from the Sonoyta Valley, Sonora during the survey of the boundary between (USNM 59910), the other from Black Mexico and the United States. No other Butte, Baja California (USNM 60855). deer with similar characteristics were col- Pieces of skin trimmed from the hide as it lected at or near that locality. was made into a rug comprised the only In the same publication Mearns (1897) material representing the type specimen in described the "Burro deer or desert mule the National Museum until 1902 when the deer," Dorcelaphus hemionus eremicus rug was acquired from Anita McGee based on a male (USNM 63403, the type) (Poole & Schantz 1942). collected in 1895 by W. J. McGee in the In 1901, Merriam described Odocoileus Sierra Seri, Sonora, Mexico. McGee did hemionus canus based on a male (USNM not keep the skull, but processed the hide 99361) from Sierra en Medio, Chihuahua, as a deer skin rug. Mearns (1897) de- Mexico. Merriam (1901) distinguished O. scribed the subspecies as pale gray in color h. canus from the western subspecies O. h. with short pelage, a dark dorsal stripe, pal- eremicus solely on the basis of antler con- er forehead markings, and wide, heavy ant- formation. The Sierra en Medio lies only 40 VOLUME 113, NUMBER 1 321 km southwest of the type locality for Dor- ulations of desert mule deer. Only Cowan celaphus crooki in New Mexico. (1936, 1961) and Longhurst & Chattin
In his list of big game of North America, (1941) attempted to quantify differences be- and replacing Dorcelaphus with the correct tween deer within the ranges of eremicus senior synonym Odocoileus, Seton (1898: and crooki. Cowan's (1936) interspecific 286) included O. h. eremicus as a subspe- cranial distinctions were based on only four cies of mule deer, but maintained the eremicus skulls; of the two additional er- "Crook black-tailed deer" as O. crooki. Ly- emicus specimens from Mexico that he dekker (1915) also listed this animal as a used to differentiate external characteristics, black-tailed deer, O. columbianus crooki. one was from Tiburon Island, Sonora, Following Merriam's (1901) description, O. which Cowan (1961) later considered to be h. canus was used for the desert mule deer a different subspecies (O. h. sheldoni). in West Texas, Arizona, New Mexico, and Cowan (1961) based his differentiation of northcentral Mexico (Seton 1909, Lantz eremicus on measurements of only one 1910, Lydekker 1915, Bailey 1931, Cowan male and one female specimen, which may 1936, Cahalane 1939, Dalquest 1953). have been previously described (Cowan Several mammalogists believed that 1936, Longhurst & Chattin 1941). Long- crooki was based on a hybrid between hurst & Chattin (1941) added descriptions Coues white-tailed deer {p. virginianus of pelage variations to differentiate eremi- couesi) and a desert mule deer but lacked cus, but they had only one crooki skin and known hybrids for comparison (Lydekker three skulls for comparison. Cowan's 1898, Seton 1929, Bailey 1931, O'Conner (1936:236) measurements of eremicus from 1939). Goldman & Kellogg (1939) exam- southwestern Arizona and California are ined the holotype of O. h. canus and other within the normal variation of crooki re- specimens from the Sierra en Medio, Chi- ported by Hoffmeister (1986). Hall (1981) huahua, along with the holotype of Dorce- continued to treat western Sonoran mule laphus crooki, and concluded that the type deer as a separate subspecies (O. h. eremi- of crooki was an unusual specimen of mule cus), but provided no supporting informa- deer and not a hybrid. Because the name tion. Hoffmeister (1962) found mule deer crooki antedates canus, they adopted O. h. from southern Arizona and northern Sonora crooki as the correct name for desert mule within 80 km of the type locality of O. h. deer in the north-central states of Mexico eremicus (Sierra Seri, Sonora) to be suffi- and adjacent Arizona, New Mexico, and ciently similar to warrant treating eremicus Texas. Hoffmeister (1962) also re-examined as a synonym of crooki. Hoffmeister the type of crooki and compared it with (1986), while remarking that he could not specimens of O. virginianus and O. hem- confirm that O. h. eremicus was a synonym ionus. He interpreted the specimen as sim- of O. h. crooki, still implied that western ply a mule deer with some features that Sonoran mule deer were not distinguishable were intermediate with or shared by white- from those farther east. The purpose of this tailed deer. study is to confirm hybrid status of the type Based on the type specimen from Sierra specimen for O. h. crooki and clarify sub- Seri, the range of O. h. eremicus was des- specific nomenclature for desert mule deer. ignated somewhat arbitrarily as western So- nora, southwestern Arizona, and extreme Material and Methods southeastern California (Mearns 1907:210).
Hoffmeister (1962) listed O. h. eremicus as I re-examine the type specimen of O. h. a synonym of O. h. crooki because he did crooki to compare and contrast its qualita- not consider western Sonoran mule deer tive and quantitative characters with corre- (burro deer) distinguishable from other pop- sponding features of mule deer, white-tailed 322 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON deer, and their hybrids from southern Ari- and other characteristics are available for zona, southern New Mexico, and adjacent mule deer X white-tailed deer hybrids, in- Mexico. Published data from previous com- cluding both captive produced and geneti- parisons are supplemented by measure- cally confirmed, wild-taken animals. Char- ments (in millimeters) from female O. h. acteristics of known hybrids produced in crooki (n = 12), female O. v. couesi (n = captivity provide a morphological basis for 17), the type specimen of Dorcelaphus assessing suspected hybrids between mule crooki, and a known O. h. crooki X O. v. and white-tailed deer encountered in the couesi F, hybrid. The type is an adult fe- wild. The primary features used to distin- male, thus I included only adult females guish between desert mule deer and Coues (>2 years old), as determined from tooth white-tailed deer concern the length and wear and replacement (Robinette et al. color pattern of the tail; size, position, and 1957, Severinghaus 1949), for comparison. color of hair tuft of the metatarsal gland,
Cranial measurements (Table 1 ) include the depth of the lacrimal pit, and dimensions of six used by Hoffmeister (1962, 1986) in ad- the body and cranium (Table 1). My eval- dition to depth of lacrimal fossa (deter- uation of the holotype of crooki based on mined as either shallow, deep, or "no de- these features follows. cision" by Hoffmeister 1962). All cranial Metatarsal glands. —Metatarsal glands measurements were taken with a metric dial of desert mule deer are positioned high on caliper and recorded to the nearest 0.1 mm. the metatarsus, exceed 75 mm in length, Values for paired measurements (e.g., and are circumscribed with brown hair (Ca- length of right and left nasals) are averages. ton 1877, Hoffmeister 1986). Those of
Elsewhere in this report, I use the term hy- Coues white-tailed deer are positioned be- brid to refer to only verified O. h. crooki X low the midpoint of the metatarsus, mea-
O. v. couesi Fi hybrids, unless otherwise sure 25 mm or less in length, and are noted. rimmed by white hairs (Quay 1971, Hoff- meister 1986). Unlike either parent, all Review of Characters known F, hybrids have metatarsal glands that are intermediate in length, location, and Hoffmeister (1962) reviewed character- appearance (Table 2). As affirmed by istics useful for distinguishing O. virgini- Mearns (1907) and Bailey (1931), the po- anus and O. hemionus in Arizona in his sition of the metatarsal gland in the holo- evaluation of the type of crooki. These in- type of crooki is intermediate in compari- cluded cranial and external measurements son to its location in white-tailed and mule
(including size of metatarsal gland) along deer (Fig. 2). It is nearly identical in loca- with qualitative descriptions of the type of tion, length, and appearance to metatarsal antler, lacrimal pit, color pattern of the tail, glands of F! hybrids produced in captivity and color and position of the metatarsal (Day 1980) and to wild mule X white-tailed gland. He relied heavily on size because de- deer hybrids whose status was confirmed sert mule deer are larger than Coues white- genetically (Wishart 1980). tailed deer. Nevertheless, Hoffmeister did The metatarsal glands of the type of not have adults of known hybrids available crooki (right = 34 mm, left = 42 mm) are to evaluate their size characteristics when longer than those of white-tailed deer (25 deciding on the taxonomic status of the type mm or less) and well below the range for of crooki. Although acknowledging Ni- desert mule deer (75-150 mm; Table 1). chol's (1938) success in producing hybrids Day's (1980) measurements of the metatar- in captivity, Hoffmeister (1962:52) tended sal glands of two adult hybrids born in cap- to discount the occurrence of hybrids in the tivity were 50 mm for a female and 73 mm wild. Today, however, data on dimensions for a male. Wishart (1980) reported lengths VOLUME 113, NUMBER 1 323 of glands from two wild-taken mule X couesi, but at or exceeding the upper limit white-tailed deer hybrids (status confirmed for desert mule deer (Table 1). The color by electrophoresis) as 50 mm for a female pattern of the tail of the type of crooki re- and 62 mm for a male. Halloran & Kennedy sembles the tails of captive-born hybrids, (1949) and Lang (1957:14) gave lengths of which Day (1980) described as "dark red- metatarsal glands of adult female desert dish-brown or reddish-black above with mule deer from southern New Mexico that white beneath and along the borders" (Fig. averaged 2 to 3 times the length of the 3d). Tails of some subspecies of mule deer gland in the holotype of crooki. (e.g., O. h. fuliginatus) commonly have a
Anderson et al. (1964) included the co- dark dorsal surface; however, this pattern is rona of hair (circumglandular hair tuft) in rare in southern Arizonan and New Mexi- their measurements of the metatarsal glands can populations. Of 349 desert mule deer of 431 adult female mule deer from the observed in southeastern Arizona during Sacramento and Guadalupe mountains of January 1998, no adults had dark tails re- southern New Mexico. Metatarsal gland sembling the type of crooki. However, in lengths of females from the Sacramento areas of sympatry with white-tailed deer, Mountains (identified as O. h. hemionus) two fawns seen in the company of female averaged 138 mm (range, 100-190); those mule deer each had a wide, dark tail stripe. of the Guadalupe Mountains (identified as I do not know if these were hybrids or pure 0. h. crooki), 131 mm (range, 90-190). The mule deer fawns; yet hybrid fawns are typ- shortest (90 mm) is considerably longer ically seen in the company of mule deer than the longest circumglandular hair-tuft (Wishart 1980, Kay & Boe 1992), implying measurement (70 mm) on the type of that the usual hybrid cross is between an crooki. aggressive white-tailed buck and a mule Mearns (1897, 1907) described the hairs deer doe. surrounding the metatarsal gland of the type Length of hind foot. —Hoffmeister (1962, of crooki as "sooty at the base and white 1986) allowed that the length of hind foot apically." On examination, these hairs are (400 mm) of the crooki type is more char- "sooty at the base;" however, while pale, acteristic of white-tailed deer. Based on the they are not white apically (Fig. 2c). Instead data at hand (Table 1), 400 mm is at the they are nearly the same pale color as the upper extreme of length of hind foot for remainder of the leg, which Mearns (1897, Coues white-tailed deer and at the lower ex-
1907) described as "cream-buff, except treme for desert mule deer. G. I. Day's (in where new clay colored hair is coming in litt.) measurement of the length of hind foot on the anterior border." Metatarsal glands in a captive-born hybrid is 405 mm. of hybrids produced in captivity are either Total length. —The total length of the circumscribed with white hairs (G. I. Day, type of crooki (1440 mm; Mearns 1897) in litt.) or the hairs match the brown col- and that of a female hybrid (1549 mm; G. oration of their mule deer parent (J. C. I. Day, in litt.) is within the range for desert Haigh, in litt.). mule deer, but longer than normal for Coues
Tail. —Mearns (1897:2) described the tail white-tailed deer (Table 1). The ratio of tail of crooki as "colored much as in D. col- length to total length for the type of crooki umbianus, but has a longer terminal switch; is 7.4 X which is at the upper extreme for upper side and extremity of tail all black, Coues white-tailed deer; however, the ratio lower side white medially, and naked to- in Day's female hybrid is 8.4 X and within wards the base" (Fig. 3c). He gave its ver- the normal range for desert mule deer (Ta- tebral length as 195 mm, which is longer ble 1). The range of ratios of tail length to than that of a female hybrid (184 mm; G. total length in Coues white-tailed deer is
1. Day, in litt.) and in the range of O. v. 5.6 X -7.5 X, whereas the normal range of 324 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
Table 1. —Comparison of Coues white-tailed deer (Odocoileus virginianus couesi), desert mule deer (O. hemionus crooki), and an O. virginianus couesi X O. hemionus crooki F, hybrid, and the type specimen of Dorcelaphus crooki. All measurements (in millimeters) are from adult females two years of age or older. Characteristics of the type of Dorcelaphus crooki from Mearns (1907); cranial measurements from A. L. Gardner
(in litt.). Measurements of known hybrid (UA 22358) supplemented by morphological information from G. I.
Day (in litt.). Sample sizes in parentheses.
Interpretation Character O. v. couesi F, hybrid Type of crooki O. h. crooki of crooki type
Metatarsal gland: Location Below midpoint Intermediate Intermediate At or above mid- Hybrid of shanka point of shanka Length Usually <25 a 50 34R, 42Lm 75-150a Hybrid 14-23 (4) b 102-121 (4)<-' 18 (l) d X = 132 e Length hair tuft 70 90-190 (431)' Hybrid Color hair tuft Whiter Brown or white Pale brown Buff or Brown; Hybrid never whiteag Tail:
Length 215-260 (3) h 184 195 170-228 (5) h Mule deer or Hybrid
>188 3 145-180 (8) a d c 170-230 (9) 152-191 (5) 165-229 (8) 1 127-185 (8>i Dorsal color White border, dull Dark reddish- Black White with black Hybrid cinnamon 11 blackd terminal brush h Gray, reddish- Like white- No white border, brown, grayish tailed deerk usually without brown or almost midband 3 black3
Length of hindfoot 387-390 (3) h 405 400 430-464 (5) h Hybrid <404 a 380-490 (448)'
e X = 409 406-445 (5)c d 3 332^105 (18) 406-475 (8) h Total length 1410-1450 (3) h 1549 1440 1370-1570 (5) Mule deer or 1230-1420 (18)d 1346-1549 (5) c Hybrid
1397-1702 (8) 1 1430-1582 (8)i
h Ratio of tail to total 5.6 X-6.6X (3) h 8.4 X 7.4 X 6.0 X-8.9 X (5) Hybrid d m length 5.9 X-7.5 X (9) 8.2 X-10.5 x m c 6.4 X-7.3 X 8.1 X-10.2 X (5) X = 5.7 X e X = 8.0 X (8)' X = 10.1 x e Depth of lacrimal 3.0-5.7 (4) n 5.3 5.9R, 6.3L 6.4-11.2 (ll) n Hybrid
fossa Shallow 3 *1 Deep3h m h Length of ear <172 188 190 190-193 (2) Mule deer d c 145-170 (18) 184.2-203.2 (5) 175-220 (8) 3 194.7-209.6 (8) 1 n Basilar length 205-216 (4) n 222 239-265 (7) Hybrid 190.9-216.2 (13)° 229° 230-246 (12) 3 n Length of nasals 71.3-79.8 (5) n 72.0 79.1-95.6 (8) 60.4-76.7 (13)° 80.7° 79-95 (4) h Mule deer 56.4-61.0 (3) a 62.5-81.7 (12) 3 n n Orbital width 55.3-59.7 (5) 63 65.7-79.8 (10) Hybrid 54.5-56.0 (4) a 64.2° 68.0-81.0 (12) 3 VOLUME 113, NUMBER 1 325
Table 1. —Continued.
Interpretation Character O. v. couesi Fj hybrid Type of crooki O. h. crooki of crooki type
51.5-62.2 (13)° 51.4-60.8 (2)p
n Zygomatic width 87.8-97.7 (6) n 96.3 99.1-113.7 (9) Mule deer 90-97 (4)a 102.1" 100-110 (13)a 89.4-100.4 (11)°
96.6-100.7 (2)p n Length of upper P-M 63.5-67.2 (5) 70.5 72.1-82.6(11)" Mule deer toothrow 61.1-69.6(13)° 76.4" 80-89 (4) h 63.4-66.8 (4) a 70.1-85.5 (12) a n Length of lower P-M 72.5-76.2 (3) n 80.7 80.9-100.5 (10) Mule deer toothrow 66.1-77.6 (11)° 87.2" 87-97 (4) h a 72.1-74.4 (2) a 82.4-97.4 (12) Usual topographic 1231-21541 above 1800 <1400 r White-tailed elevation (m) deer
a Hoffmeister 1986. b Quay 1971. c Halloran & Kennedy 1949. d G. I. Day, in litt. e Lang 1959.
'Anderson et al. 1964. ^Caton 1877. h Mearns 1907.
1 Cowan 1961.
1 J. C. Truett, in litt. k Nichols 1938. 'Bailey 1931. m Hoffmeister 1962. n This study.
A. L. Gardner, in litt.
p Krausman et al. 1978.
*> Anthony & Smith 1977. r Krausman 1978.
ratios in desert mule deer is 7.7X-10.5X fossa is diagnostic; it is deeper (6.4-11.2
(Mearns 1907, Hoffmeister 1962, G. I. Day, mm) and larger in desert mule deer than in in litt.). An exception is a desert mule deer Coues white-tailed deer (3.0-5.7 mm). from west of El Paso, Texas (Mearns 1907), Hoffmeister's (1962:49) "no decision" on with an unusually long tail (228 mm) and the depth of the fossa in the type specimen short total length (1307 mm) yielding a ra- of crooki was because both fossae are fe- tio of 6.0 X. nestrate. However, the floor of each fossa is Length of ear. —Total length of ear for clearly evident and the depth of the right the type specimen of O. h. crooki ( 1 90 mm) fossa measures 5.9 and the left fossa, 6.3. is within the normal range for desert mule These measurements are intermediate be- deer and is longer than that of a white-tailed tween the ranges of lacrimal-fossa depths of deer (Table 1). The length of ear for two the two species and confirms Wishart's adult F, hybrids (O. h. crooki X O. v. coue- (1980) observation for known mule X si) was 188 mm (female), and 209 mm white-tailed deer hybrids. Depth of lacrimal (male), also within the normal range for fossa in another hybrid doe measures 5.3 mule deer (G. I. Day, in litt.). (Table 1), which is near the upper range of Lacrimal fossa. —Depth of the lacrimal that for white-tailed deer. — 1 ' — «c
326 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
>> =: = __ Q o< -r t -1- 1 — —O- s fc <: -d >s g aj -0 • — °o 3 O u ^ c cs (N 3 r3 -^- sC O 3 X ON O U u U C/3 —— ^ "1 —— -5 X ^d _ 0^ CD 3 ON ON tt O On ON On ON O CD S3 3 3 a •— JZ -3 c u c/3 £t/j £ GO sfl •_ • -o C/3 5^ 3 ] c 3 Q c/3 k _u O S3 03 £ -S2 <£ •n|- — r--(N — m N m rt m ft, --—5 *> 3 •a k. CO £ _ t4— a UC/3 ~i E OJ = ^ !s 3 O E <: a U^ "5* cfl d C»3 s £ X c5 ^ ~ &5 ~ >, s 3 < X -3 ^ JJ *^ .j i: Fig. 2. Metatarsal glands of: a) desert mule deer {Odocoileus hemionus crooki [=eremicus])\ b) Coues white- tailed deer (O. virginianus couesi); c) holotype of (Dorcelaphus crooki, USNM 20572/35752); d) known F, hybrid (O. h. crooki X O. v. couesi). Photographs by J. R. Heffelfinger (a & b), L. M. Snyder (c), and G. I. Day (d). Fig. 3. Tails of: a) desert mule deer (Odocoileus hemionus crooki [ = eremicus]); b) Coues white-tailed deer (O. virginianus couesi); c) holotype of (Dorcelaphus crooki, USNM 20572/35752); d) known F, hybrid (O. h. crooki X O. v. couesi). Photographs by J. R. Heffelfinger (a & b), L. M. Snyder (c), and G. I. Day (d). 328 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Cranial measurements. —Hoffmeister's these four measurements from a male mule (1962, 1986) decision that the type of X white-tailed deer hybrid (as determined crooki was a mule deer pivoted on the use from metatarsal gland characteristics) were of skull measurements. Adult desert mule within the normal range for mule deer. Wis- deer in the region of the type locality of hart (1980) used five cranial measurements crooki are much larger and may weigh in an analysis of a male and a female wild- twice as much as Coues white-tailed deer taken hybrid (both confirmed by electro- resulting in some cranial dimensions being phoresis) in Alberta. All measurements fell interspecifically diagnostic. Hoffmeister within the normal range for mule deer with (1962:48) provided generalized interspecif- the exception of post-palatal width of the ic limits, but not actual ranges of the six female, which was within the range for measurements (basilar length of Hensel, white-tailed deer. length of nasals, orbital width, zygomatic Type locality.—Mearns (1897, 1907:190) breadth, and length of both upper and lower collected the type of Dorcelaphus crooki on molariform toothrows) he used in separat- 9 June 1892, on the summit of "Emory ing female mule and white-tailed deer. He Peak of the Dog Mountains, where I noted said all six measurements of the type spec- its range as from 1500 to 1868 meters." imen for crooki were within the lower range Mearns (1907:87) was camped at "Dog of these measurements for desert mule deer. Spring . . . [which] is about 2 kilometers . .. However, I found that orbital width and north of [Boundary] Monument No. 55 .. . basilar length were below the correspond- near the south extremity of the Dog Moun- ing ranges for this measurement for mule tains, a rugged range of which Emory Peak, deer; the remaining four measurements having an altitude of 1868 meters (6129 were within the normal range. The discrep- feet), is the highest." Dog Spring (31°21'N, ancy between Hoffmeister's (1962) mea- 108°19'W) appears on several old maps as surement of the nasals of the type of crooki "Ojo del Perro." The Dog Mountains are and the longer measurement in Table 1 known today as the Alamo Hueco Moun- credited to A. L. Gardner results from Hoff- tains. Apparently, the name Emory Peak meister's measurement equaling the shortest does not appear on any topographic map of distance from the frontal-nasal suture to the the area produced from 1881 through 1983 proximal margin of its anterior border be- (C. Kollen, pers. comra.). A Department of tween medial and lateral anterior projec- the Interior, General Land Office map dated tions. Gardner's measurement is the average 1903 shows an "Emory Sp." northwest of of the greatest distance between anterior Ojo del Perro, in the general vicinity of and posterior points of right and left nasals; Pierce Peak (31°27'N, 108°20'W). As the right nasal measures 80.0 mm and the Pierce Peak is drained to the north by Emo- left, 81.5 mm. Hoffmeister's (1986) princi- ry Canyon and 1 1 km northwest of Dog pal components analysis using 1 1 cranial Spring, it is most likely the same peak iden- measurements grouped measurements of tified by Mearns as Emory Peak. The ele- the type of crooki with those of mule deer. vation of Pierce Peak is given today as Cranial measurements of an adult captive- 1877 m; however, a map of the area dated born hybrid doe are either intermediate or 1942 has the elevation as 6149 ft (1874 m). within the normal range for mule deer (Ta- Mearns' (1907) elevation of 1868 m for ble 1). Emory Peak is nearly equivalent; further- Cowan (1962) reported four cranial di- more, Pierce Peak is covered with alligator mensions (width of nasals, interorbital juniper {Juniperus deppeana), which width, palatal width, and postpalatal width) matches his description of the area. used to differentiate mule deer and white- The elevation at which the type of crooki tailed deer in Alberta, Canada. Three of was collected, presumed to be above 1800 VOLUME 113, NUMBER 1 329 m, is above the normal upper elevational am's name O. h. canus was used for the limit for desert mule deer (1400; Krausman desert mule deer. Goldman & Kellogg 1978, McCulloch 1972). Coues white-tailed (1939), having noted that a mule deer (0. deer, however, often occur in highest den- h. peninsulae) from lower Baja California, sities between 1230 and 2150 m elevation Mexico, had a tail color pattern similar to (Anthony & Smith 1977). Both species are that of the type of crooki and reasoning that present in the Alamo Hueco Mountains, but their animal could not be a hybrid because white-tailed deer are less common today white-tailed deer were not in Baja Califor- than they were earlier in this century (Bai- nia, concluded that the type of crooki was ley 1931; Raught 1967; A. Hurt, pers. an abnormal specimen of mule deer. Hoff- comm.). meister (1962, 1986) acknowledged that the type of crooki was intermediate in some Discussion features, but believed the preponderance of evidence, particularly of size, supported his In the original description of Dorcela- assessment that the animal was simply an phus crooki, Mearns (1897:3) said "The abnormal mule deer. He also said that the skull has very nearly the same conforma- metatarsal gland may be small or indistinct tion as that of D. columbianus [black-tailed in some populations of mule deer else- deer], the lacrimal fossa being deeper than where, citing Hershkovitz's (1958:538) ob- in the Virginia deer, but shallower than in servation that two mule deer from lower the mule deer. The same intermediate con- Baja California, Mexico, had poorly-devel- dition obtains with respect to the vomer, in oped glandular tissue underlying well-de- the relationships of the nasal and premax- fined metatarsal hair tufts. Nevertheless, the illary bones, in the form and size of the type specimen of crooki has shortened teeth; and, in short, the whole animal ap- metatarsal glands and circumglandular tufts pears to be a compromise between the char- unlike those of any known mule deer, but acteristics of the white-tailed and mule consistent in size, form, and position with deer." those of known hybrids (Tables 1 & 2). In 1907, Mearns referred to this taxon as Hoffmeister (1962:52), in his statement Odocoileus crooki and explained naming "The few 'hybrids' that I have been able to the deer for General George Crook. Mearns track down either prove to be clearly O. also received a specimen of a 2-year-old hemionus or O. virginianus," implied that buck shot in the vicinity of Bill Williams hybridization between these species proba- Mountain, Arizona, in 1884 by a member bly did not occur in the wild. of General Crook's hunting party. Mearns White-tailed deer X mule deer hybrids believed this specimen also represented his are known to have been produced in cap- new species of black-tailed deer. I have not tivity as early as 1865 (Gray 1972). Other examined the male to verify its hybrid sta- examples of hybridization have been doc- tus because its identity has no bearing on umented at captive facilities in Arizona (Ni- the status of the name crooki because it is chol 1938, Day 1980), Colorado (Spraker not the type. Mearns (1907:187) reported et al. 1997), Illinois (Caton 1877), Texas the length of metatarsal gland on this spec- (Derr 1990), Wyoming (Guiroy et al. 1991, imen as 13 mm, which is typical of a white- E. S. Williams, in litt.), and Alberta, Canada tailed deer (Table 1). (Lingle 1992, W D. Wishart, in litt.). For several decades following Mearns' Whitehead (1972) reported white-tailed X (1897) description of Dorcelaphus crooki, black-tailed deer hybrids produced in cap- several authorities suggested that the type tivity in Tennessee. was a hybrid (Lydekker 1898, Seton 1929, Hybridization between white-tailed deer Bailey 1931, O'Conner 1939) and Merri- and mule deer has been documented genet- 330 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON ically or on the basis of metatarsal gland ever, many characteristics from the type morphology in the wild in Arizona (Day specimen for crooki fall outside the normal 1964, P. A. Dratch, in litt., J. A. Holcomb, range of variation for mule deer, and are in litt.), Montana (Cronin 1991), Texas consistent with those of known white-tailed (Carr et al. 1986, Stubblefield et al. 1986, X mule deer hybrids. The overwhelming Derr 1990, Ballinger et al. 1992), Washing- concordance of morphologic evidence in ton (Gavin & May 1988), Wyoming (Kay this comprehensive analysis reveals the ho- & Boe 1992), and in Alberta (Wishart lotype of Dorcelaphus crooki as a hybrid 1980) and British Columbia (Cowan 1962), between Coues white-tailed deer and desert Canada. White-tailed deer and mule deer mule deer. This, then, has serious repercus- are sympatric in the vicinity of the type lo- sions for the current scientific name for de- cality of crooki (Hoffmeister 1962; A. Hurt, sert mule deer. A scientific name based on pers. comm.), and hybrids are documented a type specimen later found to be a hybrid from adjacent areas in Arizona and Texas. is invalid and can not be used for either of The only genetic tests that will differen- the parental species even if it has priority tiate white-tailed deer and mule deer are over all other available names (ICZN 1985: electrophoresis of albumin (Scribner et al. Art. 23 [h]). This reanalysis clarifies the 1984) and erythrocyte acid phosphatase (P. long-standing confusion regarding the sta- A. Dratch, in litt.), and isoelectric focusing tus of this contentious type specimen and of muscle esterase (Oates et al. 1979). All therefore the correct scientific name for this of these analyses require fresh or frozen taxon. The oldest available name for desert samples. No molecular markers are cur- mule deer (formerly known as O. h. crooki rently known that will differentiate these and O. h. canus) is Odocoileus hemionus species from skin samples from museum eremicus Mearns, 1897; an abbreviated specimens (P. A. Dratch, in litt.). synonymy follows: Known hybrids are large and some body (length of ear and total length) and cranial Odocoileus hemionus eremicus (Mearns) measurements (zygomatic breadth, length of nasals, and upper and lower molariform Dorcelaphus crooki Mearns, 1897:2; un- toothrows) are within the normal range for available name because it is based on a hybrid. desert mule deer (G. I. Day, in litt.). The phenomenon of heterosis (hybrid vigor) in Dorcelaphus hemionus eremicus Mearns, "Sierra Seri, F, hybrids is well known in cervids (Krzy- 1897:4; type locality near California, in arid winski 1993, Tate et al. 1997). The deer the Gulf of the most farming industry has capitalized on heter- portion of Sonora, Mexico." osis by crossing the phenotypically diverse, Odocoileus hemionus canus Merriam, but presumed conspecific, red deer and wa- 1901:560; type locality "Sierra en Media, piti (Cervus elaphus). F, hybrids show Chihuahua, Mexico." higher and faster weight gains making them Odocoileus hemionus crooki: Goldman & more profitable than either purebred paren- Kellogg, 1939:507; name combination. tal stock (Pearse 1993). Variability in over- all size of F, phenotypes means that most Acknowledgments cranial and body measurements are poor choices for evaluating hybrid status in deer I am very much indebted to A. L. Gard- (Cowan 1962; Day 1980, in litt.; Wishart ner, USGS Patuxent Wildlife Research Cen- 1980). ter, Biological Survey Unit, National Mu- Any mule deer may have an abnormal seum of Natural History (formerly called tail, metatarsal gland, lacrimal fossa, cranial the United States National Museum), for measurements, or length of hind foot. How- measurements of specimens including the VOLUME 113, NUMBER 1 331 type of Dorcelaphus crooki and for tech- deer in southeastern Arizona. —Ecological Monographs 47:255-277. nical advice and significant assistance with Bailey, V. O. 1931. Mammals of New Mexico.—North L. Snyder of the Na- the manuscript. M. American Fauna 53:1-412. tional Museum of Natural History provided Ballinger, S. W., L. H. Blankenship, J. W. Bickham, & photographs of the type specimen of D. S. M. Carr. 1992. Allozyme and mitochondrial crooki. Thanks are extended to V. C. DNA analysis of a hybrid zone between white- tailed deer and mule deer (Odocoileus) in West Bleich, D. E. Brown, M. A. Cronin, P. R. Texas.—Biochemical Genetics 30:1-1 1. Krausman, R. M. Lee, and J. R. Purdue for Cahalane, V. H. 1939. Mammals of the Chiricahua their review of an earlier draft of this man- Mountains, Cochise County, Arizona.—Journal uscript. F. J. Abarca, D. E. Brown, S. Gal- of Mammalogy 20:418-440. lina, R. S. Henry, D. Humphreys, A. Mar- Carr, S. M., S. W. Ballinger, J. N. Derr, L. H. Blan- kenship, & J. W. Bickham. 1986. Mitochondrial tinez, E. Mellink, B. L. Morrison, J. Rom- DNA analysis of hybridization between sym- ero, A. Santos, F. Tapia, M. Weber, and D. patric white-tailed deer and mule deer in West L. Weybright provided important informa- Texas.—Proceedings of the National Academy tion regarding the current range of desert of Sciences 83:9576-9580. mule deer. Arizona Game and Fish Depart- Caton, J. D. 1877. The antelope and deer of America. ment Wildlife Managers collected data on Forest and Stream Publishing Company, New York, 426 pp. tail color pattern during routine deer sur- Cowan, I. M. 1936. Distribution and variation in deer veys. J. C. deVos provided assistance with (genus Odocoileus) of the Pacific Coastal Re- photographs and M. Alderson generated the gion of North America.—California Fish and map of the distribution of desert mule deer. Game 22:155-246. N. B. Carmony provided insightful com- . 1961. What and where are the mule and black-tailed deer? Pp. 334-359 in W. P. Taylor, ments regarding subspeciation. I thank D. ed., The deer of North America. Stackpole E. Brown, P. K. Devers, Y. Petryszyn, M. Company, Harrisburg, Pennsylvania, 668 pp. T. Pruss, and especially R. S. White for al- . 1962. Hybridization between the black-tail lowing access to specimens and for osteo- deer and the white-tail deer.—Journal of Mam- logical assistance. G. I. Day, P. R. Kraus- malogy 43:539-541. Cronin, M. A. 1991. Mitochondrial and nuclear genetic man, C. Y. McCulloch, and J. C. Truett relationships between deer (Odocoileus spp.) in shared morphological data from their earlier western North America.—Canadian Journal of R. work. A. Fisher, A. Hurt, C. Kollen, C. Zoology 69:1270-1279. LeQuiev, E. M. Rominger, and C. G. Dalquest, W. W. 1953. Mammals of the Mexican State Schmitt provided details on the Dog/Alamo of San Luis Potosi.—Louisiana State University Hueco mountains of southwestern New Studies, Biological Science Series l:(vi) + 233 Mexico. Thanks are also extended to E. pp. Day, G. I. 1964. An investigation of white-tailed deer Mellink for providing the Spanish version (Odocoileus virginianus couesi) forage relation- of the abstract. Production costs for this ships in the Chiricahua Mountains. Unpublished manuscript were paid by the Federal Aid in M.S. thesis, University of Arizona, Tucson, 101 Wildlife Restoration Act, W-53-M and W- pp. 78-R, of the Arizona Game and Fish De- . 1980. Characteristics and measurements of captive hybrid deer in Arizona. The South- partment. — western Naturalist 25:434-438. Derr, J. N. 1990. Genetic interactions between two spe- Literature Cited cies of North American deer, Odocoileus vir- ginianus and O. hemionus. Unpublished Ph.D. Anderson, A. E., L. G. Frary, & R. H. Stewart. 1964. dissertation, Texas A&M University, College A comparison of three morphological attributes Station, 1 1 1 pp. of mule deer from the Guadalupe and Sacra- Gavin, T A., & B. May. 1988. Taxonomic status and mento mountains, New Mexico.—Journal of genetic purity of Columbian white-tailed Mammalogy 45:48-53. deer. —Journal of Wildlife Management 52:1- Anthony, F. G., & N. S. Smith. 1977. Ecological re- 10. lationships between mule deer and white-tailed Goldman, E. A., & R. Kellogg. 1939. Status of the — 332 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON name Dorcelaphus crooki Mearns.—Journal of the family Cervidae living and extinct. Rowland Mammalogy 20:507. Ward, Limited, London, 329 pp. Gray, A. P. 1972. Mammalian hybrids: a check-list -. 1915. Catalogue of the ungulate mammals in with bibliography. Commonwealth Agricultural the British Museum. British Museum (Natural Bureaux, Farnham Royal, England, 262 pp. History) 4:1-439. Guiroy, D. C, E. S. Williams, R. Yanagihara, & D. C. McCulloch, C. Y 1972. Deer foods and brush control Gajdusek. 1991. Immunolocalization of scrapie in southern Arizona.—Journal of the Arizona amyloid (PrP27-30) in chronic wasting disease Academy of Science 7:113-119. of Rocky Mountain elk and hybrids of captive Mearns, E. A. 1897. Preliminary diagnosis of new mule deer and white-tailed deer. —Neuroscience mammals of the genera Mephitis, Dorcelaphus, Letters 126:195-198. and Dicotyles from the Mexican border of the Hall, E. R. 1981. The mammals of North America, 2nd United States. —Proceedings of the United States National edition. John Wiley and Sons, New York, 2:vi Museum 20:467-471. . 1907. Mammals of the Mexican boundary of + 601-1181 + 90 pp. the United States. Bulletin of the United Halloran, A. F, & C. A. Kennedy. 1949. External mea- — States National Museum 56:1-530. surements of the mule deer, Odocoileus h. Merriam, C. H. 1901. Seven new mammals from Mex- crooki. —Journal of Mammalogy 30:76-77. ico, including a new genus of rodents.—Pro- Hershkovitz, P. 1958. The metatarsal glands in white- ceedings of the Washington Academy of Sci- tailed deer and related forms of the Neotropical ences 3:559-563. region.—Mammalia 22:537-546. Morrison, B., B. Muller-Using, & M. Cotera. 1992. Hoffmeister, D. F 1962. The kinds of deer, Odocoileus, The translocation of mule deer in Nuevo Leon, in Arizona.—The American Midland Naturalist Mexico. Pp. 153-154 in B. Bobek, K. Perza- 67:45-64. nowski, and W. Regelin, eds., Transactions of . 1986. Mammals of Arizona. University of Ar- the 18th International Union of Game Biologists izona Press and the Arizona Game and Fish De- Congress, Krakow, 620 pp. partment, Tucson, xx + 602 pp. Nichol, A. A. 1938. Experimental feeding of deer. ICZN. 1985. International code of zoological nomen- University of Arizona Agricultural Experiment clature, 3rd edition. International Trust for Zoo- Station, Technical Bulletin 75:1-39. logical Nomenclature, London, xx + 338 pp. Oates, D. W, N. L. Dent, & K. A. Pearson. 1979. Kay, C. E., & E. Boe. 1992. Hybrids of white-tailed Differentiation of white-tailed and mule deer and mule deer in western Wyoming.—Great tissue by isoelectric focusing.—Wildlife Society Basin Naturalist 52:290-292. Bulletin 7:113-116. Krausman, P. R. 1978. Forage relationships between O'Conner, J. 1939. Game in the desert. Derrydale two deer species in Big Bend National Park, Press, New York, 167 pp. Texas.—Journal of Wildlife Management 42: Pearse, A. J. 1993. The recent status of deer farming 101-107. in New Zealand. Pp. 401-413 in N. Ohtaishi Krzywinski, A. 1993. Hybridization of milu stags with and H.-I. Sheng, eds., Deer of China. Elsevier, red deer hinds using the imprinting phenome- Amsterdam, 418 pp. non. Pp. 242-246 in N. Ohtaishi and H.-I Poole, A. J., & V. S. Shantz. 1942. Catalog of the type Sheng, eds., Deer of China. Elsevier, Amster- specimens of mammals in the United States Na- dam, 418 pp. tional Museum, including the Biological Sur- Lang, E. M. 1957. Deer of New Mexico. New Mexico veys collection. —Bulletin of the United States Department of Game and Fish, Santa Fe, 41 pp. National Museum 178:xiv + 1-705. Lantz, D. E. 1910. Raising deer and other large game Quay, W B. 1971. Geographic variation in the meta- animals in the United States. —United States tarsal "gland" of the white-tailed deer (Odo- Department of Agriculture, Biological Survey coileus virginianus).—Journal of Mammalogy Bulletin 36:1-59. 52:1-11. Leopold, A. S. 1959. Wildlife of Mexico: the game Raught, R. W 1967. White-tailed deer. Pp. 52-60 in birds and mammals. University of California New Mexico Wildlife Management. New Mex- Press, Berkeley, 568 pp. ico Department of Game and Fish, Santa Fe, Lingle, S. 1992. Escape gaits of white-tailed deer, mule 250 pp. deer and their hybrids: gaits observed and pat- Robinette, W L., D. A. Jones, G. Rodgers, & J. S. terns of limb coordination.—Behaviour 122: Gashwiler. 1957. Notes on tooth development 154-181. and wear for Rocky Mountain mule deer. Longhurst, W. M., & J. E. Chattin. 1941. The burro Journal of Wildlife Management 21:134-153. deer.—California Fish and Game 27:2-12. Scribner, K. T, R. J. Warren, & S. L. Beasom. 1984. Lydekker, R. 1898. The deer of all lands: a history of Electrophoretic identification of white-tailed — — VOLUME 113, NUMBER 1 333 and mule deer feces: a preliminary assess- Appendix 1 ment.—Journal of Wildlife Management 48: Specimens Examined 656-658. With the exception of the two type specimens of Seton, E. T. 1898. A list of big game of North Amer- mule deer (both males) listed below, specimens used ica.—Forest and Stream (October) pp. 285-286. in this analysis, including the type of Dorcelaphus . 1909. Life histories of northern animals. Chas. crooki, are all adult females (>2 years), and deposited Scribner's Sons, New York, 1:1-673. in the following institutions: Arizona State University . 1929. Lives of game animals. C. T. Branford (ASU); Arizona Game and Fish Department, Tucson Company, Boston, 412 pp. (AGFD); University of Arizona (UA); National Mu- Severinghaus, C. W. 1949. Tooth development and seum of Natural History (USNM). A. L. Gardner mea- wear as criteria of age in white-tailed deer. sured the USNM specimens; I measured all others. Journal of Wildlife Management 13:195-216. Odocoileus hemionus crooki [=eremicus] (14). Spraker, T. R., et al. 1997. Spongiform encephalopathy United States. Arizona: Yavapai Co., 8 km NE Horse- in free-ranging mule deer {Odocoileus hemion- shoe Dam (ASU 643); Maricopa Co., 3 km E Horse- us), white-tailed deer (Odocoileus virginianus) shoe Dam (ASU 637); Pinal Co., N side of Canyon and Rocky Mountain elk (Cervus elaphus nel- Lake (ASU 643), Picacho Mountains (UA 24418- soni) in northcentral Colorado. —Journal of 24420, 24429, 24430, 24436, 24478, 25299, 25308). Wildlife Diseases 33:1-6. Mexico. Chihuahua: Sierra en Medio (USNM 99361, Stubblefield, S. S., R. J. Warren, & B. R. Murphy. type of O. h. canus). Sonora: Sierra Seri (USNM 1986. Hybridization of free-ranging white-tailed 63403, type of O. h. eremicus). and mule deer in Texas.—Journal of Wildlife Odocoileus virginianus couesi (17). —United States. Management 50:688-690. Arizona: Cochise Co., Chiricahua Mountains (UA Tate, M. L., G. J. Goosen, G. Patene, A. J. Pearse, K. 20340, 20346); Graham Co., Blue River (USNM M. McEwan, & P. F. Fennessy. 1997. Genetic 32115); Pima Co., Santa Rita Mountains (AGFD 10, analysis Pere David's X red deer interspecies of UA 23304), Baboquivari Mountains (AGFD 53201); hybrids.—Journal of Heredity 88:361-365. Santa Cruz Co., Santa Rita Mountains (USNM 1-26 Wallmo, O. C. 1981. Distribution and habits. Pp. 202931). New Mexico: Catron Co., Mogollon Moun- in O. C. Wallmo, ed., Mule and black-tailed tains (USNM 148574); Grant Co., 32 km W Silver deer of North America. Wildlife Management City (USNM 286685), head of Mimbres River (USNM Institute and University of Nebraska Press, Lin- 147476); Hidalgo Co., near Cloverdale (USNM coln, 605 pp. 35748). Mexico. Chihuahua: Colonia Garcia (USNM Whitehead, Jr., C. J. 1972. A preliminary report on 99347, 99350); Sonora: San Luis Mountains (USNM white-tailed and black-tailed deer crossbreeding 36320); E side of San Luis Mountains (USNM 35751, studies in Tennessee.—Proceedings of the An- 37085); Pozo de Luis (USNM 59229). nual Conference of the Southeastern Associa- Odocoileus hemionus crooki [ = eremicus] X O. vir- tion of Game and Fish Commissions 25:65-69. ginianus couesi hybrids (2).—Arizona: Pima Co., Uni- Wishart, W. D. 1980. Hybrids of white-tailed and mule versity of Arizona Captive Facilities (UA 22358). New deer in Alberta. —Journal of Mammalogy 61: Mexico: Hidalgo Co., Summit of the Dog Mountains 716-720. (USNM 20572/35752, type of Dorcelaphus crooki).