Paleobiology, Biogeography, and Systematics of the Black-Footed Ferret, Mustela Nigripes (Audubon and Bachman), 1851

Home , Elaine Anderson, Mustelinae, Polecat, Steppe polecat

Great Basin Naturalist Memoirs

Volume 8 The Black-footed Ferret Article 3

5-1-1986

Paleobiology, biogeography, and systematics of the black-footed ferret, Mustela nigripes (Audubon and Bachman), 1851

Elaine Anderson 730 Magnolia Street, Denver, Colorado 80220

Steven C. Forrest Department of Biological Sciences, Idaho State University, Pocatello, Idaho 83209, and Biota Research and Consulting Inc., Box 2705, Jackson, Wyoming 83001

Tim W. Clark Department of Biological Sciences, Idaho State University, Pocatello, Idaho 83209, and Biota Research and Consulting Inc., Box 2705, Jackson, Wyoming 83001

Louise Richardson Department of Biological Sciences, Idaho State University, Pocatello, Idaho 83209, and Biota Research and Consulting Inc., Box 2705, Jackson, Wyoming 83001

Follow this and additional works at: https://scholarsarchive.byu.edu/gbnm

Recommended Citation Anderson, Elaine; Forrest, Steven C.; Clark, Tim W.; and Richardson, Louise (1986) "Paleobiology, biogeography, and systematics of the black-footed ferret, Mustela nigripes (Audubon and Bachman), 1851," Great Basin Naturalist Memoirs: Vol. 8 , Article 3. Available at: https://scholarsarchive.byu.edu/gbnm/vol8/iss1/3

This Article is brought to you for free and open access by the Western North American Naturalist Publications at BYU ScholarsArchive. It has been accepted for inclusion in Great Basin Naturalist Memoirs by an authorized editor of BYU ScholarsArchive. For more information, please contact [email protected], [email protected]. PALEOBIOLOGY, BIOGEOGRAPHY, AND SYSTEMATICS OF THE BLACK-FOOTED FERRET, MUSTELA NIGRIPES (AUDUBON AND BACHMAN), 1851

Elaine Anderson', Steven C. Forrest", Tim W. Clark," and Louise Richardson"

Abstract. — Extensive literature review and 48 mammal collections containing recent specimens of the endangered ferret black-footed {Mustela nigripes) are used to characterize historic distribution of the species. Specimens (n = 120) were measured from eight collections to characterize black-footed ferret morphology and variation. Twenty-one Pleistocene and Holocene faunas in North America show ferrets dating to 100,000 yr B.P. Recent specimens (n = 412) indicate close association with prairie the dog {Cynomys spp. ) and suggest ferrets may have been less rare than previously thought. At least 103 (25%) of all specimens were taken by federal predator and rodent control agents, and males outnumber females in collections 2.04:1. Average and extreme measurement for external, cranial, and postcranial dimensions are tabulated. Ferrets show a high degree of sexual dimoqihism, with discriminant analysis correctly classifying 95% of all specimens to sex. Ferrets also exhibit north-south clinal variation in size, but they do not appear to exhibit variation based on species of Cynomys associate. The taxonomic relationship among ferrets and close relatives is described.

The black-footed ferret (Mustela nigripes) is a medium-sized musteline that is listed as ^IS^IT^^ endangered throughout its former range and currently receives full protection under the U.S. Endangered Species Act of 1973 (16 use 1531 et. seq.). Endemic to North Amer- ica, black-footed ferrets formerly occupied an extensive range from the Great Plains of Canada to intermontane regions of the interior Rocky Mountains and southwestern

United States. The species is currently known from only one population restricted to an ap- pro.ximately 150 sq km area in northwestern

Wyoming (Fig. 1). Decline of the black-footed ferret over the last 50 years is attributed to the often systematic eradication of its principal prey and associate, the prairie dog {Cijnomys spp. ), which is often viewed as an agricultural pest throughout the West. Prairie dogs are semifossorial colonial rodents (Sciuridae) that offer an abundant source of prey and burrows for ferret shelter. Because black-footed ferrets are primarily nocturnal and spend much of their time un- derground, they seldom were observed in the Fig. 1. Historic range of the black-footed ferret (shaded area) compared with the current known range wild by naturalists until recent technologies, (dot). specifically the high-intensity portable spot- light, made observation possible. Few details of the species biology were known until a Dakota, was studied from 1964 to 1974. Prior small population in Mellette County, South to that time information on distribution and

'730 Magnolia Street, Denver, Colorado 80220. and Consulting Inc. Box 2705, Jackson, Wyoming ^Department of Biological Sciences, Idaho State University, Pocatello, Idaho S3209, and Biota Research , 83001.

11 12 Great Basin Naturalist Memoirs No. 8 specimens of ferrets were collected sporadi- CU— Cornell University Division of Biological Sciences, Ithaca, New York cally by commercial trappers, museum collec- DMNH— Denver Museum of Natural History, Denver, tors, or federal and state rodent and predator Colorado* control agents of the U.S. Fish and Wildlife FMNH— Field Museum of Natural History, Chicago Service (formerly the Biological Survey [BSC] HM— Hastings Museum, Hastings, Nebraska and Bureau of Sport, Fisheries, and Wildlife ISU — Iowa State University, Ames KSU — Kansas State University, Manhattan [BSFW]). Specimens are therefore few and KUMNH— University of Kansas, Lawrence* scattered among many collections. MCZ— Museum of Comparative Zoology, Harvard Uni- Records of M. nigripes specimens and sight versity, Cambridge, Massachusetts* reports have been compiled for some states, MDFWP— Montana Department of Fish, Wildlife and Parks, Bozeman* but no comprehensive record of black-footed MHM— Minnilusa Pioneer Historical Museum, Rapid exists ferret distribution based on specimens City, South Dakota other than Hall (1981). Some authors have MSU— Montana State University, Bozeman included measurements from limited sam- NDSHS— North Dakota State Historical Society Mu- seum, Bismarck ples, but no systematic analysis based on a NGFP— Nebraska Game, Fish, and Parks, Lincoln large sample has been made. The present NMC— National Museum of Natural Sciences, Ottawa, study is based on a comprehensive examina- Ontario tion and analysis of black-footed ferret re- NSCM — Northwestern State College, Alva, Oklahoma mains and literature and describes the pale- NYZ—New York Zoological Society, Bronx, New York NZP National Zoological Park, Washington, D.C. obiology, distribution, and skeletal mor- — OSLf —Oklahoma State University, Stillwater phometry of M. nigripes. OU— University of Oklahoma, Norman PAT— Patuxent Wildlife Research Center, Laurel, Maryland Materials and Methods ROM—Royal Ontario Museum, Toronto SDNHM — San Diego Natural History Museum, San Sixty-eight collections mammal were con- Diego, California tacted and 48 of them reported having M. SNMH— Saskatchewan Museum of Natural History, nigripes in their collections. Of these, eight Regina collections were examined and measured. SYR— State University of New York, Syracuse SZCM — State Zoological Collection, Munich, German Collection data were supplemented by a thor- Federal Republic ough hterature review. Evidence of ferrets UCB— University of California, Berkeley was confirmed either by the presence in mu- UCM— University of Colorado Museum, Boulder* seums of specimens (skins, skeletal material) UMMZ— University of Michigan Museum of Zoology, Ann Arbor of M. nigripes or by observations of ferrets in UMMNH—James Ford Bell Museum of Natural His- hand reported in the literature biologists by tory, University of Minnesota, Minneapolis familiar with the species. Some literature re- UND— University of North Dakota, Grand Forks ports, therefore, include live-captured or UNSM— University of Nebraska State Museum, Lincoln University Department of Zool- killed animals that were not collected or pre- USD— of South Dakota, ogy, Vermillion served as museum specimens. Sight reports USNM — United States National Museum, Washington, or secondary sources, however authentic, DC* were not included. UW— University of Wyoming, Laramie. Collections containing black-footed ferrets UWZM — University of Wisconsin Zoological Museum, Madison are listed below. Asterisks denote collections WGF—Wyoming Game and Fish Department, from which specimens were measured. Cheyenne W^ H. Over Museum, University of South Da- AMNH—American Museum of Natural History, New WHO— York* kota, Vermillion YPM — Peabody Museum, Yale University, New Haven, ANSP— Academy of Natural Sciences, Philadelphia Connecticut AUG— Augustana College, Sioux Falls, South Dakota ZSP Zoological Society of Philadelphia BMS— Buffalo Museum of Science, Buffalo, New York — BNP— Badlands National Park, Interior, South Dakota BSC— Biological Services Collection, Fort Collins, Colo- Record localities are listed in Table 6 as they rado* appeared on specimen labels or in the litera- CDOW— Colorado Division of Wildlife, Denver ture, with any comments or clarifying notes CMNH— Carnegie Mu.seum of Natural History, Pitts- burg included in the text or remarks. Specimen CSU—Colorado State University, Fort C'ollins label data were organized by collection date 1986 Anderson et al.: Biogeography and Systematics 13

LM'

LM' LC-M2

LM|tr-^LM|

WM,tr i,\ WM|tr pt- LM |tr yM

Wl3-l3

WC-C

Wp4.p4 Wp4-p4

POP POP

Fig. 2. Skull and mandible of black-footed ferret (Ad. S , Baca County, Colorado. DMNH 2248) showing measurements taken. A, Lateral view of skull. B, Lateral view of mandible. C, Occlusal view of ?*. D, Occlusal view of M'. E, Occlusal view of Mj. F, Dorsal view of skull. G, Ventral view of skull. For symbols see Materials and Methods. and state or province of collection. Localities skulls, (72 of known sex), 17 skeletons, and 55 (where known) were plotted on maps using fossil (Pleistocene to Holocene) specimens. In dark circles for precise locations and open addition, 19 skulls and one skeleton of the circles where location was known only to Siberian polecat (M. eversmanni) , a possible county. Asiatic conspecific of the black-footed ferret, Specimens with one exception were mea- were also measured. Data on external measured by E.A. These included 120 recent surements were taken directly from skin tags 14 Great Basin Naturalist Memoirs No. 8 and were supplemented by field measure- 21. Breadth of protocone of P'* (WP*pc). Greatest transverse width from the buccal border of the tooth to ments of live-caught known adult ferrets from the edge of the protocone. at the extant population Meeteetse, Wyo- 22. Upper molar breadth (WM'). Greatest transverse ming, from 1982 to 1984. width M'. Measurements of skeletal material were 23. Upper molar length (LM^nner). Greatest anterior- made with vernier calipers to the nearest 1/10 posterior length of the inner lobe of M . 24. Length of M, (LMj). Greatest anterior-posterior mm. Material was separated by state, species length of Ml measured on the lingual side. in locality (from of prairie dog the collected 2.5. Trignoid length of Mi (LMitr). From the posterior the literature), and sex, if known. Figure 2 edge of the protoconid to the anterior edge of the shows points between which cranial measure- tooth. 26, 27. Breadths of Mi (WMi tr, WMi tal). Greatest width ments were taken (after Anderson 1970). Cra- of the trigonid measured across the protoconid- nial included: measurements taken metaconid; greatest width of the talonid measured across the hypoconid-entoconid. 1. Condylobasal length (CBL). The least distance from a 28. Palatal breadth at canines (PB C-C). Width of palate line connecting the posteriormost parts of the oc- between canines. cipital condyles to the anteriormost parts of the 29. Basioccipital breadth (B OB). Breadth of basioccipital premaxillae. taken at midpoint between bullae. 2. Basilar length of Hensel (BL). Least distance from a line connecting the anterior border of the foramen Postcranial measurements taken included: magnum to the posterior margin of the first upper incisors. Humerus 3. Rostral breadth (WC-C). Width across the rostrum Total length (TL). Greatest distance from the greater above the canines. tuberosity to the medial epicondyle. 4. Bimolar breadth (WP^-P^). Greatest width across the Proximal breadth (PB). Greatest width across the hind cheek teeth measured at the posterior margin greater and lesser tuberosities.

of P^ and the anterior margin of M . Least shaft breadth (LSB). Least diameter of shaft. 5. Interorbital breadth (INB). Least distance across the Distal breadth (DB). Greatest width across the medial frontal bones at the fronto-maxillary suture. and lateral epicondyles. 6. Postorbital breadth (POP). Greatest width across the Ulna postorbital processes. Total length (TL). Greatest distance from the top of the 7. Postorbital constriction breadth (POC). Least width olecranon to the styloid process. across the frontal bones behind the postorbital Breadth olecranon process (B 01 Pr). Maximum width processes. of the olecranon. 8. Mastoid breadth (MW). Greatest width across the mastoid processes perpendicular to the long axis of Radius the skull. Total length (TL). Greatest distance between the head 9. Mandible length (LJAW). Total length from the sym- and the styloid process. physis at the alveolus of Ii to the most distant edge Proximal breadth (PB). Maximum breadth across the of the condyle. head. 10. Mandible height (H). From the lower border to the tip Distal breadth (DP). Maximum width of the distal end. of the coronoid process. Femur 11. 12. Ramus depth (DP3.4, DM1.2). Depth of the jaw Total length (TL). Greatest distance from the head to between P3.4 and M1.2 measured from the level of the medial epicondyle. the alveoli to the lower border. Proximal breadth (PB). Maximum breadth between the 13. Maxillary tooth row length (LC-M'). Least distance greater trochanter and the head. from the anterior border of the canine at the alveo- Least shaft breadth (LSB). Least diameter of the shaft. lus to the posterior border of M ' at the alveolus. Distal breadth (DB). Greatest distance across the 14. Mandibular tooth row length (LC-M2). Least distance condyles. from the anterior border of the canine at the alveolus to the posterior border of Mj at the alveolus. Tibia 15. Incisor breadth (Wl^-I^). Least width from the buccal Total length (TL). Greatest distance between the lat- side of right I^ to the buccal side of left I^. eral condyle and the medial malleolus. 16. Canine length (LC). The least distance between the Proximal breadth (PB). Maximum breadth between the anterior and posterior edges of the canine at the medial and lateral condyles. level of the alveolus. Distal breadth (DB). Maximum width across the distal 17. Canine breadth (WC). Transverse width of the canine end. at the level of the alveolus. Fibula 18. 19. Premolar length (LP^ LP^). Least distance from Length (L). Total length between the lateral condyle the anterior to the posterior edges of the premo- and the lateral malleolus. lars measured on the buccal side in the plane of the tooth row. Calcaneum 20. Premolar breadth (WP^). Transverse width of P^ mea- Length (L). Total length between the calcaneal sured at the center of the cusp. atuberositv and the cuboid facet. 1986 Anderson et al.: Biogeography and Systematics 15

Astragalus Length (L). Greatest perpendicular length of the bone. Baculum Length (L). Greatest length of the bone from the proxi- M. nigripes mal end to the base of the curve. Recent Specimens were placed in tentative age classes by the following criteria: (1) juvenile: cranial sutures open, deciduous dentition present, but permanent teeth beginning to erupt, epiphyses of long bones not fused; (2) young adult: internasal, nasomaxillary, ba-

' sisphenoid, and basioccipital sutures fused ^ I I =f but not obliterated, permanent dentition fully erupted except for upper canines, teeth un- worn or only slightly worn, epiphyses of long M. nigripes bones fused, but sutures still visible; (3) adult: Pleistocene/ all cranial sutures obliterated, permanent Holocene dentition fully erupted, well-developed sagit- tal crest especially on males, epiphyseal sutures obliterated.

Statistical Methods

' ' a' I Analyses were a I I I performed on Hewlett I I 6.05 6.65 7.25 7.85 8.45 9.05 Packard 3000 computer using the Statistical 6.35 6.95 7.55 8.15 8.85 9.35 Package for the Social Sciences (SPSS), including Discriminant Analysis and One-way Fig. 3. Frequency histogram of length of Mi for Re- Analysis of Variance (ANOVA). Linear dis- cent, Pleistocene, and Holocene specimens. criminant analysis was performed between sexes using standardized measurements and Diagnosis. — Mustela nigripes is a mink-sized confined to specimens of known sex. Juveniles mustelid weighing 645-1125 g. Upper parts were ommitted from the analysis to avoid allo- yellowish buff, occasionally whitish, espe- metric variation. Ranges, means, and stan- cially on the face and venter; feet black; black dard deviations were calculated for both sexes mask across the eyes, particularly well de- of M. nigripes . Scattergrams and frequency fined in young animals; tail black tipped. Skull diagrams were used to describe relationships is relatively short and broad; mastoid process between fossil and recent material and inter- is notably angular (Hillman and Clark 1980). species comparisons. A second linear discrim- Closely resembles M. eversmanni, the steppe inant analysis was performed on standardized ferret of Eurasia. Differs from M. putorius, cranial measurements of male and female fer- the European ferret, and M. vison , the Amer- rets to identify groups based on geographic ican mink, in being light colored with black variation and subgenera of prairie dog associ- markings; the latter two species are uniformly ate. One-way analysis of variance (ANOVA) dark colored, and M. p. furo, the domestic was used to explore further variation of indi- ferret, is uniformly light colored, often al- vidual variables with regard to geographic binistic. clinal variation. Morphometry Description Data on external measurements for Recent taken directly from skin tags Mustela nigripes (Audubon & Bachman) material were and are supplemented with field measure- Putorius nigripes Audubon & Bachman 1851: 297. Type ments of live-caught juvenile and adult ferrets locality Ft. Laramie, Goshen Co., Wyoming. Meeteetse from 1982 to 1984. Average Mustela nigripes Miller 1912: 102. First use of binomial. from measurements No subspecies are recognized. ( ± S. D.) and extreme external 16 Great Basin Naturalist Memoirs No.

4.0

3.5

3.0 WM, talonid

2.5

2.0

1.8 — —

1986 Anderson et al. : Biogeography and Systematics 17

5.2 a a 5.0 DD

4.5 a DD

DDD D So

4.0 - DHHO(9 DDD LM'in %ODD m^ 3.5 [Pod & o oo ooo o OOOO' oor A OOA^OA ^OO A o o

—I —I 4.8 5.0 6.0 6.5 7.0 7.5 8.0 WM I

Fig. 5. Relationship between the length of M 'inner lobe and the width of M' for Mustela nigripes (circles), M. eversmanni (triangles), and M. vison (squares) for Recent (open symbols) and Pleistocene (solid symbols) material.

Table 1. Mandibular and dental dimensions for Pleistocene specimens of M. nigripes. For symbols used in Column 1 see Cranial Measurements under Materials and Methods. 18 Great Basin Naturalist Memoirs No.

5.5

5.0

4.5

WP^pc 4.0-

3.5

3.0 1986 Anderson ETAL.:BiOGEOGRAPHY AND Systematics 19

Table 2. Cranial and dental dimensions for M. nigripes (Recent). . .

20 Great Basin Naturalist Memoirs No. 8

I2 set back of Ij and I3. The canines are rela- those of Recent specimens. Table 3 gives tively large and slightly curved. The anterior postcranial dimensions of Recent M. nigripes premolars (P""^, P2.4) are double-rooted, rela- Compared with mink, the limb bones of fer- tively short and broad, and single-cusped. rets tend to be more rugose and show less The upper carnassial (P^) is trenchant with a curvature, but it is difficult to separate the two relatively small protocone; the width of the species when only limb bones are available.

tooth across the protocone is less than that of The baculum of M. nigripes is similar to that

mink (Fig. 22). P^ is longer than the width of of mink in having the distal end hooked M\ The upper molar (M') has the characteris- sharply backward. In young animals the prox- tic hourglass shape of the Mustelinae, but the imal end is a simple, laterally flattened base

inner lobe is not as expanded as that of mink that with age develops a collar and becomes

(Fig. 5). There is no trace of a metaconid on quite rugose. The ventral groove extends the lower carnassial (Mj) and the trigonid is more than half the length of the shaft (Burt longer than the talonid; ferrets have a nar- 1960). Eight bacula were examined and mea-

rower talonid than do mink (Fig. 4). M2 is sured; the Pleistocene specimen from Isleta relatively small, and circular in shape. Fig- Cave (Fig. 8) was identical in size and mor- ures 4, 5, and 6 show that measurements of M. phology to the Recent material. nigripes fall within the range of M. eversmanni Life history observed. In No supernumerary teeth were The black-footed ferret is a mostly noctur- a few mandibles P, (2 specimens) or M2 (4 nal, solitary carnivore. The range of the black-

specimens) was absent and the alveolus footed ferret is sympatric with that of the closed; whether the tooth had been lost and prairie dog (Cynomys) throughout North the alveolus closed during the life of the ani- America, and breeding populations of ferrets mal or whether the tooth had never erupted have only been found in association with

could not be determined. Only one specimen prairie dog colonies (Linder et al. 1972, For- (USNM 21976) had an abcessed tooth (P'). rest etal.. Black-footedferret habitat, 1985). Ruprecht (1978) noted deviations in the num- Ferrets live in the burrows made by prairie ber of teeth in M. piitorius in Poland and dogs and exploit prairie dogs as their major

Holland. With advanced age the tooth cusps food source (Sheets et al. 1972). Ferrets also become worn smooth and the canines, per- eat lagomorphs, mice (cricetids), voles (mi- haps broken earlier in life, are stubby and crotines), ground squirrels and pocket go- rounded. No studies are available on the se- phers (geomyids), birds, and insects (Hender- quence oferuption of the teeth of M. nigripes , son et al. 1969; Clark et al. 1985). nor have there been any studies on age deter- Breeding occurs in March-April. Coitus in mination by counting the annuli tooth ce- lasts from 1.5 to 3 hours, and gestation is mentum or using radiographs to determine approximately 42-45 days (Carpenter and the size of the pulp cavity of the canine. Aver- Hillman 1978). Litter sizes range 1-5 young

age and extreme dental measurements are and average 3.3-3.4/litter (Linder et al. 1972, in Table 2. shown Forrest et al.. Life history characteristics, The appendicular skeleton of the black- 1985). Juveniles first appear aboveground in

footed ferret is unspecialized and shows no late June. The sex ratio at this time is equal extreme modifications as are seen in badgers (Forrest et al.. Life history characteristics, and otters. The shafts of the limb bones are 1985). relatively straight, the proximal and distal Primary mortality sources for black-footed ends are not greatly expanded, and processes ferrets are unknown. Potential predators of are not overly developed. The calcaneum has ferrets include: badgers (Taxidea taxus), coy- a well-developed trochlear process, and the otes (Cants Uitrans), bobcats {Lynx rufus),

posterior articular surface is rounded and golden eagles (Aquila chrysaetos), great- smooth (Stains 1966). Only a few limb bones of horned owls (Bilbo virginianus), and hawks M. nigripes have been recognized in Pleis- (Henderson et al. 1969; Forrest et al. Litter tocene faunas (Little Box Elder, Jaguar, and survey, 1985). Forrest et al. (Life history Isleta caves); their measurements fall within characteristics, 1985) identified four major 1986 Anderson et al. : Biogeography and Systematics 21

Table Postcranial 3. dimensions, Recent M. nigripes. For symbols used in column 1 see Materials and Methods.

Bone 22 Great Basin Naturalist Memoirs No. 8

Table 4. Black-footed ferret remains from Pleistocene, Early Holocene, and archeological faunas and steppe ferret remains from the Pleistocene of Alaska. 1986 Anderson etal.: BiogeographyandSystematics 23

Prey species Remarks References

No Cynomys, many spp. rodents ? Id. Originally identified as M. vison, see text Getz I960, Hibbard 1970 L. Martin, pers. comm. leucurus Cynomys Grassland, warmer than today Stalker et al. 1982 No Cynomys, many spp. rodents Originally identified as M. vison Slaughter 1966 No Cynomys, many spp. rodents Age originally thought to be Sangamonian Dreeszen 1970 Corner, pers. comm. No Cynomys, rodents abundant May be "Citellus" zone in part; open prairie R. G. Corner, pers. comm. Cynomys sp. Steppe Voorhies and Corner, in press No Cynomys, many spp. rodents Articulated skull and mandible; open prairie R. G. Corner, pers. comm. Cynomys leucurus J. Burns, pers. comm. Cynomys leucurus Anderson 1968, 1974 Cynomys sp. Hager 1972 Cynomys ludovicianus ? Id. Juvenile; deciduous dentition Schultz and Howard 1935 No Cynomys, many spp. rodents Outside historic range Kurten and Anderson 1972 No Cynomys, many spp. rodents D. Walker, pers. comm. No Cynomys Outside historic range Cuilday and Adam 1967 No Cynomys Outside historic range, cool grassland C. R. Harington, pers. comm.

Cynomys gunnisoni Harris and Findley 1964

Cynomys ludovicianus Corner 1977, pers. comm.

No Cynomys, many spp. rodents Carnivore trap; outside historic range White et al. 1984

Specimen burned J. Hubbard, pers. comm. Cynomys gunnisoni W. Gillespie, ms. and pers. comm. 104 Cynomys ludovicianus 1 specimen burned, 2 found in bone cache Anderson, ms.

No Cynomys, many spp. Cool grassland. Only record of M. e. in North Anderson 1977 rodents, lagomorphs America

designation of M. ci vison. The next earliest 10 of the faunas containing ferrets. The other record of ferret may be late Illinoian/early sites did not contain prairie dogs, but various Sangamonian (Adams/Clay counties, Nebra- rodent and lagomorph species were abun- ska, exact age uncertain; the specimens from dant. At two archeological sites, Atlatl Cave the "Citellus" beds were originally thought to and Upper Plum Creek Rockshelter, burned be the same age but are now regarded as late ferret bones were found, indicating their pos- Wisconsinan in age) or Sangamonian (Medi- sible use by Paleoindians. cine Hat), about 100,000 yrs B.P. By the During the late Pleistocene the steppe fer- late Wisconsin/early Holocene (15,000-8,000 ret, M. eversmanni , ranged east to Beringia, B.P.), ferrets ranged across the Great Plains the vast unglaciated land mass that extended west to Montana (Orr Cave) and Idaho from northeastern Siberia to western Alaska. (Jaguar, Moonshiner caves) and even as far Its remains have been found in deposits near north as Yukon Territory (Old Crow). At most Fairbanks, Alaska (Anderson 1973, 1977). The sites only a few bones representing one indi- specimens, a partial skull and two mandibles, vidual have been found, but at Little Box- are characterized by large size, broad facial Elder Cave at least 40 specimens and 15 indi- region, massive postorbital processes, pro- viduals (based on left mandibles) have been nounced postorbital constriction, crowded identified. The site, in the foothills of the tooth row, and enlarged canines. Measure- Laramie Mountains, contains a large number ments exceeded those of M. eversmanni mich- of prey animals including Cynomys cf leucu- noi, the largest extant subspecies. Anderson rus (n = 77-l-). Prairie dogs have been found in (1977) described the material as a new subspe- 24 Great Basin Naturalist Memoirs No. 8 1986 Anderson et al. : Biogeography and Systematics 25

Table 5. Black-footed ferret specimens associated with ethnographically known Indian tribes.

Ethnographic 26 Great Basin Naturalist Memoirs No. 8

Table 6. Recent black-footed ferret specimen accounts by state, 1851-1984. 1986 Anderson et al. : Biogeography and Systematics 27

Meas- ured Other Citation by us Remarks

X Gunnison's prairie dogs Gunnison's prairie dogs X-mount W. S. Carlos (A. M. Alexander) Gunnison's prairie dogs O. Wright (A. M. Alexander) Gunnison's prairie dogs

X-mount Mrs. M. H. Maxwell Coues 1877 Dr. Law (through F. V. Hayden) Coues 1877

S. N. Rhoads C. K. Worthan (donated by S. N. Rhoads)

White-tailed prairie dogs C. E. Aiken C. E. Aiken Gary 1911 E. R. Warren 2,806 m elevation; Gunnison's prairie dogs E. R. Warren G. DeLong 3,126 m elevation; dead in lake (origin unk.) Warren 1910 White-tailed prairie dogs R. S. Bull Felger 1910; Warren 1910 R. S. Bull Collection, Meeker Hotel; white- tailed prairie dogs R. S. Bull Felger 1910; Warren 1910 R. S. Bull Collection, Meeker Hotel; white- tailed prairie dogs E. R. Warren C. Deardorff Standing mount

E. Sutton W. W. Davidson W. W. Davidson Standing mount J. B. Burns W. D. Holhster A. H. Burns A. H. Burns

Gunnison's prairie dogs

R. J. Niedrach

S. O. Singer S. O. Singer S. O. Singer

Gunnison's prairie dogs S. O. Singer

D. Spencer O. W. Shirley X Gunnison's prairie dogs O. W. Shirley

R. Dietrich X Road kill

Road kill A. E. Borell White-tailed prairie dogs W, Dicus White-tailed prairie dogs F. Barnes Gunnison's prairie dogs F. Barnes Gunnison's prairie dogs W. Dicus White-tailed prairie dogs W. Dicus White-tailed prairie dogs R. C. Prater X Gunnison's prairie dogs L. E. Miller X Gunnison's prairie dogs Cahalane 1954 Drowned in ditch. Cahalane 1954 Road kill Bissel 1979

X-mount L. H. Kerrick

Not listed in Ghoate et al. 1982. A. B. Baker 28 Great Basin Naturalist Memoirs No. 8

Table 6 continued.

Skel- Disposition County Sex eton Crania Skin

1885 Systematics 29 1986 Anderson et al.: Biogeography and

Meas- ured by us Remarks Other CoUecter

A. B. Baker A. B. Baker A. B. Baker A. B. Baker A. B. Baker A. B. Baker A. B. Baker A. B. Baker NZP specimen; (no accession card) from NZP Kerrick L. H. toUSNM22May

A. B. Baker A. B. Baker A. B. Baker Baker A. B. BSC Baker A. B. BSC A. B. Baker C. A. Hawkes L. W. Purington L. W. Purington NZP (no accession card), rec'd. 19FebbyUSNM NZP (no accession card), rec'd. IQFebbyUSNM NZP (no accession card), rec'd. 24 Feb by USNM NZP (no accession card), rec'd. 24 Feb by USNM NZP (no accession card), rec'd. 19 Feb by USNM NZP (no accession card). rec'd. 19 Feb by USNM

L. W. Purington A. B. Baker X X BSC W. H. Osgood

EH. Herrick (L. H. Kerrick?) 1911 NZP rec'd. 3 Apr 1909; died Nov 2

1915 X NZP rec'd. 17 Jun 1910; died 2 Jul H. Byxbe 1913 NZP rec'd. 17 Jun 1910; died 26 Nov NZP rec'd. 17 Jun 1910 Feb 1911 NZP rec'd. 17 June 1910; exchanged 2

X R. Kellogg Conrad O. X Permanent loan to MCZ 4 Jan 1949 Conard (O. Conrad?) D. Missing body mount Choate and Fleharty 1975 "lona"; X-mount

Taylor 1961 Boggess et al. 1980

1877 C. Cavilieer Coues R. Williams (donated by S. N. Rhoads) From FMNH P. Youngman, pers. comm. X-mandible L. L. Walters

Wells Parker & Skin missing Apr 1980 L. L. Walters X X L. L. Walters X ADC Reports X X X 30 1986 Anderson ETAL.iBioGEOGRAPHY AND Systematics 31

Meas- ured

Collecter i)v us Remarks 32 Great Basin Naturalist Memoirs No.

Table 6 continued.

Skel- Year Date Disposition County Site Sex eton Crania Skin 1918 Mar 22 USNM 230773 Catron Magdalena, 75 mi SW X 1925 Nov 14 YPM 1970 Bernalli( Albuquerque, 12th St. X 1929 Apr 7 ANSP 14509 Lincoln Picacho, 5 km S X X 1929 Dec 8 BSC 1210 Colfax Moreno Valley, Aqua Fria X

1930 Aug 13 KU 7146 Santa Fe Santa Fe, 13 km SW X X 1934 Oct 20 USNM 251453 McKinley Gallup X X 1937 — Unknown Cibola El Moro National Monument X

McKinley Mexican Springs

Destroyed Lovington, 17 km N

North Dakota 1986 Anderson et al.: Biogeography and Systematics 33

Meas- ured Other CoUecter Citation by us Remarks 34 1986 Anderson et al.; Biogeography and Systematics 35

Meas- ured Other by us Remarks

Linder et al H. Behrens T. Bennett 36 Great Basin Naturalist Memoirs No. 8

Table 6 continued.

Skel- Year Date Disposition County Site Sex eton Crania Skir

1905 Private Collection Baylor Seymour

1933 Dec UCM 5263 Lubbock Lubbock M 1934 Feb UCM 5287 Lubbock Lubbock 1934 Dec 21 UM 76971 Lubbock Slide, 5 km SW 1986 Anderson etal.: BiogeographyandSystematics 37

Meas- ured Other by us Remarks

D. White from ZSP. received 14 Aug 1905, died 28 Feb 1906 D. Spencer D. Spencer

junnison s prairie dogs

A. Culbertson udubon & Bachman 1851 Destoyed by 1872 (Coues 1872).

Capt. J. Gillis Coues 1877

J. Mason and C. Ruby F. Bond S. E. Piper

US Biological Survey Day & Nelson 1928 Killed 10 in predator trapping

White-tailed prairie dogs Clark 1975 Clark 1975 Clark 1975

P. Muchmore Clark 1975 Road kill; formerly in WGF collection; white- tailed prairie dogs R. W. Fautin Road kill R. W. Fautin Stolen; white-tailed prairie dogs L. Hogg X Hogg Ranch kill; white-tailed prairie dogs

J. Renner Road kill; white-tailed prairie dogs L. Richardson X White-tailed prairie dogs T. W. Clark X Starved in burrow; white-tailed prairie dogs T. W. Clark Juvenile; white-tailed prairie dogs X-carcass M. Karl Young of year; white-tailed prairie dogs X-carcass D. E. Biggins X Killed by predator; young of year; white-tailed prairie dogs X-carcass D. E. Biggins Adult, killed by predator; white-tailed prairie dogs Adult; white-tailed prairie dogs X-carcass J. Hasbrouck X-partial D. E. Biggins Partial cranium; killed by predator; juvenile; crania white-tailed prairie dogs X-mandibles V. Semonsen Killed by predator; juvenile; white-tailed prairie dogs

T. M. Campbell III Clark and Campbell 1981 X Adult; white-tailed prairie dogs Martin & Schroeder 1978 1/2 skull

J. Bridges V. Jameson Martin & Schroeder 1979 X White-tailed prairie dogs Martin & Schroeder 1979 X White-tailed prairie dogs D. Higgins Martin & Schroeder 1979 1/2 skull; white-tailed prairie dogs S. Martin Martin & Schroeder 1979 X White-tailed prairie dogs Martin & Schroeder 1979 X White-tailed prairie dogs S. Martin X White-tailed prairie dogs L. Richardson Full head (eagle); white-tailed prairie dogs T. W. Clark X Adult; white-tailed prairie dogs X Adult; white-tailed prairie dogs J. Grenier X-mandibles T. W, Clark X Subaduit; white-tailed prairie dogs dogs S. C. Forrest X Adult; white-tailed prairie L. Richardson X Subaduit; white-tailed prairie dogs dogs L. Richardson X Adult; white-tailed prairie dogs L. Lee X Adult; white-tailed prairie dogs T. Thome Trap kill; white-tailed prairie dogs T. Thome Trap kill; white-tailed prairie dogs X-mandibles T. W. Clark White-tailed prairie T. Taylor X White-tailed prairie dogs prairie dogs B. Phillips White-tailed 38 Great Basin Naturalist Memoirs No.

Table 6 continued.

Skel- Disposition Sex eton Crania Skin Saskatchewan 1986 Anderson etal.: BiogeographyandSystematics 39

Meas- ured by us Remarks

C. Pickett

H. F. Hughes H. F. Hughes

J. Prochazka -

40 Great Basin Naturalist Memoirs No.

is greatly reduced. Gilbert (1977) identified 10,843 ha of C. leucurus colonies in Rio Blanco and Moffat counties in 1977 and Bissell (1979) estimated 21,500 ha for 9 of 26 counties in C. ludovicianus range in the state in 1978.

No estimate of C. gunnisoni distribution is available. Over 247,230 ha of C. gunnisoni occupied colonies disappeared from 1945 to 1947 during epizootics of sylvatic plague (Armstrong 1972).

Kansas

Occurrence of M. nigripes in Kansas was reviewed by Choate et al. (1982). We list eight additional records, including one specimen from Decatur County and one specimen in the CU collection dated 1883 (Table 6). Addi- tional literature records include a mounted specimen from Wallace County examined by Coues (1877) supplied by L. H. Kerrick.

Fig. 10. Black-footed ferret specimens from Arizona. About 1888 another ferret from Wallace Prairie dog distribution (shaded) after Cockrum (1960). County that had resided in the National Zo- ological Park was given by Kerrick to the USNM (12299/22929). These are obviously destroyed. Eight of 10 of the most recent different specimens, but whether Kerrick was specimens (1940-1952) were collected west of associated with NZP or was the collector of the the Front Range. One mandible was found in Wallace County animals is unknown. The dis- prairie dog colony searches in Logan County position of several other animals residing at in 1977 (Bissell 1979), but, like many speci- the NZP from 1905 to 1915 is also indicated in mens found ejected from burrows by prairie Table 6. Forty-eight specimens are known for dog digging activity, it may have been under- the state (Fig. 12). ground for an undetermined length of time 18 ferrets in collected before being brought to the surface. A record Of Table 6 from 1877 for Sedgewick County listed by Armstrong to 1890, 15 were collected by A. B. Baker. Several listing (1972) was found to be a sight record only and museum labels Baker as the collecter also indicate the was col- is not listed. specimen Two specimens were found above 2800 m. lected under the auspices of the BSC, but it is not One of these (UCM 10658) was found in asso- known whether Baker was employed by ciation with C. gunnisoni in Teller County at BSC. Recent specimens include one collected 2800 m. The other specimen (UCM 10660) by hand in 1957 in Sheridan County (Taylor was found drowned in Lake Moraine, eleva- 1961) and a skull and mandible of unknown age a prairie town in tion 3125 m in El Paso County, far from any found on dog Gove prairie dog colony. This specimen and an- County in 1978 (Boggess et al. 1980). other from Grand County (DMNH 653) were Ferrets and prairie dogs historically occu- the only two specimens from Colorado not pied most of Kansas west of the Flint Hills directly associated with prairie dogs and may (Fig. 12). However, prairie dogs that occu- have represented dispersing individuals. pied an estimated 809,390 ha in Kansas in Three species of prairie dogs occur in Colo- 1903 were reduced to some 14,570 ha (98% rado: C. ludovicianus, C. leuctirus, and C. reduction) by 1973 (Choate et al. 1982).

gunnisoni. Burnett (1918) estimated that the Choate et al. (1982) feel that ". . .the outlook is three combined species occupied 5,665,720 poor that the black-footed ferret will continue

ha in the state in 1918. The area now occupied to occur in Kansas, if, indeed, any remain

by prairie dogs in the state is unknown, but it here now." Systematics 41 1986 Anderson ETAL.:BioGEOGRAPHY AND

COLORADO Armstrong (1972). from Colorado. Prairie dog distribution (shaded) after Fig. 11. Black-footed ferret specimens 42 Great Basin Naturalist Memoirs No.

Fig. 13. Black-footed ferret specimens from Montana. Prairie dog distribution (shaded) after Hall (1981).

Montana prairie dogs were reduced substantially (U.S. Specimens of the black-footed ferret from Bur. Land Mgmt. 1982). There is no estimate Montana have not been described. Forty-four of the current total area occupied by prairie specimens are known from the state (Table 6, dogs in the state. Fig. 13). Coues (1877) reported the earliest In 1984 and T. M. Campbell and SCF found specimen (now lost) from the "Milk River." two separate remains, a black-footed ferret The most recent specimen was taken in Carter skull and a mandible (MDFWP 2344a and County in 1953. Thirty two (73%) of these 2344b) on a prairie dog colony in Carter ferrets come from seven counties in the south- County, where ferrets reportedly had been eastern part of the state. An undated speci- observed in 1977 (Jobman and Anderson men (USNM 13113/21976) lists the collection 1981). From the condition of the remains and location near "Ft. Custer." Ft. Custer (in the recent occupancy history by prairie dogs

Bighorn County) was activated in 1877 and in the area, it was estimated that they were no decommissioned in 1898, so it is assumed the more than 10 years old, supporting the 1977 specimen dates from that period. sighting. Repeated searches in the area failed Prairie dogs (C. ludovicianus except for a to produce other evidence or observations of small intrusion of C. leucurus in southern living animals. Carbon County [Flath 1979]), occupy the eastern two-thirds of the state except the Nebraska three extreme northeast counties (Daniels, Black-footed ferrets from Nebraska were Roosevelt, Sheridan) north of the Missouri recorded by Fichter and Jones (1953) and River (Hall 1981). Historic distribution of Jones (1964). We list an additional six speci- prairie dogs in the Burlington Northern Rail- mens, for a total of 23 from the state (Table 6, road right-of-way showed extensive contigu- Fig. 14). The most recent specimen was a road ous areas (Flath and Clark 1986). Federal pro- kill from Dawson County in 1949. grams poisoned 2,832,860 ha of prairie dog Additional information on two specimens is and ground squirrel habitat in Montana in also available. A specimen mentioned by 1920 alone (Bell 1921). Vigorous prairie dog Coues (1877) and identified (USNM 14580) as control efforts continued on a statewide basis coming from Nebraska has no date but should until the 1950s, and in some counties areas of be about the time of the Coues report of 1877. 1986 Anderson ETAL: BioGEOGRAPHY AND Systematics 43

. 44 Great Basin Naturalist Memoirs No.

NORTH DAKOTA

Fig. 16. Black-footed ferret specimens from North Dakota. Prairie dog distribution (shaded) after Hall (1981).

McKinley County. Hubbard and Schmitt Little is known of former prairie dog (C. (1954) described the substantial role of BSC ludovicianus) distribution, although there trappers in the collection of ferret specimens were likely prairie dogs found east and north in the state. of the Missouri River. In 1920, 2,428,166 ha Cynomys ludovicianus is found in the south- were treated with poisons for prairie dogs and ern and eastern parts of the state, and C. ground squirrels in North Dakota (Bell 1921). gunnisoni is found at higher elevations in the Grondahl (1973) estimated only 2740 ha of northwest. Prairie dog area in the state de- prairie dogs remained by 1973, all west of the ". clined from an estimated 4,856,333 ha in 1919 river. Seabloom et al. (1980:) . . regard to less than 202,350 ha in 1979-1981 (Hub- sightings (of black-footed ferrets) as repre- bard and Schmitt 1984). Hubbard and senting transients rather than a viable resi-

Schmitt "assume the ferret is still a member of dent population " and cite the paucity of the state's fauna and that it could occur any- prairie dogs remaining in the southwestern " where that prairie dogs occur. part of the state.

North Dakota Oklahoma No account of ferret specimens for North Lewis and Hassein (1973) listed recent fer-

Dakota is available other than Bailey (1926). ret specimens and sightings for Oklahoma. We located nine specimens, all collected west Only four specimens are known, with one of the Missouri River (Table 6, Fig. 16). Re- additional literature reference (Table 6, Fig. cent specimens include one found in 1954 in 17). A specimen was collected in Cleveland Sioux County and a skull found in 1980 in County in 1928, and Hibbard (1934) reported southeastern Billings County. a ferret taken in Texas County in 1932. Cyno- Teddy Roosevelt described ferrets found mys ludovicianus probably occupied "mil- near his ranch in western North Dakota in the lions ' of hectares in Oklahoma at the turn of late 1800s as "that rather rare weasel-like ani- the century, including one colony 35 km long mal ... I have known one to fairly depopulate in tall grass prairie between Kingfisher Creek a prairie-dog town, it being the arch-foe of and El Reno (Lewis and Hassein 1973), but these httle rodents" (Seton 1929: 571). only 3845 ha remained in 1968 (Tyler 1968). 1986 Anderson etal.: Bioceocraphy and Systematics 45

O-i

OKLAHOMA

Fig. 17. Black-footed ferret specimens from Oklahoma. Prairie dog distribution (shaded) after Hall (1981).

Black-footed ferrets were considered extir- D. P. Stearns, ESC, captured one ferret pated in Oklahoma as of September 1980 by alive near Pine Ridge, Shannon County, on 16 the U.S. Fish and Wildlife Service, Albu- September 1923. This is undoubtedly the fer- querque, New Mexico (Jobman and Anderson ret trapped by BSC in Pine Ridge September

1981). 1923 reported by Linder et al. (1972). This

animal was sent to the NZP (11281), where it South Dakota lived until 4 November 1925. It was subse- A detailed description of ferret distribution quently catalogued into the USNM (243799). and occurrence is available for South Dakota, Linder et al. (1972) listed 43 ferrets taken by where ferrets were studied in Mellette and BSC from 1924 to 1929. Table 6 lists 8 known adjacent counties from 1964 to 1974. Hender- specimens from that period. Four specimens son et al. (1969) described ferret specimens in the SDMNH were taken in South Dakota and sight reports for South Dakota from 1889 during this period and correspond to the 3 to 1967, and additional records were dis- specimens taken in 1928 not identified by cussed in Linder et al. 1972. Table 6 includes month in the Linder et al. (1972) list and the 1 an additional 15 specimens not in those ac- specimen from 1929. Therefore we have de- counts. Ninety-nine specimens are reported, ducted them from the Linder et al. (1972) tally with 57 specimens destroyed or of unknown for those years. The remaining 4 specimens disposition (Fig. 18). from that period may also have been collected Additional notes were also made for the fol- by BSC, but insufficient data are available on lowing specimens: the collectors to verify this. Both tallies are Moon (1905) noted a "pair sold alive" (Hen- therefore included. derson et al. 1969: #6). The New York Zoolog- Rose (1973) briefly discussed the history of ical Park listed two arrivals of M. nigripes in prairie dogs in South Dakota. Tovras 24-32 October 1905, but no accession card was made km long were common in major drainages. H. to verify this transaction. A specimen that R. Wells estimated 710,935 ha of prairie dogs came from NYZP to AMNH (22894) on 1 June in the state in 1923 (Linder et al. 1972). In 1906 with no data was undoubtedly one of 1968 BSFW estimated 24,281 ha in the state, these animals. Disposition of the second ani- a reduction of 96% (Rose 1973). Linder et al. 405,000 ha mal is unknown. We therefore list AMNH (1972) presented data showing 22894 as coming from this source. were poisoned by various government agen- 46 Great Basin Natur\list Memoirs No. 8 h 1986 Anderson et al.: Biogeography and Systematics 47

Utah Only one specimen is known for Utah (Dur- rant 1952), found in 1937 south of Blanding, San Juan County (Table 6, Fig. 20). Three species of prairie dogs are found in Utah: C. leucurus, C. gunnisoni, and the endemic Utah prairie dog, C. parvidens. Cynomys parvidens is geographically disjunct and there is no evidence to suggest that M. nigripes has ever occurred with this species.

Wyoming Black-footed ferret reports from Wyoming have been discussed in Clark (1980) and Clark and Campbell (1981), including an additional

126 sight records not listed here. In all, 60 ferret remains are known from 1851 to 1984, and 24 of these come from the Meeteetse area where the known population is currently under study (Table 6, Fig. 21). Five ferrets hsted

Fig. 20. Black-footed ferret specimens from Utah. in Clark and Campbell (1981) were actually Prairie dog distribution (shaded) after Durrant (1952). from South Dakota (Garst 1954). Ferrets

WYOMING

(shaded) after T. W. Clark, personal Fig. 21 . Black-footed ferret specimens from Wyoming. Prairie dog distribution communication. 48 Great Basin Naturalist Memoirs No. ^

C. ludovicianus ^>>

SASKATCHEWAN MOHT.tU.DAK.

Fig. 22. Black-footed ferret specimens from Saskatchewan. Estimated extent of prairie dogs shown by dotted Hne.

range farther west in the state than previously Saskatchewan reported by Hall (1981). Twenty-one specimens were located in one

Ferrets occurred throughout Wyoming, ex- U.S. and four Canadian museums (Table 6). cept the mountainous northwestern corner, All of these specimens were collected in in association with C. ludovicianus in the east southern Saskatchewan with the exception of and C. leucuriis in the west. Between 1915 FMNH 8207, from Gleichen, Alberta (not and 1923, 1,120,290 ha were poisoned for mapped). Gleichen is several hundred kilo- prairie dogs (Martley 1954). An additional meters out of present prairie dog range and is 445,080 ha were poisoned fiom 1923 to 1928 also disjunct from the next closest record of in Niobrara, Weston, and Campbell comities black-footed ferret in Saskatchewan. Because only, including one colony 160 km long from we have no other evidence to support ferret Indian Creek to Campbell County line (Day occmrence or recent prairie dog occurrence and Nelson 1929). Cheyenne, Wyoming, was at that latitude at this time, we regard this built on the site of a large old colony (Day and record as spurious. It is possible the skin was Nelson 1929), where a ferret specimen was picked up in fur shipments from another loca- collected in 1877 (Coues 1877). Fragmentary tion and subse(juently sold to FMNH. records of prairie dog poisoning show that Prairie dogs were not reportcxl from Canada prairie dogs have been reduced by at least until" 1927 (Soper 1938, 1944, 1946) and then

75% since 1915 (Clark 1973). Clark et al. only in the vicinit)' of Climax and \'al Marie in (1985) estimated that about 6,000 prairie dog extreme southwestern Saskatchewan. Ferret colonies (ca 90,000 ha) still exist in Wyoming, specimens were taken from 1924 to 1937 over but most are small and contain low densities of a greater geographical area (Fig. 22). Prairie prairie dogs. dogs may have been distributed at low densi- 1986 Anderson et al.: Bkk.eocraphy and Systematics 49 ties or were expanding throughout southern beled in the MCZ collection. Five of the spec- Saskatchewan and Alberta at that time and imens in this group were collected for zoos. were not recorded in l)iological surveys. Along with the Canadian evidence, additional Ground-dwelhng rodents that might provide reports outside the range of Cynomys are ferret habitat (with the exception of Sper- specimen USNM 21965 listed "from" Licks mophilus richardsonii) are absent in the area River, Missouri, and a note by Ames (1874) of ferret specimen distribution. Woodchuck listing "P. nigripes", with no evidence, in the {Marmota monax) and Frankhn's ground fauna of Minnesota. However, these reports squirrel (S. franklinii) are typically found at are far from potential range as determined by the eastern range ofCijnomys in the continen- prairie dog distribution, and we conclude tal U.S. and are found much farther north in they are erroneously placed as originating in Canada than the known distribution of prairie these locations. In the case of the Missouri dogs (Hall 1981). Rather than imply an alter- account, for example, the specimen could nate habitat for the black-footed ferret in have been taken on the Kansas plains and Canada, the distribution of ferret specimens subsequently ascribed by the collector to his more likely suggests the former range of home location. No place name for Lick's River Cynomys. The fossil history of Cymmiys in in Missouri could be found. Alberta goes back at least one million years. At Medicine Hat, Cynomys spp. has been found Summary in Wisconsin-age deposits and C. leucunis has We list 412 specimens in Table 6. The cur- been identified in the Sangamonian and mid- rent deposition is known for 310 of them. The dle Wisconsinan faunas (Stalker et al. 1982). largest number of state records (99) and extant Cynomys leiicurus has been at found at Janu- specimens (50) are from South Dakota. (late Wisconsin, Burns personal ary Cave J. Twenty-one specimens are noted from Can- ludovicianus communication), and C. was ada. Only 6 specimens were collected outside fauna (Storer recognized in the Hand Hills of known prairie dog range, although the asso- although this identification has been 1975), ciation of some of the Canadian specimens is questioned Burns, personal communica- (J. uncertain. Of the 412 records, at least 103 tion). Cynomys has not been reported from (25%) were taken by federal predator and ro- any Pleistocene fauna in Saskatchewan. It is dent control agents. The number taken by possible that intensive agriculture in the museum collectors is unknown but probably Prairie Provinces eliminated prairie dogs in is also significant. At least 41 animals (10%) many places before they could be recorded. were captured alive and held by individuals or Prairie dogs totaled only 503 ha in 1971 and zoos. are currently found only near Val Marie (Ker- Specimens collected by year are given in win and Scheelhaase 1971). Ferrets are con- Figure 23 (n = 318). The highest collection fig- sidered extirpated in Canada by the Commit- ures date from the 1920s. This peak corre- tee on the Status of Endangered Wildlife in sponds to the period in which the BSFW was Canada, 1978 (Thornback and Jenkins 1982). entering numerous agreements with state ex- tension services in the West to control prairie Additional reports dogs and carrying out large-scale poisoning Sixteen additional specimens are catalogued campaigns (Day and Nelson 1929, Linder et in museums with little or no identifying data al. 1972, Hubbard and Schmitt 1984). Be- (Table 6). Some of these may be the speci- cause ferrets never have been of economic mens that are "unknowns" from other loca- value, many specimens that were taken up to tions. Some dates of acquisition can be this time probably were destroyed and never guessed from catalogue numbers, but this is reported. Elsewhere, changing land use sig- not reliable. nificantly reduced potential ferret habitat and MCZ 14947, labeled as received in 1862, contributed to ferret decline. In several eastern prairie states (Kansas, Nebraska, Okla- was collected by F. J. Thompson, who was the collector of record for Abilene, Taylor homa, Texas), 65% of all specimens collected County, Texas, in 1882 (Coues 1882). This date prior to 1910. The early demise of ferrets attrib- specimen was lost and may have been misla- in these states is probably directly 50 Great Basin Naturalist Memoirs No.

1880 1910 1920 1930 1940 1950 I960 1970 1980

UJ 1986 Anderson etal.: Bioceocraphy andSystematics 51

20 -

AUG SEP OCT NOV DEC

Fig. 24. Collection of black-footed ferret specimens (n = 234) by month based on all records (1851-1984).

telids (King 1975). The 200 ferret specimens of known sex in Table 3 (137 males and 67 Males females) show a sex ratio of 2.04M:1F. Since sex ratios at birth are 1:1 (Forrest et al., Life

history characteristics , 1985), it seems likely

that this collection bias is similar to trap biases

seen for other mustelids, and is not a result of a skewed adult sex ratio. Trap biases in mustelids are a result of males having larger activity areas and longer movements (and therefore more encounters with traps or haz- ards) and being less trap-shy (King 1975, Pow- ell 1979). Black-footed ferret males have

larger activity areas (Biggins et al. 1985,

Richardson et al. in preparation), which further supports this theory.

MORPHOMETRIC VARIATION Sexual Dimorphism

Adult females averaged 93% of male body length for both museum- and field-measured groups and were 68% of males in body weight. Skull length for females averages 93% of that of males based on CBL. Five variables were chosen by stepwise maximizing of Wilks' lambda for cranial measurements as the best discriminators of sex: CBL, LC, POC, INB, and WM'. The results of the cranial discriminant analysis produced excellent discrimination between classes (Fig. 25). Coefficients for known specimens not 52 Great Basin Naturalist Memoirs No. 8

^WYOMING- SOUTH DAKOTA-NEBRASKA

£ 1986 Anderson ETAL.; Biogeography and System atics 53

Table 7. Discriminant classification of male and female black-footed ferrets from four localities showing number of members from each location correctly classified. 54 Great Basin Naturalist Memoirs No.

A A Group centroids

DISCRIMINANT SCORE

Fig. 29. Histogram of discriminant scores from a discriminant analysis between M. nigripes specimens taken from black-tailed prairie dog range (dark shading) and white-tailed prairie dog range (light shading).

variables (INB, WBC, WMjtr, LC-M\ WC, nignpes and WMjtal) were chosen by stepwise maxi- eversmonni mizing of Wilk's lambda, which discriminated between white-tailed and black-tailed prey putorius groups, only 53.3% of the white-tailed group frenata were placed correctly in that category, indicating a erminea high degree of overlap between groups (Fig. 29). This analysis suggests that no nivalis morphometric variation in black-footed ferrets vison occurs based on the species of prairie dog they are found to associate with. However, other differences may exist that involve eco- Fig. 30. Single linkage dendrogram using generalized logical or behavioral characteristics that could distances between species centroids based on a consensus taxonomically separate these groups but are ofMustela males and females (after Youngman 1982). not reflected in morphometric analyses.

Ferrets and Their Relatives ferent densities, have different behavior patterns, and are geographically separated, it The genus Miistela includes weasels (sub- might be expected that ferret differentiation genus Mustela), mink (subgenera Lutreola may have evolved with each subgenus of and Vison, see Youngman 1982), ferrets and prairie dog, which could be reflected in mor- polecats, (subgenus Putorius, European phological differences. For example, prairie workers often use Putorius as a generic name), dogs show similar external sizes and dimen- and South American weasels (subgenus sions among species (Hall 1981) but may differ Grammogale). "Ferret" and "polecat" are along similar latitudinal gradients. Size differ- interchangeable common names, though ences could be reflected in the size of burrow polecat is generally used for the Old World openings used and weight of the animal, species. Based on single linkage dendrograms which could in turn affect the size or confor- derived from morphometric variables, mation of ferrets found with them. Youngman (1982) suggested that the polecats

Because of north-south clinal variation and M. putorius, M. everstiuinni , and M. ni- sexual variation among ferrets, comparisons gripes, form a natural group distinct from the were made only between male ferrets from weasels and M. vison. Figurt^ 30 shows the the range of black-tailed prairie dog (n=^5()) phylogenetic relationships of some of the spe- and ferrets from the range of white-tailed cies in this group. These highly efficient small prairie dog (n = 15) subgenera from the two carnivores range in size from the tiny least

regions closest to the geographic center of weasel (M. nivalis rixosa , wt 38-63 gm), the ferret range (Wyoming-South Dakota-Ne- smallest living carnivore, to the Siberian or braska and Colorado-Kansas). Although six steppe polecat (M. eversmanni, wt to 2050 1986 Anderson et al. : Biogeography and Systematics 55

Fig. 31. Photograph.s of A/, nitiripcs (A), M. visou (B), M. eversmanni (C), and M. ptitorius furo (D).

gm). All of them have a long lithe body, short appearance, their skulls and teeth are similar legs, a long low braincase, and short powerful and can be confused. Table 9 shows some jaws equipped with elongated bladelike car- differences between them. nassials (P , Mi), sharp canines, and three The domestic ferret (M. putorius furo) was premolars in each jaw half (two in the lower bred in captivity as early as the fourth century jaw of the South American weasel, M. B.C. for use in controHing rodents and driving africana). Primarily Holarctic in distribution, rabbits from their burrows (Nowak and Par- weasels and ferrets are terrestrial and mink adiso 1983). It is also kept as a pet. Leonardo semiaquatic. About 15 extant species are rec- da Vinci's famous painting "The Lady with a ognized. Weasel" actually depicts the domestic fer- Of these, only four, M. nigripes, M. evers- ret (Kowalski 1976). Its distribution is now manni, M. putorius, and M. vison, concern us worldwide in captivity. Coloration is gener- here. Table 8 compares the four species and ally pale yellow or whitish (often albino) with Figure 31 illustrates them. Although mink no black or dark markings. Escaped domestic and ferrets differ markedly from each other in ferrets have been mistaken for black-footed 56 Great Basin Naturalist Memoirs No. 8

Table 8. Comparisons between Mustcla nigripes, M. eversmanni, M. ptitoriits, and M 1986 Anderson etal: Bioceography and Systematics 57 ferrets (Choate et al. 1982), but they are en- Detailed studies are still lacking for M. ev- tirely different in appearance (Fig. 31). ersmanni, and so far there have not been any studies on genetic variation Polecats probably arose in Europe in the between the two

species, so the question of Af . Villafranchian (3-4 mil yrs B.P.). The earliest eversmanni and M. nigripes conspecificity known species, Stromers polecat (M. stro- remains unre- solved. meri), ranged from the late Villafranchian to the middle Pleistocene, when it was replaced Another taxonomic problem in the ferrets is by the modern species. Though smaller in the recognition of subspecies. No subspecies size, Stromers polecat was closely allied to of A/, nigripes have ever been named, and our the European polecat (A/, putorius) and was studies do not show any taxonomically signifi- probably ancestral to both the European pole- cant geographic variations between samples. cat and the steppe polecat (M. eversmanni). Two or perhaps three subspecies of A/, putori- These two polecats have been considered con- ous are recognized based on slight differences specific by some workers, but studies by Rus- in size and color. Seventeen subspecies of Af. sian mammalogists (Stroganov 1962) have eversmanni have been described, eight of shown them to be distinct, well-defined spe- them from Siberia. Strogonov (1962:370) said, cies that differ in size, coloration, and habitat. "The Siberian polecat shows more geographi- Although their ranges overlap in Hungary, cal variation than the European polecat, this Romania, and southern European Russia, being manifested in changes in fur structure they are nowhere truly sympatric, being sepa- and in dimensions of body, skull and claws." rated by different habitat preferences (Corbet Whether all of these subspecies are valid or 1966). Hybrids occur only under exceptional merely represent oversplitting is unknown. circumstances. Unlike M. nigripes, the Of the three species of polecats, M. evers- far the largest geographic range, steppe polecat is not closely associated with nmnni has by any one species of rodent and feeds on susliks extending from Hungary to far eastern Asia {Spermophilus spp.), marmots, hamsters and across the broad band of steppes, forest 50° 60° voles; in winter, pikas (Ochotona spp.) are a steppes, and semideserts between and major food source in some areas. Rodent bur- N latitude. rows, especially those of susliks, are often ex- The historic range of M. nigripes included propriated by polecats for shelter and dens, the Great Plains and mountain valleys. This though they may dig their own. Miistela ev- was a relatively homogeneous environment ersmanni is valued as an exterminator of ro- without major geographic barriers. However, dents and for its fur, which is, however, of Endler (1977) points out that there is no evi- lower quality than that of M. putorius. Al- dence that allopatry is necessary for differenti- though M. eversmanni is not considered to be ation. Gradation within a continuous range endangered, hunting the animal in Siberia is (parapatry) is very common, as is pointed out prohibited. by the north-south differentiation demon- strated for Af. nigripes in this paper. Addi- Mustela eversmanni and A/, nigripes are tional specimens from the northern and closely related, and their possible con- southern extremes of the range would proba- specificity has been noted by several workers bly demonstrate more strongly this clinal vari- (see Youngman 1982 for references). Al- ation. Whether geographic isolation, for ex- though their size and coloration are similar, ample, in South Park, Colorado (USNM and analysis shows only slight differences in 247073), would eventually have resulted in cranial and dental measurements (Figs. 20, distinct subspecies will, of course, never be 21, 22), Anderson (1977) considers them sepa- known. rate entities. Ferrets entered North America from Beringia, and then ad- That the two species are closely related cannot be Siberia, spread across doubted, but until detailed comparative and statistical vanced southward through icefree corridors studies are made on the large collections of Mustela evers- to the Great Plains. Kalela (cited in Kurten manni in Soviet institutions, these data are compared 1957) noted that between 1880 and 1940, M. with the information already compiled on Mustela ni- putorius extended its range in Finland from gripes, and behavioral and chromosomal studies are un- north to central Os- dertaken on both species, I regard them as distinct. the Karelian Isthmus 58 Great Basin Naturalist Memoirs No. 8

trobothnia and west to the Gulf of Bothnia at a tection (Schonewald-Cox et al. 1985), which is rate of 7.5 km annually or 750 kni/century. This not the case for this species.

rate is probably applicable for ferrets spreading The possibility that the steppe ferret and the across Siberia into the New World in the Pleis- black-footed ferret are representatives of a tocene, when conditions were favorable. single holarctic species exploiting similar ecological niches in the New World and Old has Discussion been suggested. This in no way diminishes the unique position the black-footed ferret Our evidence supports the contention of holds in the prairie ecosystems of this conti- others (e.g., Linder et al. 1972, Hubbard and nent. It does suggest that options that might Schmitt 1984) that black-footed ferrets were draw on M. eversmanni to assist in recovery probably common historically. We have lo- efforts for the endangered M. nigripes should cated physical remains or verified reports of be further explored. ferrets from 128 of 513 counties (25%) within the historic range o(Cynomys . A conservative Acknowledgments estimate is that 41,000,000 ha of western grasslands were occupied by prairie dogs in Many individuals and organizations made the early part of this century. Using the For- this study possible. We are particularly grate- rest et al. {Life history characteristics, 1985) ful to organizations that support our field work population density estimate of one ferret per at Meeteetse and allowed us time to complete 40-60 ha, habitat may have been available in this manuscript: the Wildlife Preservation the past to support as many as 500,000- Trust International, New York Zoological So- 1,000,000 black-footed ferrets, if this habitat cietv-Wildlife Conservation International, were fully occupied by ferrets. and'the World Wildlife Fund— U.S. In addi- Although the Canadian specimens cast tion to the many curators of collections who so some doubt on the nearly obligate association kindly responded to our inquiries, several between ferrets and prairie dogs, it is almost were exceptionally helpful: Carron Meaney, certain that alternate habitats do not provide Betsy Webb of the Denver Museum of Natu- adequate resources to support ferrets in the ral History; Phil Youngman of the National long term. If ferrets were living in habitats Museum of Natural Sciences, Ottawa; Nor- other than prairie dog colonies in Canada, man Slade of the University of Kansas;

then they should still be extant there; yet the Charles Smart of the Academy of Natural Sci- last specimen was taken in 1937, about the ences, Philadelphia; Horace F. Quick of the time remnant prairie dogs in Canada were University of Colorado; R. George Corner of being eliminated by expansion of agriculture. the University of Nebraska State Museum; D. Geographic variation in a species has impli- E. Flath of the Montana Department of Fish, cations for any recovery program involving Wildlife, and Parks; W. G. Gillespie of the reintroduction of animals into areas where University of Arizona; and David Baron of the they have been extirpated. It would not be Saskatchewan Museum of Natural History. prudent to attempt such reintroductions us- We would also like to thank the curators who ing animals that differ greatly from those that allowed FA to measure collections in their originally occurred in the reintroduction area. care. However, with black-footed ferrets there Mark Shumar of Idaho State University, seems to be little habitat-related variation, provided invaluable assistance with SPSS and reintroductions should prove successful statistical procedures. Denise Casey provided in any geographic area with any prairie dog art and technical figures. The manuscript was species serving as prey, provided sufficient criticalK' reviewed by John L. Paradiso, habitat still remains to support the ferrets and Da\id M. Armstrong, and Steven Minta. We their prey. With regard to clinal or other geo- greatl) thank all of these organizations and graphic variation, our analyses suggest that a individuals. case can be made for morphometric variation within this species, although the usefulness of LiTEiuTURE Cited this argument seems limited to the case where Ai.DOl's, S. E. 1940. Notes on a black-foott-d ferret raised

numerous populations are competing for pro- in eaptivity. J. Mammal. 21:23-26. 1986 Anderson et al.: Bioceography and Systematics 59

Alexander. A. M. 1932. Control, not extermination of Gary, M. 1911. A biological survey of Colorado. North

Cijnoinys ludovicianus arizonensis. J. Mammal. American Fauna 33:1-256. 13:302. Cheatheam, L K 1977. Density and distribution of A, E. 1874. Ames, Mammalia of Minnesota. Minnesota black-tailed prairie dogs in Texas. Texas J. Sci. Acad. Bull. 1:69. 29:33-40. Anderson, E. 1968. Fauna of the Little Box Elder Cave, Choate, J R , E. K. Bocgess, and F R Henderson. Converse County, Wyoming: the Carnivores. 1982. History and status of the black-footed ferret Univ. Colorado Stud. Ser. Earth Sci. 6:1-59. in Kansas. Trans. Kansas Acad. Sci. 85:121-132. 1970. Quaternary evolution of the genus Martes Choate, J. R.. and E. D. FLEHARTi', 1975. Synopsis of (Carnivora, Mustelidae). Acta Zool. Fennica native. Recent mammals of Ellis Co., Kansas. Oc- 130:1-132. cas. Papers Mus. Texas Tech. Univ. 37:1-80. 1973. Ferret from the Pleistocene of central Clark, T. W. 1973a. A field study of the ecology and Alaska. Mammal. 54:778-779. J. ethology of the white-tailed prairie dog (C. leucu- 1974. A survey of the late Pleistocene and rus) with a model oiCynomys evolution. Unpub- Holocene fauna of Wyoming. In M. Wilson, ed.. lished dissertation. University of Wisconsin, Applied geology and archaeology: the Holocene Madison. 215 pp. history of Wyoming. Geol. Surv. Wyoming Rept. 1973b. Prairie dogs and black-footed ferrets in Inv. 10:78-87. Wyoming. Pages 88-101 in R. L. Linder and C. 1977. Pleistocene Mustelidae (Mammalia, Car- N. Hillman, eds., Proc. Black-footed Ferret and nivora) from Fairbanks, Alaska. Bull. Mus. Comp. Prairie Dog Workshop, South Dakota State Uni- Zool. 148:1-21. versity, Brookings. In preparation. The fauna of Upper Plum Creek 1975. Some relationships between prairie dogs, Canyon rockshelter (5LA2158), Las Animas black-footed ferrets, paleo-Indians, and ethno- County, Colorado. graphically known tribes. Plains Anthrop. Armstrong, D. M. 1972. Distribution of mammals in 20:71-74. Colorado. Monogr. Univ. Kansas Museum of Nat. 1980. A listing of reports on black-footed ferrets in Hist. 3:1-415. Wyoming (1851-1977). Northwest Sci. 54:47-54.

Audubon. J J , and J Bachman. 1851. The viviparous 1986. Some guidelines for management of the quadrupeds of North America. V. G. Audubon, black-footed ferret. Great Basin Nat. Mem. 8: New York. Vol. 2. 334 pp. 160-168. Bailey, V. 1905. Biological survey of Texas. North Ameri- Clark, T. W., and T. M. Campbell. 1981. Additional can Fauna 25:1-226. black-footed ferret {Mustela nigripes) reports from 1926. Biological survey of North Dakota. North Wyoming. Great Basin Nat. 41:360-361. American Fauna 49:171-173. Clark, T W., L. Richardson, S. G Forrest. T. M Camp- 1932. Mammals of New Mexico. North American bell III, D. E. Casey, and K A Fagerstone. Fauna 53:1-412. 1985. Black-footed ferret prey base. Pages Bell, W. R. 1921. Death to the rodents. USDA Yearbook 7.1-7.14 in S. Anderson and D. Inkley, eds., 1920:421-438. Black-footed Ferret Workshop Proc, Laramie,

Biggins, D E , M. H. Schroeder, S C. Forrest, and L. Wyoming, September 18-19, 1984. Wyoming Richardson. 1985. Movements and habitat rela- Game and Fish Publ., Cheyenne. tions of radio-tagged black-footed ferrets. Pages CoCKRUM, E. L. 1960. The recent mammals of Arizona, 11.1-11.17 in S. Anderson and D. Inkley, eds.. their taxonomy and distribution. University ofAri- Black-footed Ferret Workshop Proc, Laramie, zona Press, Tucson. 276 pp. Wyoming, September 18-19, 1984. Wyoming Corbet. G. B. 1966. The terrestrial mammals of western Game and Fish Publ., Cheyenne. Europe. London, G. T. FoulisandCo. Ltd. 264 pp. BISSELL. S. J. 1979. Black-footed ferret verification and Corner, R. G. 1977. A late Pleistocene-Holocene verte- habitat inventory, June 17, 1977-September 30, brate fauna from Red Willow County, Nebraska. 1978. Unpublished report, Colorado Div. Wildl., Trans. Nebraska Acad. Sci. 4:77-93. Denver. 15 pp. CouES, E. 1874. Wanted. American Sportsman 5(9): 1, BoGGESS, E. K.. F R. Henderson, and J R. Choate. Nov. 28, 1874. 1980. A black-footed ferret from Kansas. J. Mam- mal. 61:571. 1877. Fur-bearing animals: a monograph of North of Interior. USGS Bolton, H E. 1916. Spanish exploration in the South- American Mustelidae. Dept. west, 1542-1706. Charles Scribner & Sons, New Misc. Publ. 8:149-153. Texas. Amer. Nat. York. 217 pp. 1882. The black-footed ferret in Burnett, W L 1918. Rodents of Colorado in their economic 16:1009. relation. Office of State Entom. Circ. 25:1-31 Crabb. W O.. and G W Watson. 1950. Black-footed 31:99. Burt, W H 1960. BaculaofNorth American mammals. Misc. ferret in Montana. J. Mammal. Publ. Univ. Michigan Mus. Zool. 113:1-75. D.-KY, AM.. AND A P. Nelson. 1929. Wild life conserva- control in Wyoming under the leadership Cahalane, V. H 19.54. Status of the black-footed ferret. J. tion and Mammal. 35:418-424. of the United States Biological Survey. Joint Publ. Biol. Surv., Wyoming State Game & Fish Carpenter, J W, andC N. Hillman. 1978. Husbandry, re- U.S. production and veterinary care of captive ferrets. Dept., Wyoming State Ext. Serv., Wyoming Amer. Assoc. Zoo Vet. Ann. Proc, Knoxville, Tenn. Dept. Agric, Wyoming State Hist. Res. and Publ. 1978:36-47. Div. 26 pp. 60 Great Basin Naturalist Memoirs No. 8

Dreeszen, v. H. 1970. The stratigraphic framework of Guilday, J E., and E K. Adam. 1967. Small mammal Pleistocene glacial and periglacial deposits in the remains from Jaguar Cave, Lemhi County, Idaho.

central plains. Pages 9-22 in W. Dort and J. K. Tebiwa 10:26-36. Jones, eds.. Pleistocene and Recent environments Hager, M R 1972. A late Wisconsin-Recent vertebrate of the Central Great Plains. University of Kansas fauna from Chimney Rock, Animal Trap, Larimer Press, Lawrence. County, Colorado. Contrib. Geol. 11:63-71.

Durrani, S. D. 1952. Mammals of Utah. University of Hall. E R 1981. The mammals of North America, 2d ed.

Kansas, Lawrence, Publ. Mas. Nat. Hist. 6: J. Wiley & Sons, New York. Vol. 2:601-1181. 1-549. Halloran. a. F. 1964. The mammals of Navajoland. Endler.J. A. 1977. Geographic variation, speciation, and Navajo Tribal Museum, Window Rock, Arizona. clines. Monogr. Population Biology, Princeton 23 pp.

University Press 10:1-246. H.\rris, a. H . and J S Findley. 1964. Pleistocene-Re- of the cent fauna ofthe Isleta caves, Bernalillo County, Fighter, E.. and J. K. Jones. 1953. The occurrence

black-footed ferret in Nebraska. J. Mammal. New Mexico. Amer. J. Sci. 262:114-120. 34:385-389. Hassien, F D 1976. A search for black-footed ferrets in Flath, D. L. 1979. Status of the white-tailed prairie dog the Oklahoma Panhandle and adjacent area and an in Montana. Proc. Montana Acad. Sci. 38:63-67. ecological study of black-tailed prairie dogs in Flath, D. L., andT W. Clark 1986. Historic status of Texas County, Oklahoma. Unpublished thesis, black-footed ferret habitat in Montana. Great Oklahoma State University, Stillwater. Ill pp.

Basin Nat. Mem. 8:63-71. Henderson. F R . P. F Springer, and R Adrian. 1969. Forrest, S. C, T. W. Clark, D. E. Biggins, L. Richard- The black-footed ferret in South Dakota. South son, K. A. Fagerstone, and T M Campbell 111. Dakota Dept. Game, Fish, Parks, Tech. Bull. No. 1985. Life history characteristics of the genus 4:1-37. Mustela, with special reference to the black-footed Hibbard, C. W. 1934. The occurrence oi Erethizon epix-

ferret, M. nigripes . Pages 23. 1-23. 14 in S. Ander- anthum bruneri and Mustela nigripes in Kansas. J. son and D. Inkley, eds.. Black-footed Ferret Mammal. 15:70-71. Workshop Proc, Laramie, Wyoming, September 1970. Pleistocene mammalian local faunas from 18-19, 1984. Wyoming Game and Fish Publ., the Great Plains and central lowland provinces of Cheyenne. the United States. Pages 395-433 in W. Dort and

, , envi- Forrest, S C , T W Clark, L Richardson, T M Camp- J. K. Jones, Jr. eds. Pleistocene and Recent bell III. 1985. Black-footed ferret habitat: some ronments ofthe Central Great Plains, University management and reintroduction considerations. of Kansas Press, Lawrence. Wyoming Bur. Land Mgmt. Wildl. Tech. Bull. Hillman, C. N., and T. W. Clark. 1980. Mustela ni- No. 2, Cheyenne. 49 pp. gripes. Mammalian Species 126:1-3. Forrest, S. C, L. Richardson. T. W. Clark, and T M. Hoffmann, R. S., P L. Wright, and F. E. Newby. 1969. Campbell 111. 1985. Litter survey and mark/re- The distribution of some mammals in Montana. I.

capture of black-footed ferrets at Meeteetse, Mammals other than bats. J. Mammal. 50:579- July-October 1984. Unpublished report to Wyo- 604. ming Game and Fish, Wildlife Preservation Trust, HOMOLKA, C. L. 1964. Our rarest mammal? Audubon New York Zoological Society and World Wildlife 66:244-246. Fund. 12 pp. Hoogland. J. L. 1981. The evolution of white-tailed and Fortenbery, D K 1971. Unpublished report to Div. black-tailed prairie dogs (Sciuridai': ('yiioDUjs leu- Wildl. Serv. Region II, Albuquerque, N. Me.x. curtis and C. ludovicianus). Ecology 62:252-272.

Garst. W E 1954. Black-footed ferret in South Dakota. Hubbard, J, P., and C, G. Schmitt. 1984, The black-

J. Mammal. 35:594. footed ferret in New Mexico. Unpublished report Getz. L. L 1960. Middle Pleistocene carnivores from to New Mexico Bur. Land Mgmt., Santa Fe. 118

southvyestern Kansas. J. Mammal. 41:361-365. pp. Gilbert, G. L 1977. Inventory of potential black-footed Jobman. W. G.. and M. E. Anderson. 1981. Potential ferret habitat in the White River Resource Area, present range ofthe black-footed ferret as of Janu-

Colorado. U.S. Bur. Land Mgmt. Coop. Educ. ary 1, 1981. Unpublished report, U.S. Fish and

Publ. No. 1, Craig Dist., Colo. 21 pp. Wildlife Serv., Pierre, South Dakota. 65 pp. Gillespie. W. B. In preparation. Vertebrate remains from Johnson, D R 1969. Returns ofthe American Fur Com-

Atlatl Cave, Chaco Canyon, New Mexico. pany, 1835-1839. J. Manuual. 50:836-839.

, 1964. of Gray, J. E. 1865. Revision of the genera and species of Jones. J K Jr Distribution and taxonomy mam- Mustelidae contained in the British Museum. mals of Nebraska. Universitv of Kansas Publ. Mus. Proc. oftheZool. Soc. London 1865:110. Nat. Hist. 16:1-356. Grinnell, G B 1895. The story of the Indian. D. Apple- Kkuwin, L, andC. G. Scheelhaase. 1971. Present status ton Co., New York. pp. of the black-tailed prairie dog in Saskatchewan. 1896. Range ofthe black-footed ferret. Forest and Blue Jay 29:35-37. Stream 47:84. Kinc;, C M 1975. The si'x ratio of trapped weasels Grondahl, C R 1973. Status ofthe black-tailed prairie dog (Mustela nivalis). Mammal Review 5:1-6.

and black-footed ferret in North Dakota. Pages 44-47 King, J 1955. Social behavior, social organization and tnR. L. LinderandC. N. Hillnian, eds., Proc. Black- population d\ iiamics in a black-tailed prairie dog footed Ferret and Prairie Dog Workshop, South Da- town in the iilack Hills of South Dakota. C(mtrib. kota State University, Brookings. Lab. Vert. Biol. L'niversity of Michigan 67:1-123. 1986 Anderson et al. : Biogeocrapiiy and Systematics 61

KowALSKl, K. 1976. Mammals, an outline of theriology. SCHMIDLY, D J. 1977. The mammals of Trans-Pecos Translated and published for the Smithsonian In- Texas. Texas A&M University Press, College Sta- stitution and the National Science Foundation, tion. 225 pp. Washington, D. C. 617 pp. SCIIONEWALD-COX, C. , W. M J, BaYLESS. AND J. KURTEN. B 1957. Mammal migrations, Cenozoic stratig- SCHONEWALD. 1985. Cranial morphometry of raphy and the age of Peking Man and the australo- Pacific Coast Elk (Cervus elaphus). J. Mammal. pithecines. J. Paleontol. 31:215-257. 66:63-74. KuRTEN. B., AND E. ANDERSON 1972. The sediments and SCHULTZ, C B , AND E B HOWARD 1935. The fauna of fauna of Jaguar Cave: the fauna. Tebiwa 15:21-45. Burnet Cave, Guadalupe Mountains, New Mex- Lewis. J C. 1973. Additional records of black-footed fer- ico. Proc. Acad. Nat. Sci. Philadelphia 87: rets in Oklahoma. Southwest Nat. 18:350. 273-298.

Lewis, C , and F Hassein 1973. Status of prairie dog J Seabloom.R W , M G McKenna, and R. D.Crawford. and surveys for black-footed ferrets in Oklahoma. 1980. Recent records of mammals from southwest- Pages 60-75 in R. L. Linder and C. N. Hillman, ern North Dakota. Prairie Nat. 12:119-123. eds., Proc. Black-footed Ferret and Prairie Dog Seton, E T 1929. Lives of game animals. Pages 275-298 Workshop, South Dakota State University, Brook- in Vol. 4, Pt. 1. Doubleday, Doran Co., Garden ings. City, New York.

LiNDER, R. L , R. B Dahlgren, andC N Hillman 1972. Sheets, R. G., R. L. Linder, and R B. Dahlgren. 1972. Black-footed ferret-prairie dog interrelationships. Food habits of two litters of black-footed ferrets in Pages 22-37 in Symposium on Rare and Endan- South Dakota. Amer. Midi. Nat. 87:249-251. gered Wildlife S.W. U.S., 22-23 September Slaughter, B H. 1966. The Moore Pit local fauna, Pleis- 1972, Albuquerque, New Mexico, New Mexico tocene of Texas. Paleontol. 40:78-91. Dept. Game and Fish, Santa Fe. J. SOPER, D. 1938. Discovery, habitat and distribution of Lock, R A 1973. Status of the black-footed ferret and J. the black-tailed prairie dog in western Canada. black-tailed prairie dog in Nebraska. Pages 44-47 J. Mammal. 19:290-300. in R. L. Linder and C. N. Hillman, eds., Proc. Black-footed Ferret and Prairie Dog Workshop, 1944. Further data on the black-tailed prairie dog in 25:47-48. South Dakota State University, Brookings. western Canada. J. Mammal. LOVAAS, A L. 1973. Prairie dogs and black-footed ferrets 1946. Mammals ofthe northern Great Plains along

in the National Parks. Pages 139-148 in R. L. the International Boundary in Canada. J. Mam- Linder and C. N. Hillman, eds., Proc. Black- mal. 27:127-153. footed Ferret and Prairie Dog Workshop, South Soule, M. 1980. Thresholds for survival: maintaining Dakota State University, Brookings. fitness and evolutionary potential. Pages 151-169 Conservation Martin, S. J 1983. Additional records of black-footed in M. Soule and B. Wilcox, eds.. ferrets in Wyoming. Southwest. Nat. 28:95. biology: an evolutionary ecological perspective. organized, cooperative Sinauer Associates, Sunderland, Massachusetts. Martley, H. J. 1954. History of predator and rodent control (1915-1953). Unpub- Stains, H. G. 1976. Calcanea of members ofthe Musteli- lished report, U.S. Fish Wildl. Serv., Casper, dae. Parti, Mustelinae. Bull. So. California Acad. Wyoming. 32 pp. of Sciences 75:237-248. 1912. List of North American land Miller, G S , Jr. Stalker, A MacS, C. S Churcher, and R. S. Hill. 1982. 1911. mammals in the U.S. National Museum, Ice age deposits and animals from the southwest- Bull. U.S. Nad. Mus. 79:1-455. ern part ofthe Great Plains of Canada. Misc. Rept. opening of Morgan, D. L. 1953. Jedediah Smith and the 31, Geol. Surv. Canada, Wall Chart. University of Nebraska Press, Lincoln. the West. Pleistocene prairie dog in south-cen- Storer, J. E 1975. 458 pp. tral Alberta. Blue Jay 33:247. Moon, G F 1905. What have I got? Trappers World 1:4. Stroganov, S U 1962. Carnivorous mammals of Siberia. Nelson, E. W. 1919. Annual report of Chief of Bureau of Israeli Program for Scientific Translation, Biological Survey. Pages 275-298 m USDA Ann. Jerusalem, 1969. 522 pp. Rept. Dept. Agric. for year ended June 1919. Taylor, D 1961. Notes on a recent collection of a black- NowAK. R. M., AND J. L. Paradiso. 1983. Walker's mam- footed ferret. Trans. Kansas Acad. Sci. 64:41. mals of the world. 4th ed. Johns Hopkins Press, 349-351 in Thornbach, , AND M. JENKINS. 1982. Pages Baltimore, Maryland. Vol. 2:992-993. J The lUCN Mammal Red Data Book, Part 1. Powell, R. A. 1979. Mustelid spacing patterns: variation Gresham Press, Surrey, United Kingdom. on a theme by Mustela. Zeit. Tierpsychol. 50; Forrest. T. M. 153-165. Thorne, E T, M. H Schroeder, S C. Richardson, D. E. Biggins, L. R. Progulske, D R 1969. Observations of a penned, wild- Campbell III, L Williams. 50: Hanebury, D. Belitsky, and E. S. captured black-footed ferret. J. Mammal. and care of black- 619-621. 1985. Capture, immobifization 9.1-9.8 in S. Dakota. footed ferrets for research. Pages Rose, B J 1973. History of prairie dogs in South Anderson and D. Inkley, eds.. Black-footed Fer- Pages 76-77 in R. L. Linder and C. N. Hillman, Proc, Laramie, Wyoming, Sep- eds., Proc. Black-footed Ferret and Prairie Dog ret Workshop 18-19, 1984. Wyoming Game and Fish Workshop, South Dakota State University, Brook- tember Publ., Cheyenne. ings. black-footed ferret fi-om Texas. Ruprecht, a L. 1978. Dentition variations in the com- True, F. W 1885. A mon polecat in Poland. ActaTheriol. 23:239-245. Amer. Nat. 19:720. 62 Great Basin Naturalist Memoirs No.

associates VooRHiES. , In press.. Tyler, J D 1968. Distribution and vertebrate M R and R G Corner. Small of the black-tailed prairie dog in Oklahoma. Un- mammals with boreal affinities in late Pleistocene published dissertation. University of Oklahoma, (Rancholabrean) deposits of eastern and central Norman. 85 pp. Nebraska. Trans. Tertiary-Quaternary '83. U.S. Bureau Land Management. 1982. Black-tailed Warren, E R 1910. The mammals of Colorado. G. P. prairie dog control/management in Phillips Re- Putnam and Sons, New York. 300 pp.

source Area. Kept. Bur. Land Mgt., Lewistown White. J A , H G McDonald, E Anderson, and J M. Dist., Lewistown, Montana. 63 pp. SoiSET. 1984. Lava blisters as carnivore traps. Velich, R. 1961. Notes on mammals from Nebraska and Spec. Publ. Carnegie Mus. Nat. Hist. 8:241-256. P., F. southwestern Iowa. J. Mammal. 42:92-94. Young, S. and A. Halloran, 1952. Arizona speci-

Vereshchagin, N K., and G F Baryshnikov 1984. mens of the black-footed ferret. J. Mammal. Quaternary mammalian extinctions in northern 33:251. Eurasia. Pages 483-516 in P. S. Martin and R. G. YOUNGMAN, P M 1982. Distribution and systematics of Klein, eds., Quaternary extinctions — a prehistor- the European mink, Mustela hitreola Linnaeus ic revolution. University ofArizona Press, Tucson. 1761. ActaZool. Fennica 166:1-48.