Mamatian Species No. 37, pp. 1-6, 5 figs. Canis lupus. By 1. David Mech Published 2 May 1974 by The American Society of Mammalogists Canis lupus Linnaeus, 1758 Gray Wolf Conis lupus Linnaeus, 1758:39. Type locality, Sweden, Canis Iyenon ‘Schreber, 1775zpl, 89. Type locality Quebec, ‘Quebec, by restiction (Goldman, 195738). Conis Lupus-Criseus Sabine, in Franklin, 1825:651, not Griseus Boddaert, 1708. Canis “LupussAléus Sabine, in Franklin, 1828:655, not albus ery, 1792, Canis nubilus Soy, in Long, 1823-169, Type locality Engineer Cantonment, approximately 2 mi. N'Ft, Calhoun, Wash ington County, Nebraska, Canis pallines Sykes, 1831:101, Type locality Deccan, Tn Canis hodophilax Temminck, 1833:28% Type locality Hondo, Japan. Canis tariabilis Wied-Neuwied, 1841:95. ‘Clark, near Stanton, Merce® County, Lupus renal, Wagner, WH SGT Tope any Eavoe Tupus laniger Hodgoon, 1841-474. Type locality “Tibet,” but possibly little Tibet in Kashmir Canis chanco Gray, 1863:98. ‘Type locality, Chinese Tartar. Canis niger Sclater 1874p. 78, not niger Kere, 1792 Canis pambasileus Eliot, 1905.19. Type locality Susitna River, ‘egion of Mt. MeKiley, Alaska. Lupus fitchneri Matschic, 1908:158. Type locality Siningfo, Kansu, China. Lams aranarenss Matchie, 1908199. Tope locality Kerner, Lupus tschiliensis Matschie, 1908:160. Type locality Coast of ‘Chili, China apes atiscus Nok, (2) 191145. Type Tcaity Chlyshman lacies, Altai Canis tundarum Miller, 1912-1. ‘Alaska CONTEXT AND CONTENT. Order Carnivora, Family Canidae, The genus, Canis includes eight species. “Approx smately 24 New World and eight Old World subspecies of C. Tipus’ are zecognized, the number depending on authorities accepted. For summary, sce Mech (1970) for {ull synonymy, See Pocock (1935), Goldman (1944), Ellerman end Mortison Scott 17961), Novikow (1956), and Hall and Kelson (1959) Inpus Linnaeus, 1758:89, see above (eynonyme are flavus Kerr, niger Hermann, communis Dwigubels, orientalis Wagner, majar Ogerien, minor Ogéron, deitanus Cabrera, signatus Cabrera, lyeaon Trouessart, alticus Noak,ialicus Altobello, and. urjak.Bolkay). Type cay Fon Type locality Point Barrow, C1, fycaon Schreber, 1775:pl. 89, see above (cynonyms are ‘canadensis Blainvile and ungevensis Comeat) CL. albus Kets, 1792:137. Type locality Jenivea eastern USSR (synonyms Domaniewski) 1. carspestris Dwigubski, 1904:10. Type locality between Black Sea and Caspian’ Kirghizia, to Yenesei River (syn- ‘onyine are desertorum Bogdanoy, cubanensis Ognev). fe turacanensis Ogney, and dyboushit CL, mabilus Say, 18232108, soe above (a synonym is earibilis Wied-Newwied) C.1, occidentalis Richardson, 1829:60, ‘Type locality Simpson, Mackensie, Canada (synonyms a Richardson) 1. pallipes Sykes, 1831:101, see above UW hodophilax Temminck, 1839284, see above (synonyms ‘are hodopylax (sic) Temminck and japonicus Nehring) 1, juscus, Richardson, in Beechey, 1839:5. ‘Type locality Columbia River below the Dalles, between Ozegon and Washington, USA’ (a synonym is gigas Townsend) 1. chanco Gray, 1863:94, see above (synonyms are lanier Hodgson, niger Selates, filchneri Matschie, Karanorensis Matschic, tschliensis Matschi, and coreants Abe) CL griseoalbus Baied, 1858-108. Type locality Cumberland stiete Richardson, ater as House, Saskatchewan, Canada (synonyms are Lupus. Griseds Sabine, and knightit Anderson). .1, pambasilens Elliot, 1905:79, se. above. CLL. findrarum Miller, 1912:1, sce above. Vi, bailey? Nelson “and Goldman, 1929-165. Colonia Garcia, Chihuabus, Mexico. . 1 hattat Kishida, 1981:75." Type locality Sapporo, Hok: lkaido, Japan (rex Pocock is a synonym). ©, Lcratsodon Hall, 1952:420,, Type locality Taheie Canal, ‘Nootka’ Sound, Vancouver island, B.C, Canad 6. arabs Pocock, 184:65, Type locity €. Lito Pocock, 1935:689, Type locality “Cape York, on atin Bay, NW. Greenland 1. arctos Pocock, 1935:682. "Type locality "Melville Island, © ©, "M, Allen. and Barbour, 1937:280. Type locality “Newfoundland, Canada.” CL Grremonus Goldman, 1987:51. “Type locality “Red Lodge, a ‘Type locality ‘Carton County, southwestern’ Montana.” USA. |. fabraderius Goldman, 1987:38. Type locality “vicinity of ‘Fore Chimo, Quebec, Canada.” C. 1. ligon’ Goldman, 1957:39- Type locality “head of Duncan Canal, Kupreancf Island, Alexander Archipelago, Alabk USA, . L, mogellonensis Goldman, 1987343. Type locality “S.A. eee, 10" miles northwest of Luna, Catron County, Now Mexico,” USA, CL monsirabilis, Goldman, 1937:42. Type Jo ‘south of Rankin, Upton County, Texas,” USA C. Lyoungt Goldman, 1937.40. Type locality “Hats Draw, north. slope of Blue Mountains, 20) miles northwest of Monticello, Son Juan County, Utah,” USA, 1, aloes Goldman, 1941:109.. Type leeality Kachemak Bay, ‘Kenai Peninsula, Alaska, USA, .1, columbianus Goldman, 1941:110. Type locality Wistaria, Neside Ootsa Lake, B.C. Canada. 6.1 ionias Gpldnan, 1912. Type lcaiy head of ‘Canada ©. c. “10 miles Schule Lake, Reews L bernard) Anerson, 19452388. Type loality Cape Kelle, SW Hanke land, Franklin, Canada C. 1 mackensi Anderson, 1943308. “Type locality Imani, Mlackenaie, Canada (3 Pint Bathucst Inlet, Tupus-dtbue Sabine) ¢. L”mannings Anderson, 1943:392 Type locality Hantech ast Foxo Basin, west side Bain Island, Franklin DIAGNOSIS AND GENERAL CHARACTERS. Largest member (igure 1) of the Canidae except for certain breeds ft domestic dogs (Cans familias}. Adult females weigh from 18.10 38 kg (10 to 120 Ib) and measure 137 to. 152'm (45 0 60 it) in total Length: and males 20 to 80 kg (45 to 175 1b) Sand 127 to Lot m (50 to 65 ft), depending on subspecies, Fer tong and varying in col from pare hie through med tray and brown to coal black; usually grizzled gray. Generally Fesembling domestic German shepherd or husky in bead and Body configuration but distinguishable from them by having ‘orbital angle figure 2) of 40° 0 45° as compared with 53° 1 fa dogs hin 941) ad Baving lng apex: ana almost spherical tympanic bullae as. compared ‘with smaller, compressed, and slightly crumpled bullae in dogs.” Distingui able from coyote (Canis fatrans) by having larger sie, broader Snout, flatively shorter eats, and proportionately smaller brsin case. Canis rufus of eastern Texas and Louisiana is similar to Gonis lupus, being intermediate in many. characters hetween ‘woll and cojote (Nowak, 1970). See "Remarks" Further die- tinctions among these four closely velated and similae animals ‘were detailed by Lawrence and Bossert (1967) DISTRIBUTION. Formerly throughout Northern Hemi sphere north of 20° N latitade (figure 8) in all habitats and Topography except deserts and high mountain tops. DeliverateFicore 1 fetermination has restricted present range in North America (Gee figure 3), and in the Old World to castern Europe, the Soviet Union, China, and northern Indie. Remnant populations live in western Europe and Scandinavia, FOSSIL RECORD. A closely related Pleistocene species was found in Cumberland Cave, Maryland (Gidley, 1913) FORM. Guard hairs are 60 10 100 mm long, (120 to 150 mm in_mane) ‘with imbricate.seales that are, elongate in Drosimal region of hair, ctenate medially, and fattened die ally: underfur bas coronal seales (Adorjan and. Kolenosky, 1960). "Dorsal hairs are generally longer and darker than thore of venters a group of stiff hairs urrounds the preceudal gland fon dorsal side of tal about 70 mm from bace. Hildebrand Ficure 2. Anterior view of wolf skull to show the orbital angle, which is 40° to 5° as opposed to more than S3° in dogs (fom Thin, 1941). MAMMALIAN SPECIES NO. 37 Adult Canis tnpus, photo by the author. (1952) has described details of the integument in the Canidae, Shedding occurs in ate spring ‘The front foot has ive tees, including 2, short one with a ddew claw proaimad frem other four: the hind foot hes four toes. Limb posture is digitigrade; the chest is narrow and keellike with forelimbs scemingly pressed into chest, and elbows turned inward and paws outward (Young, 1944: Ijin, 1981). The legs are moderately long. Dentition 13/3, ¢ 1/1, p 4/4 my 2/3, total 42; Canines ate about 26 mm long and carnacsials are wel de ‘eloped, "The cranium is elongate and tapering anteriorly, with Jong. jaws (Figure 5), Por further details on skull and’ teeth sce Goldman 11944). ‘The baculum is pointed and has a ventral groove. The especially song and large masseter muscles allow powerful bite ‘Atkins and Dillon (2971) have compared the brain to that of ether anid ‘The simple stomach can hold 7 10 9 kg (15 to 20 1b) of food. “The liver is relatively large, varying in males from 07 to 19 kg (1.6 to 42 Ib) and. averaging 12 kg (26 Ib) and Jn females 68 to 82 kg (1.5 to 18 Ib) (Makridin, 1962). Except for minor differences noted in anatomy ie similar to that of the domestic dog. ‘are 10 mamaae (Goldman, 1944) FUNCTION, Fine undefur and long guard hairs com serve a high proportion of body heat enebling wlves to fone tion in temperateres Tower than “i0* Freese travel st sual rate of 8 kim per hour and s running gait of $5 to 70 km ct hour is facilitated by the animal's long legs and powerful Keg muscles (Mech, 1910), The extended rostrum provider abundant surface forthe olfactory organ, allowing the’ wall to detect odors of prey at distances up 10 24 km (15 mi) under evorableconditfons (Mech, 1966). Digestion of foed may take place within afew hours (Sakriin, 1959)" Howling and scent Inarking vis urine posts, feces, an srateking ae common and tnay function in teritory maintenance, although direct evidence {or this hypothesis lacking. Growing. with a frequency of S80 to 490 ls, and barking, at 320 to 904 He (Tembrock, 1963), fr both at ttes manifestations of theeat, ‘The howling of three ult males was described as follows: “(The howls] a com tinuous sound from half second to IL seconds i Tenaths Tt onsnts of 4 fundamental frequency which may lie between 150 and 780 eycles per second (Ti, and up to 12 harmonically related overiones. Most of the time, the pitch remains constantMAMMALIAN SPECIES NO. 37 NN Ficune 3. North circumpolar projection of Holarctic Region to show the original range of Cans lupus and the following subspecies: 1. C.l. monstrabilis, 2. C. 1. baileyi, 3. C. L mmogollonenss, 4. C. 1. lycaon, 5. Cl, nubilis, 6. Gx. younel, Te Gol. tremotus, 8 C.1 juscus, 9G. labradorius, 10. Co hudsonicus, 11. C1. griscoalbus, 12 C. 1. occidentalis, 13, €.1 columbianis, 18.°C. FE crassodan, 15. C. i ligoni, 16. C. l. beo thucus, 17. C1. manning, 18. C. 1, bernard, 19. C, 1. mackensit 20. €.1. tundraram, 21. C.{, pambasileus, 22. C. 1. lees, 23 G., arctoe, 2h Cc. orion, 25" CL. albus’ 26. C21 lupucy 27 C. ‘campesris, 8. C. 1. chonco, 29. C. pallipes, 30. C. L arabs, S1.C. 1. hatte and 2. C1, hadephilay or varies smoothly, and may change direction as many at four Ot five times. Tote intensity does not greatly vary throughout (Theberge and Falls, 1967), Howling by one pack of wolves ‘may stimulate howling in adjacent packs. REPRODUCTION AND ONTOGENY. After courtship that may last from days to months, welves copulate daring an estrus of Sto 7 days. Blood may’ flow from the vulva for a few days to a few weeks before estrus. The receptive. petiod may be anytime from January in low latitades to April in high Taitudes. Many courtship attempts are made, but few succeed (Rabb et al, 1967). Courtship may take place between adult members of packs or between lone welves that pair during the ‘ting season. Copulation involves typical canid coupling. in ‘which the bulbous base of the penis locks into the’ vaginal Sphincter, and the tie may last for more than 30 minutes. Tn dogs. and presumably woltes, ejaculation occurs intermittent]y ‘during the tie. Definite mate preferences are shown, but not all courtship or copalation by members of 2 pair is directed the mate Maturing females may come into heat 2 weeks Tater than animals that have bred hefore. (Rausch, 1967). In Alaska, mature females shed an average of 73 ova and implant 65 embryos, and newly maturing females, Gl ova and. 34 embryos (Rausch, 1967). Gestation laste’ 63. days, and an average of six young (extiemes, one to 11) are born blind and Helpless, usually in a hole in the ground, but often in a rock crevie, hollow log, overturned stump, or ether place. of quict nd shelter. ‘The same den is sometimes used year after Year ‘The female usually stays near the young for at least 2 months, Ficuns 4 Range of the gray wolf in North America prior to sctilement by Europeans and reduced range today (darker shading in the north and in Mexico). An area in western Montana and adjacent parts of Idaho| and Wyoming. where populations are perhaps becoming reestablished 1s shown with question mer, vile the male and other pack members hunt and feed both the Female and the pups. Eyes open at day TI to 15, most milk teeth are present by about week 3, and weaning takes place at bout week 3. After about week 8, the pups are moved to. ground nest, where they romp and play over an area of, up to nacre Ub hectare) an acen known as-a “rendezvous site" ‘The UPS may spend up to 3 weeks at one site but ae then shifted 4s far away as 8 kim (9 mi) to another. Probably depending on the degree of development of paps, they may continue this hehavior even through ‘winter, although pups in good physical condition will join adult members of pack in their travel, fas early as October, at which time they may weigh 27 ka (60 ity “and be almost of adult sige, Adalt teeth replace deciduous teeth between weeks I6 and 26 (Schonberner, 1965). Epi Biota caege cate abot month 12 Hausch 196) jalves may gain sexual maturity in year 2 but often do not breed until 3) In populations unexploited by man, only about (60% of the adalt females bread (Pilot etal, 1969), whereas in“ exploited populations 907% may breed (Rausch, 1967). Wolves may live 16 years (Young, 1944), but 10 years is an old age for individuals in the wild ECOLOGY. | Wolves orginally occupied moet habitats in the Nerthers Hemisphere: “They are. predators oo. lags ‘animals primarily, including deer (Odotoleus). soon or Od World elk (dices ales), ted deer or wapiti or New World lle \Gersus claphas), carlo (Reangijer warendus), bison (Bison Bison), "musk-oxen (Oxibus “mschatu) mountain sheep. (Ovi), and. mountain goats” (Oreamnes americanus) Beaver (Castor canadensis) isthe smallet consistent prey reported, although almost any species of animal may be eaten Including any type of domestic animal, Most analyses of wolf Fredotion on wild species have shown thet young, eld, and ftherwise inferior members of prey populations constitute’ most af the animals killed by wolves tsimmaried by Mech, 1970 see alo Mech and Frenzey 1971), Less than 8% of attacks by ‘®pack of 15 wolves on moose were succeafl an Iee oval Michigan, where the major prey is moose (Mech, 1966). Chaves Fanging fvom 100 m to 5 kon aze the rule. Wolves frst attack the rump of larger prey, but the heady shoulders fankty or romp of deer. Hametsinging has aot been documented. with wild prey. Usually all ofthe prey is eten except for the larger ned and unk’ of hide" Average Will rates vary frm one der per wll per 18 daye (Mech and Frenael, 1971) to oneFicone 5. | Skull of Canis fupus tundrarum from Wahoo Lake, Alasks (after Bee and Hall, 1956:169). moose per wolf per 45 days (Mech, 1966), with corresponding fverage estimated consumption rates being 25 kp (50 1b) of Aer per woll per day to-6.3 ke (139 Ib) of moore per wolf per day. Remains of kis often supply food to such scavengers Is ravens (Corvus coraz), bald ‘eagles (Haliaeetus levee gephalas),foxee ele), and ether eal bide and aml avens frequently follow’ wolf packs for miles, evident search of ea food ® " ‘According to Mech (1970) who summarized the reports of parasites in wolves, the following have been recorded Species of flukes (Erematoda), 21 epecies of tapeworms (C toda), 24 species of oundworms (Nematoda), three spec Of spiny-headed worms (Acanthocephala), two apecies of lie, fone species of flea, seven species of ticks, ne species of tongue worm, and one species of mange mite. Probably the most important parasite of the wolf is Echinococcus granulosus, the hnydatid tapeworm, the larval stage of which can infect man, raves known forthe woll include rabies, distemper, athetis, cancer, and miscellaneous other disorders (Mech, 1970) Few animals compete successfully with the” wolf, except for man and perhaps the cougar (Felis concolor). ‘There is some evidence (Mech, 1970) that high densities of wolves may reduce populations of the coyate (Canis latrans), wolverine (Galo gudo), and lynx (Lynx canadensis). ‘The age structuze of wolves in an unexploited population in Ontario. was 35% pups, 40% yearlings, and 256 adults (Pimlott et al, 1969), whereas in an exploited population in Alaska it was 42% pups, 29% yearlings, and. 30% adults (Rausch, 1967). Mech (1970) has caleulated that survival MAMMALIAN SPECIES NO. 37 rates of wales in unexplited populations are 6 to 4396 for ‘ups from birth to frst winter, 38% from fr to second winter Sind aboot 8070 somally for adults. Momaliy factors incl GBeseh peraien Staaten, Untraperi, ante, ines 3 prey, and exploitation and’ perenation by human being. ‘Wolves sel great deal saualy more a night but aon ring the day ifthe temperatate is cool (Mcsby 1970)- Thee types of movements cam be distingwished (1) travels within me sange, (2) dispersal ond (5) migrations. Home Tange tines vary from 150 kon {SO ni") im Minnesota (Mock aad Prenat 1971) to 13000 in? VSi00" mt) in Alaska (Bark holder, 1959), Daily travels within home range vary from lometers per day to up to 1 (33 miles). A siopersing roll trpeled a suaightline distance of 206 hm (129m te E months (Mech and Fremac, 1971). Tn tundce areas, wolves follow migrating caibow hers and thot snnsally travel more than 0 (100 mi one way, rom tundra to tai and back els, 1968). ‘The fates that individ) migrating wolves dave been Enown to teael ts 350 km (225 mi) Rust, 1973) Studies in progress in Minnesota (Mech, unpublished) show {family groupe of packs vecupy exclusive Home ranges, and that Joe volves cover much larger acu, siting about amongst these, and being chased by the resident packs Highest known ‘atural density of wold populations anywhere on a yeararound basis is one wef per 26m! (10 mi) om the average Pit, 1967). ew other species have ad auch a diversity of relationships with mans has the wat. Evidently ary humans tamed wolves and domesticated em, eventually selectively breeding them and finally developing the domestic. dog (Canis Jamiliars) from thea At present, the tring of wales for pets contines, 4 proces not feu if considerable time ts devoted toy and Well farms now sll pups for $100 to $325 each, Most of exe Animals end'up in os Because ofthe wolfe habit of killing ‘domestic livestock, the species was exterminated by bourties Felioning, and government contol programs in more than 95% tthe ates of the a8 contiguous United. Staten, in. much of Mexico dhe sete farming aren of Canads, in, "mox of ‘reste Europe, and in much of eastern Europe aid the Soviet, Union” In the latter two areas concened programs of exter rmination ‘cominue.(Pullianen. i965). although there i some fidence that oficial policy toward the wll in the USSR is hanging (Osbanin, DTD)” Va provinces stil 4 bounties on wolves, alhough in most of the areas litle Alamage to lvetock i done, The well i informally conidered in danger of extinction in Mexico. To the United States outside ot Alaska, 'six of the eight subspeies that originally occurred at Jeast i pare inthe county are ow considered eatlnet there, (CT enon and C.L tremotus are offically on the Secretary ofthe Imeror' list of endangered mals). Te posillty of the occurrence of some C2 nabils in, Minnesta has been faised by Mech and Frenael (971). Im Alaska, there are Festrcions on taking wolves in some’ areay, and boonies on them in others Harper, 1970). Wolves are deyally protected in Michigan, where ell remnant population exit in Wise ‘Onin, where thers may be afew individuals, and in Tele Royale National Park, where the population numbers 20 to 90 (Mech 1060; Jordan trey 1967)" Inthe Superior National Forest of Minnesota, woltes ace protected by federal amy lthoigh wate lave allow almost unrestricted aking. and in-some parts of Minnesota tate contol programs effering $50. per” animal Eiled are sn effect Several hundred wolves are sill prevent in the nave with an estimated 300 to 900 living in Superior National Forest (Mech, 1973). ‘Wolf far is used for parkas, mukluks, and rags. Rew wolf pelt from foreated atest sell Tor 813 10 $100: depending on wality and calor Arete wolves bring higher prices than those from taher regions. Wolves are hunted for sport in parts of ‘Aaska and Canada, usually ith an aerate searching for the animals ‘on frozen lakes open tundra. Trapping wolves for fur and bounty romaine Tucative for some Indians, Eskimos, anda few whites ‘The main methods of studying wolves have been natural observation in open areas aril tacking and observation in winter, livetrapping via stel trope and snares with locks, ond {ageing with radiocollas. For anesthetzing wildeaught wolves, a combined intramslar injection of 30 me of pheneylidine iydrochloride and 25 mg of promazine hydrochloride hes been ‘gece (Seal and Ericheon, 190; Mech and Frenec, 197). Cnsusing has proved difiealt because of low density and ex. tensive ateas involved. Tn limited areas, extensive aerial obser tations in winter have allowed reasonably accurate esimates (ech, 1965, 1975; Pimlott etal 1969)MAMMALIAN SPECIES NO. 27 sun BEHAVIOR. The wal i sal ail, sesly lanctioning in at ate basically family groups (Mech, 1010). Packs ually contain 5 to 8 members Bot packs of up 10 36’ have been reported. (Rausch, 161).” Pacha rebel Logether hy strong affectional tee that develop, are reinforced, an become fixed in pups during the frst 8 monte of ie, of By couraip dao ben tm ne mae ale Order 18 mutmtained inthe pack by a dominance hierarchy with the fale male dominant fo the adult female and pups, the female dlomiaant tothe pups, and leat vader among the pups Ta larser packs a mule onder and female order develope among arate, Bat the leaders almost always a tale, known a8 the Me male in most interactions between two wolves, one greets the axher by demonstrating its dominance, while the other Indicates Submission (Sehenkel, 1947)" The dominant posture includes sTsitPlegged stance, ears erect and forwady lips pulled forward, duane beoling, and al verial The tecth maybe baredy and there may be growling. There are tro typer of submisan, tetive "and postive. "Active submistion ie les intense and involves drawing back of the lips and cor, raid thrusting out at the tongue, lowering of the hind. quaers and paling of the! til alongside or hetween the lege. ‘The entire body is ‘sully lowered and the more is pointed up toward the mouth othe dominant animal and may touch i This fv often ae- Campanied by lining and wnat. in panive sarin, is'more Intense the wolf vols onto ts back and. draws In its paws toward its body, and often urinates, According to Schenket (1967), is behavior develops fom the elimination posture of young pups in which the. adult Ticks the perineal Seay stimulating and ingesting the ase of the pup. Active deep fom a seen postr os whieh tovthem the Ustsl method St weaning and feeding af pups for the fist of thet Ie. Social anki ‘huge ie probably estalihed noe Hoy ening bat fb pup ted by Mech om thelr patents, fought sviouay at the age of 20 day, establined thee eat tht toy aad ene loi ater tha. "Sica Infractions an dome lemonsrations occur every day throughout the year” bot {ntenefy during the breeding seaton. At tat tie some sit in'rank may take ple, but theee ony occur amg a members." Only rarely is intrapack fighting resorted ‘Courtship between previcusly mated wolts_ may take place several mont belore breeding. (Cusler. 1950) or Just, {Tew days belore the breeding season (Mech, unpublished, auth coat oa een deal of ing, png, ead rains ant rohing al wasting and octal ply with Considerable ody contact Either aie or female may” Shum or tek afoss the bak 9 te her“ engl, feceptive, ill Hit her tall and_dapay her valva." Mounting nay take place from the side ntl ut is som oriented fom elind, with the male grasping the female around tnd iniering his penis, and thrusting. unt the bulbous ‘base ‘ot his pens fs locked ebind the vaginal sphincter. While the pai sil Toeked together by their gestale the mae dismount, Tite hind Tog over his partner's back and trou his body away from her. The pete then remain coupled, back ts back, {eras longue 30 minutes, daring which cjactlationvorurs many “There are thece main methods uf communication among volves: (2) howiing and other vocaliationts (2) visual di: bays including postures ond positions of tatious body pat, Especially the faces and (3) seentmerking. Tt fa krows that indisidual wolges may have distinctive hl (Tueberge snd Hale 867)-"The bowling of pure sully Kihei fonsierable yapping, wheres the pte of adults int lower. One of the functions of howling isin assembling the bck, and. enitory advertisement is suspected as anaher. {larce barking, whining, and growling ae her. voalzatione Body postions'and facial expressions show social status pr ieseribed above. Inguinal presentation and siting weived in satus, demonstrations, but probably alo Serve in individu. identifies. Seemsmarking (Kleiman, 1960) involves deposition of urine andor feces on conspicuous objects tong tale and often includes thea often att eis specvtated ing is a means of advertising ter ‘ory, but thie has not been proved. ‘Wolves are good swimmers and do not hesitate to wade or sovim across ves and Takes: they sometimes follow prey into water even in winter (Pimlet eal, 1960). Wolves resto their sides uovally, although they Sometimes rest on their al months abdomen; when sleeping deeply, they lie curled up with nose beneath ial’ Psy “nvles chasing, ambushing! and, mock fighting, and these activities may help cach pup. determine it Status among its peers, Behavior of wolves has been ie pack (Schenkel, 1987. Rabb ot al 1961) ig Observation (Murie, 1544), and through ‘the use In winter (Mesh, 19662 Jordan et al, 1967; Haber, 1973). "Extensive studies comparing the behavior of wolves in captivity with that of other canids are being eondueted by Fox “1972 GENETICS, To date no differences in karyotypes have been found between the ‘wolf and the domestic dog or the coyote (Hungerford and Snyder, 1966), o the red wolf (Nowak, 1970). According. to Hou’ and Benivachke (1967), both dog. and coyote have 29 pairs of chromosomes, with the autowomes Alescribed as “acrocentrics or teleocentrics” and the gex chromo- Somes ae “submetaceatrie™ for the X and “minute” for the Y. fake oot a mit tei fr the Yn the dog Tia 1941)" crossed a wolf with a black mongeel sheep dog and then made various Types of crosses for four generations, totaling 101 individuals, all of which were fertile, TEARS (0 ee a ne scr it ie i a eka ihe tat a's a ea Sane Se Goteestere auras tame ant ES By oman any fe ocean sate a Send gl fa I Coeur ee Soca a ered Ah ration of this account came ‘ldlife Fund and the World Wildlife Fund LITERATURE CITED A. S, and G. B, Kolenosky. 1969. A manual for identification of huirs of selected Ontario mammals. Ontario: Dept. Lands and Foreas Res, Rept. (Wildlife) Wel 54, Allen, €.M, and T. Barbour, 1987. ‘The Newfoundland well ‘Jour. Mammal. 18:220-234, Andorin, KuM. 1913,” Summary of the large wolves of Canada, with descriptions of three new Arctic races. ou. Mammal. 24:586-395, Atkins, D. L,, and L, S, Dillon. 1971, Evolution of the care ‘bellum fa the genus Canis. Jour. Mammal. §2:96-107 Baird, SF. "1858, "Mammals. In Reports of Explorations and Surveys ... Washington, D.C. vol 8 pt. 1, 764 pp Bee, J. W., and E.R Hall. 1956. "Mammals of 'Nomthern Alaska” 2. Mise. Publ, Mus, Nat. Hist, Unis. Kansas :1-308, Beechey, F. W. 1839. The zoology of Capt. Beechey's voy: age’... 1825-28, Mammalia by J. Richardson. London, 180 po. Rurkholder, BL. 1959. Movements and behavior of wolf Financial support le re from the Welder pack In Alaska. Jour, Wildlife Mgt. 2321-11, Gig ols, 9. Araie with Harper and Bros, NY. SOL Ellermaoy J. Rand, C. $. Montson-Seot. 1951. Checklist ‘of Paluearctic and Indian mamta. . British Museum (Nat. Hist), London, 810 p. Eliot, D. G, 1905. A checklist of mammals of the Noxth ‘American continent, the West Indies, and the neighboring ‘eas, Field Columbian Mus, Publ. 105, Zool. Sex. 6:1- 761, Fox, M.W. 1972. The behavior of wolves, dogs, and related ‘canis. Harper and Row, New York, 220 pp Franklin, Jobn. 1823. Narrative of a joumey to the shores ‘of the polar sea, . -. John Murray, London, 768 pp. 50 pls, maps. Gidley, J. 'W. 1913. Preliminary report on 2 recently dite covered Pleistocene cave-deposit neat Cumberland, Mary. land. Proc. US. Nat. Mus. 46:03-102. Goldman, E, A. 1937. ‘The wolves of North America, Jour Mammal. 1837-48= 1941, Three new wolves from North America, Proc. Biol Soe. Washington 54:109-114 = 1944,""The "wolves of North America. Classification of wolves, The Amer, Wildl, Int. Washington, D.C., part 2 Pp, 387-636, Grayr I.E.” 1863, Notice of the chanco or golden wolf (Canis hanco) from Chinese Tartary. Proc. Zool. Soc. London, Bs abt, G:C, 1972. Eight years of wolf research at McKinley hak Aleka pp 79,3838 (Apu pp WH, 5,558 Hal, ‘ER! 1992. Remarks onthe affinities ofthe mammalign 12 of Vancouver Island, British Columbia, with, de Scrinions of new saspectee. Unit: Calor Publ Zoek 1959. ‘The mammals of Nosth ‘Ameria. "The Ronald Press, New York i+ 547-1088, Harper, J 1970. Wolf management in Alaska. Pp, 24.27, Tn (3. E. Jorgensen, CE. Faulkner, and L. D. Mecb, eds) Proceedings “of ''Symposiuin, on’ Wolf Management in North America U. & Bureau of Spor Fisheries and Wid. Iie, Minneapalis 30 Hildebrand, Me 1982.” The integument in Canidae. Jour Mannmal. $8:419-428, Hodgson, By H. 1847. Description of the wild ses (dsinus olyden) and wolf of Tibet (Lupus laniger). Caleta, Tour, Nat. Hise 7:87 Hou, TG, and K-Benizece, 1967. chionosomes, Isfolios 20,21. Hungerford, D. A, and R. 1. Snyder. 1966, Chromosomes of 1 European woll (Canis lupus) and of a hactean camel ‘camels Batrianus). Mammal. Chromosome, News. 20 2, Thin, 1981, An atlas of mammalian Woltsdog genetics. Jour. Genetics 42 9-414 Jolicgeur, P. 1959. Multivariate geographical variation in the Wolf Canis fupus L. Evolution 13:283-209. Jordan, PLA, P.-C. Shelton and D. L- Allen. 1967, Tumovers, and sovial structure ‘of the Tale population. Aner. Zool, 7:235-252 Keleal, J.-P. 1968.” The caribou. Monogr. 3:1-340, Kishidev 1951, [Tite amlowns paper iaad x preprint from aumber of journal that was never printed). Lansahis 3125) 72-75, Kleiman, Devra, 1966. Scent marking in the Canidae, Symp, Zaal, See. London, 18:167-17T Narbers, Royale. wall fanadian Wildlife Serv Kuyt, E. 1972. "Food habits of wolves on barren-ground Caribou range. Canadian Wildlife Serv, Rept. Ser, 21 136, Lawrence, Barbara, and W. H. Bosser. 1967. Multiple char ‘eter analysis of Canis lupus, latrans, and familiaris, with 4 discussion of the relationships of Conis niger. Amer Fook 1228-292, Linnaeus, C- 1758." Systema Naturae ed. 10, vo, 1, 823 pp, Macidin, VB. "1950" "Material po biologit valka v tundsah ‘enetskogonatsionalnoge.okruga. Zool, “Zhur, 39:1719- 1728 (cited from Pulliainen, 1965), ~ 1962. [The wolf in the Yamal north). TA13-1417 ‘aranslation by Peter Lent) Matschie, B.- 1908-- Chinesivche Siugeticee Tilchner nach China und Tibet 131-282, ple, viewxiv. Berlin. Mock, L. D."1966.""The wolves of Tele Royal. Bark Serv. Fauna Ser, 111-210, 1970, ‘The woll, Na. York. $89 pp. — 1973. “Wolf numbers in the Superior National Forest of Minnesota, USDA Forest Ser. Res. Papen, NC-97:1-10, (North Central Forest Exp. Sta., St. Paul, Minnesota) Mech, L. D. and Frenzel, L. Dz Je. (ed.), 1971. Ecological Studies of the. timber “wolf in northeastern Minnosota USDA Forest Sere. Res. Pap, NC-S2:1-62. North Cental Forest Exp. Sta, St. Paul, Minnesota) Miller, G. Su. Jr, 1912. ‘The names of the large wolves of forthern and western North America. Smithsonian’ Mic Call, $905) 1-5, Marie, A. 1948.. The wolves of Mount MeKinley. U. $. Nat Park Serv, Fauna Ser. 5:1-238 Nelson, E. W..'and E. A. Goldman. 1929. A new wolf from Mexico. Jour. Mammal. 10:165-166. Noak, T. 1611. Fichse und Walfe des Altai, Zoo 35:457-475, L. Salvi, Uppsala, Zoe, Zhwe, 4 In_ Expedition Zool Bot. ‘Ergebn. pb. U.S. Nat 121 Hist. Press (Doubleday), New Anz ‘MAMMALIAN SPECIES NO. 37 Novikov, G. A. 1956, Carnivorous mammals of the fauna of the USSR, Zool, Inet. Acad. Sci. USS, Moscow, 284 pp. (The Israel Program’ ior Seientitie tions, Jerussem, 1962) Nowak, RM. 1970. Report on the red. wolf Wildlife News, pp. 82-94 Oshanin, S. 1971, ‘What will happen to the wolf? Soviet Lite, pp, 52- Paradiso, Jr L- 1968. Notes on recently collected specimens ‘of cast Texas canids, with comments on the speciation ‘and taxonomy of the ved wolf, Amer. Midland Nat. 80: Detenders 529-54, Pimlott, D. H. 1967. Wolf predation and ungulate popu Tations. Amer. Zool, 7:267°278 Pimlott, D. HJ. A, Shannon, and C. B. Kolenosky. 1969, ‘The ecology of the timber wolf in Algonquin Park. On. tarlo Dept. Lands Forests Res. Rept. (Wildlife) 871-92. Pocock, RL 1934, Preliminary disgroses of some new races ‘of South Avablan Mammals. Ann. Mag. Nat. Hist. ser. 10, 141636, 1935, "he pp. 647-686, nen, E, 1965. Studies on the wolf (Canis lupus 1.) in Finland, Ann, Zools Fenn, 2:215-259, Rabb, C. B, J. H. Woolpy, and B. E. Ginsburg ‘elationships in a group of captive wolves, 405-31. 3 of Canis lupus. Proc. Zool. Soe. London, Pull 1967. Social ‘Amer. Zool Rausch, R. A, 1967. Some aspests of the population eeslogy fof wolves, Alaska, Amer. Zool. 7:253-265. Richardson, J. 1629. Fauna Boreali Americana, Josiah Fle. ‘cher, Norwich (England), part 1, 300 pp. Rutter, R. J, and D. H. Pilot 1968, "The world of the ‘wolf. J.B. Lippinestt Co, Philadelphia and New York, 202 pp. Say, T. "1823, Account of an expedition... under the com: ‘mand of Maj. SH. Long (E. James, compiler). Longman, Horst, Rees, Orme, and. Broser, London, vol. 1, 503 pbs vol, 2, 482 pp. Schenkel, K. 1947. Expression studies of wolves. Behaviour 1:81129 (unpublished translation from German by Agnes Klassen) 1967. Submission: ite features and function in the wolf and dog.” Amer. Zool. 7:319-329. Schonberner, D. 1965. Observations on the reproductive bi ‘logy of the woll. Z. Saugeterk. 2O(3) 71-178. (trans lated from German by 8. Van Zyll de Jong) Schteber, JC. D. von 1775. Die Siugthiere'».. . Erlangen, pl 8 Selater, P. L. 1874, On the black wolf of Thibet, Proc. Zoo ‘See. London, pp. 631-6 Seal, U. Sy and A. W. Erickson, 1969. Phencyclidine hydso ‘chloride immobilization of the Carnivora and other mam mals. Fed. Proc. (Symp. Lab, Anim. Ancathes) 28:1410- is. Sykes, W. H. 1831. Catalogue of the Mammalia of Dukhun ‘Deecan) ; with observations on their habits, ete, and characters of new species. Proc. Zool. Soc. London, pp. 99-105 Tembrock, ©, 1968. Acoustic behavior of mammals, Pp. 75 "783, in Acoustic behevior of animals (He Busnel, ed). El Sevier Publishing Co. Landon, 983 pp. Temminck, C, J. 1638." Over de Kennis en de. Verbreiding der Zoogdieren van Japan. Tijdschr. Natuarl, Geschied Physiol. 5:288. ‘Theberge, J. By and J. B, Falls. 1967. Howling as a means ‘communication in 'tmber wolves, “Amer. Zool 538 Wagner, A. J. 1861. Die Raubthiere, Supplementband, Ab thelling” 2:1-558 (in Schteber, Die Saugthiere in’ Ab: Dildungen nach der Natur) ied.Newwied, Maximillian. 1841. Reise das Innere Nord Amerika in den Jahren 1832 bis 1834 J. Hoelscher, Cob lena, 2 vole, 687 pp “+ map. Young, 8. 194d. The wolves of Noth America. The Amer. Wildlife Insi., Washington, D. C, part 1, 386 pp. 733 ‘The principal editor of this account was S. AnoEnso%. LD. Mrcn, Minwesora Freie Sraniox, Paruxst Winpuire Reseancit Csarer, US. Buagav of Sport Fisitetes aNp Witouire,(starise aponrss: Nowrit CenmaL. Forest Exreat DENT STATION, FARWELL AVE, St, PAUL, MiNnesorA 55101)