MAMMALIAN SPECIES wo. 120, pp. 1-8, 4 es Castor canadensis. By Stephen H. Jenkins and Peter E. Busher Published 8 June 1979 by the American Society of Mammalogists Castor canadensis Kuhl, 1820 North American Beaver Castor canadensis Kuk, 1820:68. Type locality Hudson Bay, Can- ‘da. Castor ubauratus Taylor (1912:167), Type locality Grayson, Sen Toaguin River, c Castor caecator Bangs (19133513). Type loc George, Newfoundland. CONTEXT AND CONTENT. Order Rodentia, Family Castoridee. The family contains one genus with two species, C canadensis and C. fiber. the European beaver. The following 23 Subspecies of C. canadensis are recognized by Hell and Kelson (1958) .c- acadicus V. Bailey and Doutt, 1942.87. Type locality Nep- ‘niga River, New Brunswick. . es batleyi Neon, 1927:125, Type locality Humboldt River, & ‘ni above Winnemucca, Humboldt Cor, Nevada 19163439. Type locality Beluga River, Cook Ss near Bay St . €. eaecator Bangs (1913:513). See above, -€. canadensis Kuhl, 1820:68. See above €.°C carolinensis Rhoads, 1896:420. Type locality Dan River, ‘neat Danbury, Stokes Co., North Carolina . cc concisor Warren and Hall, 1980:358. Type locality Monu: iment Creck. SW of Monument, El Paso Co,, Colorado. C.ecduchesnet Durrant and Crane, 1948:413, Type locality Du- ‘chesne River, $600 f.,10;mi. NW Duchesne, Duchesne Co., Utah Cc. frondator Mearns, 1897:502. Type locality Rio San Pedro, ‘Sonore, near Mon. 98, Mexican boundary Cc. idoneus Jewett and Hall, 1940387. Type locality Foley Creek, tuibatary to Nehalem River, Tillamook Co., Oregon. C. e-labradorensis V. Bailey and Doutt, 1942:86. Type locality 5 mi, above Grand Falls, Hamilton River, Labrader. Cc. leucadontus Gray, 1869:298, Type locality Vancouver Island, British Columbia ipacifcus Rhoads a synonym). C. ¢. mexicanus V.. Bailey, 1913:191. Type locality Ruidoso Creek, 6 mi. below Rutloso, Lineain Co., New Mexico. C. ¢. michiganensis V" Bailey, 1913:192. Type locality Tahqua- ‘menaw River, mi. above falls, Luce Co, Michigan Cc. missouriensis V. Bailey, 191933. Type locality Apple Creek, C. ¢. phacus Heller 1909:250. Type locality Pleasent Bay, Ad- ‘miralty Island, Alas. C. c. repentinus Goldman, 1992:266, Type locality Bright Angel c Creek, 4000 ft, Grand Canyon, Anzona «rastralis Durrant and Crane, 1948611. Type locality Red Butte Canyon, 5000 f., Fort Douglas, Utah. €. sagitiatus Benson, 1933320. ‘Type Tocalty Indianpoint Creek, 3200 ft, 16 mi. NE Barkerille, Britsh Columbia C. e- thastensis Taylor (1916:433). Type locality Cassel, Hot ‘Creek, near Pit River, Shasta Co., California. CG. e. subauratus Taylor (1912:167). See above 6. tarlort Davis, 1999:273. Type locality Big Wood River, near Bellevue, Idaho, ce. texensis V. Bailey, 1905:122, Creek, Texas Type locality Cummings G. eagcator and C. subauratus were relegated to subspecifc tank by Allen (1942) and Grinnell (1933), respectively DIAGNOSIS. C. canadensis closely resembles C. fiber of Eurasia in external appearance. They diferin chromosonte num- ber: C. canadensis, 2N = 40; C. fiber, 2N = 48 (Lavrov and Or- lov, 1923). There are alco consistent differences in cranial mor- phology (Lavrov and Orlov, 1978; Trossyfisk, 1973), GENERAL CHARACTERS. Figure 1 shows external ap- pearance, and figure 2 the tail. Typical adult sizes are: total Tength, 1600 to 1200-mm: til length, 258 to 325 mm: tall width, 0 to 200 mm; hind foot length, 156 to 205 ma; ear length, 23 0 20 mm; greatest length of skull 121 to 146 mon aygomatic width, 87 to 108 mam (Grinnell et al, 1937; Osborn, 1953); weight, TL to 26 keg (Grinnell et a., 1937’ Leege and Williams, 1967; Nov Kowaki, 1967; Aleksiuk and Cowan, 19684). Maximum reported Weights are 87 to 39 kg. but these occur rarely (Geianell eal, 1984; Schorger, 1953). There is a prominent depression in the basioceiptal. Dental formula is 1/1 © 0, p Ul m 88, total 20 The eyes have a nicttating membrane. cars and nose are val: vilar, and ips close behind incisors. Exch limb has five digits, Hind feet are webbed between digits and the second digit of each is split. Front feet are small and al digits are clawed. Underfur is dense and lead-gray in color. Guard hair is long and coarse. Color may range from yellowish-brown to black, with reddish: brown being most common. Tall Is Nattened dorsoventally, scaled, and relatively hairless Ite black in young animals but becomes lighter with age DISTRIBUTION. Castor canadensis occurs naturally in streams, ponds, and the margins of large lakes (Shelton, 1966) throughout North America, except for the arctic tundra, penin~ ular Floride, and southwestern deserts (Sgure 3). Boundaries between subspecies are uncertain because of large-scale planting and restocking operation, and so are not shown 5. C. canadensis has been introduced into Eurasia, and may even be cut-competing C. fiber in some arees (Lahti and Helminen, 14), FOSSIL RECORD. The earliest known representative of the family Castoridae is from the early Oligocene of Natrona ‘County, Wyoming (Emzy, 1972). The evolutionary relationships of the Castoridae with other extant families of zadents are not Ficune: 1 External view of Castor canadensis. Photo from American Museum of Natucal History photo fies,Fioune 2. Tail of beaver. American Museum of Natural History photo files. Seale represents about 100 mm. known (Wood, 1959; Wahler, 1972), although Fischer (1972) showed that the placenta of C. canadensis is generally sciuro- tmorph in form. The gents Castor apparently arose in the Plio- ene, and is fst represented by fossie from Bobmnerze von Mel- chingen, Germany: Fresno Co., California: and. perhaps Shani, China (Siron, 1985) Speciation within the genus during the Pleistocene is not fully understood (Fichter, 1972), but these medium-sized beavers covxisted with much lager forme (Caso roides in North America, Trogontherium in Eurasia) until about 10,000 years B.P. (Martin, 1967. FORM. Female beavers have four pectoral mammse. Ex- cept during lactation, there is no external sexual’ dimonphism, ‘Osborn (1953) pictured the baculum, and Osborn (1938) and Bond (0956) gave bacular measurements, The skull ie robust, with Aistinct pit inthe basiocepital, a small slit-ikeinfraorbital es 4 dorsoventrally broadened jugal, and a tubular extension ofthe balla enclosing the long external auditory meatus (Sgure 4). Bond (21956) found no sexual dimorphism in cranial measurements, The cheek teeth are siongly hypsodon with molasiform premolars. ‘The incisors are evergrowing and prominent. Osborn (1969) de- seribed their structure in detail, Robertson and Shadle (1954) described skeletal changes with age, particularly epiphyseal con- Salidation and ankylosiss Carlson and Welker (1976) characterized the tal, with emphasis on musculature and innervation Central nervous system and brain structure were described by Piller (1959s) and Carlson and Welker (1976), Pier (1959) described the anatomy of the hypothalamus. Since the epigotis the trachea opens ony into the nares (Coles, 1970). the palate (Coles, 1970). The of other rodents except for the in the stomach. The caecum is large, but not as large in relation to body size as in guinea pigs and rabbits (Currier et a, 1960), Aleksile and Cowan (19690) gave heart and kidney weights for 0- 3 yearold beavers. MeKean and Carlton (1977) reported mean heart weightbody weight ratio for adutsized beavers of 0.22%, aan ungsualy low value for a mammal. "There ion in size and shape of beaver ovaries 3 during estrus of pregnancy (Provost, 1962}: The maximum size reported by Provost (1962) was 15 by 25 mm. Corpora lutea are 6 t0 8 mm in diameter and prominent: ‘corpora albiantia are also prominent and remain visible for at Teast one year after they are formed (Provost, 1962). Fischer (971) described the steuctare and development of the placenta, land noted the existence of an endometrial papilla which forms on the mesometral wall of the uterus just before choroallantoic con- taet. This structure is apparently unique to, beavers. Osborn (1953) and Conaway (1958) gave testes. and seminal vesicle ‘weights of males of various ages at various stages in the repro Guctive cycle. Conaway (1938) described the histology ofthe uter- ‘us masculinus, which is quite variable in shape and size but not ts large a that of C. fiber, The urogenital system opens into a ‘Common pouch (cloaca) with the scent glands and rectum. Srend- Sen (1978) described anal and castor glands, which are present in oth sexes. FUNCTION, Lactation and milk composition were de- scribed by Zurowski et al. (1974) for C. fiber. The lower pair of ripples yield more milk than the upper pair. Duration of lactation MAMMALIAN SPECIES 120 Ficuae 3. Distribution of Castor canadensis in North Americe. Modified after Hal and Kelson (1959) by extension into the d ‘ages ofthe east slopes of the Sierra Nevada mountains of centr California and extreme westera Nevada (Richardson, 1958; per- sonal observations), and ino the Alaska Peninsula (Mi. S. Boyce, personal communication). Isolated populations in peninsular Florida dN. Layne, personal communication) and San Diego County. California (Bond, 1977) are not shown. is a leat 90 days, and milk composition remains the same from day 10 to day 90. Carlson and Welker (1976) found that somatic sensory ce- rebral neocortex projections from the upper lip and hands of bea ‘ers are more prominent than in other rodents, whereas projec- tions from the tail and vibrissae are similar in telative Size those of other rodents. Miller (1967, 1970) described aspects of nerve function which are adaptive for cold temperatures. In pat Ueular, he showed that peripheral nerves function at lower tem peratures than interior nerves and that caudal nerves contine to Conduct impulses at —5.0°C in some animals (ive of his nine subjects ‘Kitts ee af. (1958) and Stevenson et al. (1959) gave data on normal blood chemistry. Clausen and Ersland (1968, 1970) de- Sctibed in detail the effects of diving on the blood chemistry of fiber. Beavers can remain underwater for at least 15 minutes, though Ito 2 minutes is more usual (rving and Orr. 1938) ving (1987 found that cessation of respiration for | to 2 minutes follows Infation of the lungs. During this pered, blood pressure drops, ‘muscle blood flow drops, and blood flowin the brain increases. ‘These responses also occur in terrestrial maramals during a= physiation, but are more pronounced in beavers and other diving rammals(rving, 1937). Interestingly. total oxygen storage pacity in beavers is about the same relative to body weight as hhuman beings, and much less thaa in harbor seals (MeKean and Carton, 1977. Postgastric fermentation by caecal microorganiems enables beavers to assimilate shout 30% of the cellulose in their diet (Carrer et al, 1960; Hoover and Clarke, 1972). Efficiency of digestion is futher increased by caecotrophy, the periodic inges- tion of material produced in the caecum. This fecal material is {quite different in color and form from the usual feces (Wilsson, 197), Food pascage time for the beaver gut is atleast 60 hours (Curses et al, 1960), Maintenance metabolism of beavers is reported to be about 64 kealkgiday (Pearson, 1960) The tail functions in thermore: lation (Coles, 1966), fat storage (Aleksiuk, 19702), and. com- ‘munication (ace Behavior section Aleksiok and Cowan (19695) investigated seasonal changes in hormonal status, metabolim, and growth, In beavers from the arctic (Mackensle Delta) there was @ marked depression in thyroid activity and growth during ‘inter; these changes were less apparent in beavers from a temMAMMALIAN SPECIES 120 Fioune 4. Skull of Castor canadensis, Univ. of Nevada Mu- seum of Biology no. T165. Drawn by C. Hewitt. Scale represents 50mm. perate climate (Celifornia) held under the same environmental ‘conditions in the lab in British Columbia, ONTOGENY AND REPRODUCTION. Beavers repro- dduce once a year, generally n January or Februery (Hodgdon and Hunt, 1953: Bergerud and Miller, 1977). The gestation period ts bout 107 days (Wilson, 1971). Birth usually occurs in May or June (Shadle, 1930; Bradt, 1939: Hodgdon and Hunt, 1953; Ox: bom, 1953; Bergerud and Miler, 1977), but occasionally as early as February (Miller, 1948) or a late as November (Thomas, 1943 Cook and Maunton, 1958). Osborn (1958) found no evidence that young females breed later than alder ones, though earlier workers (Grinnell etal, 1937) had suggested this. Limited date suggest 3 that timing, and duration of the breeding season may vary with Iatitade ot habitat (Thomason and Jacobson, 1978). Most studies report mean ite sizes between three and four. (Of 215 pregnant or recently post-partum females examined by Hedgdon and Hunt (1953), Osborn (1953), and Brenner (1968), 51% contained three of four embryos or placental sears and 965 contsined six or less (see Hodgdon, 1949, for methods). Litter ‘ice varies markedly with habitat, even within a limited geogeaph- Iclarea (Yeager and Rutherford, 1957; Pearson, 1960; Rutherford 1964). The most important proximate factors infuencing liter size appear to be quantity and quality of available food and se- verity of winter weather. Litter size i alo positively correlated ‘with weight of the mother (Pearson, 1960: Boyce, 1974) OF the few studies in which age was estimated by degree of dental de velopment rather than weight (Novakowski, 1965; Henry and Bookhout, 1969: Boyce, 1974, only Henry and Bookhost’= showed a consistent inereate of litter size with age. C. Aber has “maller litters than C. canadensis where the two species are sym= pitric in Finland (Wilsson, 1971; Lal and Heleninen, 1974) Bergerud and Miller (1977) reported data on prenatal growth Beavers are born fully furred, with eyes at least partially open tnd incisors erupted (Bradt, 1989; Guenther, 1948). Individual Birth weights are inversely related to litter ste, and average 340 to 690 g (range of within tier means of one litter reported by Shadle, 1980, and three reported by Bradt, 1999 ‘Bailey (1927) stated that beavers are weaned at about 6 weeks 102 months, although they begin to eat sold food somewhat ea Tier. Zurowsii et al- (1974) showed that lactation lasts at least 3 months in C- fiber, and suggested thatthe same may be rue for C. canadensis. Aleksiuk and Cowan (19692, 19680) studied ‘growth of beavers inthe wild and in captivity, Beavers from the Miachencie Delta grew only daring summer, beavers from Cali fornin grew yearround In thelr sample of arctic beavers, growth, ally stopped at age 3.5. However, duration of growth may hetween populations in the same geographic area (Boyce, 197%, so broad generalizations about growth patterns are not pos sible at present There are no reliable eports of beavers becoming sexually mature in the first winter after bieth, but both sexes may be mature in their second winter, at age'1.5 (Larson, 1967; Henry and Bookhout, 1969). Henry and Bookhout (1969) found that 40% of LS-year-old females and 8996 of older females trapped in ‘ortheastern Obio during early February had ovaries with corpora Iutea. In interior Alaska and northern Canatla, sexual maturity tay he delayed until age 2.5 or even later for virtually all females land most males (Novakowski, 1969; Boyce, 1974). Age of eexual maturity depends on colony composition: in his study in Wood Buffalo National Park, Novakowski (1965) found that the only pregnant 2.5-year-ld females @ of 21) were trapped in colonies Tncking older females, Larson (1967), Henry and Bookhout 1969), and Boyce (1974), working in Maryland, Ohio, and Alaska, respectively. collected data on the age structure of beaver populations. Age was deter mined according to degree of dental development of trapped sens (van Nostrand and Stephenson, 1964: Larson and van nd, 1968). There were marked differences among the pop- ‘lation samples: for example, frequency of kits (0.5 tol yeat old fat the time tumples were taken) varied from 14% to 42%, and of kets plus yeatings from 40% to 589%. Some ofthese differences can be attributed to differences in intensity of expletation uf the populaticns by trappers. This relationship was most cleatly dem» tnstrated by Boyer (197%), who compared trapped and untrapped populations on two nearby river systems in interior Alaska. Ber- {gerd and Mille (1977) presented survivorship curve for 8 pop Slaton in Newfoundland which they presumed was stationary: Jannval mortality rate was approximately 30% for all age classes ‘The longevity record fora wild beaver is 20.5 to 21 years, but few animals ive beyond 10 years (Lareon, 1967) ECOLOGY. Like many rodents, beavers build elaborate nests and burrows, and store food for winter use, Theie ability to {ut trees is unique, and enables them to build mud and wood lodges" surrounded by open water and watertight dame even in fact-lowing streams, Lodges usually have two or more under: water entrances and a chamber afew inches above water level (Grinnell taf, 1987). Stephenson (196) found that temperatires inside an Ontario beaver lodge in winter were both higher and Tess variable than external ar temperatures. Temperatures inthe lodge deereased when beavers left. Dame may be very Tong and cctise linge areas of land to he flooded. Wilson (1971) and Hart- ‘man (1975) showed that beavers are stimulated to build dams by the sound of running water. Berry (1928) gave a detiled descrip-4 tion of eaver-made canals a less familiar example of their a {eration of habitat than dams or lodges. Warren (1927) presented humerous examples uf the variety of sizes and shapes of beaver dams, lodges, and canals. “The fundamental unit of a beaver population is the colony, which typically consists of four to eight related individual (Bradt, 1938; Bergerud and Miller. 1977) occupying a pond oF section of ‘more (Bergerud and Miller, 1977) ot less (Warner, 1976) ‘exclusively. Adult females appeat to be more sedentary than ‘adult males (Townsend, 1953: Lee, 1968; Bergerud and Miler, 1977. Beavers disperse at about two years of age (Bredt, 1938 ‘Townsend, 1953; Aleksiuk, 1968), or later in some populations (Novakowshi, 1965; Bergerud and Miller. 1977). Dispersal move- Libby, 1957; Hibbard, 1958; Leoge, 1968, with 110 kim being the m ince reported (Hibbard. 1958). The density (of colonies) ‘Sikm* (Voigt er al. 1976), but more typical values favorable habitat are 0.4 to O.d/km* (Aleksiuk, 1968; Vogt e l., 1976; Bergerud and Miler, 1977, rs ae "choosy generalist” herbivores (terminology of 1969). They eat the leaves, twigs, and bark of most species of woody plants which grow near water, as well as many Afferent kinds of herbaceous plans, especially aquatics. Despite this gonerality, beavers are usually quite selective. For example, Tenkins (1974 found that 16 of IT tee genera present ats beaver pond in central Massachusetts were cut over a two-year period, but 6 genera accounted for more than 90% af all trees cut. Where pen (Populus spp), willows (Salix spp. and conifers (espe= cially Pinas spp.) aze the only woody species present in their habitat, beavers strongly prefer aspen and willow. Hall (1960) suguesied that aspen is prefered to willow, although willow more closely approximates a tenewable resouree for beavers than as- pen. However, beavers thrive even in the absence of aspen and willow (Chabreck, 1958; Jenkins, 1975) During summer, beavers rely largely on herbaceous vegeta tion (Chabreck, 1958; Northcott, 1971) or wilow leaves and twigs (Aleksiuk, 197H)), In northern populations, primarily during fall, when a food cache of br builtin the water near a lodge or bank burrow. Materials from this cache are eaten throughout the winter, although beavers wil continue to go ashore to cut fresh trees as ong as they can break through the ie at the edge of their pond. There is some evidence that branches of certain tee species are added to eaches not for future food use but simply to hold the eache in place (Slough, 1978). Slough’s study emphasized the pitfalls of relying solely'on survey ofcut stumps as a messure of food preference. Where fvallable, the lesby Toots, thizomes, and runners of water Mies (uphar and Nymphaea) may be an important source of winter food (Nash, 1951; Hakala, 1952; Gibson, 1957; Shelton, 1966). Most work indicates that beavers either prefer small tees (Nixon and Ely, 1969; Kienaler, 1971) or show no selectivity by size (Gibson, 1957; Henry, 1967). Jenkins (1974) demonstrated igeater selectivity by size at greater distances from shore. Trees ger than about 10 cm in diameter are sometimes debarked at 1c base without being felled (Chabreck, 1958), or felled and the trunks debarked in place, with only the branches eared back to the pond for addition to'the food cache (personal vbservation). ‘Availability of food, particularly aspen tees, is an important determinant of habitat suitability for Beavers (Lawrence, 1954; Rutherford, 1955) Aspen isan eary successional species in most habitat types within the beaver's geographic range, =0 fre and other disturbances which initiate secondary succession may ule timately ead to incresred beaver populations (Lawrence, 1958), Numerous physical features of lakes and streams also influence their suitability for occupation by beavers (Yeager and Ruther- ford, 1957; Slough and Sadler, 1977). Certain sites with special tepographic ot edaphic conditions may permit continuous oceu- pation by beavers. These sites provide a source of colonists for temporary occupation of patches of early successional deiduous tuees, especially aspen, created by fie or other disturbance (Slough and Sadler, 1977, Major predators on beavers are wolves, Canis lupus (Vaigh ft al., 1970), and coyotes, Canis larrans (Young and Jackson, 1951): Wolverines, Gulo fuscus (Rausch and Pearson, 1972), and bears, Ursus americanus (Hakala, 1952), may oceasionally prey fon adults; and mink, Mustela vison (Swank, 1949), on kits. Bea- ‘vers comprise ahighiy variable fraction of the diet of wolves, with {maximum of 154 reported by Voigt eal. (1976) fora population in central Ontario. Major ectoparasites include Platyprylius cas: tris, Prolabidocarpus canadensis, Schizocarpus mineaudi, Lep. tinillusvallidus, and Leodes banisi (Erickson, 194 Lawrence MAMMALIAN SPECIES 120 al, 1961; Janzen, 1963), Relatively few species of helminths parasitize beaver intestinal tacts, the most common being the trematode Stichorchis subsriquetris and the nematodes Travas- sostus americanus and Castorstrongylus castors (Erickson, 1944 Brenner, 1970). Ernst etal. (1970) described two species uf coc- cidians obtained from beaver feces, Epidemics of tularemia, spread directly by water Gellson ef al., 1942) ot by the tick Trodes banksi (Lawrence et al. 1956), have been recorded in beaver populations (also see Stenlund, 1953). Other browsing mammal, especially mouse and snowshoe hares (Nomthcut, 1964; Shelton, 1956). ace potential competitors of beavers. Damming of streams by beavers affects sh popula: tions in many ways, often beneficially (Neff, 1957; Gard, 1961; Hanson and Campbell, 1963), ut sometimes harmfully (Rupp. 1954: Knudsen, 1962). Waterfowl populations may be greatly hanced by the presence of beaver ponds (Rutherford, 1959). Bea- vers influence vegetation not only by their selective cutting of Certain tree species, but also by producing elevated water tables fand associated changes in soil chemistry (ives, 1942; Wilde et al, 1950), They may hasten or impede forest succession adjacent to occupied ponds or streams, depending on local conditions (Lawrence, 1954; Rutherford, 1955: Slough and Sadieir, 1977) Ives (1942) argued that beavers had an important role in land: seape evolution in North America ‘Some important methodological papers are those of White law and Pengelley (1954) on handling lve-trapped beavers, Miler (1564) 'on marking beavers for individual identification, Osborn (955) and Larson and Knapp (1971) on sexing ive beavers, van ‘Nostrand and Stephenson (1964) on age determination, and Bush cer (1975) and Lancia and Dodge (1977) on radlo-telemetty. Bailey (927) and Wilsson (1971) diseussed various aspects of keeping beavers in captivity. Transplanting beavers has been spectactr larly successful in establishing new populations (Harris and ALL ddous, 1946; Knudson and Hale, 1963) Patric and Webb (1953), ‘Yeager and Hill 1054), Yeager and Rutherford (1957), and others hhave discussed stocking, harvesting, and other aspects of beaver ‘management BEHAVIOR. General descriptions of beaver behavior were presented by Morgat (1868), Mill (1913), Dugmore (1914), Ware Fen (1927), and Seton (1929). More recent feld studies emphasiz- ing behavior include those of Tevis (1950), Schramm (1968), and Hodgdon and Larson (1973). Wissen (197) conducted extensive experimental studies, both in the laboratory and in the Beld, of Cifber behavior ‘Moat workers consider beavers to be monogamous. Novak 7) presented evidence that almost all colonies in his study in Ontaro contained only one pregnant female. The few tases reported by others in which tro pregnant females were present in some colonies (for example, Hammond, 1983, Warner, 1976) occurred in high density populations. Reproductive behav. Jor was described by Roth (1938), Bradt (1939, 1940), and Hediger (21970). Copulation usualy takes place inthe water, but may eceur in a lodge or burrow (Wilsson, 197). Wilson (1971-218) dia agramed a copulatory position of captive C. canadensis observed bby Hediger 1970), Aggression is characterized by hissing, grunt- ing, and tooth-sharpening (Leighton, 1933; Novakowshi, 1963: Wiisson, 1971). Nesrly all aggressive interactions observed by ‘Tevis (1950) in a New York colony were associated with compe: tition for food. Tesis (1950) considered the adult male and female te be co-dominant, but Hodgdon and. Larson (1973) reported strong dominance by the adult female over other colony members in ther observations of e Massachusetts colony. In both studies there were indications that young animals were dominated by ‘older ones, Other known social behaviors are mutual grooming, ‘ose touching, and wrestling or dancing (Tevis, 1950; Schramm, 1968; Wilsson, 197) Some instances of these latter behaviors ‘ean reasonably be interpreted play. It is important to ote that ‘social interactions among beavers, even among members of the Same colony, are relatively rarely observed, at least ouside the Todge Beavers communicate by postures, vocalization, tail-slap- ping, and scent-mounding. Leighton (1933) described seven di ferent sounds made by captive beavers, although only two oF three of these have been reported in Geld studies (Schramm, 1968; Hodgdon and Larson, 1973). Novakowski (1969) presented sound speetrographs of normal beaver vocalizations, and die- cussed the beheviors accompanying vocalizations of a laboratory colony. Hodgdon and Larson (1973) showed that kits vocaized ‘more than adults, but that most vocalizations were directed to- ward adults rather than other kite or yearlings. Tailslapping,MAMMALIAN SPECIES 120 particularly its effets on other members of «colony, was stu fn detail by Hodgdon and Larson (1973). In the colony they stud ied, the adult female tal-slapped most often. Males and females ‘of all ages were more likely to respond (by ewimaing to deep ‘water) to # talelap by a female than to one by a male. The ‘possibility of substantial variation between colonies in tal slap- ping and other social behaviors remains to be explored. There is Fairly good evidence that wilslapping funetions to warn other beavers of danger (Tevis, 1950), but additional functions, such as frightening a potential predator, are also possible. Sceni-mound- ing, in which castoreum (washed out of the castor glands by urine) ‘and possibly also enal gland secretions (Svendsen, 1978) are de- posited on ples of mud within a colony's home range, appears to Function primarily to mark teritoral boundaries (Aleka, 1968; Wilsson, 197) Locomotion inthe water is powered primarily by the webbed bind feet: the front fect are held tight against the body. Trans- verse motions of the til are used intermittently, especially when Swimming rapidly (Wilsson, 1971). On land beavers usually walk, bh they ean gallop if frightened. Tevis (1850) described various types af dives Beavers feed on shrubs such as willow by grasping a branch with @ forelimb, pulling it to the mouth, and cutting i off with the teeth. When cutting trees they stand on thei hind feet and fanaw with their incisors Construction behavior was described by Warren (1927) and. Shadle 1956). Hedgdon and Larson (1973) presented data on age snd sex differences In amount of Hime spent on ledge mainte: hance, food cache building, and dam maintenance. Wilsson (1971) ‘described the ontogeny of these (and other) behaviors in CC. fiber. Though complex behaviors, lodge and damn maintenance ‘pear to be largely innate, The formers released by rising water Ievels and appearance of holes in the sides or tof of the lodge, and the latter is released by the sound of running water (Wilsson, 197), Beavers are active fer about 12 hours a day during spring, summer, and fal, from about 1600 or 1800 hours until 0600 or (0700' hours (Busher, 1975). During the day, all members of colony rest together in a single lodge (Tevis, 1950) or in several Subgroups, each in a different lodge or hank burrow (Busher 1975). At northern latitudes during winter beavers exlbit a"free running cireadian rhythm of period length 26.25 to 28.0 b, with for without relative ceordination, depending on available light in- tensity, which in turn depends on ice and snow cover conditions” (Potvin and Bovet, 1975). GENETICS. C. canadensis has 2N = 40 chromosomes, with a fundamental numberof 80 chromosomal arms (Lavrov and Grlov, 1973). Larson and Knapp (1971) described sexual di- ‘morphism in polymorphonuclear neutrophilleucocytes, apparent- Ty resulting from inactivation of one. member of the pait of X- ‘chromosomes present in cells of females. Lavrov and Orlov (1973) ‘obtained copulatione of C. canadensis and C. fiber in captivity, bat no hybrid offspring were born. REMARKS. An enormovs number of books and papers Ihave been writen about beavers. Yeager and Hay (1955) listed cearly works, Much Gsefal but hard-toobtain information itn theser, dissertations, and reports of various governmental agen- cies concerned with wildlife management. Beavers were historically important as a primary stimulus for exploration of western North America (Cline, 1973). They were seduced to low numbers throughout much of North America bby 1900, but have been reintroduced and hecome re-established [Im most of their former range and are quite abundant in many areas (Anderton, 1968), We thank C. Hewitt for drawing fire 4,°M. Boyce for comments on the manuscript, and'S. Anderson for assistance above and beyond his duties as editor LITERATURE CITED Aleksiuk, M. 1968. Scent-mound communication, territoriali- 'y, and population regulation in beaver (Castor canadensis Kh. Jour. Mammal. 89:739-762. — 19700. “The function of the tail asa fat storage depot inthe beaver (Castor canadensis). Jour. Mammal. $1:145-148. = 19105. The seasonal food regime of arctic beavers, Ecology 51:264.270, Aleks, M., and I. MeT. Cowan, 19600. Aspects of season- ‘al energy expenditure in the beaver (Castor cmadensie Kuhl) ft the northern imit ofits distribution, Canadian Jour. Zoo aran-abl. — 19656. The winter metabolic depression in Arctic beavers (Castor canadensts Kuhl) with comparisons to California bea: vers. Canadian Jour. Zool. 47:965-979. Allen, G. M._ 1943. Extinet and vanishing. mammals of the Tyestern hemisphere with the marine species of all the ‘ceans. Spee. Publ. Amer. Comm. Int, Wildife Protection 1 avi + 620 pp. Anderson, S. 1964. Return of the beaver. Nat. Hist. 78:38 ‘«. Bailey, V._ 1905. Biological survey of Texas. N. Amer. Fauna Bsat-202. 113. Two new subspecies of North American beavers, roc. Bick, Soc. Washington 25:191-198 1919, "A new subspecies of heaver from North Dakota, Jour. ‘Mammal, 1:31-32. 1927." Beaver habits and experiments in beaver culture Tech. Bull U.S. Dept. Agr. 21:1-40. 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Mammal. 19:199-162 1539, Breeding habite of beaver. Jour. Mammal. 20:486- 488, = 1940, Note on breeding of beavers. Jour. Mammal 21:220-221, Brenner, F. J, 1964, Reproduction of the beaver in Crawford County, Penneylvani. Jour. Wildlife Mgt. 28:743~747 1970.” Observations of the helminth parasites in beavers, Jour, Mammal. 51:171-173. Basler, PE. 1975. Movements and activities of beavers, Cas- tor canadensis, on Sagehen Creek, California, MA. thesis, San Francisco State Univ., San Francisco, California. 86 pp. Carlson, Mey and W. 1. Welker, 1976. Some. morphological, physiological and behavioral specializations in North Ammer fean beavers (Castor canadensis) Brain Behe. Evo. 18:302— 326, CChabreck, R. H. 1958. Beaver-forest relationships in St. Tam- ‘many Parish, Louisiana. Jour. Wildlife Mgt. 22:179-183, Clausen, C.,and A. Ersland. 1968. The respiratory properties ‘the blood of two diving rodents the beaver and the water ‘ole. Respiratory Physial 5:350-359, 1970, Blood O, and acid-base changes inthe beaver during submersion. Reepizatory Physiol. 113104112, Cline, GG. 1974. Peter Skene Ogden and the Hudson's Bay ‘Company. Univ, Oklahoma Press, Norman. xy + 279 pp. Coles, RoW. 1966, Thermoregulation af the beaver. Ph.D. dissertation, Harvard Univ., Cambridge, Massachusetts. 8 ions in the beaver, Jour. Conaway C. 1958. 'The aterus masculinus of Castor can densi. Jour, Mammal 39:97" 108 Cool Av and E. Rr Maunton. 1954. A study of criteria for “atinaing the age of beavers, New York Fish and Came Jour 1:28-46 Currier, Ay, W. D, Kitts, and I. MoT. Cowan, 1960. Celllose igeston in the beaver (Castor canadensis). Canadian Jour Zook 38:1109-1136 Davia, W. B. 1089, "The recent mammals of Idaho. Caxton Painters, Dupmores A: R16 "FRstory of the beaver inthe Western hemisphere. WH emann, Landow iv © 225 pp.6 Durrant, S. D., and H. S. Crane, 1948. ‘Three new beavers from Utah, Publ. Mus. Nat. Hist, Univ. Kansas 1:407-17 Emry, R.}.. 1972. A new species of Agnotocastor (Rodentia, Gastordae) from the early Oligocene of Wyoming. Amer Mus. Novitates 2485: 1-7 Erickson, A.B. 1944, Parasites of beavers, with « note on Paramphistomum castori Kolo and Park, 1932, a synonym cof Stichorchissubtriquetrus, Amer. Midland Nat. 31:625~ 630, Ema, J.V.,W.L-Cooper,andM.J.Frydendall. 1970. “Eimeria ‘sprekai Yakimott, 1894, and E. causeyi sp. n. (Protozoa: Himeriidae) from the Canadian beaver, Castor canadensis, Jour. Parasit. $6:30-31 Fichter, LS. 1972. The North American beavers of the genus autor, post incisor denttion—a multivariate study. Ph.D. dissertation, Univ. Michigan, Ann Arbor. 120 pp. Fischer, T. V._ 1971, Placentation in the American beaver (Castor canadensis), Amer. Jour. Anat. 131:158-184 Gard, H. 1961._ Ellects of beaver on trout in Sagrhen Creck, California. Jour. Wildlife Mgt. 25:221-242 Gibson, GG. 1957. A study of beaver colonies in southern ‘Alaonquin Park, Ontario, with particular reference to the available food: M.A. thesis, Univ. Toronto, Toronto, Ontario, i 86 op. Goldman, EA tal 13:266-267 Grays I. E,1869. On the white-oothed American beaver ‘Ann. Nag, Nat. Hist. Series 4, 4285, Grinnell, |. 1988.” Review ofthe recent mammal fauna of Ca: Hornia. Uni. California Publ Zool 0:71-234 Grinnell J J.'S. Dison, and J. M. Linsdale. 1987, Furbear- ing mimmals of Califora. Univ. California Press, Berkeley. Vol. 2, xiv + 377-177 pp. Guenther, SE. 1948, Young beavers. Jour. Mammal. 29:419- 20 Hakala, J.B, 1952. The ife history and general ecology ofthe beaver (Castor canadensis Kuhl in interior Alaska, M.S. the sls, Univ. Alaska College. 18 pp. Hall, E. Rl, and K., Kelson. 1988-, The mammals of North ‘tverica, onaid Press Co., New York, vill {47-1083 + 1932. A new beaver fom Arizona. Jour. Mam- pp, Hall, J.C. 1960, Willow and aspen in the ecology of beaver ‘on Sagehen Creek, California. Ecology $1:484-495. Hammond, M. C. 1943, "Beaver on the Lower Souria Refuge. Your, Wildlife Mgt. 7316-321. Hanion, W.D.,and KS. Campbell. 1963. The effets of poot ‘ze and beaver activity on distribution and abundance of swarmewater fishes in « North Missouri stream. Amer. Mid- Tand Nat. 69:136-199, Harper, J. L, 1969. The role of predation in vegetational di- "versity. Brookhaven Symp. Bik 22:48-62. Harris, D. and S. E. Aldous. 1986, Beaver management in ‘the Biack Hills of South Dakota. Jour. Wildlife Mat. 10:348— 355, Hartman, A.M. 1975._ Analysis of conditions leading to the regulation of water flow by a beaver. Psychol. Rec. 25:427— 431 Hediger, H. 1970. Zum fortpflanzungaverhalten des Kana- dischen bibers (Castor fiber canadensis). Forma Functio 1970:836-351 Heller, E, 1909." The mammals. Pp. 245-264, in Birds and ‘mammals of the 1967 Alexander expedition to southeastern ‘Alaska (Grinnell, J.. et all. Unie- California Publ. Zool. SlTL-264, Henry, D. B. 1967. Age structure, productivity, and habitat ‘characteristic ofthe beaver in aortheastern Ohio. M.S. the- sis, Ohio State Univ, Columbus. 8 pp. Henry.D. By and T. A. Bookhout. 1969. Productivity of bea ‘ers in northeastern Ohio. Jour. Wildife Mgt. 38:927-£32. Hibbard, E. A. 1958." Movements of beaver transplanted in ‘North Dakota, Jour, Wildlife Mgt. 22:208-211, Hodgdon, H.E., and J. S. Larson. "1973. Some sexual dffer- fences in behavior within a colony of marked beavers (Castor canadensis). Anim. Behay. 21:147-182. odgdon, K. W.. 1949, Productivity date from placental scart in beavers, Jour, Wildife Mgt. 13:812-414. Hodgdon, K. W., and J. H. Hunt. 1958. Beaver ig Main. Nine Dept Inland Fisheries and Div, Bull 3:ix + Hoover Wis andSD. Clarke. 912 Ihetver. Jour: Nutrition 102:9-16. anagement sme, Game Fiber digestion in the MAMMALIAN SPECIES 120 Irving, L._ 1987. The respiration of beaver. Jour. Cell. Comp, Physiol. 9837-451, Irving, L-, and M.D. Orr, 1985, The diving habits of the bea- er. Science 82:569 Ives, R. L.. 1982, The beaver-meadow complex. Jour. Geo ‘morph, 5191-208, Janzen, D. H. 1963.” Observations on populations of adult bea- ‘ver-beeties, Platypeyllus castors (Platypsylidae: Coleop: tera}. Pan-Pacifc Entomol. 39:215-228, Jellison, W- Li, G. M. Kohls, W. J. Butler, and J. A Wesver. 1942.” Epizootc tlaremi in the beaver, Castor canadensis, and the contamination of stream water with Pas- teurella tularensis. Amer. Jour. Hygiene 36:168-182. Jenkins, 5. H. 1914,” Food selection by heavers. Ph.D. disser- ion, Harvard Univ., Cambridge, Massachusetts. 165 pp = 1915." Food selection by beavers: A multidimensional con- tingeney table analysis, Oecologia 21:157-1. Jewett, S. G., and E. R. Hall. "1940. Anew race of beaver from Oregon. Jour. Mammal, 2187-89 Kienaler,J. M1971. Woody plant utlzation by beaver in nat ‘rally acid and acid mine water. M.S. thesis) West Virgin Uaniy., Morgantown. 87 pp. Kits, 'W."D.,"M. C" Robertson. B. Stephenson, and I. MeT. ‘Cowan, "1958. “The normal blood chemistry of the beaver (Castor canadensis). A. Packed-cll volum rate, hemoglobin, erythrocyte diameter, counts, Canadian Jour. Zool. 36:279-283 Knudson, G, J.” 1962. "Relationship of beaver to forest ‘and wildlife in Wisconsin. Tech, Bull, Wisconsin Conser- vation Dept. 25:1-52 Kaudsen, G.., and J. B, Hale, 1965, Movements of trane- planied beavers in Wisconsin, Jour. Wildlife Mgt. 20685— 88, Kuhl, H. 1820. Beitrige2ur aoologie und vergleichenden ‘mie. Verlag der Hermannschen Buchandiung, Frankfurt am Main, Abul 131 pp. Lahti, Su and M. Helminen. 1974, The beaver Castor fiber UL.) and Castor conadenais (Kuk) in Finland. Acta Theriol Wart. Lancia, RAL, and W. B. Dodge, 1977. A telemetry system for continuously recording lodge use, nocturmal and subniv- fan activity of beaver (Castor canadensis). Proc. It. Cont Wildlife Botelemetry 1:86-82. Larson, J. S. 1967," Age structure and sexual maturity within fa western Maryland beaver (Castor canadensis) population, Jour, Marmmal, 48:408-413 Larson, J. Si, and S. J. Knapp, 1972. Sexual dimorphism in beaver neutrophils. Jour, Mammal. 82:212-215 Larson, J. S.,and F. C. van Nostrand. 1968, An evaluation of Decver aging techniques. Jour. Wildlife Mgt. 32:99-103. Lavror, L. S., and V._N. Orlow.” 1973, Karyotypes and tax- ‘gnomy of modern beavers (Castor, Casterdae, Mammalia) Zeal. Zhur, 52:734-742, dn Russian, English summary.) Lawrence, W. H. 1954. Michigan beaver populations as influ- ced by fry and leping, Pu. dateraion, Unie. Miche Ann Arbor. vit + 219 p Lawrence, W. H. LD. Fay, and S. A. Graham. 1956, re port on the Beaver die-o in Michigan. Jour. Wildlife Mg. Sostet-i6r, Lawrence, W. H., K. L. Hays, and S. A. Graham. 1961," Eetoparasites of te beaver (Castor canadensis Kul, ‘Wildlife Diseases 12:1-14. Leege. T. A, 1968, Natural movements of beavers in south- ‘astern idaho. Jour. Wildlife Mgt. 32:973-976, Leege. T. Ax. and R, M, Williams, 1967, Beaver productivity in Idaho, Jour. Wildlife Mgt. 31:326-332, Leighton, A. H. 1983, Notes on the relations of beavers to ome ‘another and to the muskrat. Jour. Mammal. 16:27-35. Libby, W. L. 1957. Observations on beaver movements in ‘Alaska. jour. Mammal 38:26 McKean, T., and C, Caron. 1972, Oxygen storage in bea ‘vers. Jour. Applied Physiol. 42:545-S47- Martin, P. S. 1967. Prehistoric overkill. Bp 75-120, in Pleis- tocene extinctions: The search for a cause (P_S. Martin and HVE. Wriaht, eds). Yale Univ. Press, New Haven. x + 453 pp Mearns, E. A. 1897, Preliminary diagnoses of new mammals ofthe genera Sciarus, Castor, Neotoma and Sigmodon from the Mexican border of the United States. Proc. U.S. Nat ‘Mus, 20:501-505,MAMMALIAN SPECIES 120 Miler, D.R. 1964. Colored plastic ear markers for beavers. ‘Jour. Wildife Mgr 28:859-Bo1. Miler, FW. 1948, Early breeding of the Texas beaver. Jour. ‘Niamimai, 29:41 Miller, L- K. 1967. Caudal nerve function a8 related to tem erature in some Alaskan mammals. Comp. Biochem. Phys fol, 21:679-686, — 1970. Temperature-dependent characteristics of peripheral nerves exposed to different thermal conditions in the same Animal. Canadian Jour. Zool, 48:75-81, Mill, EA. 1013.” In beaver world. Houghton Miffin Co., Bos ton, xiv + 228 pp Morgan, L. H. 1868. ‘The American beaver and his works. J B. Lippincot and Co., Philadelphia. xv + 380 pp Nash, J. Br 1951." An investigation of some problems of eco (gy of the beaver in northem Manitebe., Manitoba Dept. Mines and Natural Resources, Game and Fisheries Branch, 64 pp, Neff, D.'l. 1957. Ecological effects of beaver habitet aban- ‘donmient in the Colorado Rockies. Jour, Wildlife Mgt 21:30— 6 Nelson, E, W. 1927. Description of a new subspecies of bea- ‘ver, Proc. Biol. Soe. Washington 40:125-126. Nixon, C. Mand J. Ely. 1969." Foods eaten by a beaver cok ‘ony in southeast Ohio, Ohio Jour. 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Viltrery 8:115- 266.a Wood, A.E, 1959, Eocene radiation and phylogeny of the ro- ‘dents, Evolution 18:394-361. Yeager, L. E and K. G. He Dibliography on the Dept. of Game and Fish | Yeager, L. E Beaver management Drablems on western public lands. Trans. N- Amer. Wildlife tnd Nat, Resources Conf, 19:462-479 Yeager, L. E., and W. H. Rutherford. 1957, An ecological ‘bass for beaver management in the Rocky Mountain region Trans. N. Amer. Wildife and Nat. Reeources Conf. 22:260- 209) 1985, A contribution toward a MAMMALIAN SPECIES 120 Young, S, P., and H. H.T, Jackson. 1951 ‘Stackpole Co.. Harrisburg. av * 411 pp. Zarowski, W., J. Kisza, A. Kruk, and A. Roskosz. 1974. Tactation and chemical composition of milk of the European Dbeaver (Castor fiber Ly). Jour. Mammal. 55:847-850, ‘The clever covet. Primary editors for this account were SYDNEY ANDERSON and Dante F. WILLIAM SS, H. JENKINS AND P. E. BUSHER, DEPARTMENT OF BIOLOGY, Usivensity oF Nevapa, Reno, NEVADA 89557