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Factors Affecting Black Bear Reproductive Success and Cub Survival Author(s): Kenneth D. Elowe and Wendell E. Dodge Source: The Journal of Wildlife Management, Vol. 53, No. 4 (Oct., 1989), pp. 962-968 Published by: Wiley on behalf of the Wildlife Society Stable URL: http://www.jstor.org/stable/3809596 . Accessed: 10/12/2013 16:27
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BEARREPRODUCTIVE BLACK SUCCESS AND FACTORSAFFECTING CUB SURVIVAL

D. ELOWE,1 Wildlife Research Unit,University of Massachusetts, KENNETH MassachusettsCooperative Amherst,MA01003 Wildlife of Massachusetts, WENDELL E. DODGE, MassachusettsCooperative Research Unit,University Amherst,MA01003

Abstract: Data from 18 adult female black bears (Ursus americanus) from an exploited population in westernMassachusetts bearing62 cubs in 27 litters(31 M and 31 F) providedinsightsinto black bear nutrition and populationdynamics. Diet and nutritionalanalysesof food items indicated that nutritionalcondition of pregnant females affected reproductivesuccess. Twenty-six of 28 females with access to high fat and carbohydratemast diets produced cubs, but 10 of 10 females with low carbohydratefall diets failed to produce cubs. Forty-one cubs (21 M and 20 F) in 18 families were followed intensively from birth until 3 years of age. Overall mortality of cubs was 41% by 1 year and 61% by 2.5 years (subadult).Mortality was much higher for male than for female cubs, even before family breakup (62 vs. 20%by age 1, 91 vs. 35%by age 3). Most cub mortality (76%)occurred between 1.5 and 5 months of age. Human interactions,behavior of the mother, and disease influenced cub survival.Thirteen of 20 female cubs survived to adulthood,and all but 1 stayed within their natal areas. Eight of 21 male cubs survived until dispersal,but 6 of 8 were shot by hunters while dispersingat age 2.5. Only 2 of 21 male cubs survived to adulthood.There was no evidence of competition for food or space. Fall mast availability appeared to influence bear densities through the number of female cubs born. J. WILDL. MANAGE. 53(4):962-968

An assessment of the long-range reproductive potential of black bear populations is essential to evaluate factors that limit bear population densities and to keep management strategies compatible with optimum bear densities. Social order was once thought to limit bear densities through establishment of territories, but it now appears to play a minor role. Although Rogers (1977) found female bears to be territorial in Minnesota, he suggested that territoriality may enhance reproductive success and permit higher bear densities through increased foraging efficiency. In Massachusetts, female bears used definite home ranges but exhibited no territorial behavior (Elowe 1984). Home ranges of adult females overlapped, and bear density did not appear to depend on social order. Other than mutual and/or temporal avoidance and breeding interactions, little social interaction was observed. Fluctuations in recruitment of young into the population, influenced by nutritional or other factors, appears to be 1 way bear densities are naturally limited. Rogers (1976), Eiler (1981), Wathen (1983),

and Elowe (1984) suggested that black bear reproductive success was related to nutritional condition of females. Eagle and Pelton (1983) evaluated quality of seasonal bear diets in Tennessee by comparing nutritional quality of foods found in scats in spring, summer, and fall. Cub production and survival data have been compiled concurrently with other aspects of black bear ecology in several studies (Reynolds and Beecham 1980, Eiler 1981, Wathen 1983), although only LeCount (1987) has addressed specific causes of mortality in young cubs. Our study was designed to identify factors that influence black bear recruitment rates. Specific objectives were to evaluate relationships among nutritional quality of female bears' diets and physical condition, reproductive success, and cub survival after birth. The extent, timing, and causes of cub mortality were examined based on data collected from May 1980 to May 1986 in Massachusetts. E. V. Howard, Jr., and E. S. Howard assisted with field work, and the West Experiment Station Laboratory at the University of Massachusetts performed the proximate analyses of bear food items. We thank S. D. Miller, M. W. Sayre, I Present address: Maine Department of Inland and R. T. Duby for critical review of the manuFisheriesand Wildlife, Wildlife ResourceAssessment script. This study was funded, in part, by the Massachusetts Division of Fisheries and Wildlife Section, P.O. Box 1298, Bangor, ME 04401. 962

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through the Massachusetts Cooperative Wildlife Research Unit and Federal Aid in Wildlife Restoration Project W-35-R-VI-4.

STUDYAREAAND METHODS
The study area encompassed approximately 2,800 km2 in western Massachusetts (42030'N, 72045'W). Elevations were not great (<1,064 m), but terrain was steep and rugged in most places. Northern hardwood forest associations of beech (Fagus spp.), maple (Acer spp.), and hemlock (Tsuga spp.) covered higher elevations (>300 m) of the northern and western parts of the study area and oak (Quercus spp.) replaced beech in lower elevations of the eastern and southern sections. Spruce (Picea spp.)-fir (Abies spp.) associations predominated in moist areas at high elevations (Elowe 1984). Small towns were scattered throughout the study area with sections of residential housing and outlying dairy farms. Approximately 75% of the area was forested (MacConnell and Niedzwiedz 1975). We captured bears primarily with trained bear hounds (Elowe 1984) and cable foot snares. Bears were then weighed, measured, blood samples taken, and selected bears were fitted with radio collars. A first premolar tooth was removed from all bears except cubs to age by cementum analysis (Willey 1974). All females assumed to be pregnant (i.e., >3 yr old and without cubs that yr) were stalked at least every 3 days from early September until denning (20-35 data points/F/ fall) to determine individual diets. Fall foods were determined by opportunistic combinations of direct observation of feeding, gross analysis of known origin scat samples, and evidence of feeding in proximity to a female's location. Major fall foods were classified as those that were estimated to comprise >80% of the fall diet (1 Sep-denning). Samples of foods eaten by pregnant females were dried, ground, and compared using proximate nutritional analyses, which provided the dry weight percent protein, fat (ether extract), carbohydrate (nitrogen-free extract), and fiber contents. Only a limited number of samples of each food item were analyzed, so nutritional composition data were averaged with data from King and McLure (1944), Bump et al. (1947), and Short and Epps (1976). We assumed that nutrient levels of the major food item were an adequate index of overall dietary levels. Where 2 food items made up the major portion of the diet, nutrient composition was calculated from

the proportion of evidence of use. Mast crops were classified visually as scarce, good, or very abundant (Rogers 1976) while following female bears. Dens were visited the first week of February and the number, sex, weight, and health of cubs was noted. Small metal ear tags were used to identify individual cubs. In late March, dens of females with new cubs were again visited and cubs were weighed, tagged with permanent ear tags, and fitted with small, expandable transwt = 55 g) designed to break mitter collars (Tc off at approximately 2-3 months (Elowe 1987). Mothers also were weighed again to detect differences in condition. After emergence, families were intensively stalked to monitor female feeding behavior and cub behavior and mortality. During fall and spring stalking and tracking, we made every effort to observe wind direction, silence, and to ensure that bears were not disturbed. Bears were observed and/or followed closely as long as possible without disturbing (1 min-6 hr). Visual observations, in most cases (>85%), were made on bears that were unaware of our presence. If visual contact was impossible, the area was searched for feeding and habitat information and then left undisturbed. Cubs surviving to 1 year of age were fitted with larger radio collars while denned with their mother to provide data on family break-up, dispersal, and mortality.

RESULTS FallNutrition
Although many different foods were eaten throughout the fall, the diet of female bears consisted of 1 or 2 major food items (i.e., >80% of the diet), depending on availability. For example, females with access to abundant black or chokecherries (Prunus spp.) at higher elevations (>300 m) selected these over all other foods. Beech nuts were eaten occasionally but not extensively until cherries were gone in late fall or cherry production failed. Bears in lower elevations (<300 m) depended on oak and to a lesser degree, hickory (Carya spp.) mast for the bulk of their fall diet. Other minor foods were witch hazel (Hammelis virginiana), rose hips (Rosa multiflora), and pokeberries (Phytolacca americana). When fall mast crops failed, bears fed in cornfields or ate herbaceous materials (i.e., grasses). Some females traveled ?50 km to feed on corn

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of fall mast and effects on blackbear reproductive Table1. Abundance success in Massachusetts,1980-85.
Potential mothers Yr Mastavailabilitya ProTotal duced Comments

1980 1981 1982 1983 1984 1985

Red oak, beech, cherries,and hickory very abundant All mast very scarce Red oak good, but scattered;beech and cherry mast scarce Red oak good, but scattered;beech scarce;cherry mast very abundant Beech, cherry, and red oak very abundant Oak scattered;beech and cherry scarce

5 2 6 7 4 2

5 8 7 7 5 6 Unsuccessfulfemale hit by car-possibleinternal injury 2 successfulmothersfed on corn Mast failed in unsuccessfulfemale's range

a After Rogers'(1976) classification.

when mast crops failed in their normal ranges. Meat in fall diets, as indicated by the presence of hair and/or bones in seat samples, was found only 3 times in 6 years. One scat contained red squirrel (Tamiasciurus hudsonicus) hair and 2 scats contained grey squirrel (Sciurus carolinensis) hair. Abundance of fall mast crops affected reproductive success the following winter (Table 1). Digestible carbohydrates in fall diets in 1981 during a major mast failure were lower than levels in 1983 during a good mast crop year (Table 2). This had a direct effect on the female's condition during the denning period and on the number of cubs born to these females. Diet, condition, and reproductive response were similar in those females without access to fall mast in other years, also. Typical fall diets in good mast years contained 20-40% more carbohydrates and 3-15% more fats than fall diets in poor mast years, depending on elevation and species of mast available. From 1980 to 1986, 26 of 28 female bears with access to higher fat and carbohydrate mast crops produced cubs while 10 of 10 females subsisting on lower carbohydrate herbaceous fall diets were in worse physical condition and failed to produce cubs. February weights of bears that ate herbaceous diets the preceding fall were typically 25-40% lower than weights of the same individuals with access to better mast diets in other years.

Females bore 1 cub in 5 of 27 litters (18%), but these were all first litters for those females. The sex ratio in first litters was 72 M:100 F and was not significantly different from 50:50 (P > 0.10). There were 2 cubs in 37% of the litters and 3 cubs in 41%. Only 1 female (4%) had 4 cubs. Of 22 litters of > 1 cub, 3 were all male (1 with 3 M, 2 with 2 M) and 2 were all female cubs (2 with 2 F). Among 10 known first litters, 4 bears had their first litter at 3 years old, 5 at 4 years, and 1 at 5 years. Backtracking the reproductive history of other radio-collared females indicated that most females probably had their first litter at age 4.

CubSurvival
A subsample of 41 cubs (21 M and 20 F) from 18 families was followed intensively from birth until 3 years old. Overall mortality by 1 year was 41% and by 2.5 years (subadult) was 61%. However, even before family break-up, male cub mortality was higher than female cub mortality (Table 3). By age 1, 62 versus 20% of male and female cubs, respectively died and by age 3, 91% of male versus 35% of female cubs died. Thirteen of 17 mortalities to cubs <1 year old occurred between 1.5 and 5 months old. There was no mortality observed between early February and mid-March. Six of 17 first year mortalities occurred between mid-March and emergence in mid- to late April. One cub was found dead in a den after the mother and 2 remaining cubs emerged. The cub was estimated to have died weeks earlier. No wounds >3 were evident and the cause of death, although unknown, was presumed to be disease. No cub

CubProduction
Reproductive data were collected from 18 adult female black bears bearing 62 (31 M and 31 F) cubs in 27 litters (2.4/litter) from 1980 to 1986. Yearly sex ratios did not differ significantly from 50:50 (P > 0.10).

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Table 2. Nutrient success of adultfemale bears in 1981 (oak, beech, compositionof fall diets (% dry wt) and reproductive and 1983 (oak mast heavy but scattered, beech and cherrymast heavy). mast failures) cherry,and hickory
Nutrientsa Yr Bear Major fall foods Protein Fat Carbohydrate Fiber Den wt (kg) Cubs born Feb 1982

1981

1983

019 004 010 023 008 028 017 003 019 014 031 039 015 017 003

Corn Corn Herbaceous Herbaceous Herbaceous Herbaceous Herbaceous Herbaceous Red oak, corn (1:1)b Red oak Red oak Red oak Beech Cherry, beech (2:1) Cherry, beech (2:1)

10.8 10.8 16.3 16.3 16.3 16.3 16.3 16.3 8.4 5.9 5.9 5.9 10.9 6.5 6.5

3.3 3.3 4.3 4.3 4.3 4.3 4.3 4.3 8.5 13.6 13.6 13.6 17.3 8.0 8.0

80.3 80.3 40.0 40.0 40.0 40.0 40.0 40.0 70.6 61.0 61.0 61.0 30.1 49.1 49.1

2.4 2.4 25.6 25.6 25.6 25.6 25.6 25.6

10.3

18.2 18.2 18.2 26.6 16.0 16.0

65 56 47 44 32 45 36 36 75 50 70 72 52 61 55

3 3 0 0 0 0 0 0 3 2 2 3 2 1 2

a Nutritional values are means from this study, Bump et al. (1947), King and McLure (1944), and Short and Epps (1976). b Nutrient values of diets composed of >1 major food item were calculated from estimated proportion of use, because intake volumes were not known.

mortality was caused by research den work. However, all 5 cubs from 2 families died when mothers abandoned dens due to other human disturbance. One den was located near a house and the other was about 30 m from a frequently used snowmobile trail. Thus, human disturbance was the major cause of death for cubs before emergence from dens. Transmitters fitted to cubs allowed the presence or absence of cubs with the mother to be easily verified and missing cubs to be located, although the cause of death could not always be determined. Seven of 17 first-year mortalities occurred between emergence and 1 June. Causes of cub mortality in addition to those included in Table 3 that were noted between emergence and the end of June 1985 included abandoned (n = 1), falling from tree (n = 1), presumed disease (n = 1), and consumption by the mother (n = 2). It is not known whether the 2 cubs eaten by the mother (same family) were already dead before they were eaten. Hair, claws, and ear tags were found in scats next to the bed that the mother had used about 1.5 weeks after emergence in mid-April, and both cub radio collars were found in the bed. A third cub survived and remained with the female. One of 5 cubs that died between July and January was suspected to have been shot and 4 died of unknown causes. Recently, LeCount (1987) found that adult male bears and other predators were a significant source of mortality

to young cubs in Arizona, but no evidence of such mortality was found during our study. Age and/or experience of the mother appeared to influence survival of young cubs. Of 22 litters, 4 of 7 first-time mothers, and zero of 15 females that had already raised a litter, lost entire litters (2 litters abandoned, 2 unknown). Only 1 of 13 female cubs surviving to age 2.5 moved from a natal area. She moved approximately 15 km and remained in that area for 5 years. All male cubs surviving to age 2 dispersed between 30 and 200 km from their natal areas. Hunting was the major cause of mortality in male cubs > 1 year old. Only 1 subadult female was killed by hunting, but 6 of 8 subadult males were shot by hunters while dispersing at age 2.5.
Table3. Causes of mortality to 19 male and 7 female black bear cubs (fromsample of 21 M and 20 F) in Massachusetts between 1980 and 1985.
By age 1 Cause of mortality M F By age 3 M F

Abandonedin den Natural accident Disease F died Cars Hunting Handling Unknown Totals
n Remaining

3 1 3 1 1 1 3 13
8

2 2 6 4
16

1 2 3
13

6
2

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Thus only 2 of 21 male cubs versus 13 of 20 female cubs survived to adulthood.

AND MANAGEMENT DISCUSSION IMPLICATIONS

Bears used certain fall foods to the exclusion of other abundant potential food sources. Use was directed towards foods high in carbohydrates and fats and low in fiber as was found by Eagle and Pelton (1983). High carbohydrate diets facilitate fat deposition needed for winter metabolism. Nelson et al. (1983) have shown that fat stores are essentially the only sources of metabolic energy during denning and even for a short time following emergence. Although corn is an excellent source of carbohydrates, damage to cornfields was low except in areas of complete mast failures. Bears used natural mast crops over agricultural crops. Rogers (1976) noted that agricultural damage increased during years of mast failures. Bears in areas of mast failures that did not have access to cornfields were in poor condition in winter as a consequence of a low digestible carbohydrate diet of grasses in fall. In early February following a poor mast year, bears were emaciated and weights were typically 25-40% lower than weights of the same bears following an abundant mast year. Den weights of adult female bears in good condition ranged from 54 to 100 kg, and no threshold weight was evident between those females that produced cubs and those that did not. However, no female in our study that appeared emaciated or in other than good condition produced cubs. Female bears in Massachusetts reach 80-90% of their adult maximum mean weight by 4 years of age (K. D. Elowe, unpubl. data). Thus, additional growth of young adult females cannot account for either the wide variability of weights of females in good condition or the differences observed between females in poor versus good condition. Poor nutritional condition, such as a negative energy balance that might occur during severe mast failures, may cause the hypothalamus to direct nutrients away from reproduction via control of growth hormone release from the anterior pituitary (Jainudeen and Hafez 1980, Kaltenbach and Dunn 1980). Lack of nutritional support for pregnancy may cause inhibition of luteinizing hormone (LH)-releasing hormone (RH) release from the hypothalamus, restricting LH release from the anterior pituitary. Because

maintenance of the corpus luteum and its production of progesterone are dependent on LH, lower levels may cause termination of pregnancy and resorption of the conceptus through imbalanced uterine progesterone-estrogen levels (Mayer 1963, Nutting and Meyer 1963). Blood analyses have been used by some researchers to attempt to determine nutritional and reproductive condition of deer (Seal et al. 1972, Warren et al. 1982). While blood indicators of protein metabolism (i.e., blood urea nitrogen and urea/creatinine ratios) may be helpful in distinguishing effects of high or low protein diets in deer, it appears to have little significance for determining the nutritional plane of bears during winter. Because bears use triglycerides and glycerols from fat tissue as a main winter metabolic pathway (Nelson et al. 1983), triglyceride levels in the blood may indicate the presence or absence of available fat stores. However, no relationship was found between fall diets and 22 winter blood serum parameters, including triglycerides, or between females that produced cubs and potential mothers that did not produce cubs (Elowe 1987). Differential mortality of male and female cubs <1 year old was not easily explained. The difference was even more pronounced if cubs that died of abandonment (3 M and 2 F) and mother dying (1 M and 2 F), events affecting both sexes equally, were separated from the sample; males accounted for all first-year mortality (n = 9). Research handling accounted for 2 of 3 mortalities to subadult females, further indicating a disparity in natural vulnerability between young male and female bears. There was no evidence of differential parental investment towards male or female cubs that might result in differential mortality (Trivers and Willard 1973). Although nursing rates were not known, there was no evidence that male and female cubs were not in company of the mother equally. Low nutritional condition and other environmental stresses have been shown to affect males with higher metabolic rate (e.g., humans, rats [Rattus spp.], and red deer [Cervus elaphus]) more than females, possibly causing increased vulnerability to starvation and disease (Hamilton 1948, Schein 1950, Clutton-Brock et al. 1982). Clutton-Brock et al. (1985) suggest that in polygynous species where male size and fitness increases reproductive success, increased male growth rates may have been selected for

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at the cost of higher juvenile mortality. Known breeding habits of bears and our mortality data support this theory. Stomachs of male cubs found dead during our study frequently contained milk, and no deaths could be attributed to starvation. However, other factors may have affected male cub mortality indirectly. Milk is the primary food of cubs from January until at least early June. Four cubs from 2 different families that were separated from their mothers in mid-June survived (1 mother died, and the other separation was of unknown cause). Other cubs were commonly observed eating herbaceous matter as early as the first week of May, but most continued nursing to some extent into August. Jenness (1985) stated that milk composition is greatly influenced by diet. Early spring nutritional condition of lactating female bears depends, to a large extent, on fall nutrition. Thus, the presence or absence of adequate fat stores may influence the quality or quantity of milk produced and could have a significant effect on the health of cubs. Wathen (1983) reported that low fall food supplies increased cub mortality the following spring. However, in our study, the 2 years of highest cub mortality (9 of 14 and 8 of 12) followed years of very abundant mast and high cub production. Lowest cub mortality (1 of 6) followed a year when mast crops failed and only females feeding in corn produced cubs. Wathen's (1983) observation of increased cub mortality following mast failures may be correct for females in marginal condition that produce cubs. Bears in Massachusetts, however, appeared to require a high nutritional threshold before producing cubs and therefore, no data were available on cubs born to mothers in marginal or poor condition. Dispersal of male cubs after family breakup caused them to become more vulnerable to human-caused mortality. Hunting was the only cause of mortality in male cubs >1 year old. Extreme movements in unfamiliar areas made road crossings and human contact inevitable. Because of the promiscuous breeding nature of bears, however, it is improbable that a high male cub mortality rate has any significant adverse effects on population reproductive success. Black bears in Massachusetts do not appear to respond to lower population densities by increased reproduction or cub survival as suggested by Lindzey et al. (1983) in Pennsylvania.

Nutritional factors that govern reproductive rate, social structure of the Massachusetts population, dispersion of food sources, and bear movement data (Elowe 1984) indicate that competition for food or space is not an important factor in increasing reproductive success. Mast crops are usually in widespread abundance or widespread scarcity and there was no evidence of competition for fall foods at estimated density levels of bears in Massachusetts (1 bear/7 km2 [Elowe 1984]). At present mortality and exploitation levels in Massachusetts and bordering states, 13 of 20 females versus only 2 of 21 males lived to maturity. Fluctuations in fall mast crops, then, appeared to influence the density of bears through the number of female cubs born.

LITERATURE CITED
BUMP, G., R. W. DARROW,F. C. EDMINSTER, AND

W. F. CRISSEY. 1947. The ruffed grouse. N.Y. State Conserv. Dep. 915pp.
CLUTTON-BROCK, T. H., S. D. ALBON, AND F. E.

GUINESS. 1985.

Parental investment and sex

differences in juvenile mortality in birds and mammals. Nature 313:131-133.

-, F. E. GUINESS, AND S. D. ALBON. 1982. Red

deer: behavior and ecology of two sexes. Univ. of Chicago Press, Ill. 378pp. T. C., ANDM. R. PELTON.1983. Seasonal EAGLE, nutrition of black bears in the Great Smoky MountainsNational Park.int. Conf. Bear Res. and Manage.5:94-105. EILER, J. H. 1981. Reproductive biology of black bearsin the SmokyMountains of Tennessee.M.S. Thesis, Univ. Tennessee, Knoxville. 127pp. K. D. 1984. Home range, movements,and ELOWE, habitatpreferencesof blackbears (Ursus americanus) in western Massachusetts.M.S. Thesis, Univ. Massachusetts, Amherst.112pp. . 1987. Factors affecting black bear reproductive success and cub survival in Massachusetts. Ph.D. Thesis, Univ. Massachusetts,Amherst. 71pp. HAMILTON, J. B. 1948. The role of testicularsecretions as indicated by the effects of castrationin man and by studies of pathological conditions and the short lifespan associatedwith maleness. Recent Prog. Hormone Res. 3:257-322. productive failure in females. Pages 449-470 in E. S. E. Hafez, ed. Reproduction in farmanimals. Lea and Febiger, Philadelphia,Pa. R. 1985. Biochemical and nutritionalasJENNESS, pects of milk and colostrum. Pages 164-197 in B. L. Larson, ed. Lactation. Iowa State Univ. Press,Ames.
KALTENBACH, C. C., AND T. G. DUNN. 1980. En-

M. R., AND E. S. E. HAFEZ. 1980. ReJAINUDEEN,

docrinologyof reproduction.Pages 85-113 in E. S. E. Hafez, ed. Reproductionof farm animals. Lea and Febiger, Philadelphia,Pa.

This content downloaded from 71.85.136.238 on Tue, 10 Dec 2013 16:27:11 PM All use subject to JSTOR Terms and Conditions

968

* Elowe and Dodge AND CUB SURVIVAL BEARREPRODUCTION

J. Wildl. Manage. 53(4):1989

KING, T. R., AND H. E. McLURE. 1944. Chemical composition of some American wild feedstuffs. J. Agric. Res. 69:33-46. LECOUNT, A. L. 1987. Causes of black bear cub mortality. Int. Conf. Bear Res. and Manage. 7: 75-82. AND LINDZEY, J. S., G. L. ALT, C. R. MCLAUGHLIN, W. S. KORDEK. 1983. Population response of Pennsylvania black bears to hunting. Int. Conf. Bear Res. and Manage. 5:324-339. W. P., AND W. NIEDZWIEDZ. 1975. MACCONNELL, Remote sensing 20 years of change in Berkshire County, Massachusetts, 1951-1971. Mass. Agric. Exp. Stn. Res. Bull. 629. 117pp. G. 1963. Delayed nidation in rats: a method MAYER, of exploring the mechanisms of ovo-implantation. Pages 213-228 in A. C. Enders, ed. Delayed implantation. Univ. Chicago Press, Ill. NELSON,R. A., G. E. FOLK, JR., E. W. PFEIFFER, J. C. J. JONKEL, ANDD. L. STEIGER. J. CRAIGHEAD, 1983. Behavior, biochemistry, and hibernation in black, grizzly, and polar bears. Int. Conf. Bear Res. and Manage. 5:284-290. NUTTING,E. F., AND R. K. MEYER. 1963. Implantation delay, nidation, and embryonal survival in rats treated with ovarian hormones. Pages 233251 in A. C. Enders, ed. Delayed implantation. Univ. Chicago Press, Chicago, Ill. D. G., ANDJ. J. BEECHAM. 1980. Home REYNOLDS, range activities and reproduction of black bears in west-central Idaho. Int. Conf. Bear Res. and Manage. 4:181-190. L. L. 1976. Effects of mast and berry crop ROGERS, failures on survival, growth, and reproductive

success of black bears. Trans. North Am. Wildl. and Nat. Resour. Conf. 41:431-438. 1. 1977. Social relationships, movements, and population dynamics of black bears in northeastern Minnesota. Ph.D. Thesis, Univ. Minnesota, Minneapolis. 194pp. SCHEIN, M. W. 1950. The relation of sex ratio to physiological age in the wild brown rat. Am. Nat. 84:489-496. SEAL, U. S., L. J. VERME,J. J. OZOGA,AND A. W. ERICKSON.1972. Nutritional effects on thyroid activity and blood of white-tailed deer. J. Wildl. Manage. 36:1041-1052. SHORT,H. L., AND E. A. EPPS,JR. 1976. Nutrient quality and digestibility of seeds and fruits from southern forests. J. Wildl. Manage. 40:283-289. TRIVERS, R. L., AND D. E. WILLARD. 1973. Natural selection of parental ability to vary the sex ratio of offspring. Science 179:90-92. WARREN, R. J., R. L. KIRKPATRICK, A. OELSCHLAEGER, P. F. SCANLON, K. E. WEBB, JR., AND J. B.

WHELAN. 1982. Energy, protein, and seasonal influences on white-tailed deer fawn nutritional indices. J. Wildl. Manage. 46:302-312. WATHEN, W. G. 1983. Reproduction and denning of black bears in the Great Smoky Mountains. M.S. Thesis, Univ. Tennessee, Knoxville. 135pp. WILLEY, C. H. 1974. Aging black bears form first premolar tooth sections. J. Wildl. Manage. 38: 97-100.

Received 3 July 1986. Accepted 6 January 1989.

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