Palaeoecology of cave bears as evidenced by dental wear analysis: a review of methods and recent findings

Abstract

The study of dental wear was first used years ago to infer the palaeoecology of fossil mammals and in particular their diet. Results depend predominantly on the scale of the analysis used. Analyses of dental macrowear, mesowear or microwear do not provide the same type of dietary information, be it about the seasonal, annual or lifetime diet. This contribution focuses on emblematic species, cave bears (Ursidae), in particular Ursus spelaeus spelaeus. Methods used by previous researchers to infer their dietary preferences and thus their palaeoecology are reviewed and compared. This review is complemented by an analysis of several specimens of cave bears from the Goyet cave in Belgium, using dental microwear texture analysis (DMTA), a methodology widely applied for reconstructing palaeodiets. Three main conclusions are drawn here: (1) DMTA is the method that provides the most precise palaeobiological inferences; (2) during the pre-dormancy period, cave bears show dietary flexibility; (3) dental wear alone might be not sufficient to provide a complete reconstruction of the cave bear palaeodiet.     

Blood chemistry of black bears from Pennsylvania during winter dormancy

Twenty-four serum chemistries were measured in blood samples collected from 20 adult female black bears (Ursus americanus) and their offspring, including 14 yearlings and 37 cubs, in northeastern Pennsylvania during winter 1984. Four other captive adult females were bled before, during, and after they were subjected to unseasonably warm temperatures during February. Levels of serum urea nitrogen (SUN) and creatinine were lower (P less than 0.05), and iron was higher (P less than 0.05) in male cubs compared to female cubs; serum chemistries were similar (P greater than or equal to 0.05) between sexes for yearlings. Total protein, albumin and creatinine levels increased with age of bears, whereas chloride, alkaline phosphatase, potassium, inorganic phosphorus and SUN/creatinine were higher (P less than 0.05) in cubs than in yearlings and adults. The relatively high serum calcium in cubs was probably related to rapid bone development and dietary intake of calcium during winter dormancy in cubs. Low serum calcium in adults was attributed to lactation and a lack of dietary intake. Urea/creatinine ratios averaged 5.5 and 4.6 for yearling females and males, respectively, 6.3 for adult females, and 29.0 and 22.8 for female and male cubs, respectively. Levels of serum chemistries of black bears apparently are relatively stable during winter denning, when bears are without food or water and do not urinate or defecate for several months. This stability indicates that black bears are resistant to the extremes in extrinsic environmental conditions. Abnormal blood chemistry values may indicate metabolic stresses that are not being controlled by bears.(ABSTRACT TRUNCATED AT 250 WORDS)     

Behavioural Differences between Single Scandinavian Brown Bears (Ursus arctos) and Females with Dependent Young When Experimentally Approached by Humans

Carnivore-human encounters that result in human injury present a conservation and management challenge and it is therefore important to understand under what conditions such incidents occur. Females with cubs are often involved when humans are injured by brown bears Ursus arctos. In Scandinavia, this is particularly true for unarmed recreational forest users. Our aim was to document behavioural differences between single bears and females with cubs in order to develop recommendations to minimize the risk of injuries to recreational forest users. We documented the reactions of GPS-collared females with cubs and single brown bears to experimental approaches by humans to 50 m from the bear on 42 and 108 occasions, respectively. The majority of females with cubs (95%) and single bears (89%) left when approached. Bears that left were passed at shorter distances and were in more open areas than those that stayed. Both groups had similar flight initiation distances, which were longer for bears that were active at the time of the disturbance. Females with cubs selected more open habitat than single bears, also for the new site they selected following disturbance. Females with cubs, particularly active females with cubs of the year, moved greater distances and spent more time active following the approach. Females with cubs and single bears were seen or heard in 26% and 14% of the approaches, respectively. None of the bears displayed any aggressive behaviour during the approaches. Females with cubs selected more open habitat, perhaps predisposing them to encountering people that are not involved in hunting activities, which might be the primary explanation why females with cubs are most frequently involved when unarmed people are injured by bears in Scandinavia. To mitigate injury risks, one must consider factors that bring bears closer to human activity in the first place.     

The cave bear’s hibernation: reconstructing the physiology and behaviour of an extinct animal

Abstract

When studying an extinct species such as the cave bear (Ursus spelaeus ROSENMÜLLER 1794), it is possible to apply a variety of molecular biology techniques such as the study of stable isotopes or mitochondrial DNA (mDNA) to infer patterns of behaviour or physiology that would otherwise remain concealed. Throughout Europe and along time, differences in the isotopic values (δ¹³C and δ¹⁵N) of cave bears arise from environmental differences and the Pleistocene climatic evolution. The climate determines the hibernation length, during which the cave bears undergo a particular physiology that can be related to an increase in δ¹⁵N during climate cooling. In order to verify whether hibernation affected the isotopic values, we compared cave bears in different ontogenetic stages. The results show that perinatal values reflect the values for mothers during hibernation, while juveniles show differences in maternal investment. A previous study in the literature based on complete mitochondrial DNA sequences of several individuals collected from closely situated caves showed that each cave housed, almost exclusively, a single lineage of haplotypes. This pattern suggests extreme fidelity to the birth site, or homing behaviour, and that cave bears formed stable maternal social groups, at least for the purpose of hibernation. Studies of this type offer unexpected data on the palaeobiology of this extinct animal.     

Changes in the intramolecular stable carbon isotope ratios with age of the European cave bear (Ursus spelaeus)

Changes with age in the diet and metabolism of the extinct European cave bear (Ursus spelaeus) from the Divje Babe archaeological site in northwestern Slovenia were investigated by comparing the stable carbon isotope ratio of whole bone collagen with that of the peptide-bonded carboxyl carbons in collagen. These carboxyl carbons were selectively released by decarboxylation of the collagen hydrolysate with ninhydrin (2,2-dihydroxy-1,3-indanedione). The stable carbon isotope ratio increased with age for both the peptide-bonded and other carbons. Although only one-third of the carbon atoms in collagen are peptide-bonded, they account for most of the observed change with age of the stable carbon isotope ratio of the whole collagen. Stable nitrogen isotope ratios decreased with age and were negatively correlated with changes in the stable carbon isotope ratios. These observations support previous investigations on the unique metabolism of hibernating bears.     

Chronological and Isotopic data support a revision for the timing of cave bear extinction in Mediterranean Europe

Abstract

The Cave Bear, Ursus spelaeus (sensu lato), was one of many megafaunal species that became extinct during the Late Pleistocene in Europe. With new data we revisit the debate about the extinction and paleoecology of this species by presenting new chronometric, isotopic and taphonomic evidence from two Palaeolithic cave bear sites in northeastern Italy: Paina Cave and Trene Cave. Two direct radiocarbon dates on well-preserved collagen have yielded ages around 24,200–23,500 cal yr BP, which make them the latest known representatives of the species in Europe. The carbon (δ¹³C) and nitrogen (δ¹⁵N) isotopic values of bone collagen exhibit values similar to those of older cave bears from Swabian Jura and France, suggesting that the feedings preferences of cave bears remained unchanged until the disappearance of this species in Europe. Several bear remains preserved traces of human modification such as cut marks, which enables a reconstruction of the main steps of fur recovery and the butchering process. The broad range of plant types available and the favorable location of Berici Hills may have played an important role in the range expansion of cave bears and their interaction with the Paleolithic hunters settled the same area.     

Resolving temporal variation in vertebrate diets using naturally occurring stable isotopes

Assessments of temporal variation in diets are important for our understanding of the ecology of many vertebrates. Ratios of naturally occurring stable isotopes in animal tissues are a combination of the source elements and tissue specific fractionation processes, and can thus reveal dietary information. We review three different approaches that have been used to resolve temporal diet variation through analysis of stable isotopes. The most straightforward approach is to compare samples from the same type of tissue that has been sampled over time. This approach is suited to address either long or short-term dietary variation, depending on sample regime and which tissue that is sampled. Second, one can compare tissues with different metabolic rates. Since the elements in a given tissue have been assimilating during time spans specific to its metabolic rate, tissues with different metabolic rates will reflect dietary records over different periods. Third, comparisons of sections from tissues with progressive growth, such as hair, feathers, claws and teeth, will reveal temporal variation since these tissues will retain isotopic values in a chronological order. These latter two approaches are mainly suited to address questions regarding intermediate and short-term dietary variation. Knowledge of tissue specific metabolic rates, which determine the molecular turnover for a specific tissue, is of central importance for all these comparisons. Estimates of isotopic fractionation between source and measured target are important if specific hypotheses regarding the source elements are addressed. Estimates of isotopic fractionation, or at least of differences in fractionation between tissues, are necessary if different tissues are compared. We urge for more laboratory experiments aimed at improving our understanding of differential assimilation of dietary components, isotopic fractionation and metabolic routing. We further encourage more studies on reptiles and amphibians, and generally more studies utilizing multiple tissues with different turnover rates.     

Ancient DNA reveals differences in behaviour and sociality between brown bears and extinct cave bears

Ancient DNA studies have revolutionized the study of extinct species and populations, providing insights on phylogeny, phylogeography, admixture and demographic history. However, inferences on behaviour and sociality have been far less frequent. Here, we investigate the complete mitochondrial genomes of extinct Late Pleistocene cave bears and middle Holocene brown bears that each inhabited multiple geographically proximate caves in northern Spain. In cave bears, we find that, although most caves were occupied simultaneously, each cave almost exclusively contains a unique lineage of closely related haplotypes. This remarkable pattern suggests extreme fidelity to their birth site in cave bears, best described as homing behaviour, and that cave bears formed stable maternal social groups at least for hibernation. In contrast, brown bears do not show any strong association of mitochondrial lineage and cave, suggesting that these two closely related species differed in aspects of their behaviour and sociality. This difference is likely to have contributed to cave bear extinction, which occurred at a time in which competition for caves between bears and humans was likely intense and the ability to rapidly colonize new hibernation sites would have been crucial for the survival of a species so dependent on caves for hibernation as cave bears. Our study demonstrates the potential of ancient DNA to uncover patterns of behaviour and sociality in ancient species and populations, even those that went extinct many tens of thousands of years ago.     

Zinc Isotope Ratios as Indicators of Diet and Trophic Level in Arctic Marine Mammals

Carbon and nitrogen stable isotope ratios of bone collagen are an established method for dietary reconstruction, but this method is limited by the protein preservation. Zinc (Zn) is found in bioapatite and the isotopic compositions of this element constitute a very promising dietary indicator. The extent of fractionation of Zn isotopes in marine environments, however, remains unknown. We report here on the measurement of zinc, carbon and nitrogen isotopes in 47 marine mammals from the archaeological site of Arvik in the Canadian Arctic. We undertook this study to test and demonstrate the utility of Zn isotopes in recent mammal bone minerals as a dietary indicator by comparing them to other isotopic dietary tracers. We found a correlation between δ⁶⁶Zn values and trophic level for most species, with the exception of walruses, which may be caused by their large seasonal movements. δ⁶Zn values can therefore be used as a dietary indicator in marine ecosystems for both modern and recent mammals.     

The impact of metabolism on stable isotope dynamics: a theoretical framework

Stable isotope analysis is a powerful tool used for reconstructing individual life histories, identifying food-web structures and tracking flow of elemental matter through ecosystems. The mechanisms determining isotopic incorporation rates and discrimination factors are, however, poorly understood which hinders a reliable interpretation of field data when no experimental data are available. Here, we extend dynamic energy budget (DEB) theory with a limited set of new assumptions and rules in order to study the impact of metabolism on stable isotope dynamics in a mechanistic way. We calculate fluxes of stable isotopes within an organism by following fluxes of molecules involved in a limited number of macrochemical reactions: assimilation, growth but also structure turnover that is here explicitly treated. Two mechanisms are involved in the discrimination of isotopes: (i) selection of molecules occurs at the partitioning of assimilation, growth and turnover into anabolic and catabolic sub-fluxes and (ii) reshuffling of atoms occurs during transformations. Such a framework allows for isotopic routing which is known as a key, but poorly studied, mechanism. As DEB theory specifies the impact of environmental conditions and individual state on molecule fluxes, we discuss how scenario analysis within this framework could help reveal common mechanisms across taxa.     

Milk intake in blue fox (Alopex lagopus) and silver fox (Vulpes vulpes) cubs in the early suckling period

Milk intake of fox cubs (2-16 days of age; body weight, 96-649 g) in ten blue fox litters and ten silver fox litters were measured by the water isotope dilution (WID) technique following a single intraperitoneal injection of tritiated water (3HHO). Litter size varied from four to 14 in blue foxes and from three to eight in silver foxes. Silver fox cubs had higher birth weights than blue foxes. Inter-species body weights and growth rates were apparently dependent on litter size and the dam's constitution. In both species growth rate increased with age and body weight (7-35 g per day). In the cubs, the biological half-life of body water turnover (BWT) rose from 1.5 days at 2-3 days of age to 2.5 days at 13-16 days of age, although a considerable scatter was seen. The mean daily milk intake of the cubs varied with body weight, from 31 to 193 g per day, whereas daily milk intake per unit of body mass remained stable at 30-35 g per 100 g body weight. The ratio of milk intake to body weight gain varied considerably among cubs, averaging 4.5 g/g during the 3-week experimental period. In suckling fox cubs, the calculated daily intake of metabolically energy (ME) corresponded fairly with the estimated energy requirements for growth and maintenance of the young. Finally, the applicability and the accuracy of the WID technique was evaluated in ten 3-week-old fox cubs, by tube-feeding with a milk replacer for 48 h, which documented that the daily rates of milk intake and water turnover can be accurately measured in suckling fox cubs by the WID technique following a single injection of 3HHO.     

An Overview of the Ichnological and Ethological Studies in the Cave Bear Den in Urşilor Cave (Western Carpathians,Romania)

This paper presents a preliminary investigation of late Pleistocene cave bear traces from Ur?ilor Cave in the western Carpathians, Romania. The bears left thousands of traces on the walls, plateaus, and slopes of the cave interior. Some areas in the cave have been heavily trampled, leaving more than 140 hibernation beds as well as fur impressions. The footprints of cave bears are assigned to Ursichnus europaeus nov. ichnogen. and nov. ichnosp. and the cave bear-beds to Ursalveolus carpathicus nov. ichnogen. and nov. ichnosp. as behavioral traces. Tens of thousands of scratch marks on the slopes and top of a clay plateau are the result of bears moving from the hibernation area down to a stream and back. These traces reveal that the cave bears had short claws, similar to those of modern, primarily herbivorous black bears. Deep within the cave, three weathered, articulated cave bear skeletons still lie in their hibernation beds on the clay plateaus or a natural cave corner. One of these bears was a one-year-old male cub that did not survive its first hibernation; a second skeleton close was an adult female. The third, a young male, was found close to the end of the cave system. The bears of Ur?ilor Cave would have felt well protected against carnivores during their hibernation because their sleeping places were so deep within the cave. Their strategy may well have been to avoid any conflict with hyenas and lions during hibernation.     

Short-term behavioural response of polar bears (<Emphasis Type="Italic">Ursus maritimus</Emphasis>) to snowmobile disturbance

In this study the distance, at which polar bears detected and actively responded to approaching snowmobiles was measured and the behavioural response was recorded. The study was performed on Svalbard, an arctic island where human traffic has increased substantially in recent years. Fieldwork was conduced in April and/or May during the years 2003–2005. Polar bears were observed on ice with telescopes and binoculars. Undisturbed polar bears were observed continuously and their behaviours recorded, during the time when two snowmobiles moved toward the bear(s). Distances between the bear, the observer, and the approaching snowmobiles were measured using GPS positions taken on the track towards the bear. Data on the behavioural response of 20 encounters with bears were collected. On average, bears were alerted to the snowmobiles at 1,164 m. Mean distance at which the locomotive response occurred was 843 m, and there was a statistical significant difference in distance between sex and age classes [326 m (95% CI = 138–496 m) for adult males; 1,534 m (95% CI = 508–2,768 m) for adult females with cubs; 164 m (95% CI = 49–543 m) for two adult females without cubs; and 1,160 m (95% CI = 375–1,353 m) for single medium sized bears]. The responses of the polar bears to the snowmobiles were categorized according to intensity and persistence of reactions. Females with cubs and single medium sized bears tended to show more intense responses than adult males and lone adult females. Wind direction affects sound and odour transmission, and although an effect on response distance was not found, the response intensity was affected by wind direction. We conclude that female polar bears with small cubs in particular may have a greater risk to be disturbed, since they react at greater distances with amplified reactions; thus, users of snowmobiles should take particular care in areas where females with cubs are present.     

Survival and Conflict Behavior of American Black Bears after Rehabilitation

Wildlife agencies face difficult situations when orphaned or injured American black bear (Ursus americanus) cubs (<12 months old) or yearlings (≥12 and <24 months old) are captured. One option is bear rehabilitation, the care and feeding of cubs or yearlings in a semi-natural environment, followed by release. Unfortunately, the survival and movements of bears released from rehabilitation facilities are often poorly documented and the ultimate reasons for success or failure poorly understood. Our goal was to assess survival and post-release conflict of orphaned bear cubs and yearlings following release from a rehabilitation facility, Appalachian Bear Rescue (ABR), in Townsend, Tennessee, USA, from 2015–2016. We predicted that rehabilitated bears would survive at similar rates, die from similar causes, and engage in similar conflict behavior to wild conspecifics. We equipped 42 black bear cubs and yearlings from ABR with global positioning system-collars and released them in Great Smoky Mountains National Park or Cherokee National Forest, Tennessee and North Carolina, USA. Estimated annual survival using known-fate methods for all released bears was 0.93 ± 0.06 [SE]). Survival for 13 bears released as cubs was 0.64 ± 0.14, whereas none of the bears released as yearlings died within 1 year after release (n = 29). Survival of rehabilitated bears was similar to or higher than published rates for wild conspecifics. Three of 42 bears (7.1%) released from ABR engaged in conflict behavior up to 1 year following release, and those had spent time involved in conflict behavior with their mothers (e.g., approaching humans) prior to being orphaned. Despite not having the typical post-natal experience with their mothers, the bears in our study appeared to behave and survive similarly to their wild conspecifics. Rehabilitation is effective for managing orphaned or injured bears. Best survival occurred for bears released as yearlings; however, managers can maximize cub survival through fall releases when plentiful wild foods are available. © 2019 The Authors. The Journal of Wildlife Management published by Wiley Periodicals, Inc. on behalf of The Wildlife Society.     

Stable Isotope Analysis in Primatology: A Critical Review

Stable isotope analysis has become an important tool in ecology over the last 25 years. A wealth of ecological information is stored in animal tissues in the relative abundances of the stable isotopes of several elements, particularly carbon and nitrogen, because these isotopes navigate through ecological processes in predictable ways. Stable carbon and nitrogen isotopes have been measured in most primate taxonomic groups and have yielded information about dietary content, dietary variability, and habitat use. Stable isotopes have recently proven useful for addressing more fine‐grained questions about niche dynamics and anthropogenic effects on feeding ecology. Here, we discuss stable carbon and nitrogen isotope systematics and critically review the published stable carbon and nitrogen isotope data for modern primates with a focus on the problems and prospects for future stable isotope applications in primatology. Am. J. Primatol. 74:969‐989, 2012. © 2012 Wiley Periodicals, Inc.     

Upper Pleistocene Panthera leo spelaea (Goldfuss, 1810) remains from the Bilstein Caves (Sauerland Karst) and contribution to the steppe lion taphonomy,palaeobiology and sexual dimorphism

Remains of the steppe lion Panthera leo spelaea (Goldfuss) from historical digs in the Bilstein Caves of Warstein (Sauerland, NW Germany) are described. Their age seems to be from the Early Weichselian periods (Upper Pleistocene). Whereas the Bilstein cave was inhabited by cave bears at that time only a few hyena prey remains, were most likely imported into the cave entrance by hyenas. Bite and crush marks on a few bones of Bison priscus, Bos primigenius, Cervus elaphus, a rhinoceros Coelodonta antiquitatis vertebra and even several chewed cave bear bones prove the hyena presence which is similar to other caves in the Sauerland hyena den cave rich region. Additionally some larger wolves subspecies Canis lupusspelaeus bones were found, but only few Crocuta crocuta spelaea remains are present. After taphonomic comparisons to six other hyena and cave bear den caves of northern Germany, this cave can be classified as a cave bear den, which was briefly used by hyenas only for food storage or commuting or cave bear predation site in one part of the Cave. The lion material refers at least to one young adult lioness, one more adult female and two male lions; therefore, at minimum, the remains of four adult individuals are represented. The absence of juvenile lion material, in contrast to cave bear cub remains in the Bilstein Caves, proves that P. leo spelaea did not use this and all other caves in the region to raise their cubs. The bone material from the Bilstein Caves would prove the same hyena-lion antagonism conflict being recently proven for the Perick Caves, Balve Cave or Martins Cave well. Other situations in caves such as the Keppler Cave and the Bilstein Cave initially show the more complex taphonomic situation of lion remains in European caves, especially in cave bear dens, where they seem to have hunted periodically cave bears, such as it is already proven for hyenas in the Sauerland Karst and other caves of Europe.     

Comparison of Acetate Turnover in Methanogenic and Sulfate-Reducing Sediments by Radiolabeling and Stable Isotope Labeling and by Use of Specific Inhibitors: Evidence for Isotopic Exchange

Acetate turnover in the methanogenic freshwater anoxic sediments of Lake Vechten, The Netherlands, and in anoxic sediments from the Tamar Estuary, United Kingdom, and the Grosser Jasmunder Bodden, Germany, the latter two dominated by sulfate reduction, was determined. Stable isotopes and radioisotopes, inhibitors (chloroform and fluoroacetate), and methane flux were used to provide independent estimates of acetate turnover. Pore water acetate pool sizes were determined by gas chromatography with a flame ionization detector, and stable isotope-labeled acetate was determined by gas chromatography-mass spectrometry. The appearance of acetates with a different isotope labeling pattern from that initially added demonstrated that isotopic exchange occurred during methanogenic acetate metabolism. The predominant exchange processes were (i) D-H exchange in the methyl group and (ii) (sup13)C-(sup12)C exchange at the carboxyl carbon. These exchanges are most probably caused by the activity of the enzyme complex carbon monoxide dehydrogenase and subsequent methyl group dehydrogenation by tetrahydromethanopterine or a related enzyme. The methyl carbon was not subject to exchange during transformation to methane, and hence acetate with the methyl carbon labeled will provide the most reliable estimate of acetate turnover to methane. Acetate turnover rate estimates with these labels were consistent with independent estimates of acetate turnover (acetate accumulation after inhibition and methane flux). Turnover rates from either radioisotope- or stable isotope-labeled methyl carbon isotopes are, however, dependent on accurate determination of the acetate pool size. The additions of large amounts of stable isotope-labeled acetate elevate the acetate pool size, stimulating acetate consumption and causing deviation from steady-state kinetics. This can, however, be overcome by the application of a non-steady-state model. Isotopic exchange in sediments dominated by sulfate reduction was minimal.     

Seasonal and sex differences in the structure and chemical composition of adipose tissue in wild polar bears (Ursus maritimus)

Biopsy samples of adipose tissue from the upper thigh were collected in spring and in late summer/autumn from 370 wild polar bears ( Ursus maritimus ), including adult females with and without cubs, adult males and juveniles. Mean adipocyte volume was measured from all samples and chemical assays of the lipid, total protein and collagen were also performed on samples from 53 bears. Mean adipocyte volume was smaller in all specimens in spring than in late summer/autumn, but the differences were greatest for solitary adult females. The range of adipocyte volumes was much greater for adult females than for adult males, and in females only, mean adipocyte volume correlated significantly with total body mass. Therefore, adipocyte volume measurements from biopsy samples provide some information about fatness in adult females, but are worthless as an indicator of body composition in males and juveniles. In juveniles and females, but not adult males, the lipid content of the adipose tissue was up to 18% lower in autumn than in spring. The collagen content was significantly higher in autumn than in spring in all bears except females with cubs. We suggest that these differences in chemical composition arise from accumulation of water within and between the adipocytes, which would minimize tissue shrinkage, and from changes in the vascularization of the adipose tissue. These properties may be adaptations to rapid fattening and prolonged fasting and the sex differences may reflect the contrasting reproductive strategy of female and male polar bears.     

GRIZZLY BEAR DEMOGRAPHICS IN AND AROUND BANFF NATIONAL PARK AND KANANASKIS COUNTRY,ALBERTA

Abstract: The area in and around Banff National Park (BNP) in southwestern Alberta, Canada, is 1 of the most heavily used and developed areas where grizzly bears (Ursus arctos) still exist. During 1994–2002, we radiomarked and monitored 37 female and 34 male bears in this area to estimate rates of survival, reproduction, and population growth. Annual survival rates of bears other than dependent young averaged 95% for females and 81–85% for males. Although this area was largely unhunted, humans caused 75% of female mortality and 86% of male mortality. Females produced their first surviving litter at 6–12 years of age (x̄ = 8.4 years). Litters averaged 1.84 cubs spaced at 4.4-year intervals. Adult (≥6-years-old) females produced 0.24 female cubs per year and were expected to produce an average of 1.7 female cubs in their lifetime, based on rates of reproduction and survival. Cub survival was 79%, yearling survival was 91%, and survival through independence at 2.5–5.5 years of age was 72%, as no dependent young older than yearlings died. Although this is the slowest-reproducing grizzly bear population yet studied, high rates of survival seem to have enabled positive population growth (Λ = 1.04, 95% CI = 0.99–1.09), based on analyses using Leslie matrices. Current management practices, instituted in the late 1980s, focus on alleviating human-caused bear mortality. If the 1970–1980s style of management had continued, we estimated that an average of 1 more radiomarked female would have been killed each year, reducing female survival to the point that the population would have declined.     

Reproductive Ecology and Cub Survival of Florida Black Bears

Abstract: We investigated reproductive ecology and cub survival of Florida black bears (Ursus americanus floridanus) in Ocala National Forest and the adjacent residential area of Lynne, Florida, USA, 1999-2003. We documented production of 81 cubs from 39 litters. Average litter size was 2.08 ± 0.11 (SE) cubs. The mean age of first reproduction was 3.25 ± 0.27 years. Excluding females that reproduced in consecutive years due to litter loss, interlitter interval was 2.11 ± 0.11 years. The mean annual fecundity rate was 0.57 ± 0.06. We used expandable radiocollars to monitor the fate of 41 bear cubs. The probability of cubs surviving to 9 months of age was 0.46 ± 0.09 and did not differ between cohorts or study locations. The most important causes of cub mortality included infanticide and mortality caused directly or indirectly by collisions with vehicles. Our results indicate that reproductive rates of female black bears in the Ocala study area are comparable to those reported for other black bear populations from eastern United States, but cub survival rates are lower than those reported for most black bear populations. Management of Florida black bears should emphasize strategies to reduce the mortality of cubs.