Condition-dependent reproductive success in bighorn ewes

Individual bighorn ewes ( Ovis canadensis ) at 4–14 years of age were 1.5% heavier preceding years when they weaned a lamb than preceding years when their lamb died before weaning. Intra-individual differences in mass between years of successful and unsuccessful reproduction appeared independent of a ewe's multiyear average mass. Relative mass loss both in the winter before and in the winter after a given repro-ductive episode increased with reproductive success. Long-term monitoring of individual mass and reproductive success is a promising technique to study life histories in capital breeders, because it allows to partially account for differences in reproductive potential.     

Predator-induced natural selection on temperament in bighorn ewes

Temperament traits in animals may have important fitness consequences, but have received little attention from ecologists or evolutionary biologists. A few studies have linked variation in temperament with fitness, but none has measured selection on temperament traits. We estimated the strength of selection on female boldness and docility on bighorn sheep ewes, Ovis canadensis. The Ram Mountain population experienced little predation pressure during the first 25 years of study, then 2 years (1997 and 1998) of frequent predation by cougars, Puma concolor, during which adult ewe mortality almost tripled over the long-term average, to 27% a year. During years of high predation, we found moderate selection favouring bold ewes, and age-specific selection on docility. Old ewes appeared more vulnerable to predation than young ewes. In contrast, no evidence of selection on temperament traits was observed during 2 years of low predation (1996 and 1999). These results suggest predator-induced selection favouring bold and nondocile ewes. Leadership was highly correlated with age and may increase the risk of predator encounter. Leadership alone, however, could not explain the higher vulnerability of old ewes to predation. Cougar predation on bighorn sheep occurs sporadically and unpredictably, probably because individual cougars often are prey specialists. Cougar predation may have limited microevolutionary effects on temperament in bighorn sheep, because it mostly affects ewes near the end of their reproductive life span and because of potential countervailing selection on boldness and docility. Copyright 2003 Published by Elsevier Science Ltd on behalf of The Association for the Study of Animal Behaviour.      

Selfish mothers: reproductive expenditure and resource availability in bighorn ewes

We present a marginal value model explaining intraspecific andinterspecific variation of mating systems in nonterritorialungulates. The model takes into account the simultaneous effectsof spatial and temporal distribution of females, female groupsize, male-male competition, female choice, and the operationalsex ratio (i.e., the proportion of estrous females). The modelpredicts that higher numbers of females per group increasesthe average exploitation time of such groups by males. An increasein female group density, operational sex ratio, and age-specificfighting success of males are predicted to reduce the averageexploitation time of female groups, leading to roving of males(i.e., moving between female groups). In contrast, an increasein die female rejection rate of males and in the time spentby males on nonmating activities (i. e., foraging, lying down,ruminating, migrating) are predicted to increase the averageexploitation time of female groups and to favor staying behaviorof males (i.e., defending a female group over a longer periodof time). Consequently, die model predicts that young maleswill tend to be "stayers," whereas middle-aged and old malesare expected to be "rovers." Model predictions were tested widifield data collected on muskoxen Ovibos moschatus in a naturalpopulation in west Greenland. Observed correlations betweenbull exploitation times of cow groups and the six above-mentionedsocial and environmental parameters were all in die predicteddirection and statistically significant in five of die six cases.Overall, 69% of die observed variation in exploitation timeof cow groups by males was explained by die model predictions.Stepwise regression suggested that, of die six parameters, variationin sex ratio (R² = .56) and time spent on nonmating activities(R² = .35) had the largest effects on male exploitation time.Also, die observed age-specific variation in bull exploitationtime of cow groups was as predicted.     

Consistency of temperament in bighorn ewes and correlates with behaviour and life history

Individual differences in temperament may affect how animals react to novel situations, avoid predation, invest in reproduction and behave in a variety of social contexts. Little information is available, however, about individual differences in temperament for wild animals. For bighorn sheep, Ovis canadensis, ewes captured as part of a long-term study, we compared behaviour during handling to behaviour in the field and reproductive history. We considered 'bold' ewes those that were frequently trapped during the summer, and assigned to each ewe a docility index based on her behaviour during handling. Measurements of temperament for the same individual at different captures were highly consistent. Temperament was not affected by reproductive status or age, nor was it related to body mass. Correlations between behaviour at the trap and in the field were weak and mostly nonsignificant, suggesting that temperament is domain specific rather than domain general. Bold ewes tended to start reproducing earlier and have higher weaning success than shy ewes. Variability in temperamental traits in the study population could be maintained by life-history trade-offs and by yearly changes in selective pressures. Copyright 2000 The Association for the Study of Animal Behaviour.     

Early primiparity in brown bears

We documented 2 cases of unusually early primiparity in brown bearsUrsus arctos Linneaus, 1758 in an introduced population in central Austria. Two females gave birth at the age of 3 years.     

Mass‐dependent reproductive strategies in wild bighorn ewes: a quantitative genetic approach

In the Ram Mountain bighorn sheep (Ovis canadensis) population, ewes differing by more than 30% in body mass weaned lambs with an average mass difference of only 3%. Variability in adult body mass was partly due to additive genetic effects, but inheritance of weaning mass was weak. Maternal effects could obscure genetic effects in the phenotypic expression of weaning mass, particularly if they reflected strategies of maternal expenditure that varied according to ewe mass. We performed a quantitative genetic analysis to assess genetic and environmental influences on ewe mass and on maternal expenditure. We used the mean daughters/mother regression method and Derivative Free Restricted Maximum Likelihood models to estimate heritability (h²) of ewe mass and indices of maternal expenditure. We found additive genetic effects on phenotypic variation in maternal mass, in lamb mass at weaning (absolute maternal expenditure) and in weaning mass relative to maternal mass at weaning (relative maternal expenditure). Heritability suggests that maternal expenditure has the potential to evolve. The genetic correlation of ewe mass and absolute maternal expenditure was weak, while ewe mass and relative maternal expenditure were strongly negatively correlated. These results suggest additive genetic effects on mass‐dependent reproductive strategies in bighorn ewes. Mass‐dependent reproductive strategies could affect lamb survival and phenotypic variation in adult mass. As population density increased and reproduction became costlier, small females reduced maternal expenditure more than large females. Constraints on reproductive strategy imposed by variations in resource availability are therefore likely to differ according to ewe mass. A general trend for a decrease in maternal expenditure relative to maternal size in mammals suggests that size‐dependent negative maternal effects may be common.     

Population subdivision among desert bighorn sheep (Oviscanadensis) ewes revealed by mitochondrial DNA analysis

We used behavioural observations and mitochondrial DNA (mtDNA) sequence analysis to examine demographic and genetic structure within and among home-range groups of desert bighorn sheep (Oviscanadensis) ewes in the Peninsular Ranges of southern California, USA. We identified substantial genetic variation in the first 515 bp of the mtDNA control region and determined that seven haplotypes were distributed in a nonrandom fashion among these ewe subpopulations. Although a significant (P < 0.01) amount of mtDNA variation (33%) was partitioned among home-range groups, we did not find strong evidence for matrilineal substructuring within these groups. Based on analyses of molecular variance, and comparisons of behavioural associations and distances between centres of activity, we concluded that within a given home-range group, bighorn sheep ewes generally associate with other ewes based on their availability rather than their matrilineal relationships. Our results also supported the conclusion that multiple ewe subpopulations exist within the Peninsular Ranges, and that these subpopulations are the most basic demographic and genetic units.     

Determinants and consequences of interspecific body size variation in tetraphyllidean tapeworms

Tetraphyllidean cestodes are cosmopolitan, remarkably host specific, and form the most speciose and diverse group of helminths infecting elasmobranchs (sharks, skates and rays). They show substantial interspecific variation in a variety of morphological traits, including body size. Tetraphyllideans represent therefore, an ideal group in which to examine the relationship between parasite body size and abundance. The individual and combined effects of host size, environmental temperature, host habitat, host environment, host physiology, and host type (all likely correlates of parasite body size) on parasite length were assessed using general linear model analyses using data from 515 tetraphyllidean cestode species (182 species were included in analyses). The relationships between tetraphyllidean cestode length and intensity and abundance of infection were assessed using simple linear regression analyses. Due to the contrasting morphologies between shark and batoid hosts, and contrasting physiologies between sharks of the Lamnidae family and other sharks, analyses were repeated in different subsets based on host morphology and physiologies (“sharks” vs. batoids) to determine the influence of these variables on adult tetraphyllidean tapeworm body size. Results presented herein indicate that host body size, environmental temperature and host habitat are relatively important variables in models explaining interspecific variations in tetraphyllidean tapeworm length. In addition, a negative relationship between tetraphyllidean body size and intensity of infection was apparent. These results suggest that space constraints and ambient temperature, via their effects on metabolism and growth, determine adult tetraphyllidean cestode size. Consequently, a trade-off between size and numbers is possibly imposed by external forces influencing host size, hence limiting physical space or other resources available to the parasites.     

Genomic consequences of genetic rescue in an insular population of bighorn sheep (Ovis canadensis)

Genetic rescue is a management intervention whereby a small population is supplemented with individuals from other populations in an attempt to reverse the effects of inbreeding and increased genetic load. One such rescue was recently documented in the population of bighorn sheep (Ovis canadensis) within the National Bison Range wildlife refuge (Montana, USA). Here, we examine the locus-specific effects of rescue in this population using a newly developed genome-wide set of 195 microsatellite loci and first-generation linkage map. We found that the rate of introgression varied among loci and that 111 loci, 57% of those examined, deviated from patterns of neutral inheritance. The most common deviation was an excess of homozygous genotypes relative to neutral expectations, indicative of directional selection. As in previous study of this rescue, individuals with more introduced alleles had higher reproductive success and longevity. In addition, we found 30 loci, distributed throughout the genome, which seem to have individual effects on these life history traits. Although the potential for outbreeding depression is a major concern when translocating individuals between populations, we found no evidence of such effects in this population.     

Nursing behaviour of bighorn sheep: correlates of ewe age,parasitism, lamb age,birthdate and sex

The nursing behaviour of marked bighorn ewes, Ovis canadensis, and their lambs was observed in Alberta, Canada, for 2 years. Suckle duration decreased as lambs grew older. Nursing behaviour in midlactation was correlated with maternal condition; young ewes, ewes with high faecal counts of lungworm (Protostrongylus spp.) larvae, and ewes whose lambs were born late allowed shorter suckles and were less likely to nuzzle their lamb's rump during a suckle. Sons did not appear to receive more milk than daughters. Suckle duration in mid-lactation was correlated with survival for female lambs but not for males. Lambs born late were disadvantaged, probably because nutritious forage was accessible to their mothers only in early lactation.     

Many eyes or many ewes: vigilance tactics in female bighorn sheep Ovis canadensis vary according to reproductive status

In gregarious animals, there is usually a negative relationship between individual vigilance and group size. This effect of group size is generally explained by increasing probability of predator detection (the many-eyes hypothesis) and by the dilution of risk occurring in larger groups. Few studies have attempted to examine the specific implications of either hypothesis on the expected vigilance pattern of an animal. Here we examine whether reproductive status affects vigilance patterns in bighorn sheep Ovis canadensis ewes. We also test whether the observed vigilance patterns are consistent with predictions from dilution or detection models of vigilance. Although vigilance decreased with increasing group size, vigilance tactics differed between barren and lactating females. Lactating ewes relied solely on predator detection. In contrast, barren ewes benefited from both detection and dilution effects when group size increased and adjusted vigilance effort according to the proportion of lactating ewes in their group. It is generally assumed that gregariousness increases safety. Here we further show that reproductive status influenced how animals reduce predation risk and that some individuals take advantage of the vigilance effort provided by others.     

Determinants of musculoskeletal frailty and the risk of falls in old age

Neuromuscular parameters that describe locomotion are indispensable variables for the diagnosis and treatment of frailty, fall risk and osteoporosis. A scientifically-based standardized locomotor assessment should be an essential part of medical examinations in research and clinical practice. There has been no previous consensus regarding which test procedures should be included in a locomotor assessment. The goal of this article is to provide a rationale for the selection of appropriate locomotor tests in a comprehensive locomotor assessment for elderly patients. We propose that a locomotor assessment should comprise the parameters that have been proven predictive for both falls and impending disability. The parameters should be measured in the standard units of physics. Therefore, we propose the following tests for a standardized locomotor assessment: (1) Self-selected gait velocity as the single best measure of general locomotor status and a good predictor of age-related adverse events; (2) Chair rise test (timed 5 chair rises) which measures power on vertical movement and the hip surrounding muscles as the most important neuromuscular risk factor for falls and fall-related fractures; (3) Tandem standing and tandem walking to measure postural capacity (balance) to the side; (4) Timed up and go test as a global screening procedure; (5) Clinical gait analysis with special focus on regularity; and (6) At least on a research level, movement must be measured referring to the terms of physics by mechanography. Mechanography (Leonardo force plate system, Novotec Pforzheim, Germany) records the time course of ground reaction forces, velocity of the vertical movements of the center of mass and power during unrestricted physiological movements. In the mechanogram the eccentric and concentric phases of movements can be differentiated and the storage of energy in the elastic elements of the body can be examined. The kinetics of human movement is explained by mechanograms of a two-legged jump. The ground reaction forces resulting from a jump down from a height of 0,46 m are demonstrated as a performance that is representative for human coordination. One goal of this text is to underline the insights that arise if the rules of physics are applied to human movement. A deeper understanding enables us to create more effective treatments for disorders of the muscle-bone unit. Bringing physics and cybernetics into the field of osteoporosis is a great heritage of Harold Frost.     

The effects of primiparity on reproductive performance in the brown bear

We studied the effects of primiparity on litter size, offspring size, and cub loss in brown bears (Ursus arctos) in two study areas (north, south) in Sweden from 1987 to 2006. Sexually selected infanticide (SSI) has been suggested previously as a mortality factor in our study populations. Females in the south became primiparous earlier than females in the north. Primiparous females had significantly smaller litters of cubs than multiparous females. We found no evidence that primiparity was costly in terms of the interlitter interval. Primiparous mothers had a higher probability of cub loss than multiparous mothers. The probability of cub loss was analyzed separately for the pre-mating and the mating season. The probability of cub loss by primiparous females in the pre-mating season increased with both increasing population density and deteriorating food conditions, whereas the probability of cub loss during the mating season decreased with increasing age of primiparity and increased with male turnover (a variable predicting SSI). The temporal patterns of cub loss by primiparous females suggested that the critical times for reproductive success by primiparous females were the pre-mating season (from birth to shortly after leaving the den) and the mating season. Cub loss in these periods was independent and caused by different factors. Cub loss before the mating season seemed to be most influenced by food conditions, whereas that during the mating season appeared to be caused by SSI.     

Population dynamic consequences of competition within and between age classes

We investigate the population dynamics of a semivoltine species whose juvenile development takes two years to complete, and is followed by a very short reproductive adult stage. Reproduction is synchronized so at any given time the juvenile population consists of two cohorts. Coexistence of the two cohorts requires that the strength of intea-cohort competition exceeds that of inter-cohort competition, an extension of the competitive exclusion principle. The population may exhibit population cycles with low integer period. We identify two mechanisms for two-year cycles, and note that four-year cycles are a natural consequence of overcompensation. Three year cycles can occur and we discuss the mechanisms involved.     

Elaeophorosis in bighorn sheep in New Mexico

Two bighorn sheep (Ovis canadensis) in New Mexico (USA) were found to be naturally infected with Elaeophora schneideri. An adult ram examined in 1997 in the Fra Cristobal Mountains had 26 nematodes in the carotid and iliac arteries, and microfilariae were present in the skin, nasal mucosa, brain, and lungs. This ram was markedly debilitated prior to euthanasia and extensive crusty, scabby lesions were observed on its head. In 1998, a yearling ewe found dead adjacent to Watson Mountain near the Gila Wilderness area was found to have 13 nematodes present in its heart. This is the first report of E. schneideri in bighorn sheep, and we suggest that bighorn sheep are susceptible to E. schneideri infection wherever they coexist with mule deer (Odocoileus hemionus hemionus) and appropriate tabanid vectors.     

Hemorrhagic disease in bighorn sheep in Arizona

Two bighorn sheep from Arizona (USA) were submitted for necropsy. One was a Rocky Mountain bighorn (Ovis canadensis canadensis) and the other was a desert bighorn (Ovis canadensis mexicana). Both had lesions consistent with those of hemorrhagic disease (HD). Epizootic hemorrhagic disease virus (EHDV) type-2 and bluetongue virus (BTV) type-17, respectively, were isolated from the sheep tissues. To our knowledge, HD caused by either EHDV or BTV infection has not been documented previously in Arizona bighorn sheep.     

Detection probability and Pasteurellaceae surveillance in bighorn sheep

We investigated the influence of detection probability (i.e., the probability of detecting the disease or organism of interest) on the repeatability of results reported from bacterial culture tests used to demonstrate the presence of species in the Pasteurellaceae family that infect bighorn sheep (Ovis canadensis). We also estimated occupancy probabilities (i.e., the probability an individual bighorn in a herd is infected) for each cultured biovariant and examined the effects of detection probability on the number of samples needed to detect the Pasteurellaceae biovariants from within an individual sheep as well as from within a herd. We collected 5-15 samples from free-ranging bighorns in Colorado, using oropharyngeal swabs or swabs of lungs, and submitted these swabs either immediately or after 2 days for bacterial culture. We saw significant variability in results for repeated samples from each of the sheep, and detection probabilities were ≤ 0.71 for all Pasteurellaceae biovariants cultured. The delayed (≥ 2 days) sample submission reduced both the microbial diversity detected and the detection probability for the biovariants characterized when compared to samples submitted immediately. Oropharyngeal sampling had higher detection probabilities of the individual biovariants than did lung swabs, and there was a difference in the biovariants detected between oropharyngeal and lung sampling. Depending on the biovariant of interest, estimates of occupancy probabilities ranged from 0.37-0.89, and we estimated that three to >30 swab samples were necessary to obtain a 95% confidence of detecting the cultured biovariants if they were present in an individual sheep. We estimated that the optimal sample sizes to detect the observed biovariants within a sheep herd with a 95% confidence ranged from sampling two bighorns twice to sampling 40 individuals once. Detection probability impacts the results reported from bacterial cultures for Pasteurellaceae in bighorn sheep, and confounding effects of the detection process should be addressed to improve the rigor of surveillance.