Polygynous Ungulates Research Articles

Do honest signals or piercing weapons determine rank in a polygynous ungulate?

Abstract In polygynous ungulates, males are often larger than females and bear more elaborate/larger weapons. Quantifying sexual dimorphism in different traits could provide insights into species-specific evolutionary pathways of sexual selection. Concerning the combination of secondary sexual traits, we found that Himalayan tahr (Hemitragus jemlahicus) is unique among the ~20 species in the tribe Caprini, as its body mass dimorphism is ~2-fold greater than the dimorphism in horn size, whereas horn shape appears to be near-monomorphic. Whilst horns show the same growth rate in both sexes, body mass increases faster in males. Considering age variation, dominant, golden-ruffed males are also heavier than brown-ruffed, lower-ranking males. Unlike most bovids, male–male competition in tahr does not seem to have influenced weapon development, suggesting a lower importance of horns in male–male competition compared to body mass, as their unritualized combat style also suggests. Our study highlights alternative evolutionary pathways occurring in the Caprinae, where intraspecific signals involve different traits, from weapons to pelage features. Accordingly, male tahr use their ruff colour as an ‘honest’ signal of rank.

Effect of adult male sterilization on the behavior and social associations of a feral polygynous ungulate: the horse

Castration is commonly used to control the behavior of companion animals and livestock, yet there have been few longitudinal studies of its effects. Despite the ubiquity of this surgery in ridden horses, the effects of castration (termed gelding in horses) have rarely been examined in a reproductive population. We tested effects of gelding on maintenance and social behaviors of individuals pre- and post-gelding, and in comparison to intact control adult males (2 to >16 years old) in both harem and bachelor status, we then tested how gelding affected association with mares (i.e., maintenance of a harem group) compared to intact controls, and any effects on bachelor social associations. We further explored any effects on foaling rate to assess potential impacts on population growth rate. We conducted this study over four years (2017–2020) at two Herd Management Areas (HMAs) in western Utah, USA: Conger and Frisco. We conducted demographic observations year round at both HMAs to record survival and foaling rate. We additionally recorded behavioral observations at Conger HMA. In December 2017, 27 adult males from Conger (42% of adult males in the population) were gelded and returned to the range with their social groups. Due to pre-treatment observations we were able to compare stallions of known status pre- and post-treatment (harem or bachelor), as well as gelded and intact males. We had no morbidity or mortality related to the gelding surgery and all males maintained good body condition throughout the study. There was no effect of gelding on maintenance behaviors (feeding, moving, and standing). There was no effect of gelding on frequency of agonistic behavior, and a non-significant tendency for less reproductive behavior in geldings; geldings showed more affiliative and less marking behavior. Age class and/or social status were better predictors of behavior than gelding. Over time fewer geldings maintained a harem, and their harem size declined during the study. Horses that were bachelors when gelded tended to remain as bachelors, whereas intact bachelors of the same cohort mostly attained a harem. Foaling rate at Conger was reduced in the year following treatment, but then returned to pre-treatment levels. From a welfare perspective gelding is safe to use in feral horses and has minimal effects on horse behavior and social interactions in a reproductive herd. Effectiveness for population growth control would likely require a larger proportion of males in the population to be castrated for longer-term effects on foaling rate.

Contrasting selection pressure on body and weapon size in a polygynous megaherbivore.

Sexual size dimorphism (SSD) is a common morphological trait in ungulates, with polygyny considered the leading driver of larger male body mass and weapon size. However, not all polygynous species exhibit SSD, while molecular evidence has revealed a more complex relationship between paternity and mating system than originally predicted. SSD is, therefore, likely to be shaped by a range of social, ecological and physiological factors. We present the first definitive analysis of SSD in the common hippopotamus (Hippopotamus amphibius) using a unique morphological dataset collected from 2994 aged individuals. The results confirm that hippos exhibit SSD, but the mean body mass differed by only 5% between the sexes, which is rather limited compared with many other polygynous ungulates. However, jaw and canine mass are significantly greater in males than females (44% and 81% heavier, respectively), highlighting the considerable selection pressure for acquiring larger weapons. A predominantly aquatic lifestyle coupled with the physiological limitations of their foregut fermenting morphology likely restricts body size differences between the sexes. Indeed, hippos appear to be a rare example among ungulates whereby sexual selection favours increased weapon size over body mass, underlining the important role that species-specific ecology and physiology have in shaping SSD.

Linking alternative reproductive tactics and habitat selection in Northern chamois.

In polygynous ungulates, males may achieve fertilization through the use of alternative reproductive tactics (ARTs), discrete phenotypic variations evolved to maximize fitness. ARTs are often associated with different male spatial strategies during the rut, from territoriality to female‐following. Although variation in space use patterns of rutting male ungulates is known to be largely affected by the spatial distribution of females, information on the year‐round habitat selection of alternative reproductive types is scant. Here, we investigate the seasonal variation in habitat choice of a large mammal with ARTs (territoriality and nonterritoriality), the Northern chamois Rupicapra rupicapra. Global Positioning System (GPS) data on 28 adult males were collected between February 2010 and December 2013 in the Gran Paradiso National Park (Italy) and used to fit resource selection functions to explore the ART‐specific use of key topographic features, such as elevation, aspect, and slope, and vegetation phenology expressed as NDVI values. Territorial and nonterritorial chamois profoundly differed in their habitat selection not only during the rutting season. Compared to nonterritorial males, territorial males used lower elevations in summer and autumn, preferred southern slopes in spring and summer, and used steeper areas in summer but not in winter. We found no difference in seasonal selection of NDVI values between males adopting ARTs. Our results suggest that territorial males tend to occupy warmer, lower‐food‐quality habitats in late spring and summer, whereas nonterritorial males are free to follow and exploit vegetation phenology and more favorable temperatures. Different patterns of habitat selection may reflect different trade‐offs between the optimization of energy balances throughout the year and the increase of mating opportunities during the rut in males adopting alternative reproductive tactics.

Young and mature males have similar energy expenditure during the rut in a trophy-hunted population of Mediterranean mouflon

In polygynous ungulates, males invest time and energy to reproductive activities during the rut and this involvement is expected to increase with age due to different mating tactics in young versus adult males. In contrast, mating period is expected to be less costly for females for which late gestation and lactation are the most energetically demanding periods. However, empirical supports of these hypotheses through direct measures of reproductive effort are still limited in ungulate species, particularly in males. In addition, this general pattern may be modified in populations facing selective harvesting on adult males, where young males may experience less competition to mate and invest more energy during the rut. We investigated these hypotheses by studying the age- and sex-specific variations of kidney fat reserves from pre- to post-rut periods in a trophy-hunted population of Mediterranean mouflon (Ovis gmelini musimon × Ovis sp.), a polygynous dimorphic mountain ungulate. Females were found to build up energy from the pre-rut to the post-rut periods, most likely to face with the subsequent costs of gestation/lactation that occur few months later. Conversely, kidney fat mass strongly decreased similarly in both young and mature males, and with the same magnitude, suggesting a strong investment of males of all ages in this population. This might be related to the selective hunting pressure on mature males this population is facing with. This result supports the need for more empirical and comparative studies to better grasp the influence of trophy hunting on reproductive effort in male ungulates.

Weather stochasticity and alternative reproductive tactics in Northern chamois, Rupicapra rupicapra

Abstract In polygynous ungulates, male breeding success is often achieved through alternative reproductive tactics (ARTs). The maintenance of ARTs in this taxon is largely explained by condition-dependent selection, a decision rule in which individuals switch their breeding behaviour on the basis of internal factors such as ageing (status-dependent selection), or external factors such as weather conditions during the rut (environment-dependent selection). In some species, however, ARTs are fixed during the mating season, and other forms of selective pressures rather than environment-dependent selection may contribute to the evolutionary maintenance of male types within populations. Assuming female vicinity as a proxy of mating opportunity, we explored the effect of snow depth variations on the elevational overlap between females, and territorial and non-territorial male chamois, Rupicapra rupicapra, in three consecutive rutting seasons. In snow-free periods, females overlapped with non-territorial males, but not with territorial ones. With increasing snow depth, females and territorial males progressively occupied similar elevations. Our results suggest that snow cover variations may impose changes in the spatial distribution of males and females. In turn, weather stochasticity over multiple ruts, rather than within-rut environment-dependent selection, may impose variations in mating opportunities of ARTs, possibly contributing to the coexistence of different phenotypes within populations.

Tooth wear as a practical indicator of sexual differences in senescence and mastication investment in ecology studies

Abstract Teeth of many mammalian species do not grow further after they have emerged, nor are they replaced or repaired if they break or wear. These dental properties have been used to test evolutionary hypotheses on life expectancy relative to actual age (senescence, aging), as the animal is expected to die when all its dental crown tissue has been depleted due to the wearing effect of mastication. Females of polygynous mammals with small litter size are expected to live longer than males of the same species, as female reproductive fitness is maximised by having a long reproductive life. In contrast, maleś reproductive success is limited to the short period when they are in prime condition, as it is in this period that they are most successful in intra-sexual competition for mating opportunities. Consequently, it has been predicted that femaleś teeth should be more durable and so wear more slowly than maleś teeth. However, these predictions are affected by potential sex differences in the hardness of dental tissues, which have not been assessed in depth in the literature. I tested sexual differences in the hardness of enamel and dentine on a balanced, by sex and age, sample of 156 lower first molars of Scottish red deer of known age using Vickers micro-hardness indentation measures as a proxy of the resilience of the tooth to wear. My results clearly indicate that there are no sexual differences in enamel and dentine hardness, and that tooth hardness has a negligible effect on predicting tooth wear, which supports the use of tooth wear relative to body size as a convenient proxy for assessing sexual differences in senescence in polygynous ungulates. I warn on making conclusions from comparisons between populations or species in tooth wear when tooth hardness was controlled for.

Chronology of reproductive investment determines predation risk aversion in a felid-ungulate system.

Fear of predators can behaviorally mediate prey population dynamics, particularly when predation risk influences reproductive investment. However, the costs of reproductive investment may mitigate predation risk aversion relative to periods when the link between reproductive output and prey behavior is weaker.We posit that intensity of reproductive investment in ungulates may predict their response to predation risk such that the sexes increase risk exposure during biological seasons that are pivotal to reproductive success, such as the fawn‐rearing and breeding seasons for females and males, respectively.We examined the activity patterns of sympatric white‐tailed deer (Odocoileus virginianus), a sexually segregated polygynous ungulate, and Florida panthers (Puma concolor coryi) in the context of the “risky times – risky places hypothesis” and the reproductive strategy hypothesis. We compared detection rates and diel activity overlap of both species using motion‐triggered camera traps positioned on (n = 120) and off (n = 60) anthropogenic trails across five reproductive seasons.Florida panthers were nocturnal and primarily observed on‐trail providing an experimental framework with risky times and risky places. Contrary to studies in other taxa inversely correlating prey reproductive investment to predation risk, the sexes of deer were more risk prone during sex‐specific seasons associated with intense reproductive investment.Our results suggest spatiotemporally variable predation risk influences sex‐specific behavioral decision‐making in deer such that reproductive success is maximized.

Security, size, or sociality: what makes markhor (Capra falconeri) sexually segregate?

Sexual segregation, the tendency to seasonally live in groups comprised of separate sexes, is widespread in sexually dimorphic polygynous ungulates. The causes for such segregation are still being studied and debated to arrive at a universal explanation. We assessed sexual segregation in the markhor, Capra falconeri—a dimorphic mountain ungulate inhabiting a seasonal temperate environment. We observed markhor herd composition, gender-specific diet, and habitat use over 3 years and multiple seasons. We tested the predictions of the Reproductive Strategy-Predation Risk Hypothesis (RSH) that focuses on offspring security, and the Forage Selection-Sexual Dimorphism Hypothesis (FSH) based on different foraging needs between the sexes due to size dimorphism. Generalized linear models were used to identify the important variables that best explained segregation in markhor. Male and female markhor showed a tendency to segregate throughout the year outside the mating season. Female markhor were confined to cliffs with relatively less forage cover, whereas males used areas with relatively better forage away from the security of cliffs. Females also occurred at lower elevations compared to males. Social segregation prevailed during pre-parturition and continued until autumn when the offspring were older, indicating a role of other factors besides predation risk. Sexual segregation in markhor was influenced by multiple mechanisms.

Factors influencing reproductive success in male white‐tailed deer

ABSTRACTPositive relationships between age, sexually selected traits, and male reproductive success have been reported for a number of polygynous ungulates; however, relatively little is known about the factors influencing male reproductive success in ungulate species whose mating system is characterized by tending‐bond behaviors. Broad interest in the genetic consequences of selective harvest supports a greater understanding of the role of these factors as determinants of male reproductive success in important game species (e.g., white‐tailed deer [Odocoileus virginianus]), that exhibit tending‐bond behaviors. We investigated male reproductive success in white‐tailed deer across a range of sex ratios and age structures using a known population of deer housed in a 175‐ha enclosure in central Alabama, USA. We measured age, annual antler size, and annual body size of male white‐tailed deer and assigned paternity to 143 known‐age offspring during 2007–2014. Reproductive success was attributed to a high proportion of males during each of the 6 breeding seasons. Our most supported model indicated that annual body size and antler size of the individual were positively associated with annual male breeding success. The effects of annual antler size were sensitive to changes in mean male age of the herd, with antler size having the greatest effect on male reproductive success under older male age structures. Young (≤1.5 yr) males reproduced most frequently when male age structure was youngest (which correlated with female‐biased sex ratios in this population). Our results suggest that male age structure and sex ratio played a key role in establishing patterns of male reproductive success in white‐tailed deer. Management practices that encourage balanced adult sex ratios and older male age structures (e.g., Quality Deer Management) may promote a highly competitive environment where sexually selected traits are of increased importance to male breeding success. However, the ability of managers to alter herd genetics in a positive or negative direction through selective harvest is limited in white‐tailed deer because of the high proportion of reproducing males. © 2016 The Wildlife Society.

Influences of mating group composition on the behavioral time-budget of male and female Alpine Ibex (Capra ibex) during the rut.

During the rut, polygynous ungulates gather in mixed groups of individuals of different sex and age. Group social composition, which may vary on a daily basis, is likely to have strong influences on individual’s time-budget, with emerging properties at the group-level. To date, few studies have considered the influence of group composition on male and female behavioral time budget in mating groups. Focusing on a wild population of Alpine ibex, we investigated the influence of group composition (adult sex ratio, the proportion of dominant to subordinate males, and group size) on three behavioral axes obtained by Principal Components Analysis, describing male and female group time-budget. For both sexes, the first behavioral axis discerned a trade-off between grazing and standing/vigilance behavior. In females, group vigilance behavior increased with increasingly male-biased sex ratio, whereas in males, the effect of adult sex ratio on standing/vigilance behavior depended on the relative proportion of dominant males in the mating group. The second axis characterized courtship and male-male agonistic behavior in males, and moving and male-directed agonistic behavior in females. Mating group composition did not substantially influence this axis in males. However, moving and male-directed agonistic behavior increased at highly biased sex ratios (quadratic effect) in females. Finally, the third axis highlighted a trade-off between moving and lying behavior in males, and distinguished moving and female-female agonistic behavior from lying behavior in females. For males, those behaviors were influenced by a complex interaction between group size and adult sex ratio, whereas in females, moving and female-female agonistic behaviors increased in a quadratic fashion at highly biased sex ratios, and also increased with increasing group size. Our results reveal complex behavioral trade-offs depending on group composition in the Alpine ibex, and emphasize the importance of social factors in influencing behavioral time-budgets of wild ungulates during the rut.

Effects of harvest, culture, and climate on trends in size of horn‐like structures in trophy ungulates

AbstractHunting remains the cornerstone of the North American model of wildlife conservation and management. Nevertheless, research has indicated the potential for hunting to adversely influence size of horn‐like structures of some ungulates. In polygynous ungulates, mating success of males is strongly correlated with body size and size of horn‐like structures; consequently, sexual selection has favored the development of large horns and antlers. Horn‐like structures are biologically important and are of great cultural interest, both of which highlight the need to identify long‐term trends in size of those structures, and understand the underlying mechanisms responsible for such trends. We evaluated trends in horn and antler size of trophy males (individuals exhibiting exceptionally large horns or antlers) recorded from 1900 to 2008 in Records of North American Big Game, which comprised >22,000 records among 25 trophy categories encompassing the geographic extent of species occupying North America. The long‐term and broad‐scale nature of those data neutralized localized effects of climate and population dynamics, making it possible to detect meaningful changes in size of horn‐like structures among trophy males over the past century; however, ages of individual specimens were not available, which prevented us from evaluating age‐class specific changes in size. Therefore, we used a weight‐of‐evidence approach based on differences among trophy categories in life‐history characteristics, geographic distribution, morphological attributes, and harvest regimes to discriminate among competing hypotheses for explaining long‐term trends in horn and antler size of trophy ungulates, and provide directions for future research. These hypotheses were young male age structure caused by intensive harvest of males (H1), genetic change as a result of selective male harvest (H2), a sociological effect (H3), effects of climate (H4), and habitat alteration (H5). Although the number of entries per decade has increased for most trophy categories, trends in size of horn‐like structures were negative and significant for 11 of 17 antlered categories and 3 of 8 horned categories. Mean predicted declines during 1950–2008 were 1.87% and 0.68% for categories of trophy antlers and horns, respectively. Our results were not consistent with a sociological effect (H3), nutritional limitation imposed by climate (H4), or habitat alteration (H5) as potential explanations for long‐term trends in size of trophies. In contrast, our results were consistent with a harvest‐based explanation. Two of the 3 species that experienced the most conservative harvest regimes in North America (i.e., bighorn sheep [Ovis canadensis] and bison [Bison bison]) did not exhibit a significant, long‐term trend in horn size. In addition, horn size of pronghorn (Antilocapra americana), which are capable of attaining peak horn size by 2–3 years of age, increased significantly over the past century. Both of those results provide support for the intensive‐harvest hypothesis, which predicts that harvest of males has gradually shifted age structure towards younger, and thus smaller, males. The absence of a significant trend for mountain goats (Oreamnos americanus), which are difficult to accurately judge size of horns in the field, provided some support for the selective‐harvest hypothesis. One other prediction that followed from the selective‐harvest hypothesis was not supported; horned game were not more susceptible to reductions in size. A harvest‐induced reduction in age structure can increase the number of males that are harvested prior to attaining peak horn or antler size, whereas genetic change imposed by selective harvest may be less likely to occur in free‐ranging populations when other factors, such as age and nutrition, can override genetic potential for size. Long‐term trends in the size of trophy horn‐like structures provide the incentive to evaluate the appropriateness of the current harvest paradigm, wherein harvest is focused largely on males; although the lack of information on age of specimens prevented us from rigorously differentiating among causal mechanisms. Disentangling potential mechanisms underpinning long‐term trends in horn and antler size is a daunting task, but one that is worthy of additional research focused on elucidating the relative influence of nutrition and effects (both demographic and genetic) of harvest. © 2013 The Wildlife Society.

Rutting behaviour of territorial and nonterritorial male chamois: Is there a home advantage?

Males using alternative male mating tactics (AMTs) may express their mating effort in a variety of ways. In polygynous species with limited sexual dimorphism, differences in male aggressiveness may affect mating opportunities. We recorded the behaviour of 8 territorial and 7 nonterritorial male Alpine chamois Rupicapra rupicapra, a nearly monomorphic ungulate, during the 2011 rut in the Gran Paradiso National Park (Italy), to analyse differences in mating effort and mating opportunities between AMTs. The chamois showed a rich behavioural repertoire (31 behavioural patterns), with a prevalence of indirect aggression. Territorial males had higher frequency of aggressive and courtship behaviour than nonterritorial males over the early rut. Later, nonterritorials increased their mating effort, possibly because of reduced competition with dominant males. Territorial males monopolised all observed mating events. Our results support the hypothesis that chamois may assert dominance through intense aggressiveness rather than through horn size or body mass as found in other polygynous ungulates. Most important, differences in mating effort mediated by AMTs resulted in different mating opportunities; these benefits, however, are traded off against greater costs, due to higher levels of hormone metabolites and parasitism. Data on AMTs flexibility, lifetime reproductive success and survival are needed to clarify the mechanisms underlying the evolution and maintenance of AMTs within chamois populations.

Sex ratio in goitered gazelles (Gazella subgutturosa Guldenstaedt, 1780)

We studied the sex ratio of goitered gazelles in the naturally arid environment of Kazakhstan over a 6-year period. The main methods in our study were taking transect counts and focal observations. The sex ratio of adult goitered gazelles has demonstrated a female bias due to a much higher mortality of males of all ages, especially during years with unusually severe winters. This phenomenon is typical for many polygynous ungulates, as well as other gazelle species. Surprisingly, our data demonstrated monthly fluctuations in sex proportions, along with a bias shift from a female-dominant population during most of the year to a male-dominant population during spring. We discovered, though, that our data did not reflect any real changes in the sex ratio of the population but, instead, revealed the radical changes in behavior of pregnant females before giving birth—hiding from danger in thick shrubs or broken terrain rather than fleeing. As a result, we were not able to see many pregnant females in our spring samples (before birthing), and so received a male-biased population. During the rest of the year (after birthing), females returned to their usual behaviors of fleeing from danger that then gave us a female-biased sex ratio that reflected a more accurate status in sex proportions of the population. So, our results discovered seasonal sex difference in hiding behavior which led to a bias based on visibility.

The cost of trying: weak interspecific correlations among life-history components in male ungulates

Life-history trade-offs are well known in female mammals, but have seldom been quantified for males in polygynous species. I compared age-specific mass, weapon size, survival, and reproductive success of males in eight species of ungulates, and found weak interspecific correlations among life-history traits. Young males tended to have higher reproductive success in rapidly-growing than in slow-growing species, and in species where horns or antlers reached near-asymptotic size over the first few years of life. There was no clear interspecific trade-off between early reproduction and early survival. Reproductive senescence was evident in most species. Generation length, calculated as the mean age of fathers, was negatively correlated with the reproductive success of young males and positively with life expectancy of 3-year-olds, but not with early mortality. The main determinant of male reproductive success in polygynous ungulates is the ability to prevail against competing males. Consequently, the number and age structure of competitors should strongly affect an individual’s ability to reproduce, making classic trade-offs among life-history traits very context-dependent. Most fitness costs of reproduction in male ungulates likely arise from energy expenditure and injuries sustained while attempting to mate. Individual costs may be weakly correlated with fitness returns.

Association patterns and kinship in female reindeer (Rangifer tarandus) during rut

Non-random association of females has been reported in many social species, with kinship being the main mechanism in shaping these associations. Understanding the patterns of association during rut is particularly important in polygynous animals, as this may have implications for inbreeding vulnerability and genetic structuring of populations. However, these patterns are highly dynamic and may be influenced by the population sex ratio and male age structure. We examined females' association in reindeer (Rangifer tarandus), a highly polygynous ungulate, to determine whether genetically related females associate more often than non-kin during rut and whether this association was influenced by the aforementioned factors. During three rutting seasons the male age structure and the sex ratio of the herd was manipulated: two extremely female biased sex ratios, one with only adult males and one with only young males, whereas the third being closer to a more even sex ratio and male age structure. We found that genetic relatedness influenced female association in the mixed male age group, but not in the adult and young male age groups. Highly related females (relatedness ∼0.5) associated more closely than unrelated females. Further, period of the rut as well as male age structure and sex ratio had a significant effect on female association. Our study suggests that the female reindeer association during rut is not only influenced by preference for kin but also by male–male competition and male age as well.

Unbiased sex-specific survival in Alpine chamois

Many polygynous ungulates show higher mortality of males than of females, because of the intense male–male competition during the rut and the costs associated with the development of sexual-size dimorphism. In the weakly dimorphic Alpine chamois Rupicapra rupicapra the occurrence of differential sex-specific survival strategies is controversial. To date, only two studies investigated the survivorship of males and females in this species, producing conflicting results: these works, based on the use of life tables, require confirmation from researches carried out on living populations. We assessed the survival pattern of a protected Alpine chamois population in the Swiss National Park, where 116 individuals were marked and monitored over 13 years (1996–2008). We tested for sex-, age- and year-dependence of survival by means of capture-mark-resight models. Resighting probabilities were sex-dependent, and survival rates were time-dependent. Females had higher resighting probabilities (0.84) than males (0.74). All over the time periods, sex had a weak influence on survival probability (males=0.91; females=0.92) and survival rates remained surprisingly high until late age (1 year=0.90; 2–7 years=0.91; 8+ years=0.92). The growing evidence for a high adult survival and a weak differential mortality of the two sexes, together with the highly seasonal sexual-size dimorphism observed for Alpine chamois, might indicate the occurrence of a unique conservative survival strategy in both sexes and a low-risk mating strategy by males.

Age-related male reproductive effort in two mountain ungulates of contrasting sexual size dimorphism

In polygynous ungulates, the reproductive effort of adult males peaks during a short period in which feeding activities are sacrificed for mating activities. Hence, both fat reserves and body mass are predicted to decline markedly during this period. The decline is also predicted to be greater in fat reserves than in body mass because fat is catabolized before muscle, and to increase with the intensity of sexual selection. In contrast, no specific patterns are expected in females for which late gestation and lactation rather than mating are the energetically most demanding periods. We tested these hypotheses in two mountain ungulates of contrasting sexual size dimorphism (SSD): Himalayan tahr ( Hemitragus jemlahicus (H. Smith, 1826)) (SSD = 123%) and alpine chamois ( Rupicapra rupicapra (L., 1758)) (SSD = 26%). As expected, kidney fat declined more rapidly than body mass in adult males of both species. Kidney fat declined faster in adult male tahr compared with adult male chamois. There was no consistent pattern of changes in body mass or kidney fat in female tahr or female chamois. Our results suggest that adult males of species with strong SSD allocate more energy to mating than males of less dimorphic species.

Indirect measures of reproductive effort in a resource-defense polygynous ungulate: territorial defense by male guanacos

Trade-offs between reproductive effort and activities that are likely to enhance survival have been studied extensively in harem defense ungulates. However, among resource defense systems, ecological determinants of the variation in male reproductive effort are less understood. The aim of this study was to assess the extent of the effort devoted to territorial defense by male guanacos and how this is affected by predation risk. Time allocation to different activities and displays of aggressive behavior were compared between territorial and bachelor male guanacos within two populations with contrasting levels of predation risk. Territorial males devoted almost twice the time than bachelors to vigilance and moving, at the expense of foraging time. Aggressive interactions were more frequent and severe for territorial males than for bachelors, based on observation. These differences were larger during the reproductive season but remained statistically significant after this period, highlighting the extended effort towards territorial defense beyond the peak of reproductive activity. In contrast to previous observations on females of the same populations, there was no effect of predation risk level or group size on male time allocation, suggesting that under the ecological conditions of the study, intra-sexual competition entails a stronger influence on male individual behavior than predation risk level.

No inbreeding effects on body size in two captive endangered gazelles

Selection for body size is intricate, involving trade-offs between energy costs, reproductive output, foraging efficiency, and interaction with other community members. In dimorphic, polygynous ungulates, body size is highly correlated with reproductive success in both sexes. Body size has been proposed as a potential phenotypic indicator of genetic change, in wild and in captive populations. We analysed the relationship between adult body size and inbreeding in two captive populations of endangered gazelles. Two estimates of inbreeding were used: individual inbreeding coefficient ( F i ) and individual increase in inbreeding (Δ F i ). Six cranial traits and eight post-cranial bones were measured in 87 Cuvier's gazelles and 97 Mohor gazelles. The average level of individual inbreeding found for the alive population of Cuvier's gazelle was 0.236 and for Mohor gazelle 0.260. Our study demonstrated that within our populations most variability in body size is explained by sex, but we have not found any evidence of inbreeding depression in this morphological trait. Our results are surprising in the light of a widely held belief that from an evolutionary point of view close inbreeding has deleterious effects. The great diversity of factors acting on the effects of inbreeding on trait values make difficult to provide a simple framework to understand them all, hence, we suggest fitness consequences of inbreeding has to be assessed considering the conditions under which one should expect inbreeding depression.