Sparring,relative antler size,and assessment in male caribou

Summary There are two kinds of antler combats in male deer. Fighting is rare, violent, occurs between matched males, and can cause injury and death. Sparring is common, usually gentle, often occurs between unmatched males, and involves no risk of injury. We recorded 1308 sparring matches and only 6 fights between wild male woodland caribou (Rangifer tarandus caribou) in 2 years of study. In the 713 sparring matches between males with unequal antlers, the animal with smaller antlers initiated close to half of the encounters, but terminated nearly 90% of them. That was true whether a sparring match involved two adults, two yearlings, or an adult against a yearling. We argue that the best interpretation of that shift in the decisions of the smaller-antlered male is that sparring serves to assess a partner's weight and strength relative to one's own. Thus, through sparring, antlers could be used to allow a form of tactile (proprioceptive) assessment of fighting ability, which could be the basis for subsequent visual assessment at a distance. Offprint requests to: C. Barrette     

Litter reductions reveal a trade-off between offspring size and number in brown bears

Experimental manipulations have revealed positive effects of litter reduction on offspring mass in small mammals, but little is known about this trade-off in large mammals. We examined the determinants of natural litter size variation and quantified the effects of litter size, maternal characteristics, and litter composition on yearling mass using 24?years of data on marked brown bears (Ursus arctos) in Sweden. Infanticide by adult males is a major cause of cub-of-the-year mortality, leading to litter size reductions. Litter size (n?=?265) at den emergence ranged from one to four cubs (average, 2.7) and increased with maternal age. Litter size, however, appeared independent of maternal size, population density, interlitter interval, study area, or previous litter sex ratio. Yearling body mass increased with maternal body size but was independent of litter sex ratio. Litter size and yearling mass were negatively correlated, mostly because singletons were about 30?% heavier than yearlings from litters of two to four cubs. In reduced litters, survivors were on average 8?% heavier as yearlings than individuals from intact litters, suggesting that sibling competition reduces growth. Trade-offs between litter size and yearling mass in bears appear similar in magnitude to those found in small mammals.     

Disappearance and dispersal of Spermophilus elegans juveniles in relation to behavior

Summary Patterns of disappearance and dispersal of Spermophilus elegans juveniles during the first 6-weeks postemergence were compared for 1977 and 1978 and related to quantitative and qualitative changes in social interaction involving juveniles.Juvenile disappearance (emigration or mortality) and dispersal within the study site varied between the sexes within and between years. Female disappearance and dispersal were significantly greater in 1977, and male losses in 1978 significantly exceeded male losses in 1977. Greater female loss in 1977 resulted in total male — female losses being equivalent, whereas in 1978 juvenile loss was strongly biased toward males by the end of the 6-week period.Greater female loss in 1977 was attributed primarily to increased aggression between female juveniles in that year because of larger average litter size with more females per litter. Increased disappearance of males in 1978 showed no correlation with litter size or relative increase in number of males per litter. Male young interacted with individuals of several age/sex classes, and a possible behavioral influence on male disappearance was increased aggressiveness by yearling males toward juvenile males in 1978.Behavior appeared to act as a proximate factor in juvenile disappearance and dispersal, and the observed differences between how male and female juveniles interacted in 1977 and 1978 were hypothesized to reflect the operation of different selective pressures to increase individual male or female fitness.     

Proximate causes of natal dispersal in female yellow-bellied marmots, Marmota flaviventris

We investigated factors influencing natal dispersal in 231 female yearling yellow-bellied marmots (Marmota flaviventris) using comprehensive analysis of 10 years (1983-1993) of radiotelemetry and 37 years (1963-1999) of capture-mark-recapture data. Only individuals whose dispersal status was verified, primarily by radiotelemetry, were considered. Univariate analyses revealed that six of the 24 variables we studied significantly influenced dispersal: dispersal was less likely when the mother was present, amicable behavior with the mother and play behavior were more frequent, and spatial overlap was greater with the mother, with matriline females, and with other yearling females. Using both univariate and multivariate analyses, we tested several hypotheses proposed as proximate causes of dispersal. We rejected inbreeding avoidance, population density, body size, social intolerance, and kin competition as factors influencing dispersal. Instead, our results indicate that kin cooperation, expressed via cohesive behaviors and with a focus on the mother, influenced dispersal by promoting philopatry. Kin cooperation may be an underappreciated factor influencing dispersal in both social and nonsocial species.     

Antler asymmetry and immunity in reindeer

Fluctuating asymmetry (FA) measures an individual's ability to undergo identical development in bilaterally symmetrical characters and may indicate sensitivity to environmental stress. FA in ornamental characters is often positively related to parasite intensities, which are important environmental stressors. Parasites affect and are affected by several parts of the immune system, and the ability to resist parasites may be signalled via FA in ornaments. In this study we examined reindeer antlers, which show FA, demonstrated to be caused by parasite infections. We measured antler FA, immune parameters (i.e. densities of different classes of leukocytes, IgG levels and abomasal lymph node numbers) and intensity of abomasal nematodes in free-ranging 1.5-year-old male reindeer slaughtered in the early part of their rutting period. We found a relationship between parasite intensity and immune parameters suggesting that our measures of immune activity reflect density of current parasite infections. More important, these immune parameters were associated with FA in both the main beam length and numbers of antler tines. The immune parameters were, however, only weakly correlated to antler size. This indicates that FA, but not size, of antlers grown during exposure to a multitude of environmental stressors may reveal information about individual immunity that can be important for host-parasite interactions. Antler FA may therefore communicate an individual's quality during the rut in reindeer. Received: 30 March 1998 / Accepted after revision: 15 August 1998     

Natural selection and the communal rearing of pups in African wild dogs (Lycaon pictus)

Summary The contributions of adult (>18 months) and yearling (10–18 months) African wild dogs (Lycaon pictus), to raising pups were quantified at eight natal dens, where pups remain for their first three months of life.Adults between 2–6 years of age did most of the hunting, and the dominant male often made the first grab at fleeing prey. Yearlings contributed to the hunting but were reluctant to tackle large prey animals. Yearlings and breeding females had prior access to food. Yearlings and adults regurgitated comparable amounts of food, but in one pack watched at a time of food shortage, the yearlings failed to regurgitate and stole food from the pups.Dominant dogs chased predators, especially spotted hyenas (Crocuta crocuta) from the area of the den more frequently than other adults or yearlings chased predators. Mothers, particularly in the first six weeks post-partum, stayed and protected the pups when the pack hunted.There is a positive, but non-significant correlation between the number of adult helpers and the number of pups raised. However, the sex ratio bias towards males at birth suggests that male helpers (which predominate) usually increase pup survivorship. The roles of direct and indirect selection in the evolution of the helping behavior are discussed. Indirect selection has probably played an important role in the unusual post-reproductive survival of males and their helping behavior.Serengeti Wildlife Research Institute contribution number 284     

Age-related foetal sex ratio bias in Iberian red deer (<Emphasis Type="Italic">Cervus elaphus hispanicus</Emphasis>): are male calves too expensive for growing mothers?

Several theories predict offspring biases towards males or females with increasing reproductive resources of the mother to maximize reproductive returns by offspring, or as a result of prohibitive cost of the most expensive sex for young mothers or those in poor condition. This study examines foetus sex of 221 harvested hinds in a food-supplemented game estate for 10 years, according to hind age class (yearlings, subadults or adults), precise age, body mass and condition, and jaw length. A logistic model showed that hinds had a greater probability of bearing a male foetus with increasing age class, but not with any other variable. The greatest bias was found in yearling hinds. After controlling for age class and mass, jaw length was smaller in pregnant compared to non-pregnant yearlings and subadults, which suggests a trade-off between reproduction and growth. The bias towards females in yearlings increased as gestation proceeded, which suggests that the bias might be a result of selective abortion of male foetuses. Although results do not exclude an investment in males to increase number of grand-offspring, they suggest that young hinds may produce daughters as a trade-off between low energetic-cost offspring and their need to grow.Communicated by S. Krackow     

Age,experience, and enemy recognition by wild song sparrows

Summary Older female Song Sparrows (Melospiza melodia) breeding on Mandarte Island, B.C., Canada, are more often parasitized by a brood parasite, the Brown-headed Cowbird (Molothrus ater), than yearling females breeding for the first time. This may be explained if older Song Sparrows behave differently than yearlings towards searching female cowbirds, and are thus more readily recognized as potential hosts. We tested this hypothesis by measuring the responses of Song Sparrows to a stuffed mount of a female cowbird in 1982, when no cowbirds frequented the island. This mount, and a control (a female junco) were presented near the nests of wild female sparrows of known age. As predicted, adult female sparrows behaved differently towards the cowbird model from yearlings (Table 1). They did not differ in their responses to the junco. Adult males also responded differently to the cowbird and junco (Table 2), but adult males did not differ significantly from yearlings in their response to either model. When birds that responded weakly as yearlings in the absence of cowbirds were retested as 2-year-olds after the recolonization of the island by cowbirds, they responded strongly to cowbird models. Two-year-old birds not exposed to cowbirds as yearlings, were parasitized at a rate intermediate between the rate for experienced adults and that for yearlings. Yearling females were parasitized less often at the beginning of the period of breeding activity by cowbirds than at the end. All these results are consistent with our hypothesis that age-selective parasitism results from differences among age classes in the mobbing responses of Song Sparrows to cowbirds. The greater response of adult sparrows to cowbirds seems non-adaptive, because it apparently results in a loss of reproductive output through selective parasitism. We suggest that this paradox can be resolved if the response to cowbirds near the nest is an instance of a more general acquired response to all potential enemies that approach a nest.     

Macro-habitat patch structure,environmental harshness,and Marmota flaviventris

Summary Social behavior of 15 colonies of Yellow-bellied Marmots was studied at sites differing in both elevation (plant growing season length) and patch structure (density and spacing of suitable colony sites in large blocks of habitat) from 1976 through 1978. Colonies were typically composed of family groups. In low elevation colonies, offspring dispersed at the end of the juvenile year. In all high elevation areas, offspring dispersed as yearlings, and parent-offspring and sibling interactions during the juvenile year were highly amicable. In high elevation areas in continuous habitat, dispersal by yearlings occurred with no agonistic interactions. In high elevation areas with patchy habitat, however, brief but intense periods of extremely agonistic sibling and parent-offspring interactions occurred immediately prior to dispersal of yearlings.Social behavior and dispersal are uncorrelated with elevation, plant growing season length, or available foraging time. Social interaction among relatives in marmot colonies is amicable when other suitable colony sites are located nearby; in these areas, dispersal occurs without any antagonism. Agonistic behavior occurs only in areas with a patchy distribution of suitable colony sites. At these sites, dispersal occurs only after periods of extreme antagonism. Further, those offspring who do not disperse from the colony site are the ones who dominate and initiate agonistic interactions with their siblings. The large-scale structure of the habitat influences the propensity of the individual to disperse. When the animal is reluctant to disperse because of a combination of high transit difficulty and low desirability of a new site, angonistic social interactions with relatives force dispersal.     

Sociality reduces individual direct fitness in a communally breeding rodent,the colonial tuco-tuco (<Emphasis Type="Italic">Ctenomys sociabilis</Emphasis>)

In many social vertebrates, remaining in the natal group leads to at least short-term reductions in the direct fitness of philopatric animals. Among communally breeding rodents, the direct fitness costs of philopatry appear to increase as the frequency of successful natal dispersal decreases, suggesting a functional link between constraints on natal dispersal and the reproductive consequences of sociality. To explore this relationship empirically, I documented patterns of direct fitness among female colonial tuco-tucos (Ctenomys sociabilis), which are group-living subterranean rodents from southwestern Argentina. Demographic data suggest that successful natal dispersal is rare in this species, leading to the prediction that natal philopatry in C. sociabilis is associated with significant reductions in individual direct fitness. Using data obtained during 1996–2001, I compared the direct fitness of females that dispersed from their natal group and bred alone as yearlings to that of females that lived and bred in their natal group as yearlings. Philopatric yearlings reared significantly fewer young to weaning than did disperser (lone) yearlings. Although neither survival to a second breeding season nor the estimated lifetime number of pups reared to weaning differed between dispersal strategies, the annual direct fitness of group-living females was 23–40% less than expected, suggesting that philopatric animals experienced a substantial direct fitness cost by remaining in their natal group. These data yield important insights into the adaptive bases for group living in C. sociabilis and suggest that constraints on natal dispersal are an important factor favoring group living in this species.Communicated by J. Wilkinson     

Male morphology and behavior correlate with reproductive success in the American redstart (Setophaga ruticilla)

Summary In a population of American redstarts (Setophaga ruticilla), we determined whether a relationship exists between seasonal reproductive success (RS) and a variety of male and female morphological and behavioral characters including plumage and male song. Adult males differ dramatically in plumage from yearling males and also exhibit variable amounts of black on their breast (bib size). Adult males were more successful than yearlings in terms of the number of eggs laid by their females. Among adult males, those with smaller bibs (less black) had females that laid more eggs, and produced more hatchlings and more fledglings. We found no evidence to indicate that this result was a consequence of territory quality. We examined a number of features of song but none alone was a predictor of RS; however, one song feature (rare repeat song) correlated with bib size. When bib size and rare repeat song were analyzed simultaneously, both were found to relate to RS. No female features were predictors of RS, but females arriving earlier at the breeding site mated with males with smaller bibs. The evidence is consistent with the view that plumage of redstarts may be used as a basis for female choice. Offprint requests to: R.E. Lemon     

Factors influencing length of maternal care in brown bears (<Emphasis Type="Italic">Ursus arctos</Emphasis>) and its effect on offspring

Length of maternal care, i.e. the interval between successfully raised litters, is the most important factor explaining the variation in reproductive rate among brown-bear (Ursus arctos) populations. In this paper, we examine the variation in length of maternal care in radio-marked brown bears and its effect on their offspring in northern Sweden. Young stayed with their mothers for 1.4–1.5 or 2.4–2.5 (in one case 3.5) years and were weaned with body masses varying from 17 to 69 kg. The probability of yearling litters staying with their mother for a 2nd year increased with decreasing yearling body mass, and was higher for litters with two offspring than for litters with one or three to four offspring. Staying with their mothers for a 2nd year had a positive effect on mass gain in yearlings and this effect was more pronounced in litters with two than three to four offspring. Body mass of 2-year-olds was not related to age of weaning, suggesting that keeping offspring for an additional year mainly compensated for low yearling body mass. If large offspring body mass positively affects later offspring survival and reproduction, mothers may be able to optimize the length of maternal care according to the litter size and the size of their yearlings.Communicated by F. Trillmich     

Age-specific social dominance affects habitat use by breeding American redstarts (Setophaga ruticilla): a removal experiment

Summary By removing older males from their breeding territories, we tested the hypothesis that age-related dominance behavior influenced the pattern of habitat selection by breeding American redstarts Setophaga ruticilla (Aves: Parulinae). Fifteen older male redstarts removed in five experimental replicates during three breeding seasons were replaced by ten yearling and five older males; no redstart males of either age colonized the control sites during the same time periods, although two yearlings disappeared. Significantly more yearling males (67%, n=9) colonized the vacated areas than were present in the redstart population at large (26.8%, n=209). We reject the alternative hypothesis that yearling male redstarts occupy different habitats from older males because of age-related (innate) habitat preferences. Redstarts that colonized the territories made vacant by our removals (i.e., floaters) were a behaviorally heterogeneous group of animals. The presence of both yearling and older male floaters indicates that suitable habitat is limiting for this species and that intraspecific competitive interactions are important in habitat distribution, and potentially in population regulation.     

Why do female Belding's ground squirrels disperse away from food resources?

We examined the effects of food provisioning on the natal dispersal behavior of Belding's ground squirrels (Spermophilus beldingi). We provided extra food to adult and yearling females in their maternal territories during pregnancy and lactation, and to offspring of these females in their natal areas for 6 weeks after weaning. We used unprovisioned young of unprovisioned mothers as controls. Provisioning influenced the probability of dispersal from the natal area by female but not male S. beldingi. All surviving male S.␣beldingi dispersed by 55 weeks of age, regardless of whether they and their mothers received extra food. By contrast, we observed a significant trend, beginning 3 weeks after weaning and continuing through the yearling year, for a greater proportion of provisioned than control female S. beldingi to emigrate from the natal area. Competition for food did not appear to influence natal dispersal of females. However, overall population density, density of females weaning litters, and rates of aggression and vigilance among these females, were higher in provisioned than control areas, suggesting that competition for non-food resources was unusually intense in provisioned areas. We propose that juvenile female, but not juvenile male, S. beldingi may emigrate from the natal site to increase access to areas with low densities of conspecifics. Together with findings of earlier workers, our results suggest that spatial and temporal distributions of environmental resources are important influences on the dispersal behavior of female ground squirrels. Received: 28 February 1996 / Accepted after revision: 16 October 1996     

Maternal effects on post-weaning physical and social development in juvenile mountain goats (Oreamnos americanus)

Little is known about maternal effects on post-weaning development, yet they may be important because maternal care could have long-term consequences only evident when offspring approach adulthood. We have assessed the effects of maternal age, current reproduction (presence of a kid of the year) and social rank on the body mass, horn length and social rank of 1- and 2-year-old mountain goats (Oreamnos americanus). Maternal reproductive status and social rank did not affect the mass or horn length of either yearlings or 2-year-olds. Maternal age was positively correlated with yearling body mass for males but not females. We could not detect any maternal age effects on body mass of 2-year-olds. Maternal age and spring forage quality were positively correlated with horn length of yearlings of both sexes, but not of 2-year-olds. Juvenile females showed compensatory growth in mass between 1 and 2 years of age, but males did not. Neither sex showed compensatory growth in horn length. None of the maternal characteristics we examined directly affected the social rank of juveniles, which increased with body mass. Social rank in female mountain goats seems to be established early in life and maintained to adulthood. By affecting yearling development, maternal age could affect the reproductive success of males.     

Food allocation rules vary with age and experience in a cooperatively breeding parrot

Although it is known that parents can differ in their optimal resource allocation to offspring in size-structured broods, the mechanisms determining differences in allocation rules of carers are not yet clarified. In cooperatively breeding species, breeders and non-reproductive helpers often differ in their fitness payoffs of providing care and in their breeding experience. Cooperative breeders thus provide an appropriate system to examine two hypotheses originally proposed to explain differences in food allocation among parents: (i) food allocation between carers differs because of the distinct cost-benefit ratio of selective feeding (i.e. breeders and helpers are expected to differ in food allocation) and (ii) carers differ in their ability to feed selectively (i.e. differences in food allocation are expected between experienced adults and inexperienced yearlings). We compared feeding rates with which breeders, old helpers and yearling helpers provisioned nestlings of different hatching rank. The influence of experience upon food allocation was further assessed by comparing food allocation of yearlings early and late during nesting. We show that allocation rules differ between age classes because breeders and old helpers fed the youngest chicks most, whereas yearlings showed the opposite pattern. The role of experience was supported by the fact that yearlings adjusted food allocation to that observed in experienced adults during the breeding season. We thus suggest that food allocation in El Oro parakeets depends either on differential skills of adults to transfer food to the youngest chick or on their ability to recognize nestling needs.     

Birth season testosterone levels in male Verreaux’s sifaka, Propithecus verreauxi: insights into socio-demographic factors mediating seasonal testicular function

Previous research suggests that although reproduction and testicular function in wild sifaka (Propithecus verreauxi) are highly seasonal, birth season elevations in fecal testosterone (T) in transferring males indicate that social factors may be as important as climatic factors in regulating reproductive function in sifaka. This paper examines the relationship of male dispersal and social status to the patterning of birth season aggression and testicular function in P. verreauxi at Beza Mahafaly Special Reserve, Madagascar. Behavioral and hormonal data were collected from a total of 38 adult males, 15 residing in seven stable groups and 23 living in eight unstable groups, yielding 186 fecal samples and 493 focal animal hours of observation. The results suggest that birth season elevations in fecal testosterone are the consequence of social disruption resulting from male movements between groups and the particular responses of individual males to dispersal events. Hormonal responses to dispersal events appear to precede behavioral responses and occasionally register reactions not concomitantly evident in the behavioral response, and may be predictive of future events. Hormonal reactions occurred primarily in resident alpha males, suggesting that they differ in their assessment of destabilizing influences, perhaps due to different reproductive opportunities and/or investment. These findings are important for the new insights they provide into the role of androgens in mediating male dispersal, life history, and reproductive strategies, and suggest that investigations of androgen-behavior interactions in free-ranging populations can be a powerful new tool for assessing the contextual and motivational basis of social behavior. Received: 22 November 1999 / Revised: 27 April 2000 / Accepted: 15 October 2000     

Correlates of horn and antler shape in bovids and cervids

Using analyses that control for phylogeny, we examine whether the shapes of horns and antlers in ungulates are related to style of fighting, environmental factors, mating and social systems, or are simply arbitrary. Many of the predictions that relate horn shape to species' fighting tactics and to their mating and social systems are supported. Bovids with tips facing inwards are likely to wrestle with their horns, be monogamous and solitary, whereas those with tips facing out tend to be polygynous and live in large groups. Smooth horns are used for stabbing, and are found in females of polygynous species living in large groups. In female bovids, twisted horns are found in large species and are used in wrestling whereas, in male bovids, straight horns are found in solitary species. Finally, deer with more than five antler tines tend to be large and to fight by fencing. There was little support for shape of horns and antlers being related to environmental factors, nor were shapes arbitrary as might be expected if they had arisen through female choice. Body size had little effect on these results. In general, monogamous solitary bovids have straight inwardly facing horns whereas polygynous group-living species demonstrate a wide variety of horn shapes.Electronic Supplementary Material Supplementary material is available in the online version of this article at .Communicated by M. Elgar     

Copula duration and testes size in the yellow dung fly,Scathophaga stercoraria (L.) : the effects of diet,body size,and mating history

Summary Seasonal variation in mean hind tibia length and mean testes length is investigated in the yellow dung fly, Scathophaga stercoraria (L.). There is a cycle in mean hind tibia length and mean testes length over a season. The body size curve peaks later than the testes length curve, showing that there is no fixed relationship between the two variables. The causes of variation in testes size and its influence on copula duration are experimentally examined. Increasing the number of Drosophila eaten per day leads to increased mature testes length. Males with larger testes copulate for longer than males with smaller ones, and smaller males copulate for longer than do larger males. While testes shrivel with successive copulas, copula duration remains constant. The more females a male is prepared to copulate with in a day (up to five), the longer he copulates with each. The shrinkage of the testes of males collected throughout a day suggests that males copulate with an average of 4 females per day. The costs of sperm production are thus shown to have a significant influence on the copula duration.Offprint requests to: P.I. Ward at the second address     

Social structures,genetic structures and dispersal strategies in Australian rabbit (<Emphasis Type="Italic">Oryctolagus cuniculus</Emphasis>) populations

In this study, the pattern of movement of young male and female rabbits and the genetic structures present in adult male and female populations in four habitats was examined. The level of philopatry in young animals was found to vary between 18-90% for males and 32-95% for females in different populations. It was skewed, with more males dispersing than females in some but not all populations. Analysis of allozyme data using spatial autocorrelation showed that adult females from the same social group, unlike males, were significantly related in four of the five populations studied. Changes in genetic structure and rate of dispersal were measured before and during the recovery of a population that was artificially reduced in size. There were changes in the rate and distance of dispersal with density and sex. Subadults of both sexes moved further in the first year post crash (low density) than in the following years. While the level of dispersal for females was lower than that of the males for the first 3 years, thereafter (high density) both sexes showed similar, low levels of dispersal (20%). The density at which young animals switch behaviour between dispersal and philopatry differed for males and females. The level of genetic structuring in adult females was high in the precrash population, reduced in the first year post crash and undetectable in the second year. Dispersal behaviour of rabbits both affects the genetic structure of the population and changes with conditions. Over a wide range of levels of philopatry, genetic structuring is present in the adult female , but not the male population. Consequently, though genetic structuring is present, it does not lead to inbreeding. More long-distance movements are found in low-density populations, even though vacant warrens are available near birth warrens. The distances moved decreased as density increased. Calculation of the effective population size (N_e) shows that changes in dispersal distance offset changes in density, so that N_e remains constant.