Maternal rank is not correlated with cub survival in the spotted hyena, Crocuta crocuta

Reproductive success in female spotted hyenas, Crocuta crocuta,is positively correlated with social rank. High-ranking femalesare known to produce more offspring, but the effects of maternalrank on early cub survivorship have not been investigated. Cubmortality was examined over a 4-year period in one clan of wild-livingspotted hyenas in Kenya. Data were obtained for 100 cubs in63 litters produced by 27 adult females. Survivorship of cubsfrom birth through their first year was examined as a functionof litter size, sex of cubs, and maternal rank. Overall, cubmortality was high (61%). Contrary to expectation, singletoncubs did not survive better than twins, and there was no differencein survivorship between female and male cubs. High-ranking motherswere not more successful at raising twins or daughters thanwere low-ranking mothers. There was no correlation between cubmortality and maternal rank. Peaks in cub mortality coincidedwith life stage events, including mean age of arrival at a communalden, and age at which cubs began visiting kills. Documentedcauses of mortality included intraclan infanticide, disease,orphaning, predation by lions, and a mechanism of filial infanticidethat has not been previously described in this species: selectivelitter reduction by mothers via partial litter abandonment.No instances of facultative or obligate siblicide were detected.During this study, association between rank and number of cubssurviving to 1 year of age appeared to be due to differencesin reproductive output and not differential survival of cubswithin their first year.     

A comparative study of litter size and sex composition in a large dataset of callitrichine monkeys

In many birds and mammals, the size and sex composition of litters can have important downstream effects for individual offspring. Primates are model organisms for questions of cooperation and conflict, but the factors shaping interactions among same‐age siblings have been less‐studied in primates because most species bear single young. However, callitrichines (marmosets, tamarins, and lion tamarins) frequently bear litters of two or more, thereby providing the opportunity to ask whether variation in the size and sex composition of litters affects development, survival, and reproduction. To investigate these questions, we compiled a large dataset of nine species of callitrichines (n = 27,080 individuals; Callithrix geoffroyi, Callithrix jacchus, Cebuella pygmaea, Saguinus imperator, Saguinus oedipus, Leontopithecus chrysomelas, Leontopithecus chrysopygus, Leontopithecus rosalia, and Callimico goeldii) from zoo and laboratory populations spanning 80 years (1938–2018). Through this comparative approach, we found several lines of evidence that litter size and sex composition may impact fitness. Singletons have higher survivorship than litter‐born peers and they significantly outperform litter‐born individuals on two measures of reproductive performance. Further, for some species, individuals born in a mixed‐sex litter outperform isosexually‐born individuals (i.e., those born in all‐male or all‐female litters), suggesting that same‐sex competition may limit reproductive performance. We also document several interesting demographic trends. All but one species (C. pygmaea) has a male‐biased birth sex ratio with higher survivorship from birth to sexual maturity among females (although this was significant in only two species). Isosexual litters occurred at the expected frequency (with one exception: C. pygmaea), unlike other animals, where isosexual litters are typically overrepresented. Taken together, our results indicate a modest negative effect of same‐age sibling competition on reproductive output in captive callitrichines. This study also serves to illustrate the value of zoo and laboratory records for biological inquiry.     

Reproductive parameters of the common genet <Emphasis Type="Italic">Genetta genetta</Emphasis> (Linnaeus, 1758) in Southwest Europe

We used information from 148 litters belonging to 119 different wild and captive female common genets (Genetta genetta) in SW Europe over a period of almost 50 years to study certain reproductive parameters. Data were obtained from a combination of sources including carcasses, live captures, telemetry-based studies, captive genets, and reports in the literature. The circannual pattern of breeding was fairly similar in both wild and captive genets, with births occurring in all months except December, albeit with a large peak in spring and a secondary peak in autumn. The sex ratio at birth was near to 1:1, and mean litter size was 2.16 ± 0.76 cubs/litter with a range of 1–5 cubs. Litters of two cubs were the most frequent both in wild and captive settings. In captivity, we have observed the births of replacement litters and double litters; however, we did not find evidence of wild females that having two litters in the same year. We conclude that the possibility of breeding at any time of the year is facilitated by the continuous spermatogenesis of the males and by the ability of females to enter into estrus after losing a litter prematurely and to breed twice in the same year.     

Reproductive biology of captive female Eurasian lynx, Lynx lynx

Studies on wild Eurasian lynx (Lynx lynx) have revealed variation in reproduction between areas, years and individuals. In order to explore potential causes for this variation other than food supply, we analysed data from captive lynx, which provide conditions with minimal environmental variation as all were fed ad libitum. Data from 37 individual female lynx were available from 20 zoos in Norway, Sweden, Finland, Switzerland and the Czech Republic. Data on 177 reproductive events (where a male was available to the female at mating time) are presented. Of these events, 85% resulted in litters being born. Average litter size was 1.95, with a variation from 1 to 4. The mean birth date was 26th May, and sex ratio was not significantly different from parity. The probability of reproduction was related to age, with fewer litters produced by the very young (2–3-year old), and no sign of a senescence effect. However, a clear effect of senescence on litter size was evident. The captive lynx did not have higher reproductive rates than wild lynx, indicating that either factors other than food supply are driving the variation in wild lynx reproduction, or that a factor such as stress may be causing additional variation in the captive population.     

Litter size and infant survivorship in wild groups of cotton‐top tamarins (Saguinus oedipus) in Colombia

Cotton‐top tamains (Saguinus oedipus) are a critically endangered primate found only in Colombia. Efforts to conserve this species are centered on developing effective management plans that integrate biological information regarding population dynamics and factors that influence their survival. This study documented infants born to wild cotton‐top tamarin females from 1994–2008 at two distinct field sites in northern Colombia. Our studies have shown that wild cotton‐top tamarins typically give birth to one litter each year and infant survival to 6 months of age was greater in the wild than has been reported in captive colonies. However, similar to reports from captive colonies, litter size of wild cotton‐top tamarins ranges from 1–3 infants, with twin litters most common. Here we report the first occurrence of triplet litters in nearly 20 years of observing wild cotton‐top tamarin groups. Over the first 3 months of life, wild‐born infants exhibited highest mortality during the first week of life, similar to reports from captive colonies. Infant survival in the wild also increases with successive litters as it does in captivity. However, inter‐birth interval, group size, and the number of adult males in the group did not appear to influence infant survival in the wild. The value of such long‐term data from field studies aids in the information that can be used to model future population trends and develop effective conservation plans for this critically endangered primate. Am. J. Primatol. 71:707–711, 2009. © 2009 Wiley‐Liss, Inc.     

Early sibling aggression in mammals and its hormonal correlates

Early sibling aggression is a widespread phenomenon in birds. Ornithologists distinguish species with "obligate" and "facultative" siblicide. Sibling aggression was described in some mammal species: the domestic pig (Sus scrofa), the spotted hyena (Crocuta crocuta), the Eurasian lynx (Lynx lynx), and the Iberian lynx (L. par-dinus). In all of them, sibling aggression corresponds well with the "facultative" siblicide model in birds. Sibling aggression was observed at the age of 36-64 days in both lynx species. It is usually restricted to a single fight and can change the hierarchical structure and growth rate of the kittens. In the spotted hyena and the domestic pig, the frequency and intensity of aggressive interactions between siblings are strongest during the first days of postnatal ontogeny and then decrease gradually. The newborns of these species are much developed than newborn lynx kittens. Usually adult lynx females, in contrast to hyenas and pigs, try to stop sibling fights. This is probably related to the larger parental investment at the time of the fight in lynxes (a kitten's body weight is about 10% of the mother's) than in pigs (0.5%) and hyenas (1.9%). Sibling aggression in spotted hyenas could be related to the high level of androstenedione and is not related to testosterone concentration. In the Eurasian lynx, female sibs attack their littermates slightly more often than male sibs do, and sibling aggression is not testosterone-dependent. Hormones secreted by the adrenal glands may play an important role in this phenomenon. The data available so far, however, do not positively confirm the presence of hormonal trigger effects in mammal sibling aggression.     

Markedly Elevated Antibody Responses in Wild versus Captive Spotted Hyenas Show that Environmental and Ecological Factors Are Important Modulators of Immunity

Evolutionary processes have shaped the vertebrate immune system over time, but proximal mechanisms control the onset, duration, and intensity of immune responses. Based on testing of the hygiene hypothesis, it is now well known that microbial exposure is important for proper development and regulation of the immune system. However, few studies have examined the differences between wild animals in their natural environments, in which they are typically exposed to a wide array of potential pathogens, and their conspecifics living in captivity. Wild spotted hyenas (Crocuta crocuta) are regularly exposed to myriad pathogens, but there is little evidence of disease-induced mortality in wild hyena populations, suggesting that immune defenses are robust in this species. Here we assessed differences in immune defenses between wild spotted hyenas that inhabit their natural savanna environment and captive hyenas that inhabit a captive environment where pathogen control programs are implemented. Importantly, the captive population of spotted hyenas was derived directly from the wild population and has been in captivity for less than four generations. Our results show that wild hyenas have significantly higher serum antibody concentrations, including total IgG and IgM, natural antibodies, and autoantibodies than do captive hyenas; there was no difference in the bacterial killing capacity of sera collected from captive and wild hyenas. The striking differences in serum antibody concentrations observed here suggest that complementing traditional immunology studies, with comparative studies of wild animals in their natural environment may help to uncover links between environment and immune function, and facilitate progress towards answering immunological questions associated with the hygiene hypothesis.     

Birth and simultaneous rearing of two litters in a pack of captive African wild dogs (Lycaon pictus)

Two female African wild dogs (Lycaon pictus) in a pack of 10 animals at the Bronx Zoo gave birth to litters of nine and seven pups within a 2‐day period. Two pups from the litter of seven died before they were 2 weeks old, but the other 14 survived past weaning. Litter size, sex ratio, and pup developmental stages were all consistent with data obtained from other captive‐ and wild‐born litters. Both dams were very attentive to their litters, and during the pups' first 2 weeks spent >90% of their time in the dens with the pups. By 2 days after birth and during their first 2 weeks of life, the pups spent 86% of their time in a nursing position. Before parturition and during the first week post‐partum, one of the dams (DAL) was clearly dominant to the other (WHI). However, the females' dominance rank reversed 2 weeks post‐partum. On two occasions during the pups' first 2 weeks DAL stole and reared one of WHI's pups, but between Weeks 3 and 4 all of DAL's pups were stolen and WHI raised both litters as one until they were weaned. The pups began spending time out of their den at approximately 1 month of age. The timing of the births, the design of the wild dogs' management facility, and the presence of several dens in different enclosures within the facility all likely contributed to the successful rearing of the litters. The pups in both litters were very similar in size because of the short interval between births, so one litter did not have a competitive advantage over the other with respect to gaining access to the dam for milk. The facility's design helped mitigate aggression within the pack, and the presence of multiple dens enabled to dams to move the pups to different den sites and allowed the two females to stay visually and spatially apart from each other while remaining with the pack. Zoo Biol 0:1–17, 2006. © 2006 Wiley‐Liss, Inc.     

Effects of age and parity on litter size and offspring sex ratio in golden hamsters (Mesocricetus auratus)

Golden hamsters that were mated repeatedly from 55 days of age produced 6-12 litters. Litter size at birth rose between the 1st and 2nd litters, peaked on the 3rd, and declined steadily after the 5th litter. Offspring sex ratio (% male) at birth followed a similar pattern: increasing between the 1st and 2nd litters, remaining high through the 3rd, and becoming increasingly female-biased thereafter. Weaning success decreased sharply after the 6th litter and most dams failed to raise any young to weaning after the 9th litter. These sequential effects on litter size, offspring sex ratio and weaning success were also observed in females mated once at different ages, but they occurred considerably later in life, i.e. increasing parity hastened the effects of advanced age. These age- and parity-related changes in litter composition are consistent with the Trivers-Willard hypothesis that physiologically-stressed females would skew offspring sex ratios to favour daughters. However, since the observed changes in sex ratio were probably due to differential prenatal mortality, their adaptive significance is unclear.     

Age specific fecundity,litter size,and sex ratio in the chacoan peccary (Catagonus wagneri)

Ten years of data on the reproductive biology of the Chacoan peccary (Catagonus wagneri) were analyzed to determine average litter size, sex ratio, timing of births, and individual and age-specific fecundity. Data were obtained from a captive herd of chacoan peccaries located in the western Paraguayan Chaco. Births peak in the austral spring months of September, October, and November, with fewer litters born during the dry season months of June, July, and August. The average litter size was 2.4 with a sex ratio of 56:44 (M:F). There was no significant difference in litter size among individual females or among females of different ages. Finally, there was some individual variation in the age at which sows produced their first litter, but no discernible variation among sows in the average time between litters. Zoo Biol 16:301–307, 1997. © 1997 Wiley-Liss, Inc.     

Do non‐additive effects on decomposition in litter‐mix experiments result from differences in resource quality between litters?

Differences in resource quality between litter species have been postulated to explain why litter-mixtures may decompose at a different rate to that which would be predicted from single species litters (termed 'non-additive effects'). In particular, positive, non-additive effects of litter-mixing on decomposition have been explained by differences in initial nitrogen concentration between litter species. This interpretation is confounded because litter species that differ in nitrogen content also differ by a number of other resource quality attributes. Thus, to investigate whether initial nitrogen concentration does account for positive, non-additive effects of litter-mixing, we mixed grass litters that differed in initial nitrogen concentration but not species or structural plant part identity, and then followed mass loss from the litter-mixes over time. We used the litterbag technique and three grass species for which a gradient of four distinct initial nitrogen concentrations had been generated. We produced all no- to four-mix compositions of litter qualities for each species. Litter from different species was never mixed.
Contrary to what would be predicted, we found that when litters of the same species but with different initial nitrogen concentrations were mixed, that negative, non-additive effects on decomposition were generally observed. In addition, we found that once mixed, increasing litter quality richness from two to four mixtures had no significant, non-additive effect on decomposition. Litter quality composition explained little of the experimental variation when compared to litter quality richness, and different compositions generally behaved in the same manner. Our findings challenge the commonly held assumption that differences in nitrogen concentration between plant species are responsible for positive, non-additive effects of litter-mixing on decomposition.     

Reproductive performance of a laboratory colony of the deermouse, Peromyscus maniculatus gambelii.

During a 23-mo period approximately 600 litters and 2,500 offspring were produced in a laboratory colony established with wild-trapped deermice (Peromyscus maniculatus gambelii). The incidence of pregnancy and size of litters increased with domestication. Average litter size was 4.3 for all litters and 4.1 and 4.7 for litters born to wild-trapped and first generation females, respectively. Litter size increased up to the seventh sequential pregnancy for both wild-caught and first-generation females and then tapered off. In all litters more males were born than females. The departure from a 1:1 ratio was significant for the combined data and for litters of first-generation females, but not for litters of wild-caught females.     

Predation, sensitivity, and sex: why female black rhinoceroses outlive males

Among sexually dimorphic, polygynous mammals, adult femalestend to outlive males and respond more strongly to predatorsthan males. We asked whether a monomorphic, polygynous speciesvirtually immune to predation due to large size (black rhinoceros,Diceros bicornis) conforms to this pattern. Data on 193 interactionswith lions (Panthera leo) and spotted hyenas (Crocuta crocuta)in two nonhabituated populations in Namibia studied from 1991to 1993 revealed that: (1) females were more vigilant or aggressivethan males to either of the potential predators; and (2) whethersolitary or with calves, females attacked more often than males.Although solitary females tended to be more aggressive to lionsthan to hyenas, neither females with calves or males seemedto discriminate between the two carnivores. We also simulatedthe behavior of human predators (poachers) during 69 encounterswith rhinoceroses. While both sexes abandoned local sites becauseof our presence, females ran farther than males, covering upto 40 km in a day. These findings implicate a behavioral mechanismto explain why secondary sex ratios favor females - males aremore prone to human predation, a prediction consistent withdata from 12 populations throughout Africa. Black rhinocerosesappear to be an unanticipated exception to the well-establishedpattern of male-biased mortality in polygynous mammals; in theabsence of intense human predation (a recent event), male mortalityfails to exceed that of females, suggesting that intrasexualcompetition in a polygynous mammal: may not be the primary causeof unbalanced secondary sex ratios. Our results on the causesof sex differences in mortality and in responsiveness to differentpredators reinforce the relevance of behavioral ecology to conservation;such information is necessary for planning how best to minimizenegative human influences on the few remaining wild Africanrhinos.     

Effects of inbreeding on reproductive success,performance, litter size,and survival in captive red wolves (Canis rufus)

Captive‐breeding programs have been widely used in the conservation of imperiled species, but the effects of inbreeding, frequently expressed in traits related to fitness, are nearly unavoidable in small populations with few founders. Following its planned extirpation in the wild, the endangered red wolf (Canis rufus) was preserved in captivity with just 14 founders. In this study, we evaluated the captive red wolf population for relationships between inbreeding and reproductive performance and fitness. Over 30 years of managed breeding, the level of inbreeding in the captive population has increased, and litter size has declined. Inbreeding levels were lower in sire and dam wolves that reproduced than in those that did not reproduce. However, there was no difference in the inbreeding level of actual litters and predicted litters. Litter size was negatively affected by offspring and paternal levels of inbreeding, but the effect of inbreeding on offspring survival was restricted to a positive influence. There was no apparent relationship between inbreeding and method of rearing offspring. The observable effects of inbreeding in the captive red wolf population currently do not appear to be a limiting factor in the conservation of the red wolf population. Additional studies exploring the extent of the effects of inbreeding will be required as inbreeding levels increase in the captive population. Zoo Biol 29:36–49, 2010. © 2009 Wiley‐Liss, Inc.     

Reproduction of Asian chipmunks (Tamias sibiricus) in captivity

Factors affecting reproduction in captive Asian chipmunks, Tamias sibiricus, were examined by means of a survey of chipmunk breeders in Great Britain. Sixteen breeders were asked about the conditions under which their chipmunks were kept and their success in breeding them. Results covered 205 female-years of pairing. Breeding was promoted by large cage size and early weaning of young, and inhibited by the presence of other rodent species nearby and by extended photoperiod. A diet of seeds and nuts, with some fresh fruits and vegetables, appeared to be adequate for breeding. Breeding occurred in 80–91& of female-years when neither extra lighting nor other rodents were present, irrespective of cage size. Weaning age affected the occurrence of second litters in a year. Second litters were born less often (0–13&) of breeding (female-years) if young of the first litter remained with the mother 8 weeks or longer. Second litters were more frequent (14–45& of breeding female-years) if first litters were removed by 6.5 weeks of age, and this frequency increased with cage size. Litter size also increased with cage size (means ranged from 3 to 7). Overall breeding success increased with cage size and decreased with the presence of other rodents. In most colonies without other rodents near, mean success rate ranged from 3.1 to 8.5 young/female-year. In colonies with other rodents present mean success rate varied from 0 to 3.3 young/female-year. It is suggested that the mechanisms that control population size in the wild also act to inhibit or promote breeding in captivity.     

Male-biased reproduction and sex-ratio adjustment in muskrats

Summary Sex ratios of a population and of litters were sampled in muskrats in Ontario, Canada. Sex ratios of litters sampled from nests were male biased (54% male). Until weaning, no differential costs of producing and rearing male and female young were identified that could account for this greater production of males. Following weaning, however, male-biased dispersal of juveniles from their natal site and more frequent acquisition by females of these sites as breeding sites the following year suggested a greater investment by adult females in female young. Therefore, competition between female siblings for the acquisition of their natal site may be sufficient to result in the greater production of males. In addition, the simultaneous occupation of, and competition between, siblings and parents for the resources of the natal home range may not be necessary for local resource competition to result in a greater production of the dispersing sex. Greater-than-expected binomial variance in sex ratios of litters suggested that adjustment of sex-ratios occurred. However, we were unable to associate the adjustment of litter sex ratios with changes in maternal condition. The greater production of males and the predominance of monogamous associations between adults in this population may have lead to slightly greater variation in male fitness than female fitness. Therefore, a female in better-than-average condition may have benefited by producing more males. Similarly, a lower cost of producing dispersing males may allow nutritionally-stressed females to reduce their total expenditure on offspring by producing more males. Because these experiments were non-manipulative, maternal condition may not have varied sufficiently during this study to detect adjustments of litter sex ratios resulting from either of the above mechanisms acting separately, but the combined effects of small differences in matermal condition and selective pressures operating in the same direction may have resulted in the observed deviation from the binomial.     

Reproductive patterns in the domestic dog--a retrospective study of the Drever breed

The aim of this study was to examine the differences, between seasons of the year, in the distribution of matings and whelpings, litter size, pup deaths, and sex ratio in domestic dogs. Furthermore, we wanted to examine the effects of age and parity of the bitch at the time of whelping on litter size, as well as the effect of litter size on gestational length. A final aim was to investigate the fertility and frequency of whelping problems in a private kennel of Drever dogs. Data from the Swedish Kennel Club (SKK) registry for the Drever breed during 1995-2006, comprising a total of 2717 litters, were analyzed together with more detailed data from a private, professional kennel of Drevers, with a total of 285 matings and 224 whelpings, during the same time period. The most matings took place during winter, and the fewest during summer; consequently, most whelpings occurred during the winter and spring seasons. Of the 285 mated bitches, 78.6% whelped, 6.25% experienced dystocia, and 5.36% underwent Cesarean section. The pup death rate was 7.6%. The largest litters were born during spring. Litter size was negatively correlated with duration of pregnancy (r=-0.18). Each pup more than average caused a shortening of the gestation by 0.25 days, and each pup less a corresponding lengthening. Bitches giving birth to their first litter after 4 years of age produced a smaller litter than younger bitches. Litter size decreased after 5 years in all bitches. The number of born pups at the private kennel increased from the first to the third parity, then decreased. The number of registered pups increased from the first to the second parity in the SKK data and from the second to the third parity in the data from the private kennel, then decreased. Mating a bitch only once resulted in a smaller litter size. None of the studied factors had any effect on the sex ratio of the pups. There were significant differences between males in whelping rate among the mated bitches, but no difference in mean litter size, which indicates a female problem rather than a male one. Available data suggest that the domestic dog is still under considerable seasonal influence, although modified by ambient and management factors.     

WHY DO SOME SIBLINGS ATTACK EACH OTHER? COMPARATIVE ANALYSIS OF AGGRESSION IN AVIAN BROODS

In many parentally fed species, siblings compete for food not only by begging and scrambling, but also by violently attacking each other. This aggressive competition has mostly been studied in birds, where it is often combined with dominance subordination, aggressive intimidation, and siblicide. Previous experimental and theoretical studies proposed several life-history, morphological, and behavioral variables that may facilitate the evolution of broodmate aggression, and explain its taxonomic distribution. Here we apply phylogenetic comparative analyses for the first time to test the influence of five hypothesized facilitators of the evolution of broodmate aggression, analyzing 69 species in seven avian families using two quantitative measures of aggression: incidence and intensity. We show that incidence and intensity of aggression increase with long nestling periods and indirect feeding, and small brood size is associated with intense aggression. Large food parcels were not correlated with either the incidence or intensity of aggression. Our study suggests that indirect feeding, long nestling periods, and small broods, possibly in combination with other factors, have tended to favor the evolution of aggressive broodmate competition.     

Birth-sex ratios and local resource competition in roe deer, Capreolus capreolus

We investigated variation in the primary sex ratio within andbetween 14 populations of roe deer (Capreolus capreolus) inrelation to maternal body condition. The sex ratio was increasinglymale biased as average maternal body weight decreased. Thisrelationship did not vary according to the population consideredand was not affected by the litter size produced. This relationshipwas also apparent within populations. These results indicatethat, where environmental conditions are limiting, roe doestend to produce male-biased litters. Dispersal is more commonand occurs at an earlier age among male juveniles in this species,particularly as density increases and resources become increasinglyscarce. Thus, we suggest that where females experience environmentalstress, they tend to produce male kids to avoid potential futurelocal resource competition posed by female offspring. [BehavEcol 7: 461–464 (1996)]