Non-breeding Males Research Articles

When the ball is in the female’s court: How the scramble-competition mating system of the North American red squirrel has shaped male physiology and testosterone dynamics

Male reproductive success in most mammals is determined by their success in direct inter-male competition through aggression and conflict, resulting in female-defense mating systems being predominant. This is linked to male testosterone levels and its dynamics. However, in certain environments, a scramble-competition mating system has evolved, where female reproductive behavior takes precedence and male testosterone dynamics are unlikely to be related to inter-male competition. We studied the North American red squirrel (Tamiasciurus hudsonicus), a species with a well-established scramble-competition system. Using an ACTH hormonal challenge protocol as a proxy for competitive interactions, we compared the testosterone dynamics in breeding males in late winter with that in nonbreeding males in late spring in the Yukon. To gain an integrated picture of their physiological state, we also assessed changes in their stress response, body mass, energy mobilization, and indices of immune function. Testosterone levels at the base bleed were high in breeding males (2.72ng/mL) and virtually absent in non-breeding males (0.04ng/mL). Breeding males were in better condition (heavier body mass, higher hematocrit, and higher erythrocytes), had higher indices of immune function (neutrophil:lymphocyte ratio), but a similar ability to mobilize energy (glucose) compared with non-breeding males. Though total cortisol was higher in non-breeding males, free cortisol was twice as high in breeding males as their corticosteroid binding globulin levels were half as high. In response to the ACTH challenge, testosterone levels in breeding males declined 49% over the first hour and increased 36% over the next hour; in non-breeding males levels showed no change. Free cortisol increased only modestly (26% in breeding males; 23% in non-breeding males). Glucose levels changed similarly in breeding and nonbreeding males, declining for the first 30min and then increasing for the next 60min. Thus, testosterone and components of the stress axis function in a profoundly different manner in male red squirrels than in males of mammals with female-defense mating systems. There are four probable interrelated reasons for these adaptations in male red squirrels: the marginal benefits of each mating, the constraints of mate searching away from their own resource-based territories, energy mobilization in a harsh environment, and a long life span.

Harvest and group effects on pup survival in a cooperative breeder.

Recruitment in cooperative breeders can be negatively affected by changes in group size and composition. The majority of cooperative breeding studies have not evaluated human harvest; therefore, the effects of recurring annual harvest and group characteristics on survival of young are poorly understood. We evaluated how harvest and groups affect pup survival using genetic sampling and pedigrees for grey wolves in North America. We hypothesized that harvest reduces pup survival because of (i) reduced group size, (ii) increased breeder turnover and/or (iii) reduced number of female helpers. Alternatively, harvest may increase pup survival possibly due to increased per capita food availability or it could be compensatory with other forms of mortality. Harvest appeared to be additive because it reduced both pup survival and group size. In addition to harvest, turnover of breeding males and the presence of older, non-breeding males also reduced pup survival. Large groups and breeder stability increased pup survival when there was harvest, however. Inferences about the effect of harvest on recruitment require knowledge of harvest rate of young as well as the indirect effects associated with changes in group size and composition, as we show. The number of young harvested is a poor measure of the effect of harvest on recruitment in cooperative breeders.

Metabolic and affective consequences of fatherhood in male California mice

Physiological and affective condition can be modulated by the social environment and parental state in mammals. However, in species in which males assist with rearing offspring, the metabolic and affective effects of pair bonding and fatherhood on males have rarely been explored. In this study we tested the hypothesis that fathers, like mothers, experience energetic costs as well as behavioral and affective changes (e.g., depression, anxiety) associated with parenthood. We tested this hypothesis in the monogamous, biparental California mouse (Peromyscus californicus). Food intake, blood glucose and lipid levels, blood insulin and leptin levels, body composition, pain sensitivity, and depression-like behavior were compared in males from three reproductive groups: virgin males (VM, housed with another male), non-breeding males (NB, housed with a tubally ligated female), and breeding males (BM, housed with a female and their first litter). We found statistically significant (P<0.007, when modified for Adaptive False Discovery Rate) or nominally significant (0.007<P<0.05) differences among reproductive groups in relative testis mass, circulating glucose, triglyceride, and insulin concentrations, pain sensitivity, and anxiety-like behaviors. A priori contrasts indicated that VM produced significantly more fecal pellets than BM in the tail-suspension test, had significantly higher glucose levels than NB, and had significantly lower average testis masses than did NB and BM. A priori contrasts also indicated that VM had a nominally longer latency to the pain response than NB and that VM had nominally higher insulin levels than did NB. For breeding males, litter size (one to three pups) was a nominally significant positive predictor of body mass, food consumption, fat mass, and plasma leptin concentration. These results indicate that cohabitation with a female and/or fatherhood influences several metabolic, morphological, and affective measures in male California mice. Overall, the changes we observed in breeding males were minor, but stronger effects might occur in long-term breeding males and/or under more challenging environmental conditions.

Delayed Age at First Breeding and Experimental Removals Show Large Non-Breeding Surplus in Pied Flycatchers

Avian breeding populations have been shown to be regulated by territorial behaviour, often creating a surplus of non-breeding individuals. However, most evidence is of a male non-breeder surplus, whereas for a surplus to actually buffer a population both non-breeding males and females should be present. Here, we provide descriptive and experimental evidence for the existence of a population buffer consisting of mostly male and potentially also female Pied Flycatchers using nest box areas. First we show that local recruits often do not breed in their first year, with 23% of all recruiting males observed breeding in their first year, and 51% of females. When accounting for mortality in the years prior to observed first breeding, we estimate that only 9% of all first-year males breed locally, and 29% of first-year females. Similar percentages of first-year flycatchers skipping breeding have been observed in other study populations. We show that in the year of new establishment of our nest box plots, most kno...

An immunohistochemical study on the distribution of vasotocin neurons in the brain of two weakly electric fish, Gymnotus omarorum and Brachyhypopomus gauderio.

Hypothalamic nonapeptides (arginin vasotocin-vasopressin, oxytocin-isotocin) are known to modulate social behaviors across vertebrates. The neuroanatomical conservation of nonapeptide systems enables the use of novel vertebrate model species to identify general strategies of their functional mechanisms. We present a detailed immunohistochemical description of vasotocin (AVT) cell populations and their projections in two species of weakly electric fish with different social structure, Gymnotus omarorum and Brachyhypopomus gauderio. Strong behavioral, pharmacological, and electrophysiological evidence support that AVT modulation of electric behavior differs between the gregarious B. gauderio and the solitary G. omarorum. This functional diversity does not necessarily depend on anatomical differences of AVT neurons. To test this, we focus on interspecific comparisons of the AVT system in basal non-breeding males along the brain. G. omarorum and B. gauderio showed similar AVT somata sizes and comparable distributions of AVT somata and fibers. Interestingly, AVT fibers project to areas related to the control of social behavior and electromotor displays in both species. We found that no gross anatomical differences in the organization of the AVT system account for functional differences between species, which rather shall depend on the pattern of activation of neurons embedded in the same basic anatomical organization of the AVT system.

The diet of the Antarctic fur seal Arctocephalus gazella at the South Orkney Islands in ten consecutive years

To improve the knowledge on the Antarctic fur seal foraging behavior, evaluate whether changes in its diet are associated with changes in prey availability, and evaluate whether fisheries had negative impacts on Antarctic fur seal populations, a total of 1359 scats of non-breeding males were collected in ten consecutive sampling periods between 1994 and 2003 at Laurie Island, South Orkney Islands. Antarctic krill was the most numerous prey throughout the sampling period followed in importance by fish prey. Antarctic krill also predominated by reconstructed mass, except during 1994 and 1998 when penguins were the most important prey, and during 1995 when fish dominated. Among fish, demersal-benthic species (mainly Gobionotothen gibberifrons, Chaenocephalus aceratus and Chionodraco rastrospinosus) dominated the diet except in 1997, 1998, 1999 and 2002 when myctophids (mainly Gymnoscopelus nicholsi and Electrona antarctica) were the most important fish prey. In seasons when fish dominated the diet, the diet was primarily comprised of demersal species. Although our results are in overall agreement with previous studies, the contribution to the diet of the main preys changed throughout the sampling period, probably in parallel with changes in prey abundance. According to historical results on fish consumption, the fisheries carried out in the past have had negative impacts on seal populations from the South Orkney Islands.

Non-Breeding Eusocial Mole-Rats Produce Viable Sperm--Spermiogram and Functional Testicular Morphology of Fukomys anselli.

Ansell’s mole-rats (Fukomys anselli) are subterranean rodents living in families composed of about 20 members with a single breeding pair and their non-breeding offspring. Most of them remain with their parents for their lifetime and help to maintain and defend the natal burrow system, forage, and care for younger siblings. Since incest avoidance is based on individual recognition (and not on social suppression) we expect that non-breeders produce viable sperm spontaneously. We compared the sperm of breeding and non-breeding males, obtained by electroejaculation and found no significant differences in sperm parameters between both groups. Here, we used electroejaculation to obtain semen for the first time in a subterranean mammal. Spermiogram analysis revealed no significant differences in sperm parameters between breeders and non-breeders. We found significantly larger testes (measured on autopsies and on living animals per ultrasonography) of breeders compared to non-breeders (with body mass having a significant effect). There were no marked histological differences between breeding and non-breeding males, and the relative area occupied by Leydig cells and seminiferous tubules on histological sections, respectively, was not significantly different between both groups. The seminiferous epithelium and to a lesser degree the interstitial testicular tissue are characterized by lesions (vacuolar degenerations), however, this feature does not hinder fertilization even in advanced stages of life. The continuous production of viable sperm also in sexually abstinent non-breeders might be best understood in light of the mating and social system of Fukomys anselli, and the potential to found a new family following an unpredictable and rare encounter with an unfamiliar female (“provoked or induced dispersal”). Apparently, the non-breeders do not reproduce because they do not copulate but not because they would be physiologically infertile. The significantly increased testes volume of breeding males (compared to non-breeders) is in agreement with previously found higher testosterone levels of breeders.

Consequences of Fatherhood in the Biparental California Mouse (Peromyscus californicus): Locomotor Performance, Metabolic Rate, and Organ Masses.

Although effects of motherhood on mothers have been well documented in mammals, the effects of fatherhood on fathers are not well known. We evaluated effects of being a father on key metabolic and performance measures in the California mouse, Peromyscus californicus. California mice are genetically monogamous in the wild, and fathers show similar parental behavior to mothers, with the exception of lactation. To investigate the impact of fatherhood on fathers, focal males were paired with an intact female (breeding males), a tubally ligated female (nonbreeding males), or another male (virgins). Starting 3-5 d after the birth of each breeding pair's first litter, males were tested for locomotor performance (maximum sprint speed, treadmill endurance), basal metabolic rate (BMR), and maximum oxygen consumption ([Formula: see text]). At the end of the 11-d test period, mice were euthanized, hematocrit was determined, and organs were weighed. Speed, endurance, and [Formula: see text] were significantly repeatable between two replicate measurement days but did not differ among groups, nor did BMR. Breeding males had significantly larger hind limb muscles than did nonbreeding males, whereas virgin males had heavier subcutaneous fat pads than did nonbreeding and breeding males. Several correlations were observed at the level of individual variation (residuals from ANCOVA models), including positive correlations for endurance with [Formula: see text], [Formula: see text] with testes mass, and some of the digestion-related organs with each other. These results indicate that fatherhood may not have pronounced performance, metabolic, or morphological effects on fathers, at least under standard laboratory conditions and across a single breeding cycle.

Size does not matter for male Marbled Salamanders (Ambystomaopacum)

Understanding the phenotypic attributes that contribute to variance in mating and reproductive success is crucial in the study of evolution by sexual selection. In many animals, body size is an important trait because larger individuals enjoy greater fitness due to the ability to secure more mates and produce more offspring. Among males, this outcome is largely mediated by greater success in competition with rival males and (or) advantages in attractiveness to females. Here we tested the hypothesis that large male Marbled Salamanders (Ambystoma opacum (Gravenhorst, 1807)) mate with more females and produce more offspring than small males. In experimental breeding groups, we included males chosen specifically to represent a range of sizes. After gravid females mated and nested freely, we collected egg clutches and genotyped all adults and samples of hatchlings with highly variable microsatellite markers to assign paternity. Size had little effect on male mating and reproductive success. Breeding males were not bigger than nonbreeding males, mates of polyandrous females were not smaller than those of monogamous females, and there was no evidence for positive assortative mating by size. Although body size did not matter for male Marbled Salamanders, we documented considerable fitness variation and discuss alternative traits that could be undergoing sexual selection.

Sexual aggression by intruders in hooded crow Corvus cornix.

The hooded crow Corvus cornix is a west Palaearctic, solitary nesting, monogamous corvid. In the breeding season, populations are characterized by a social organization wherein breeding pairs are territorial and non-breeding individuals, called floaters, live in flocks. During a study of the breeding ecology of the hooded crow, conducted in a protected flooded area, we monitored nests with video cameras. We recorded two separate incidents when intruders attacked a female at the nest. We believe that she remained in the nest in order to prevent the strangers cannibalizing the nestlings by mantling over the brood. The spatio-temporal occurrence of these attacks suggests that the observed behaviour is intraspecific sexual aggression wherein non-breeding males mounted an immobilized female.Electronic supplementary materialThe online version of this article (doi:10.1007/s10211-015-0222-z) contains supplementary material, which is available to authorized users.

Non-invasive administration of 17β-estradiol rapidly increases aggressive behavior in non-breeding, but not breeding, male song sparrows

17β-Estradiol (E2) acts in the brain via genomic and non-genomic mechanisms to influence physiology and behavior. There is seasonal plasticity in the mechanisms by which E2 activates aggression, and non-genomic mechanisms appear to predominate during the non-breeding season. Male song sparrows (Melospiza melodia) display E2-dependent territorial aggression throughout the year. Field studies show that song sparrow aggression during a territorial intrusion is similar in the non-breeding and breeding seasons, but aggression after an intrusion ends differs seasonally. Non-breeding males stop behaving aggressively within minutes whereas breeding males remain aggressive for hours. We hypothesize that this seasonal plasticity in the persistence of aggression relates to seasonal plasticity in E2 signaling. We used a non-invasive route of E2 administration to compare the non-genomic (within 20min) effects of E2 on aggressive behavior in captive non-breeding and breeding season males. E2 rapidly increased barrier contacts (attacks) during an intrusion by 173% in non-breeding season males only. Given that these effects were observed within 20min of E2 administration, they likely occurred via a non-genomic mechanism of action. The present data, taken together with past work, suggest that environmental cues associated with the non-breeding season influence the molecular mechanisms through which E2 influences behavior. In song sparrows, transient expression of aggressive behavior during the non-breeding season is highly adaptive: it minimizes energy expenditure and maximizes the amount of time available for foraging. In all, these data suggest the intriguing possibility that aggression in the non-breeding season may be activated by a non-genomic E2 mechanism due to the fitness benefits associated with rapid and transient expression of aggression.

Effect of RFRP-3 on reproduction is sex- and developmental status-dependent in the striped hamster (Cricetulus barabensis)

RFamide-related peptides (RFRPs) are orthologous to gonadotropin-inhibitory hormone (GnIH) inhibiting gonadotropin release. There are only two RFRP sequences (RFRP-1 and RFRP-3) encoded in rodents. RFRP-3, which was considered as a hypothetical inhibitor on GnRH, shows a stimulatory effect on the male Syrian and male Siberian hamster in short days. As a dominant rodent pest in northern China farmland, the striped hamster (Cricetulus barabensis) has higher reproductive activities and could act as a model to study the mechanism of reproduction. However, the effect of RFRP-3 on the reproductive activity for the striped hamster is less understood. In the study, we cloned 643 bp RFRP cDNA from the striped hamster hypothalamus, which contained an ORF of 570 bp encoding two RFamide-related peptide (RFRP) sequences: SPAPANKVPHSAANLPLRF-NH2 (C. barabensis RFRP-1) and TLSRVPSLPQRF-NH2 (C. barabensis RFRP-3). We also investigated the expression variation of RFRP mRNA and GnRH mRNA in the hypothalamus from hamsters with different developmental statuses (7-week-, 13-week- and 1.5-year-olds) using FQ-PCR, in which the 13-week-old female individuals were in estrous. The striped hamsters that are 7 weeks and 1.5 years old are non-breeding individuals, and those that are 13-week hamsters have breeding phenomena. The highest hypothalamus RFRP mRNA level was found in breeding males as compared to non-breeding males. Conversely, the lowest RFRP mRNA level in the hypothalamus was observed in breeding females, with no significant level when the breeding females were compared to the 7-week-old individuals. Additionally, the investigation of GnRH expression level showed a declining expression trend across the developmental stages (7-week-, 13-week- and 1.5-year-olds) in both sexes. Significant negative and positive relationships were detected in the 13-week estrous female (r=-0.997, P=0.035) and the 13-week male (r=0.998, P=0.029) striped hamsters respectively, which suggest that RFRP-3 has inhibitory and stimulatory effects on female and male adults respectively. Our results suggest that the effects of RFRP-3 on reproduction are sex- and developmental status-dependent in the striped hamster.

The Presence of Sexually Dimorphic Submandibular Glands inTaricha granulosa, the Rough-Skinned Newt (Salamandridae)

We investigate whether Taricha granulosa, the rough-skinned newt, possesses pheromone-producing courtship glands in the submandibular dermis. The hypothesis that these newts have a pheromone-producing mental gland repeatedly appears in the literature based on a small histological study and on courtship behavior in which males press their chins on females' nares. We identify a sexually dimorphic granular gland in the submandibular region of male newts. Its secretion is granular and PAS (periodic acid-Schiff)-positive, in contrast to the PAS-negative secretion of typical granular glands. This gland type is found only in males and only in the anterior submandibular region. Compared to glands evaluated from non-breeding males, the glands from animals collected in the breeding season are hypertrophied. This sexually dimorphic granular gland is distinct from the mental gland found in plethodontid salamanders; the plethodontid mental gland is an aggregate of individual glands, whereas the granular glands in T. granulosa are not clustered together to form a distinct unit. Our results support the hypothesis that males are secreting pheromones from submandibular glands during courtship in T. granulosa.

Hormonal regulation of vasotocin receptor mRNA in a seasonally breeding songbird

Behaviors associated with breeding are seasonally modulated in a variety of species. These changes in behavior are mediated by sex steroids, levels of which likewise vary with season. The effects of androgens on behaviors associated with breeding may in turn be partly mediated by the nonapeptides vasopressin (VP) and oxytocin (OT) in mammals, and vasotocin (VT) in birds. The effects of testosterone (T) on production of these neuropeptides have been well-studied; however, the regulation of VT receptors by T is not well understood. In this study, we investigated steroid-dependent regulation of VT receptor (VTR) mRNA in a seasonally breeding songbird, the white-throated sparrow (Zonotrichia albicollis). We focused on VTR subtypes that have been most strongly implicated in social behavior: V1a and oxytocin-like receptor (OTR). Using in situ hybridization, we show that T-treatment of non-breeding males altered V1a and OTR mRNA expression in several regions associated with seasonal reproductive behaviors. For example, T-treatment increased V1a mRNA expression in the medial preoptic area, bed nucleus of the stria terminalis, and ventromedial hypothalamus. T-treatment also affected both V1a and OTR mRNA expression in nuclei of the song system; some of these effects depended on the presence or absence of a chromosomal rearrangement that affects singing behavior, plasma T, and VT immunolabeling in this species. Overall, our results strengthen evidence that VT helps mediate the behavioral effects of T in songbirds, and suggest that the chromosomal rearrangement in this species may affect the sensitivity of the VT system to seasonal changes in T.

Arginine Vasotocin (AVT) Triggers Courtship Behavior Without Exposure to External Stimuli and Modulates the Olfactory Response of MaleHynobius leechiiSalamanders

To study the function of arginine vasotocin (AVT) in a specific courtship behavior (body undulation) in male Hynobius leechii, we injected various doses of AVT or an AVT V1a or V2 receptor antagonist into breeding and non-breeding males. After these injections, we placed the males alone or with breeding females in Petri dishes and measured the incidence and frequency of body undulation. Additionally, to test whether AVT modulates the olfactory response of males, we exposed breeding males that were injected with AVT to female odors and measured the same response. Both breeding and non-breeding males intraperitoneally injected with 50 or 100 µg of AVT exhibited body undulation. Additionally, breeding males intraperitoneally injected with 50 or 100 µg of AVT exhibited an increased frequency of body undulation when exposed to female odors. The intraperitoneal injection of 5, 25, or 50 µg of the AVT V1a or V2 receptor antagonist did not significantly decrease the incidence or frequency of body undulation of 100 µg AVT-injected breeding males. However, in a post-hoc experiment, the breeding males that were subcutaneously injected with 100 µg of the AVT V1a receptor antagonist exhibited a significant decrease in the frequency of body undulation, induced by exposure to females. Additionally, a central injection of 1 µg of AVT into the brain induced body undulation in breeding males. Our results show that AVT not only induces a specific courtship behavior in male H. leechii via AVT V1a receptors but also modulates the olfactory response.

A case of natural queen succession in a captive colony of naked mole-rats, Heterocephalus glaber

Naked mole-rats occur in large colonies where usually a single queen monopolizes reproduction. Queen succession occurs from within usually as a result of aggressive encounters with subordinate females that queue for reproductive succession following colony instability, which inevitably results in death of either the queen or the challenging conspecific. We monitored a queen succession following the death of the breeding male in a colony of the naked mole-rat, Heterocephalus glaber, prior to, during as well as after replacement of the original breeding female. The response of the pituitary luteotrophs was investigated in the non-reproductive females during this period of instability by the administration of endogenous gonadotrophin releasing hormone (GnRH) and evaluating the subsequent luteinising hormone (LH) response in the blood. Larger and older non-breeding females engaged in aggressive encounters that culminated in death. The new breeding successor which arose from within the colony was a large female who continued to procreate. The six non-breeding females that were killed during reproductive takeover were larger and older females which exhibited elevated basal circulating LH concentrations as well as increased pituitary sensitivity as measured by the amount of releasable LH to an exogenous GnRH challenge. By contrast, non-breeding females that survived the succession were smaller and younger animals with reduced basal and GnRH challenged LH concentrations. Likewise, five non-breeding males which were heavier and older than those non-breeding males which survived were killed. These animals did not, however, show elevated basal or exogenous GnRH challenged LH concentrations when compared to the surviving males. The non-breeding animals of both sexes which survived the reproductive takeover event represented individuals which posed a minimal threat to the new successor and hence promoted the continuation of the marked reproductive skew that is prevalent in this highly inbred colonial subterranean hystricomorph.

Effect of reproductive status on hypothalamic–pituitary–adrenal (HPA) activity and reactivity in male California mice (Peromyscus californicus)

Previous studies indicate that reproductive condition can alter stress response and glucocorticoid release. Although the functional significance of hypothalamic–pituitary–adrenal (HPA) axis modulation by breeding condition is not fully understood, one possible explanation is the behavior hypothesis, which states that an animal's need to express parental behavior may be driving modulation of the HPA axis. This possibility is consistent with findings of blunted activity and reactivity of the HPA axis in lactating female mammals; however, effects of reproductive status on HPA function have not been well characterized in male mammals that express parental behavior. Therefore, we tested this hypothesis in the monogamous and biparental California mouse. Several aspects of HPA activity were compared in males from three reproductive conditions: virgin males (housed with another male), non-breeding males (housed with a tubally ligated female), and first-time fathers (housed with a female and their first litter of pups). In light of the behavior hypothesis we predicted that new fathers would differ from virgin and non-breeding males in several aspects of HPA function and corticosterone (CORT) output: decreased amplitude of the diurnal rhythm in CORT, a blunted CORT increase following predator-odor stress, increased sensitivity to glucocorticoid negative feedback, and/or a blunted CORT response to pharmacological stimulation. In addition, we predicted that first-time fathers would be more resistant to CORT-induced suppression of testosterone secretion, as testosterone is important for paternal behavior in this species. We found that virgin males, non-breeding males and first-time fathers did not display any CORT differences in diurnal rhythm, response to a predator-odor stressor, or response to pharmacological suppression or stimulation. Additionally, there were no differences in circulating testosterone concentrations. Adrenal mass was, however, significantly lower in new fathers than in virgin or non-breeding males. These results suggest that the behavior hypothesis does not explain HPA function across reproductive conditions in male California mice.

EFFECTS OF SOCIAL AND EXTRA-PAIR MATING ON SEXUAL SELECTION IN BLUE TITS (CYANISTES CAERULEUS)

The contribution of extra-pair paternity (EPP) to sexual selection has received considerable attention, particularly in socially monogamous species. However, the importance of EPP remains difficult to assess quantitatively, especially when many extra-pair young have unknown sires. Here, we combine measurements of the opportunity for selection (I), the opportunity for sexual selection (IS ), and the strength of selection on mating success (Bateman gradient, βSS ) with a novel simulation of random mating tailored to the specific mating system of the blue tit (Cyanistes caeruleus). In a population where social polygyny and EPP are common, the opportunity for sexual selection was significantly stronger and Bateman gradients significantly steeper for resident males than for females. In general, success with the social mate(s) contributed most to variation in male reproductive success. Effects of EPP were small, but significantly higher than expected under random mating. We used sibship analysis to estimate the number of unknown sires in our population. Under the assumption that the unknown sires are nonbreeding males, EPP reduced the variance in and the strength of selection on mating success, a possibility that hitherto has not been considered.

Linking predator–prey interactions with exposure to a trophically transmitted parasite usingPCR‐based analyses

Parasite transmission is determined by the rate of contact between a susceptible host and an infective stage and susceptibility to infection given an exposure event. Attempts to measure levels of variation in exposure in natural populations can be especially challenging. The level of exposure to a major class of parasites, trophically transmitted parasites, can be estimated by investigating the host's feeding behaviour. Since the parasites rely on the ingestion of infective intermediate hosts for transmission, the potential for exposure to infection is inherently linked to the definitive host's feeding ecology. Here, we combined epidemiological data and molecular analyses (polymerase chain reaction) of the diet of the definitive host, the white-footed mouse (Peromyscus leucopus), to investigate temporal and individual heterogeneities in exposure to infection. Our results show that the consumption of cricket intermediate hosts accounted for much of the variation in infection; mice that had consumed crickets were four times more likely to become infected than animals that tested negative for cricket DNA. In particular, pregnant female hosts were three times more likely to consume crickets, which corresponded to a threefold increase in infection compared with nonpregnant females. Interestingly, males in breeding condition had a higher rate of infection even though breeding males were just as likely to test positive for cricket consumption as nonbreeding males. These results suggest that while heterogeneity in host diet served as a strong predictor of exposure risk, differential susceptibility to infection may also play a key role, particularly among male hosts. By combining PCR analyses with epidemiological data, we revealed temporal variation in exposure through prey consumption and identified potentially important individual heterogeneities in parasite transmission.

Seasonal and sex differences in the hippocampus of a wild rodent

Studies across and within species suggest that hippocampus size is sexually dimorphic in polygamous species, but not in monogamous species. Although hippocampal volume varies with sex, season and mating system, few studies have simultaneously tested for sex and seasonal differences. Here, we test for sex and seasonal differences in the hippocampal volume of wild Richardson's ground squirrels (Urocitellus richardsonii), a polygamous species that lives in matrilineal, kin-based social groups and has profound sex differences in behavior. Based on the behavior and ecology of this species, we predicted that males would have a significantly larger hippocampus than females and that the hippocampus would be largest in males during the breeding season. Analyses of both absolute and relative volumes of the hippocampus yielded a significant difference between the sexes and seasons as well as an interaction between the two such that non-breeding males have significantly larger hippocampal volumes than breeding males or females from either season. Dentate gyrus, CA1 and CA3 subfield volumes were generally larger in the non-breeding season and in males, but no significant interaction effects were detected. This sex and seasonal variation in hippocampal volume is likely the result of their social organization and male-only food caching behavior during the non-breeding season. The demonstration of a sex and seasonal variation in hippocampal volume suggests that Richardson's ground squirrel may be a useful model for understanding hippocampal plasticity within a natural context.