Mink with Divergent Activity Levels have Divergent Reproductive Strategies

Stable individual differences in activity levels within populations have been linked to differences in reproductive rate or parental care in several species, including American mink (Neovison vison). Fur‐farmed mink are good models for studying such effects because they yield large sample sizes and readily allow investigations into maternal behaviour, reproductive success, offspring performance and the relationships between these factors. On farms, very inactive individuals generally have smaller litters, and this held true in our study populations. We tested two competing hypotheses to explain this: (1) inactive individuals are failing to cope with a challenging environment and experiencing chronic stress and/or depression‐like ‘apathy’; this predicts female‐skewed litters, poorer maternal care, higher infant mortality and poorer infant growth and (2) inactive individuals do not have reduced fitness but instead employ an alternative adaptive reproductive strategy, trading off offspring quantity for quality; this predicts enhanced maternal care, reduced infant mortality and enhanced infant growth. Inactive females’ kits, especially their sons, grew faster than active females’, even after statistically controlling for litter size; and by 21 d, inactive and active dams’ litters no longer differed in total biomass, despite the former’s smaller litter sizes. In kit retrieval tests, inactive females were faster than active dams to reach their sons (as well as more likely to contact their sons than their daughters: a bias towards male kits not evident in the active dams). Furthermore, kit growth rates and dam latencies to touch them co‐varied, suggesting the existence of consistent differences in maternal style across inactive and active dams. Hypothesis 2 was thus supported: inactive females favour offspring quality over quantity, investing more resources in fewer kits, particularly males. This potentially boosts their sons’ adult fitness. More broadly for laboratory‐based studies, possible ‘captivity effects’ on the fitness correlates of activity and other personality traits are discussed.     

Courtship in two species of periodical cicadas, Magicicada septendecim and Magicicada cassini

Courtship behaviour of two species of periodical cicadas, Magicicada septendecim and M. cassini, was studied in the field during the 1970, 1973, and 1974 emergences of these insects. In areas where both species were courting there were differences in both male and female courtship patterns, both in acoustic and behavioural components. Experiments with models showed that male M. septendecim were more likely to court crude models of females than were M. cassini males. When females were ‘courted’ with models that could imitate some of male courtship, they were more receptive when the models' ‘songs’ were those of conspecific males. Acoustic differences between species are probably used by females in mate selection, maintaining species separation even in areas where the two species overlap in both space and time.     

Influence of Litter Gender Composition on Subsequent Maternal Behaviour and Maternal Aggression in Female House Mice

It has been suggested that the sex-dependent pattern of mother-pup interaction that occurs early in infancy can affect some aspects of the animal's behaviour repertoire in adulthood. The aim of this study was to investigate the effects of litter gender composition (LGC) on subsequent maternal behaviour and aggression in the female house mouse. Albino mouse litters were reduced at birth to six pups according to two conditions: FM (3 females and 3 males) and FF (all females). At weaning (day 21), all mice were rehoused in unisexual groups. At adulthood (70–90 days) the females were mated and litters culled at birth to 6 pups (3 females and 3 males). On days 2, 4, and 6 after delivery the females were tested for maternal behaviour (10 min) and retrieval of pups. On day 6, at the end of the recording of maternal behaviour, the females were also tested for maternal aggression towards a strange adult male conspecific (5-min exposure). In absence of differences in maternal behaviour scores, FM females showed shorter latencies for retrieval of the first pup on postnatal days 2 and 4. Maternal aggression was also significantly affected by the two conditions. FM females showed higher scores of both duration and frequency of aggressive grooming, defensive upright postures, fleeing and of the number of attacks. FF females spent more time self-grooming, crouching, and on nest. These results support the hypothesis that a series of factors, such as olfactory, tactile, and acoustic stimulation provided by the pups, the variation of maternal behaviour depending upon LGC, and the qualitative and/or quantitative variation in social interaction with the opposite sex in infancy, contribute to the broad interindividual plasticity to cope in different environmental situations.     

Learning enhances female control over reproductive investment in the Japanese quail

The adaptive significance of learning is supported by studies showing its positive effects on mating behaviour, but they rarely go beyond fertilization success. Here we studied how learning contributes to qualitative reproductive investment, by testing the hypothesis that mating in the context that predicts male appearance has positive effects on female reproductive investment compared with unsignalled mating. Using Japanese quail (Coturnix japonica), we found that effects of mating in the context predicting mating opportunity depend on female body condition and receptivity, while the outcome of unexpected mating depends on male behaviour. In particular, among females mated with the familiar male in the context predicting that he will appear, female condition positively affected the number of fertilized eggs and egg mass and more receptive females tended to produce more sons. Additionally, conditioned females laid heavier eggs for daughters than for sons. In contrast, in females that were mated unexpectedly and with a novel male, the number of fertilized eggs was highly dependent on male behaviour and was negatively related to maternal body condition. Egg mass was not related to body condition, and there were no indications of sex allocation. This is, to our knowledge, the first study demonstrating how female body condition and behaviour interact with the context of mating in shaping maternal reproductive investment.     

Social Behaviour on the Wing in the False Vampire,Megaderma lyra

Social interaction occurs in bats. In a group of 10 Megaderma lyra (seven female and two male adults, as well as one female juvenile) held in captivity, two stereotyped flying behaviour patterns —the ‘grumbling flight’ and the ‘song flight’ — were observed and studied. The ‘grumbling flight’ is a social interaction in flight between at least two Megaderma lyra in which ‘grumble sequences’ are emitted. This behaviour is triggered by stress or arising aggression, and presumably attempts to avoid agonistic interactions with dangerous physical contact. The song flight was exclusively displayed by the dominant male bat and only directed at the non-lactating female members of the group, with a preference to alien females. This behaviour is composed of three behavioural stages, each accompanied by a specific ‘song strophe’. The song flight presumably aims at bonding the females to the male. During the grumbling flight and the song flight, M. lyra emits communication sounds in the ultrasonic range. The sounds consist of simple elements (FM_down, FM_up, CF), and are similar to types of sounds emitted for echolocation by various bat species.     

Strategic Investment in Sperm Removal Behaviour in a Bushcricket

Multiple mating by females is widespread and generates sperm competition among the ejaculates of rival males over fertilization. One way in which males can avoid or reduce sperm competition is by displacing or removing previous males’ sperm from female sperm stores. An apparent example of this occurs in the bushcricket Metaplastes ornatus. Males perform a specialised sperm removal behaviour (SRB), using their highly-derived subgenital plate, with which they remove sperm from the female’s spermatheca during the early phases of mating before transferring a spermatophore of their own. Here we investigated whether males strategically invest in SRB according to the amount of previously stored sperm present in females. Each male was tested twice, once with a female containing sperm (‘filled’ condition) and once with a female from whom most previously deposited sperm had recently been removed by another male (‘emptied’ condition). For comparison, a separate group of males was paired with virgin females. Males did not discriminate between non-virgin females in the ‘emptied’ or ‘filled’ conditions in terms of their investment in SRB, suggesting they may not able to perceive the amount of sperm present in the female’s spermatheca. By contrast, male investment in SRB was significantly reduced in pairings with virgin females, indicating that males are sensitive to some aspect of a female’s mating status. Our results thus suggest that males modulate SRB in response to female-mediated cues, possibly chemical cues left by previous males, which would not be present on virgin but would be on non-virgin females.     

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.     

Behaviour and fecundity of female rats mated with preferred or non-preferred males

Sexually experienced female rats were allowed to choose between two inaccessible males. Approximately half of the females developed a consistent preference, and the most ‘discriminating’ females were mated with either their preferred or their non-preferred male. Females without a decided preference were mated with one of the two males randomly selected by the experimenter. The pairs of discriminating females and preferred males had higher levels of non-sexual contacts, solicitations and sexual activity than the pairs of discriminating females and non-preferred males. Specifically, the former pairings were characterized by higher levels of non-sexual contact and affiliative behaviour (crawl over and under responses) and solicitation responses (ear-wiggling and darting) by the female and shorter latencies for intromissions and ejaculations than the pairings with a non-preferred male. The pairs that included non-discriminating females behaved more like the pairs with a discriminating female and a preferred male than those with a discriminating female and a non-preferred male. Even so, the number of females delivering litters and the size of the litters did not differ between the different experimental pairings. These data suggest that discriminating famale rat is not conferred and obvious reproductive advantage over a less discriminating female.     

Evidence for Sex-Biased Behavioral Maternal Investment in the Gray Mouse Lemur (Microcebus murinus)

To study sex-differential allocation of maternal behavior in Microcebus murinus, I recorded behavioral patterns on 21 litters from parturition to the weaning period. After a pregnancy of 61.5 ± 0.9 days, females may produce from one to four young per litter. Litters were weaned in 40 days by mothers. Behavioral observations at the beginning of the nocturnal activity period demonstrated that close contacts between mothers and infants were more frequent in multiparous mothers than in primiparous ones (p < .01). Time in close contact with offspring aged >15 days old compared to contacts with neonates is significantly lower only in single-sex litters and is more marked for all-female litters (p < .01). Mother's approaches toward mixed litters or all-male litters were always significantly greater than approaches toward all-female litters (p = .04). However, mother's approaches within mixed sex litters were not biased toward either sex (p = .7). Males in a litter may be interpreted as a stimulator of maternal behavior. Similarly, using retrieving tests of 15-min duration, a significant maternal preference for male neonates is evident. Latency to first retrieval is significantly shorter in multiparous females than in primiparous ones (p < .05) independent of size and sex ratio of the litter. For multiparous females only, male neonates were chosen first for retrieval more often than females (p < .05). Finally, the calls of young played a stimulator effect on maternal retrieving (p < .001). Accordingly multiparous mothers exhibit more interest in their young, which appears to be biased toward male neonates.     

Sexual conflicts in spotted hyenas: male and female mating tactics and their reproductive outcome with respect to age,social status and tenure

We investigated the reproductive outcomes of male and female mating tactics in the spotted hyena, Crocuta crocuta, a female-dominated social carnivore with high maternal investment, an absence of paternal care and female control over copulation. Paternity was determined using microsatellite profiling of 236 offspring in 171 litters from three clans. We found little evidence that male tactics that sought to coerce or monopolize females were successful. Polyandry and sperm competition appeared to counter effectively pre-copulatory male tactics, such as harassment, monopolization and other tactics, such as infanticide, that were against the evolutionary interests of females, and may have contributed to the stability of the male dominance hierarchy, which operated as a social queue. At least 39% of 54 females mated multiply, and 35% of 75 twin litters were fathered by two sires. Polyandry may also serve to ensure fertilization, compensate for an initial poor-quality mate or ensure fertilization by genetically compatible mates. Female mate choice matched observed patterns of affiliative male-female behaviour, indicating that affiliative behaviour is a successful male mating tactic, and was consistent with the idea that male tenure may serve as an index of male quality, although male fertility may decline with extreme old age.     

Maternal Aggression and Litter Size in the Female House Mouse

It has been commonly argued that, in house mice, female post-partum fighting against a male intruder functions to protect the offspring from infanticide. The aim of this study was to test the hypothesis that maternal aggression is actually related to pup defence and, specifically, according to parental investment theory, that its intensity should increase with litter size. 60 nulliparous albino female mice were mated and randomly assigned to four experimental groups in which litters were culled at birth to 0, 4, 8, or 12 pups, respectively. On day 8 after delivery all females were tested for maternal aggression against a stranger adult male conspecific (5-min exposure). No aggression occurred in the group in which all pups had been removed. In the other groups, the proportion of females displaying overt aggression increased with litter size. Several scores of female agonistic behaviour (proportion of females displaying overt aggression, total attacking time, frequency of tail rattling) were significantly higher for the females rearing 8 and 12 pups than for the females rearing 4 pups. Aggressive behaviour of females rearing 12 pups was not significantly higher than that of females rearing 8 pups. No male committed infanticide. These results support the hypothesis that rodent maternal aggression is strictly related to offspring defence and are consistent with the theoretical prediction that, the costs of the defence being equal and the gain in fitness increasing with litter size, the intensity of maternal defence of the young should increase with their number.     

Coping with social tension: Sex differences in the effect of food provision to small rhesus monkey groups

In previous research, male chimpanzees (Pan troglodytes) were found to be more ‘conciliatory’ than females, in that after aggressive incidents males more often engaged in socially positive contact with former opponents. The first part of the present paper presents similar results for a large breeding troop of rhesus monkeys (Macaca mulatta). To investigate further the possible sex difference in postconflict behaviour, a series of experiments was carried out on small isosexual groups of young rhesus monkeys, three male and three female groups. Food competition was created by providing either a single apple piece (Monopoly test) or a handful of small pieces (Equality test). Observations lasted for 30 min, and behaviour in tests without food provisioning was also recorded. The predicted response to food was an increase in aggression followed by restorative behaviour, such as grooming. In Equality tests both sexes showed similar responses, i.e. increased aggression and decreased grooming and cohesiveness. In Monopoly tests, however, their responses differed. After the aggression increase, which occurred in all groups, males showed a significant increase in grooming and cohesiveness, whereas females showed the (non-significant) opposite trend. Another sex difference was that changes in grooming and aggression frequencies were related to the rank order in female groups, but not in male groups. Since there was no evidence for a direct causal connection between grooming and aggression, a new model is introduced which links grooming behaviour to the type of food provision rather than to the social disturbance by aggression. According to this model inequality in food distribution causes social tensions. Males actively try to reduce these tensions, whereas females do not. Some alternative explanations are also discussed.     

The influence of female presence and seasonality on the home range size and activity patterns of male domestic cats in Brazil’s Atlantic Forest

The factors that affect the home range size of domestic cats (Felis silvestris catus) in a semi-domesticated condition (i.e., cats that receive shelter and food from humans but are free to move in the wider natural environment) is not completely understood. Here, using radio telemetry, we present the first assessment of the home range size and activity patterns of male domestic cats living in the insular environment of Brazil’s Atlantic Forest, and examine the extent to which these were influenced by the presence (‘resident males with females’: RMF group) or absence (‘single males’: SM group) of females in the residences of their owners. Daily and seasonal activity patterns have also been characterized. Home ranges of the RMF were markedly smaller than that of the SM and exhibited less overlap with the home ranges of other males than did those of the SM. Male cats were most active at twilight and evening (44.68 ± 1.07) and least active in the afternoon (28.76 ± 2.79). RMF presented nocturnal activity greater than SM, especially in the dry season. Our data indicate that presence of females is an important factor in shaping these parameters of domestic male cats.     

The effects of litter-size variation on the development of play behaviour in the domestic cat: litters of one and two

In many litter-bearing species, litter-mates interact with each other in a variety of ways during early life. In the domestic cat, Felis catus, social play is a prominent form of interaction between litter-mates. The present study investigated how the lack of a litter-mate affected the development of kitten play and of the mother-kitten relationship. The subjects were 14 litters of domestic kittens living with their mothers in large indoor cages. Seven litters contained two male kittens, and the other seven contained single male kittens. Social interactions within the families were observed from day 22 to day 83 after birth. Single kittens played a more active part in maintaining close proximity to their mothers and directed more playful behaviour at them than did kittens with a sibling. Although single-kitten mothers avoided their offspring more than did mothers of litters of two, they also directed much higher levels of play behaviour at them. Despite the marked difference in the mother-kitten play relationship in the two litter types, single kittens experienced quantitatively less social play than did kittens with siblings. As the kittens grew older, single-kitten mothers showed higher levels of aggression towards their young than did mothers of litters of two.     

Diet Quality Affects Mate Choice in Domestic Female Canary Serinus canaria

Mate choice is the result of intra‐ and intersexual selection. Even though a consensus between females is often observed in favour of some males, different constraints can affect their choice. Context (early learning, mate‐choice copying, prior experience, etc) and/or condition (genetics, body condition, age, etc) can deflect the choice of a female from its ‘standard of beauty’. In domestic canaries Serinus canaria, females display their acceptance of a male by producing a particular sexual behaviour called ‘copulation solicitation display’ (CSD). Previous studies have shown that the number and intensity of these CSDs are more important when females listen to sexy ‘A’ phrases with a rapid tempo (A16) in comparison with A phrases with a slow tempo (A8). In this study, we assess the relationship between food quality (highly or poorly diversified food during a short period) and female choosiness towards these two types of phrases. We confirm that females discriminate between A16 and A8. Nevertheless, the difference between the number of sexual responses towards A16 and A8 is weaker among females with a poorly diversified food diet in comparison with females with a highly diversified food diet. This suggests that even a short‐term modification of condition could decrease females’ selectivity towards high‐value stimuli and increase their response rates towards low‐value signals.     

Paternal and maternal behaviour in the Mongolian gerbil

Twenty-seven pairs of Mongolian gerbils (Meriones unguiculatus) were observed rearing fifty-five litters in undisturbed conditions. Parental activities were performed by both parents. Males spent more time in the nest but females showed more sniffing and licking of pups and nest building. There was no difference between the parents in the amount of body contact with the pups. The frequency of occurrence of each of these activities changed with the age of the pups. These and other measures of behaviour are discussed in relation to the changing stimulation from the young and the willingness of the female to allow the male to interact with the litter.     

Male aggression,limited female choice and the ontogeny of mating behaviour in the flesh fly Sarcophaga crassipalpis

Previous work has shown that male flesh flies (Sarcophaga crassipalpis Macquart) exhibit an ontogeny of behaviour from eclosion through sexual maturity that includes extensive changes in the expression of aggressive, non‐aggressive interactive and non‐interactive behaviours. To determine how the presence of a female flesh fly influences the manifestation of these behaviours, male flesh flies of different ages post‐eclosion are paired with same‐age females and their behaviours are monitored in a simple arena during a 50‐min observation period. All flies are socially isolated until pairing. Although the levels of expression of aggressive and non‐aggressive interactive behaviours are depressed relative to previous findings in male‐opponent pairs, the ontogeny of aggression still occurs as indicated by a significant increase, with age, in the agonistic behaviour ‘hold’. Similar to male‐opponent pairs and individual males, the performance by males of the non‐interactive behaviours ‘walking’ and ‘standing’ diminishes, whereas ‘upside‐down’ increases with age. By contrast, ‘grooming’ shows a significant age‐related decline. No courtship behaviours are observed in the males, although the aggressive behaviour ‘hold’ is a significant transition to mating. Females show no obvious courtship or rejection behaviours, although the significant increase in ‘upside‐down’ with age could possibly be a behavioural gateway to mating. The results of this study indicate that extensive age‐related changes encompassing the entire behavioural repertoire are intrinsic to male flesh flies and persist under a variety of different social contexts.     

Alternative Male Spawning Tactics and Acoustic Signals in the Plainfin Midshipman Fish Porichthys notatus Girard (Teleostei,Batrachoididae)

The plainfin midshipman Porichthys notatus has two male reproductive morphs, ‘Type I’ and ‘Type II’, which are distinguishable by their physical traits alone. Type I males are eight times larger in body mass than Type II males and have a six-fold larger relative sonic (vocal) muscle mass than Type II males. In contrast, the testicles of Type II males are seven times larger than those of Type I males. This study demonstrates morph-specific patterns of reproduction, including acoustic signals, for Type I and II males. Field censuses of nests showed that only Type 1 males maintained nests. Type II males and females transiently appeared in these nests in association with each other. Infra-red video and hydrophone recordings in aquaria showed that Type I males maintained nests and readily vocalized. Long-duration ‘hums’ and sequences of short-duration ‘grunts’ were produced during advertisement and agonistic contexts, respectively. Humming Type I males attracted females to their nests, pair-spawned, and then guarded egg clutches alone. By contrast, Type II males neither acoustically courted females nor maintained available nest sites, but rather ‘sneak-’ or ‘satellite-spawned’ at the nests of Type I males. Type II males infrequently produced low amplitude, short duration grunts that were similar in spectral, temporal and amplitude characteristics to the grunts of females. Type II males appear to be obligate sexual parasites of the nest-building, mate-calling, and egg-guarding Type I males. The dimorphic body and vocal muscle traits of the two male morphs in the plainfin midshipman are thus paralleled by a divergence in their reproductive tactics and the properties of their acoustic signals.     

Postpartum copulatory and maternal behaviour in Norway rats under seminatural conditions

Virgin female rats were mated at a freely cycling oestrus, and then again with the same males at the subsequent postpartum oestrus. Copulatory behaviour at each mating was compared and the temporal patterning of the female's mating and maternal behaviour was analysed. Postpartum females were found to conserve the number of ejaculatory series received, yet the series were shorter, resulting in a shorter time being occupied, by this mating. During mating, females spent little time with their litters, and then mainly in the post-ejaculatory intervals. Ejaculatory series were rarely disrupted by nesting. The postpartum female rat seems to engage in behavioural time-sharing.     

Mining and exploitation of natural mineral deposits by the desert tortoise, Gopherus agassizii

Six female desert tortoises, Gopherus agassizii, were observed to consume the soil of a ‘limelayer’ on an exposed site, or to excavate through the overburden where the layer was not exposed. The calcium content of soil from the layer mined was significantly higher than that of adjacent surface soil. All geophagous episodes involved reproductively mature females during the nesting season, a period in which the females were in potential calcium stress. To our knowledge this is the first report of mining and long-term exploitation of natural mineral deposits by a lower vertebrate and may represent behaviour at least as complicated as that seen in many ungulates.