Paternal kin bias in the agonistic interventions of adult female rhesus macaques (Macaca mulatta)

When agonistic interventions are nepotistic, individuals are expected to side more often with kin but less often against kin in comparison with non-kin. As yet, however, few mammal studies have been in a position to test the validity of this assertion with respect to paternal relatedness. We therefore used molecular genetic kinship testing to assess whether adult female rhesus macaques (Macaca mulatta) from the free-ranging colony of Cayo Santiago (Puerto Rico) bias their interventions in ongoing dyadic aggressive interactions towards maternal and paternal half-sisters compared with unrelated females. It turned out that females supported maternal half-sisters significantly more often than paternal half-sisters or non-kin regardless of the costs associated with such interventions. Similarly, females targeted maternal half-sisters significantly less often than non-kin when this was associated with high costs. Unrelated females provided significantly higher mean rates of both high- and low-cost support to each other than did paternal half-sisters. However, females targeted paternal half-sisters significantly less often than non-kin when targeting was at low cost, suggesting that females refrain from intervening against paternal half-sisters. Our data confirm the general view that coalition formation in female mammals is a function of both the level of maternal relatedness and of the costs of intervention. The patterns of coalition formation among paternal kin were found to be more complex, and may also differ across species, but clear evidence for paternal kin discrimination was observed in female rhesus as predicted by kin selection theory.     

Vocal alliances in Chacma baboons (<Emphasis Type="Italic">Papio hamadryas ursinus</Emphasis>)

Theory predicts that females in species with matrilineal dominance hierarchies should use nepotistic support systems to maintain their family’s rank. Female Old World monkeys, however, form alliances against other females at surprisingly low rates. Nonetheless, in many species, females utter threat vocalizations when observing others’ disputes, suggesting that these vocalizations may function as ‘vocal alliances’. We describe a playback experiment testing the efficacy of vocal alliances in free-ranging female baboons. Subjects were played the same female’s threat-grunts under three separate conditions: after being threatened by the signaller’s close relative to mimic kin support, after being threatened by a female maternally unrelated to the signaller to mimic non-kin support, and after a friendly interaction with the signaller’s close relative as a control. Subjects responded more strongly to the playback and avoided the signaller and her matrilineal relatives for a longer period of time in kin support trials than in either non-kin support or no aggression trials. In contrast, there was no difference in subjects’ behaviour between non-kin support and no aggression trials. These results corroborate observational data showing that vocal support occurs at a higher rate than physical support in female baboons, and that kin are more likely to provide vocal support than non-kin. We conclude that vocal support plays a similar role as physical support in the alliances of female baboons.     

O brother,where art thou? The varying influence of older siblings in rank acquisition by female baboons

Males in sexually dimorphic species like baboons appear to have surprisingly little influence on the reproduction and dominance ranks of their female kin, even though they could potentially increase their fitness by helping their relatives improve their ranks. Male baboons are able to dominate females several years before they emigrate, but their presence has no effect on relatives’ dominance ranks, at least when female kin are present. As a result, females usually acquire ranks within their matriline, above their older sisters. We describe the process of rank acquisition among orphaned and non-orphaned juvenile and adolescent females in a group of free-ranging baboons. Orphaned females were significantly more likely than non-orphaned females to acquire unexpected ranks. Orphaned females with older sisters often acquired ranks within the matriline, but below their older sisters’. Orphaned females with older brothers were likely to rise in rank above their matriline. Females’ interventions on behalf of younger sisters always supported the existing female dominance hierarchy, while males’ interventions tended to act against it. Similarly, in playback experiments, females appeared to be willing to support their younger sister only in disputes with lower-ranking females. In contrast, males appeared to be willing to support their sister even in disputes with higher-ranking females. Fraternal support enables females to improve their dominance ranks, but only if their mothers have died. It remains a puzzle why males have so little influence on their female relatives’ ranks when female kin are present, and so much when they are absent.     

Benefits of kin association: related and familiar zebrafish larvae (<Emphasis Type="Italic">Danio rerio</Emphasis>) show improved growth

Zebrafish (Danio rerio) larvae prefer the olfactory cues of kin to non-kin. We examined the potential benefits of kin preference by comparing growth rate, shoaling, and aggressive behavior in juvenile zebrafish housed in groups of either familiar kin or unfamiliar non-kin. Over an observation period of 5 days, the animals grew 33% more in kin groups; however, neither shoaling nor the frequency of aggressive interactions was different in groups of related versus unrelated individuals. Shoaling behavior increased with increasing observation time and increasing age, while aggressive behavior remained the same. We conclude that associating with kin probably creates a less stressful environment that allows for higher growth rates, which can lead to higher direct fitness based on increased survival and earlier reproduction. Kin recognition leading to kin-structured groups may therefore be under positive selection.     

Genetic relatedness and eusociality: parasite-mediated selection on the genetic composition of groups

Summary Contrary to the expectations of kin selection theory, intracolony relatedness in eusocial insects is often low. We examined the idea that associations of low relatedness (high genetic variability) may be advantageous because of negative frequency-dependent selection on common host phenotypes by rapidly evolving parasites and pathogens. Using the natural host-parasite system of the bumble bee Bombus terrestris and its intestinal trypanosome Crithidia bombi, we investigated the transmission properties of parasites in host groups. Within naturally infested nests and in artificially constructed groups of workers, prevalence of infestation increased with time of exposure (Table 1). The susceptibility of isolated groups of workers to the parasites to which they were exposed differed with identity and natural infestation of their nest of origin (Table 2). In addition, those workers that were related to the individual introducing an infection to their group were more likely to become infested than were unrelated workers (Table 3). Although the bumble bee workers in experimental boxes appeared to differ in behavior toward kin and non-kin, making more physical contacts with kin, we found no discernible relationship between number of physical contacts and prevalence of infestation in a group. Therefore, we conclude that differences in parasite transmission reflected interactions among different host and parasite phenotypes. This system thus demonstrates the factors necessary for negative frequency-dependent selection by parasites on common host phenotypes - variability for susceptibility and infectiousness in host and parasite populations and similarity for these traits among related individuals. If, as we show here, high genetic relatedness within groups enhances parasite transmission, kin directed altruism may increase the risk of contracting parasites and infectious diseases. Therefore, parasites and pathogens may be an important force moderating the genetic structure of social groups. Offprint requests to: J.A. Shykoff at the present address     

The kin recognition system of carpenter ants (Camponotus spp.)

Summary Hamilton's (1964) hypothesis linking haplodiploidy and eusociality in the Hymenoptera could be reconciled with the occurrence of polygyny and multiple insemination if workers are able to distinguish full (3/4 related) sisters from other familiar matri- and patrilines within the colony, and direct altruistic behavior toward them preferentially. We examined this possibility in small genetically mixed nests of the carpenter ant Camponotus floridanus, formed by the transfer of worker pupae from two unrelated source colonies. In 120 h of observation on 12 queenright and 12 queenless nests, more than 15,000 behavioral interactions were recorded. Workers antennated familiar nonkin significantly more frequently than familiar sisters. However, they failed to discriminate consistently between kin and non-kin in food exchanges and grooming. Aggressive behavior was occasionally observed in some queenless nests, but almost never in the presence of a queen. When aggression did occur, it was directed significantly more often toward non-kin. Though related adult workers did not cooperate preferentially, the biases in antennation and aggression do indicate an ability to discriminate familiar kin from familiar nonkin, which may be employed in other contexts such as the rearing of reproductive brood.     

Shoaling decisions in three-spined sticklebacks (<Emphasis Type="Italic">Gasterosteus aculeatus</Emphasis>)—familiarity,kinship and inbreeding

Shoaling with familiar kin is a well-known phenomenon. It has been described both for adult and for fry of three-spined sticklebacks (Gasterosteus aculeatus). However, evidence of preference for kin independent of familiarity is scarce. Furthermore, inbreeding effects have not been studied for shoaling preferences and there is a lack of studies about changes in individuals’ ability to recognise kin during different phases of life history. We gave inbred and outbred, nonreproductive sticklebacks of different age the choice to shoal with a group of familiar siblings vs unfamiliar non-siblings and with unfamiliar siblings vs unfamiliar non-siblings. Subadult sticklebacks preferred to shoal with familiar kin over unfamiliar non-kin, but inbred and outbred individuals had similar preferences. When given the choice between unfamiliar siblings and unfamiliar non-sibs, adult outbred fish preferred the shoal of kin, while inbred fish behaved indifferently. Body characteristics of the group such as body mass, standard length and condition of its members did not significantly explain the shoaling preferences for kin. Thus, sticklebacks were capable of recognising familiar and unfamiliar kin. The latter capability was lost by inbreeding.     

Kin recognition by juvenile predatory mites: prior association or phenotype matching?

Kin recognition, the biased treatment of conspecific individuals based on genetic relatedness, is a widespread phenomenon in animals. The most common mechanisms mediating kin recognition are prior association (familiarity) and phenotype matching. Recognition based on prior association allows identifying familiar individuals. Recognition based on phenotype matching is an extension of prior-association-based recognition and allows identifying familiar and unfamiliar individuals due to a shared phenotypic trait. I investigated which of the two mechanisms is used by cannibalistic juvenile predatory mites, Phytoseiulus persimilis. Protonymphs that were associated with either siblings or non-kin discriminated familiar and unfamiliar larvae and preferentially cannibalized the latter irrespective of genetic relatedness. In contrast, despite previous association with either siblings or non-kin, protonymphs did not discriminate unfamiliar sibling and unfamiliar non-kin larvae. Association in the larval stage therefore mediated kin recognition based on familiarity, but not phenotype matching in cannibalistic P. persimilis protonymphs. Furthermore, in the presence of a familiar prey individual, sibling cannibalism occurred significantly sooner than non-kin cannibalism. This quick sibling cannibalism may have been the consequence of preferential association of siblings and/or may indicate the occurrence of an alternative cannibal phenotype. I discuss the adaptive significance of prior-association-based recognition for P. persimilis juveniles and emphasize the ability of P. persimilis to use multiple recognition mechanisms in dependence of the ontogeny and the ecological context.     

Do kin always make better neighbours?: The effects of territory quality

Summary The effects of territory quality on kin-biased territorial defence behaviour and fitness of juvenile rainbow trout (Oncorhynchus mykiss) were examined by manipulating food and predation risk levels in an artificial stream channel. Groups of related (full sibling) or unrelated fry were observed in the channel under one of four treatments: (1) high food-low predator (high territory quality); (2) high food-high predator; (3) low food-low predator; and (4) low food-high predator (low territory quality). Fish within kin groups always initiated fewer aggressive interactions, defended smaller territories, exhibited higher proportions of threat type territorial defence behaviours (as opposed to overtly aggressive behaviours) and had a higher mean weight increase than non-kin groups. Within both kin and non-kin groups, decreasing territory quality significantly increased the frequency of aggressive interactions, the size of territories and decreased growth. It is argued that while kin-biased territorial defence behaviours are always present, decreased availability of resources and increased predation risk serve to reduce the magnitude of this kin bias. However, even in low-quality territories, the benefit of associating with kin may increase the probability of overwintering survival (increased fitness) of the young fish by increasing body size.     

Philopatry, male presence and grooming reciprocation among female primates: a comparative perspective

Competition for food and a safe location in a group are considered to be the main determinants of variation in social relationships among female primates. The effect of the presence of males is usually overlooked, however. Here we put forward two hypotheses connecting the (relative) number of males in a group, a statistic measuring the strength of positive relationships among females (the degree of reciprocity of grooming) and female residence. Under the first hypothesis, we assumed that philopatric females suffer especially from competition for males. Because females of these species are restrained to their native group and thus do not move to groups where better conditions (i.e. more males) prevail, a higher socionomic sex ratio would imply reduced competition and this should be reflected in better relationships among females (i.e. in a higher degree of reciprocation). The second hypothesis is based on the contention of Wrangham (1987) that groups with more males are better able to defend large food sources: in these groups female relationships would suffer less from within-group competition for food. We therefore also expected a positive correlation between the absolute number of males and grooming reciprocation. To test the hypotheses, matrices of grooming interactions were collected by going through the primatological literature. For ten female-resident and four female-transfer primate species, the degree of grooming reciprocation was quantified using a specially adapted matrix statistic. The results favoured the first (competition for males) hypothesis. The degree of grooming reciprocation among females was positively correlated with socionomic sex ratio, but neither with group size nor with the absolute number of males. Correlations between the degree of reciprocity and sex ratio were confirmed at three levels: within the same group at different points in time, between groups of the same species, and between species. In addition, the degree of reciprocation increased with sex ratio more strongly in typical single/variable male than in multi-male species. This we interpreted as a direct consequence of the lower social availability of males for females in multi-male groups than single-male groups. In turn, this effect may be due to males competing for females in multi-male groups. Received: 28 May 1997 / Accepted after revision: 9 November 1997     

The predatory mite Phytoseiulus persimilis manipulates imprinting among offspring through egg placement

A crucial question with respect to imprinting is how animals ensure that kin imprint on kin but not on non-kin. Imprinting takes place in a sensitive ontogenetic phase, usually in an early period of life or when offspring are produced, at which time the recipient imprints on the first referents met. In the predatory mite Phytoseiulus persimilis, imprinting among immature individuals happens in the larval stage immediately after hatching. I tested the hypothesis that adult P. persimilis females manipulate offspring imprinting by influencing the likelihood of encounters among recipients and referents via egg placement and egg aggregation. I conducted two experiments, one of which addressed imprinting and cannibalism, and the other addressed egg placement and egg aggregation. The imprinting experiment suggests that larvae imprint on any conspecific individual met in a sensitive ontogenetic phase and later on treat this individual as kin, irrespective of relatedness. After molting to protonymphs, imprinted individuals preferentially cannibalized unfamiliar to familiar larvae. Irrespective of familiarity, kin were cannibalized earlier than non-kin, suggesting the involvement of self-referent phenotype matching. The egg-placement experiment provides evidence that females adjust the aggregation level of their own eggs according to the degree of relatedness to present eggs from a previously ovipositing female. Both experiments in concert suggest that egg placement is a maternal strategy influencing imprinting among immature individuals. Apart from avoiding kin cannibalism, egg placement and imprinting by larvae may have relevance to other behaviors influenced by kin recognition, such as mate choice, prey-patch choice and dispersal.Communicated by L. Simmons     

Kinship and food availability influence cannibalism tendency in early-instar wolf spiders (Araneae: Lycosidae)

For potentially cannibalistic animals such as spiders, the ability to recognize and avoid kin and/or preferentially cannibalize non-relatives would permit exploiting conspecifics as prey while minimizing loss of inclusive fitness. We investigated the effects of relatedness and availability of alternative food on cannibalism tendency in pairs of juvenile Hogna helluo (Walckenaer), a North American wolf spider (Araneae: Lycosidae). For second-instar spiderlings (dispersing stage), cannibalism was more likely among pairs of non-sibs than pairs of sibs and, interestingly, was also more likely when other prey were available. We found no evidence of increased cannibalism in pairings involving broods of greatest average size disparity, indicating that size differences are unlikely to explain differences in cannibalism tendency. Additionally, the relative number of deaths from cannibalism or other causes did not increase with increasing risk of starvation. For third-instar spiderlings, which had lived independently of their mother and sibs following dispersal, cannibalism rates were very high in all treatments and there were no significant effects of relatedness or food availability. Our results suggest that spiders with predominantly solitary lifestyles may bias cannibalism toward non-kin during the juvenile associative period, and that this effect is lost in the subsequent instar. Results are discussed in the context of several potential mechanisms that might result in differential cannibalism.Communicated by M. Elgar     

Infant handling and mortality in yellow baboons (<Emphasis Type="Italic">Papio cynocephalus</Emphasis>): evidence for female reproductive competition?

This study examines negative and positive infant handling behavior in 24 free-ranging yellow baboon infants (Papio cynocephalus) studied over a 5-year period in Mikumi National Park, Tanzania. We test predictions of the female reproductive competition hypothesis to explain patterns of infant handling behavior by adult females (excluding the infants mother) in relation to observed cases of infant mortality by age 3 months (25% of infants in this study). Results show that: (1) low-ranking infants received more negative infant handling than high-ranking infants; conversely high-ranking infants received more positive infant handling; (2) female kin engaged in higher levels of positive infant handling than did non-kin, whereas non-kin showed higher levels of negative infant handling; (3) rates of negative infant handling varied by season, with high levels at the onset of the rainy season; and (4) high level of negative infant handling was a significant predictor of infant mortality by age 3 months (infant rank and sex did not predict survival). We discuss how the occurrence and interpretation of infant handling behavior in the literature has likely been confused by different definitions of this behavior, as well as differences in the socio-ecological context in which this behavior occurs.Communicated by C. Nunn     

Communal nesting, kinship, and maternal success in a social primate

Communal nesting, where several mothers regularly pool and cooperatively rear offspring, is unusual in mammals. This type of crèching behavior is especially rare among primates, with the notable exceptions of humans, some nocturnal strepsirrhines, and—as we show in this study—black-and-white ruffed lemurs (Varecia variegata). Here, we combine data on nesting behavior, genetic relatedness, and infant survivorship to describe variation in ruffed lemur infant care and to examine the potential benefits of ruffed lemur communal breeding. Reproductive events were rare, and females produced litters (synchronously) only once in 6 years of observation. We show that not all mothers participate in communal crèches, but those that did had greater maternal success; communal breeders spent more time feeding and their offspring were more likely to survive. Although cooperating mothers were often related, females also cooperated with non-kin, and those who shared infant care responsibilities had greater maternal success than mothers who did not participate. If there is indeed a causal link between maternal cooperation and reproductive success, this unusual behavior, like that of human communal rearing, may have evolved via some combination of kin selection and mutualism.     

Signal use by collared lizards, Crotaphytus collaris: the effects of familiarity and threat

Many lizard species use lateral compressions of the body during agonistic encounters. I investigated the signal value of the frequency at which these displays are presented and how that rate is affected by familiarity and threat. The response of resident collared lizards (Crotaphytus collaris) to tethered strangers, neighbors on the correct boundary, and neighbors displaced to the opposite boundary, was recorded by counting the number of lateral displays directed at the intruder and the number of subsequent acts of escalated aggression. There was no relationship between size asymmetry of the opponent and the rate of lateral display or aggression, nor was there a relationship between body size and the rate of lateral display or aggression. However, there was a high positive correlation between lateral display rate and aggression, suggesting that the rate of lateral displays is a conventional signal of motivation to attack. The highest rates of display and aggression were directed toward displaced neighbors, somewhat less toward strangers, and the least toward neighbors at the correct boundary. The ratio of aggressive acts to lateral displays followed the same pattern, presumably because the perceived threat to the resident decreases in the same order. Taken together these data suggest that collared lizards are able to assess the threat of an opponent and signal motivation to respond aggressively towards that opponent.Communicated by S. Downes     

Kin discrimination in the spotted hyena (<Emphasis Type="Italic">Crocuta crocuta</Emphasis>): nepotism among siblings

We examined patterns of affiliation, association, and aggression to inquire whether spotted hyenas (Crocuta crocuta) can distinguish among various groups of maternal and paternal siblings. If so, and if these animals conform to predictions of kin selection theory, then behavioral interactions among hyenas should vary with relatedness. We also considered familiarity-based recognition and phenotype matching as mechanisms hyenas might use to recognize kin. Patterns of affiliative behavior indicated that hyenas favored full-sibling littermates over half-sibling littermates or any other group of half-siblings. Rates of dyadic aggression generally did not vary with kinship. Hyenas associated more closely with half-sibling littermates than with non-littermate half-siblings, and hyenas affiliated more with maternal half-siblings than with paternal half-siblings, suggesting that familiarity-based cues might mediate discrimination among these sibling classes. In addition, operation of a phenotype-matching mechanism was suggested by the preference hyenas demonstrated during affiliative interactions for full- over half-sibling littermates, and by their lack of preference in these interactions for half-sibling littermates over non-littermate half-siblings. Phenotype matching was also suggested by our observation that paternal half-siblings cooperated more, and fought less, than did non-kin. Our data indicate that hyenas can discriminate among various types of siblings, that their social behavior conforms to predictions of kin selection theory, and that they recognize kin using mechanisms of both familiarity and phenotype matching.Communicated by S. Alberts     

Sperm exchange in a simultaneous hermaphrodite

Many simultaneously hermaphroditic animals cross-inseminate when copulating, even though unilateral sperm exchange is functionally possible. The question is why reciprocity is the rule. This study addresses sperm exchange in a hermaphroditic planarian flatworm, Dugesia polychroa, in which self-fertilisation does not occur. We investigated the availability of self sperm for inseminating a partner, how self sperm affects the likelihood of sperm donation, and whether sperm donation depends on reciprocation by the partner. The amount of self sperm depended on body size, duration of isolation and experimental date, and animals that had more self sperm were more likely to mate. Depletion of allosperm (sperm received from partners) played only an indirect role, suggesting that animals copulate more to donate sperm rather than to replenish allosperm reserves. Among 60 copulating pairs, reciprocal insemination was more common (63.3%) than unilateral transfer (18.3%). A surprising 18.3% did not exchange sperm in either direction. Individuals were more likely to behave like their partners, resulting in more symmetrical matings (either reciprocity or no sperm exchange) than asymmetrical matings (unilateral sperm donation). The amount of self sperm donated during a copulation depended mainly on the amount available before the copulation and not on the amount received from the partner. The results suggest that conditional sperm exchange, or sperm trading, takes place in D. polychroa. Received: 31 May 1997 / Accepted after revision: 1 December 1997     

Effects of kinship on territorial conflicts among groups of lions,<Emphasis Type="Italic"> Panthera leo</Emphasis>

Inclusive fitness theory predicts that cost of tolerant behaviour during competitive interactions is lower for relatives than for nonrelatives. Many studies have examined the effect of relatedness on behaviour within social groups. In contrast, kin selection acting among groups has received less attention. The genetic structure of African lion (Panthera leo) populations creates a strong possibility that kin selection among groups modifies behaviour during group conflicts. We used playback experiments and genetic data to investigate the importance of relatedness during simulated territorial disputes in lions. However, we found no effect of relatedness on territorial behaviour. Degree of relatedness did not affect the decision to approach simulated intrusions, nor did it affect the behaviour during approaches. The decision to approach was instead affected by position within the territory and consecutive playback number (a measure of habituation). For playbacks that did elicit an approach, the speed of response was not detectably affected by relatedness, but was affected by odds (the ratio of residents to intruders), number of intruders, number of bouts, presence of cubs, position within the territory, temperature and playback number. Although responses were unaffected by relatedness, it remains possible that other aspects of behaviour during natural encounters among prides are affected by kin selection.Communicated by L. Sterck     

Estimation of temporary emigration in male toads

Male boreal toads (Bufo boreas) are thought to return to the breeding site every year but, if absent in a particular year, will be more likely to return the following year. Using Pollock's robust design we estimated temporary emigration (the probability a male toad is absent from a breeding site in a given year) at three locations in Colorado, USA: two in Rocky Mountain National Park and one in Chaffee County. We present data that suggest that not all male toads return to the breeding site every year. Our analyses indicate that temporary emigration varies by site and time (for example, from 1992 to 1998, the probability of temporary emigration ranged from 10% to 29% and from 3% to 95% at Lost Lake and Kettle Tarn, respectively). Although the results provide weak evidence that males are more likely to return after a year's hiatus, a general pattern of state-dependent temporary emigration was not supported. We also hypothesized relationships between temporary emigration and a number of weather variables. While some competitive models included weather covariates, imprecise and variable estimates of the effects of these covariates precluded fully defining their impact on temporary emigration.     

Strategic use of allopreening in family-living wrens

A prominent interaction in the lives of social mammals is allogrooming. Individuals allogroom strategically and preferentially, grooming high-quality individuals that control access to resources. This results in distinct patterns of allogrooming within social groups, such that some individuals are groomed more than the others, some dyads express symmetry in allogrooming, and others exchange allogrooming for other benefits. Allopreening, the avian equivalent of allogrooming, occurs commonly in group-living birds, providing the opportunity to test whether social birds also use allopreening strategically in their social relationships. I examined this hypothesis in family-living buff-breasted wrens (Cantorchilus leucotis) by examining allopreening initiation, reciprocation and rate during interactions involving breeding adults and offspring. Buff-breasted wrens exhibit a nuclear family structure in which pairs display long-term partnerships, and juveniles delay dispersing from natal territories for about 10 months. Allopreening was symmetrical between breeding partners: males and females who initiated and reciprocated allopreening of partners with similar frequency, and within reciprocated bouts, allopreening was time-matched. Pairs in which allopreening was not observed were more likely to divorce, but allopreening patterns did not change in successive years of partnerships. Parent–offspring allopreening, by contrast, was asymmetrical in pattern; parents initiated allopreening more than their offspring, and reciprocation never occurred. The different patterns of allopreening observed within buff-breasted wren families suggest the strategic use of allopreening, possibly for the maintenance of long-term partnerships and in exchange for social benefits from offspring.