Food sharing by sibling vs nonsibling spiny mice (Acomys cahirinus)

Summary In comparison to paired unfamiliar unrelated age-mates, pairs of familiar littermate (sibling) spiny mice (Acomys cahirinus) were more active in exploring a novel environment containing limited food resources and engaged in a greater degree of cooperative feeding or food sharing. When animals born in successive litters were tested, however, familiar siblings did not differ from unfamiliar nonsiblings on measures of food sharing even though subsequent data indicate that such siblings could recognize one another. Young animals were found to feed for longer periods of time than their older cagemate regardless of whether the latter animal was a familiar sibling or an unfamiliar nonsibling.     

Incest avoidance and bonding between siblings in gerbils

Summary Social and sexual preferences for siblings versus non-siblings have been investigated in Mongolian gerbils (Meriones unguiculatus) in a series of laboratory and outdoor enclosure (100 m²) studies. When first social contact took place at pairing in adulthood full siblings and nonsiblings did not differ with respect to onset of reproduction. Correspondingly, both siblings and non-siblings paired at weaning age had a delayed onset of reproduction. Thus, no effect could be established in either case, related to the individuals' relatedness, but the importance of social exposure was clear. Non-siblings paired when 2 months old had a delayed onset of reproduction, which did not differ statistically from the onset in siblings, raised together, and housed pairwise since the same age. When sexually mature animals were paired at 3 months of age, siblings were found to start reproducing later than non-siblings (Ågren 1981). Following an initial delay in both siblings and nonsiblings paired when 5 months old, all non-sibling pairs reproduced before any sibling pair. At pair formation, at least the females in both groups were known to be sexually immature. These results indicate that for sibling-bonding to occur social exposure may be delayed until the animals are old enough, or somatically developed, to mature sexually. Thus, social exposure before weaning age during the period for primary socialization seem not to be crucial, nor the actual maturational state, sexually, in older females. To study the stability of the social bonds between siblings and non-siblings, animals kept pairwise in the laboratory were released in large outdoor enclosures. Pairs brought together before the age of sexual maturity did not remain stable, whether or not the former cagemates were siblings. Non-sibling pairs formed after this age, which had also reproduced successfully in the laboratory, did remain stable. Social and sexual interactions were studied within and between two mixed-sex sibling groups of four. The sibling groups had the possibility to establish group territories before being exposed to each other. Although the sibling groups remained stable, the females still preferred non-sibling males as sexual partners. However, socially dominant brothers copulated more frequently than subordinant brothers. This rank difference was not evident in non-siblings.     

The effects of age and relatedness on mating patterns in thornbug treehoppers: inbreeding avoidance or inbreeding tolerance?

The social environment of many species includes synchronous maturation of siblings in family groups, followed by limited dispersal of adults from their natal site. Under these conditions, females may experience high encounter rates with same-age siblings during mate searching, increasing their risk of inbreeding. If inbreeding depression occurs, mating with a sibling is often considered maladaptive; however, in some contexts, the inclusive fitness benefits of inbreeding may outweigh the costs, favoring females that tolerate some level of inbreeding depression. We evaluated mating patterns in the treehopper Umbonia crassicornis, a semelparous species in which females encounter same-age siblings during mate searching. A female U. crassicornis that mates with a brother suffers from inbreeding depression. We used a free-choice mating design that offered females simultaneous mating opportunities with three groups of males: siblings, same-age nonsiblings, and older nonsiblings. These groups represent the types of males typically encountered by females during mate searching. Our goal was to assess whether mating patterns were influenced by inbreeding avoidance by evaluating two hypotheses: kin discrimination and age-based mating (older males cannot be siblings in this species). There was no difference in the proportions of females mating with siblings vs nonsiblings, suggesting an absence of kin discrimination. However, females mated with a greater proportion of older vs younger males. Given that females do not avoid siblings as mates despite a cost to inbreeding, our results provide a possible example of inbreeding tolerance. We also discuss some factors that may have contributed to the mating advantage of older males.     

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     

Ecology of kin and nonkin larval interactions in Tribolium beetles

Summary The larvae of flour beetles Tribolium castaneum and T. confusum were reared in two kinds of groups: full siblings and unrelated individuats. These kin and nonkin groups were reared in open cultures, in which emigration was permitted (both species) and in closed cultures, in which emigration was prohibited (only T. confusum). We measured larval development time and survivorship, weight of pupae, and time of larval emigration from open cultures. The effects of age structure were investigated by establishing open cultures of larvae of uniform age (larvae hatched from eggs laid within 72 h) and cultures of larvae of variable age (eggs laid within 240 h). In closed cultures of siblings, T. confusum larvae pupated on an average 2.2 days earlier than larvae reared in nonsibling groups. In T. castaneum, more small and medium size and fewer large size larvae emigrated from groups of siblings compared to groups of nonsiblings. Males that emigrated and pupated remained with their sibs for a shorter time than did similar males raised with unrelated larvae. In T. castaneum, age structure variation reduced the sibs tendency to migrate, but did not influence interactions among unrelated larvae. The genetical effects of kinship and the ecological effects of age structure were shown to affect the interactions of Tribolium larvae reared in groups. Reducing the similarity between individuals, either genetically or demographically (using mixed broods or mixed age cohorts), changed the pattern of larval interactions. Upon occasion, the effects of kin interactions may well be the mechanical consequences of the coexistence of similar individuals rather than the effects of altruistic behavior.     

Do newborn domestic rabbits Oryctolagus cuniculus compete for thermally advantageous positions in the litter huddle?

Competition among mammalian siblings for scarce resources can be severe. Whereas research to date has focused on competition for the mother’s milk, the young of many (particularly altricial) species might also be expected to compete for thermally favorable positions within the nest, den, or litter huddle. We investigated this in newborn pups of the European rabbit Oryctolagus cuniculus, a species in which the altricial young are not brooded by the mother, and in which competition for milk is severe. In eight unculled litters (N = 86 pups) of a domestic chinchilla strain, we calculated huddling indexes for individual pups on postnatal days 2–5 as a measure of the degree of insulation they received from littermates. Pups maintained almost constant physical contact with the litter huddle. They performed brief but frequent rooting and climbing behaviors, which usually improved their huddling index, interspersed with longer periods of quiescence during which their huddling index declined. As expected, we found a significant positive relation between pups’ mean huddling index and body temperature. Unexpectedly, however, we did not find a relation between huddling index and pups’ birth weight, survival, milk intake, or efficiency of converting milk to body mass. We conclude that rather than competing for thermally advantageous positions within the huddle newborn rabbits share out thermally advantageous positions as they move in a continual dynamic flow through it. Thus, in newborn rabbits, competition for the mother’s milk exists alongside mutual “cooperative” benefits of littermate presence. This contribution is part of the special issue “Sibling competition and cooperation in mammals” (guest editors: Robyn Hudson and Fritz Trillmich).     

A field test of the effects of familiarity and relatedness on social associations and reproduction in prairie voles

Inbreeding depression is a well-documented phenomenon. In animals, one means of avoiding the costs of inbreeding is through the recognition and avoidance of kin as mates. Prairie voles (Microtus ochrogaster) are short-lived, socially monogamous rodents that demonstrate inbreeding depression in the laboratory. Field data indicate that pair formation in nature is opportunistic but pairing among close relatives seems uncommon. We examined the role of relatedness and familiarity on prairie vole social associations and reproduction by placing adult voles into 0.1-ha enclosures with familiar siblings, unfamiliar siblings, and unrelated, unfamiliar conspecifics. Live-trapping data indicated that indices of social pair bonding were random with respect to relatedness and familiarity. Among females whose litters were sired by a single male, litters were significantly more likely to be sired by unfamiliar than familiar males, but the number of litters sired by males that were unrelated to their partner was not different from the number of litters sired by males that were related to their partner. Additionally, females that produced offspring with familiar siblings were significantly more likely to have litters with multiple paternity than females not producing offspring with familiar siblings. However, multiple paternity was not influenced by relatedness of sires. Finally, older individuals were more likely to produce offspring with each other than with younger individuals. Our findings suggest that prior association is a more important mechanism of inbreeding avoidance than phenotype matching for prairie voles mating under ecologically relevant conditions.     

Food delivery and sibling competition in experimentally even-aged broods of the cattle egret

Summary The age of nestlings was made even in 12 nests of cattle egrets (Bubulcus ibis) by exchanging young chicks or eggs in a breeding colory in Japan. The growth of chicks in such synchronously hatching broods (SHBs) grew almost as fast as first-hatched chicks in 38 control (asynchronously hatching) broods (AHBs). Four last-hatched chicks in AHBs but no chicks in SHBs died of starvation. The frequency of parental nest-visits with food, that of begging food by chicks, and food mass eaten by chicks were greater in SHBs than in AHBs during the first half of the nestling period and similar thereafter. Dominance rank within each SHB was formed through ritualistic fights among siblings. It was correlated with neither growth rates nor winning ratios in food contests, but in some SHBs the subordinate chicks were attacked more frequently by dominant siblings during food contests than in AHBs. The most subordinate chick in one SHB died as a direct result of such attacks. Sibling aggressions were more frequent in SHBs than in AHBs.     

Seasonal decline in intracolony genetic relatedness in eastern tent caterpillars: implications for social evolution

Summary Genetic relatedness in social insect colonies may vary spatially or temporally as a result of changes in colony membership due to immigration or to variation in patterns of maternity and paternity. We estimated relatedness for eastern tent caterpillars (Malacosoma americanum) in laboratory colonies derived from egg masses using multilocus genotypic data derived from electrophoresis. This estimate is compared with estimates obtained from colony samples taken in the field at four intervals spanning the larval developmental season. We found that average intracolony relatedness is close to 0.5 initially but declines through the developmental season due to colony merging, showing that caterpillars do not discriminate between siblings and nonsiblings in order to preserve colony family structure. Using the intracolony values together with relatedness values for higher levels of population structure, we estimated the effective mean number of simple families represented in single colonies through the season. The overall effective number of families per tent increased from one at the time of eclosion to 1.3 by the end of the season. Average intracolony relatedness remained relatively high despite the occurrence of colony merging, apparently as a result of the low density of tents on most trees, combined with high relatedness within the original colonies. Thus, high intracolony relatedness is maintained in M. americanum populations through the effects of adult dispersal, mating, and oviposition patterns, rather than through behavioral discrimination mechanisms of the larvae. These findings underscore the importance of considering the causes of temporal variation in genetic relatedness as well as the consequences for the indirect component of inclusive fitness. Correspondence to: J.T. Costa     

How does salinity tolerance influence the distributions of <Emphasis Type="Italic">Brachionus plicatilis</Emphasis> sibling species?

Environmental salinity is important in defining Brachionus plicatilis sibling species distributions. However, while salinity influences distributions, sibling species often co-exist. Three different mechanisms potentially account for the partial co-occurrence of sibling species: (1) siblings have differing salinity tolerances that partially overlap; (2) siblings physiological tolerances may be commonly broad, but relatively small differences in tolerances differentiate distributions via interactions e.g. competition; or (3) siblings distributions may be influenced by physical factors other than salinity. Here, we assess the extent of salinity tolerance in three B. plicatilis sibling species (B. plicatilis 6TUR, B. plicatilis IOM and B. rotundiformis 6TOS) by measuring population growth rate (μ, day⁻¹) and egg development time in response to salinity (5–60‰) and salinity fluctuations (≤ Δ40‰). Sibling species were identified by analysis of the mitochondrial COI gene, and salinity responses were compared by regression analysis. Responses differed significantly between siblings, although the broad trends were similar. Positive growth occurred at all salinities, and highest growth rates ranged between 0.93 and 1.08 day⁻¹ at 16–18‰. Rapid changes in salinity reduced growth rates, but net mortality occurred only in one treatment (100% mortality on transfer from 10 to 40‰). Egg development time was largely invariant with salinity except for B. plicatilis IOM and where rotifers were transferred from 30 to 60‰. We indicate that several siblings are similarly euryhaline and tolerate salinity fluctuations. Undoubtedly, wide tolerances in B. plicatilis are adaptations to ephemeral and seasonally variable habitats. Given common broad salinity tolerances, it is unlikely that the differential distributions of sibling species are a direct result of physiological constraints. Instead, we illustrate using a simple model that subtle differences in physiological tolerances may have important impacts on interactions between sibling species, which may in turn influence distributions.     

An indirect test of inbreeding depression in the termites Reticulitermes flavipes and Reticulitermes virginicus

We analyzed tandem-running pairs of the termites Reticulitermes flavipes and Reticulitermes virginicus utilizing 13 and 12 microsatellite loci, respectively. Newly formed pairs in both species were significantly related to one another, but this average relatedness was considerably higher in R. flavipes (0.130 vs 0.060). These average relatedness levels resulted from some tandem pairs forming between nestmate termites: more than one quarter of all R. flavipes pairs (26.1%) met this criterion, while this was the case for only about one of every 20 R. virginicus pairs (5.1%). The likelihood that termites paired with siblings was inversely related to the inferred dispersal ability of the two species. F _ST, measured over identical spatial scales, was significantly higher in R. flavipes (0.034) than in R. virginicus (0.008). A comparison in R. flavipes of the observed proportion of nestmate pairs observed during tandem running vs the proportion found in established colonies revealed a significant excess of close relatives when pairs were first formed. There are two possible causes of this discrepancy: inbreeding depression (ID) may eliminate inbred colonies early in development, or related pairs may part late in the tandem-running phase or after it is completed. The latter explanation of inbreeding avoidance implies either historical or contemporary ID, and these results therefore suggest that, either directly or indirectly, ID could be a more potent force in the evolution of termite mating systems than is generally appreciated. This work was funded by grants from the United States Department of Agriculture National Research Initiative Competitive Grants Program (nos. 00-35302-9377 and 2002-35302-12490).     

Secondary sex ratios do not support maternal manipulation: extensive data from laboratory colonies of spiny mice (Muridae: Acomys)

Spiny mice of the genus Acomys (Muridae) represent a very suitable mammalian model for studying factors influencing the secondary sex ratio (SSR). The maternal effort in these rodents is extremely biased in favour of the prenatal period and, therefore, maternal manipulation of the SSR is potentially more advantageous. We studied the SSR in four populations/species of spiny mice kept in family groups consisting of two closely related females, one non-relative male and their descendants. The groups were established from founding animals aged about 3 months (maturing age) and were allowed to breed freely for several months. Each litter was sexed after birth, and relevant data were thoroughly recorded. Altogether, data were collected on 1684 litters: 189 of Acomys sp. from Iran, 203 of A. cilicicus, 875 of A. cahirinus, and 417 of A. dimidiatus. We recorded the sex of 4048 newborns of which 1995 were males and 2053 were females. The overall sex ratio was close to 1:1 (49.2%). Generalized linear mixed models and/or generalized linear models were constructed to evaluate the effect of four life history and eight social variables on the sex ratio. No consistent effects of these variables on the sex ratio were found and, interestingly, none of the variables associated with maternal life history had any effect on the sex ratio. Three factors associated with group composition (i.e. the number of immature males, the number of immature females and the number of breeding females) did have significant effects on the sex ratio, but these effects were not consistent across the studied species. In conclusion, our evaluation of this large dataset revealed that the sex ratio in spiny mice is surprisingly stable.     

Transmission of food preferences in spiny mice (Acomys cahirinus) via nose-mouth interaction between mothers and weanlings

Summary Two laboratory experiments assessed the role of nose-to-head and nose-mouth investigation in the transmission of food preferences from Acomys cahirinus mothers to their offspring. Pups spent more time nosing and licking the nose-mouth area of mothers who ate a novel food (fresh potato or carrot) than of mothers who ate no novel food. A greater amount of the less-preferred novel food (potato) was subsequently eaten by pups whose mothers had eaten potato than pups whose mothers had eaten either carrot or familiar laboratory chow only. Mothers maintained a stereotypic posture during nose-mouth investigation, holding still with mouth slightly open, providing both a source of specific food cues and a context in which pups could become familiar with them. These data suggest an active role by Acomys mothers in the development of offspring food preferences.     

Kinship, familiarity and social status modulate social learning about “micropredators” (biting flies) in deer mice

Animals can learn to recognize and respond to dangerous, threatening factors through either individual or social learning, whereby an individual learns and acquires the defensive behaviors and avoidance responses of another. Here we show that kinship, familiarity, and relative dominance of the interacting individuals affect social learning of defensive responses to micropredators in deer mice, Peromyscus maniculatus. Brief exposure of individual male deer mice to biting flies (stable fly, Stomoxys calcitrans) induced a decrease in pain sensitivity, or analgesia, and active self-burying avoidance responses. The defensive analgesic responses and their fear/anxiety/stress associated correlates facilitate the display of behavioral avoidance responses. Preparatory analgesia and avoidance responses were evident 1–3 days later when the mice were exposed to biting flies that were altered to be incapable of biting but were not displayed to similar-sized non-biting house flies. These anticipatory avoidance and analgesic responses to biting flies were also acquired through social learning—without direct individual aversive experience with biting flies. Fly-naive mice (observers) that witnessed other mice (demonstrators) being attacked by biting flies but themselves were not bitten did not display any avoidance responses. However, when exposed 1–3 days later to altered flies with biting mouth parts removed, the observers displayed socially acquired analgesic and self-burying avoidance responses. Observers whose demonstrators were either kin (siblings) and, or members of a familiar pair (kin or non-kin) displayed enhanced social learning of defensive responses. Also, within the familiar pairs, social status affected learning with subordinate observers displaying better social learning than dominant observers. These findings indicate that kinship, familiarity and social status modulate social learning of defensive responses to, and the recognition of, dangerous and threatening stimuli, likely including that of predators.Communicated by G. Wilkinson     

Sex differences in the ontogeny of social behavior in pikas: possible relationships to dispersal and territoriality

Summary A laboratory study on the ontogeny of social behavior in pikas (Ochotona princeps), an alpine lagomorph, was conducted to determine the role of early relationships between adult females and young and among siblings in the development of territorial and dispersal behaviors. Sex differences during development were examined because field studies have reported greater dispersal distances in young females than young males. At birth, females were significantly heavier than males. There were no sex differences in nursing frequency until after the 2nd week of age, when males initiated more nursing attempts than females. By the end of the weaning period (weeks 5 and 6), adult females became non-interactive with young, but aggression of young toward littermates and the mother increased until the eighth week. At this time, young males outweighed their sibling females. Young were dominant over their mothers by the age of 5 weeks, and young males were dominant over their sibling females.Sex differences were observed in aggression, scent-marking, exploratory activity, and submissive vocalizations, with higher rates in young males, except for submissive vocalizations, which were higher in females (Table 2). Vocalizations and scent-marking behavior increased over time, and were positively correlated with interaction rates.These data support the hypothesis that female young disperse farther than male young largely as the result of unsuccessful competition with male siblings for available territories close to the birthplance. A dispersal strategy for pkkas is proposed.     

Male mating speed promote hybridization in the Rana lessonae–Rana esculenta waterfrog system

Hybridization is a widespread phenomenon in many vertebrate groups. Prezygotic isolating mechanisms, probably caused by selection against hybrids with reduced fitness, reduce the likelihood of such events. Although hybrid-reduced fitness relatively to parental species is common, hybridization can also be beneficial, and hybrids sometimes outperform the pure species type. In this study, we examined two potential processes, Hubbs’s principle and male–male competition, which could enhance hybridization in the waterfrog complex and thus explain the proportion of heterospecific pairs collected in a natural pond. Firstly, by collecting 791 frogs in the field to study pair and chorus composition, we showed that in a mixed Rana lessonae–Rana esculenta population, the scarcity of hybrid R. esculenta males did not account for the proportion of heterospecific pairs: indeed, when examining pairing composition in six different choruses, we found that hybrid males were always under-represented and that R. esculenta females were found paired with R. lessonae males. Secondly, we investigated experimentally whether or nor male–male competition mechanism could explain pair formation in waterfrogs. Our mating speed experiment highlights mechanisms that could explain heterospecific pairs in a context of promiscuous mating where scramble competition was intense. To measure the rapidity with which a male grasps a female, we placed males in a grid cage with a female, and the dynamics of pair formation was monitored. R. esculenta males showed a lower pairing success than R. lessonae males as a smaller proportion of them amplexed females, and more time was needed for them to get amplexed. Thus, a less adaptative mechanism than female mate choice may also explain the mating pattern observed in waterfrog species.     

Sibling competition in guinea pigs (<Emphasis Type="Italic">Cavia aperea</Emphasis> f. <Emphasis Type="Italic">porcellus</Emphasis>): scrambling for mother’s teats is stressful

Lactation is the most energy-intense period in the life of a female mammal. This can cause severe conflict between mother and offspring over the duration of lactation but also between siblings over the amount of milk each pup gets from its mother. Thus, competitive interactions between siblings are expected, and competition is likely to increase with litter size, particularly in species where the number of offspring exceeds the number of teats. We studied sibling competition in the domestic guinea pig (Cavia aperea f. porcellus), which has two teats, but frequently bears litters of up to five pups. By cross-fostering we created non-competition (control) litters with two pups and competition litters with four pups and observed nursing behaviour on days 5, 10, 15 and 20 postpartum. Pups of larger litters had lower growth rates, indicating increased competition among siblings in these litters. Pups of larger litters had to wait longer for access to a teat and spent less time suckling than pups of smaller litters but ate more solid food instead. Additionally, we manipulated the individual short-term need of pups by separating half of the pups of each litter for 2 h from their mothers before observation. Within a litter, hungry pups achieved access to milk faster and spent more time suckling than non-hungry pups. Pups competed mostly by scramble competition. Aggressive interactions occurred only in large litters. Pups of large litters had higher cortisol levels than pups in small litters. These effects decreased with age as pups became increasingly independent of maternal milk. Pup behaviour appears to fit better with models of scramble competition than with those of honest signalling. This contribution is part of the special issue “Sibling competition and cooperation in mammals” (guest editors: Robyn Hudson and Fritz Trillmich).     

Differential aggressiveness between fire salamander (<Emphasis Type="Italic">Salamandra infraimmaculata</Emphasis>) larvae covaries with their genetic similarity

Responding differentially to kin and non-kin is known to be adaptive in many species. One example is the inclusive fitness benefits of reducing aggression toward closer relatives. Little is known, however, about the ability of animals to assess differential degrees of genetic relatedness and to respond accordingly with differential levels of aggression. In the present study, we tested whether aggressiveness between body mass-matched pairs of fire salamander (Salamandra infraimmaculata) larvae covaried with the genetic similarity between them. We quantified aggressiveness at three levels of genetic similarity by selecting pairs within and across pools from recently genotyped populations. We also assessed aggression between pairs of siblings. Aggression and associated injuries decreased as genetic similarity increased across the groups. These findings suggest that cannibalistic salamanders can assess their degree of genetic relatedness to conspecifics and vary their behavioral responses depending on the degree of similarity between them along a genetic relatedness continuum.     

Parent–offspring resource allocation in domestic pigs

Behavioural research on domestic pigs has included parent–offspring conflict, sibling competition, and the use of signals which influence resource allocation. In this paper, we review key sow–piglet behavioural studies and discuss their relevance to resource allocation theory. Sibling competition begins in the uterus and continues after birth, as piglets compete directly for access to the sow’s teats. This competition is made more severe by a unique dentition, which newborn piglets use to lacerate the faces of siblings during teat disputes. Competition often leads to the death of some littermates, especially those of low birth weight. Piglets also compete indirectly for milk, apparently by stimulating milk production at the teats that they habitually use at the expense of milk production by other teats. The complex nursing behavior of the sow appears designed to prevent the more vigorous piglets from monopolizing the milk. Sows give vocal signals which both attract piglets to suckle and synchronize their behavior during nursing episodes. Piglets give loud vocal signals when separated from the sow; calls which vary in intensity and appear to be honest signals of need. Udder massage by piglets may also serve as an honest signal of need. Parent–offspring conflict has been demonstrated experimentally in pigs. Specifically, when given the opportunity to control contact with their piglets, sows nurse less frequently, provide less milk, and lose less weight during lactation than sows that cannot control their level of contact. Because of this interesting natural history and because they are so amenable to experimentation, domestic pigs provide a rich system for testing ideas drawn from resource allocation theory. This contribution is part of the special issue “Sibling competition and cooperation in mammals” (guest editors: Robyn Hudson and Fritz Trillmich).     

Recognising one’s enemies: a functional approach to risk assessment by prey

Little has been done to compare the relative importance of various mechanisms through which prey assess the potential risk from natural enemies. We used predator-naive spider mites (Tetranychus urticae, Tetranychidae) to (1) compare the responses of prey to chemical cues from enemy and non-enemy species and (2) investigate the source of these cues. In the laboratory, we observed the distribution of T. urticae in response to cues from nine mite species, including (1) predators of spider mites, (2) predators/parasites of other animals, and (3) fungivores/pollen-feeders. When given a choice over 24 h, spider mites foraged and oviposited in fewer numbers on leaf discs that were previously exposed to predatory or parasitic mites (including species incapable of attacking spider mites) than on clean leaf discs (unexposed to mites). Interestingly, previous exposure of leaf arenas to fungivores and pollen-feeders had no significant effect on spider mite distribution. We then observed the response of T. urticae to cues from two species of predator that had been reared on a diet of either spider mites or pollen. T. urticae showed stronger avoidance of leaf discs that were previously exposed to spider-mite-fed predators than of discs exposed to pollen-fed predators. Nevertheless, for one predator species (Amblyseius andersoni), T. urticae still preferred to forage and oviposit on clean (unexposed) discs than on discs exposed to pollen-fed predators. Protein-derived metabolic wastes of predatory or parasitic mites may provide a general cue about potential predation risk for T. urticae. However, T. urticae also avoided areas exposed to pollen-fed predators, suggesting there may be other sources of enemy recognition by the spider mites. We discuss the ecological and evolutionary mechanisms that may influence the scope of information through which animals assess predation risk. Received: 11 January 1999 / Received in revised form: 25 October 1999 / Accepted: 20 November 1999