Responses to threat by female bobucks,<Emphasis Type="Italic"> Trichosurus caninus</Emphasis>, during different stages of offspring development

The costs of reproduction are widely recognised as a major selective force in the evolution of various behavioural and life-history characteristics. In particular, the behaviour of reproductively active animals is likely to change when breeding increases risk of predation. We investigated the effect of an experimentally derived threat on the vigilance and escape behaviour of female Trichosurus caninus with and without dependent offspring and at different stages of offspring development. Females with offspring showed a heightened response to the threat in comparison to females without offspring. In addition, females with offspring displayed a stronger response at earlier stages of their offsprings development.Communicated by P. Bednekoff     

Biparental defence in a spider mite (Acari: Tetranychidae) infesting Sasa bamboo

Summary Field observations on the nesting pattern of Schizotetranychus celarius (Banks) (Acari: Tetranychidae) revealed that many individuals live gregariously in a united nest. Behavioural experiments recorded on video tape recorder and 16 mm film showed that adult males and females can effectively defend their nest and offspring from a phytoseiid predator, larvae of Typhlodromus bambusae Ehara (Acari: Phytoseiidae), through counterattacks. The male spider mite often killed the predator's larva and succeeded in defending offspring and nest completely. Comparison of the behaviour patterns recorded between a nest owner parent S. celarius and two kinds of intruders (predators and conspecific males or females) revealed conspicuous differences. I suggest that there is a kind of biparental care in S. celarius, and that the life-pattern of this species therefore fits the definition of subsociality originally framed for insects.     

Differential effects of offspring condition-dependent signals on maternal care regulation in the European earwig

Parent–offspring conflict theory predicts the evolution of offspring solicitation signals that can influence the amount and/or the duration of parental investment. Short-term effects of offspring solicitation signals on parental food provisioning have been widely demonstrated, but persistent effects of offspring signals on the maintenance of parental care have been rarely studied. Also, the relation between the amount of care provided to the brood and how it is distributed among individual offspring within a brood is not well enough understood. Here, we investigated in the European earwig (Forficula auricularia) the effects of offspring condition-dependent chemical signals on the maintenance of maternal care among broods and the distribution of maternal food within broods. Mothers were isolated from their brood for 3 days and continuously exposed to chemical signals extracted from broods of experimentally manipulated nutritional state. After re-introducing mothers to their brood, a range of maternal behaviours were quantified. We found that earwig mothers groomed their offspring significantly more after exposure to chemical extract from high-food brood in comparison with mothers exposed to extract from low-food brood, which in turn displayed significantly more aggressive behaviour. Furthermore, we manipulated offspring individual nutritional condition within the brood to evaluate the effect of offspring state on the within-brood food distribution. Within broods, poorly fed individuals received significantly more food than well-fed individuals, probably due to scramble competition. These results show that earwig nymphs express multi-component condition-dependent signals and behaviours differentially affecting maternal care provisioned to the brood and the distribution of care within broods.     

Biparental investment and reproductive success in a subsocial desert beetle: the role of maternal effort

Parastizopus armaticeps is a nocturnal subsocial detritivorous desert tenebrionid that produces very few offspring per brood. The two environmental factors that constrain reproduction, rapid sand desiccation rate and food scarcity, are countered by biparental effort. Males dig and extend breeding burrows, maintaining their moisture level; females forage on the surface at night for high-quality detritus, the larval food. This was shown to be a scarce and unpredictable resource for which there is high competition. When food was supplemented in a field experiment, offspring number and survivorship doubled and burrow failure due to desiccation dropped from approximately half, the typical failure rate for unsupplemented burrows, to zero. Food supplementation did not, however, increase larval foodstore size and there was no difference in the size of the offspring produced. Supplemented females reallocated their time, foraging less and digging more with the male. This change in maternal behaviour patterns resulted in deeper burrows which remained moist longer, thus extending the larval production period. Female foraging efficiency, particularly food retrieval speed, determined how much time females could allocate to digging, consequently increasing the reproductive success of the pair. Burrow depth and sand moisture level at the burrow base were the major correlates of reproductive success, but the scarcity and unpredictability of high-quality food on the surface and the competition for this resource influenced the number of offspring indirectly through their effect on female behaviour. Received: 29 November 1996 / Accepted after revision: 7 December 1997     

Learning and natal host influence host preference,handling time and sex allocation behaviour in a pupal parasitoid

The host choice and sex allocation decisions of a foraging female parasitoid will have an enormous influence on the life-history characteristics of her offspring. The pteromalid Pachycrepoideus vindemiae is a generalist idiobiont pupal parasitoid of many species of cyclorrhaphous Diptera. Wasps reared in Musca domestica were larger, had higher attack rates and greater male mating success than those reared in Drosophila melanogaster. In no-choice situations, na?ve female P. vindemiae took significantly less time to accept hosts conspecific with their natal host. Parasitoids that emerged from M. domestica pupae spent similar amounts of time ovipositing in both D. melanogaster and M. domestica. Those parasitoids that had emerged from D. melanogaster spent significantly longer attacking M. domestica pupae. The host choice behaviour of female P. vindemiae was influenced by an interaction between natal host and experience. Female P. vindemiae reared in M. domestica only showed a preference among hosts when allowed to gain experience attacking M. domestica, preferentially attacking that species. Similarly, female parasitoids reared on D. melanogaster only showed a preference among hosts when allowed to gain experience attacking D. melanogaster, again preferentially attacking that species. Wasp natal host also influenced sex allocation behaviour. While wasps from both hosts oviposited more females in the larger host, M. domestica, wasps that emerged from M. domestica had significantly more male-biased offspring sex ratios. These results indicate the importance of learning and natal host size in determining P. vindemiae attack rates, mating success, host preference and sex allocation behaviour, all critical components of parasitoid fitness. Electronic Publication     

Heritable and experiential effects on boldness in a tropical poeciliid

Consistent differences in human behaviour are often explained with reference to personality traits. Recent evidence suggests that similar traits are widespread across the entire animal kingdom and that they may have substantial fitness consequences. One of the major components of personality is the shyness–boldness continuum. Little is known about the relative contributions of genes and the environment in the development of boldness in wild animal populations. Here, we bred wild-caught fish (Brachyraphis episcopi) collected from regions of high- and low-predation pressure, reared their offspring in the laboratory under varying conditions and tested boldness utilising an open-field paradigm. First-generation laboratory-reared fish showed similar behaviour to their wild parents suggesting that boldness has a heritable component. In addition, repeated chasing with a net increased boldness in both high- and low-predation offspring, showing that boldness is also heavily influenced by life experiences. Differences between males and females were also sustained in the laboratory-reared generation indicating that sex differences in boldness are also heritable. We discuss these results with reference to the potential underlying genetic and hormonal mechanisms as well as the environmental influences that may be responsible for expression of boldness in wild animals.     

Territorial counter-singing in male sac-winged bats (<Emphasis Type="Italic">Saccopteryx bilineata</Emphasis>): low-frequency songs trigger a stronger response

In many animal species, individuals compete for resources but avoid escalated conflicts by threat displays, i.e. a mutual signalling behaviour that enables the opponents to predict the outcome of the conflict without the necessity of actual fighting. For example, territory holders may use acoustic signals to communicate not only their own identity and the borders of their territory but also their competitive quality, fighting ability and motivation. Here, we show that male sac-winged bats, Saccopteryx bilineata, adjust their vocal territorial displays according to the fundamental frequency of territorial songs of their opponents. In playback experiments with territorial males, low-frequency stimuli elicited a higher territorial song rate and length than high-frequency stimuli. Male S. bilineata that sing more often and with lower fundamental frequencies have been shown to sire more offspring than their competitors. Fundamental frequency of territorial songs, hence, may reveal male quality and, consequently, the resulting threat posed to competing males. We argue that this is reflected in the increased response of competitors to low-frequency territorial songs shown here. Such competitive signalling behaviour has been shown in a few mammal species like red deer and baboons but, thus far, not in bats. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Paternity covaries with laying and hatching order in the collared flycatcher Ficedula albicollis

Females in most bird species engage in extra-pair copulations. Although this behaviour is widespread, benefits for females of doing so are less understood. The most widely accepted hypothesis is that they improve their previous choice of social partner and gain genetic benefits for their offspring. Some evidence for this comes from studies that find that extra-pair young (EPY) have greater fitness than their half-sibs. However, this might be also caused by maternal, non-genetic effect, a possibility that remains largely untested. Here we test whether EPY are laid in larger eggs or eggs laid early in the laying sequence in the collared flycatcher (Ficedula albicollis). The size of eggs bearing EPY and within-pair young (WPY) did not differ, however, EPY were laid in early eggs and consequently hatched earlier than WPY. As hatching asynchrony is a strong determinant of offspring size and survival in many species, including collared flycatcher, our results suggest that a caution is needed when paternal genetic effects are to be inferred from comparison of naturally occurring half-sibs.     

Reduced nest defence intensity and improved breeding success in terns as responses to removal of non-native American mink

Lack of parental experience or differences in reproductive effort may lead to variation in nest defence behaviour among individuals in a prey population. In this experimental study, we analysed nest defence behaviour using a model of an American mink, Mustela vison, a non-native predator, at colonies of arctic terns, Sterna paradisaea, in two large areas where mink had been removed and two comparable control areas with mink in the south-western archipelago of Finland, Baltic Sea, in June 2000. Furthermore, we recorded breeding success of arctic terns in the same four areas during 1998–2001. Arctic terns took higher risks in nest defence in control areas and in a short-term (mink-free for 2 years) removal area than in the long-term (mink-free for 8 years) removal area. Thus, colonies with recent experience of mink were more active in defending their offspring. The breeding success of arctic terns was significantly higher in mink-removal areas than in control areas. We conclude that arctic terns modify their nest defence behaviour in the presence of mink. However, they cannot defend their nests sufficiently against this mainly nocturnal predator, since their breeding success is reduced in areas where mink are present.Communicated by J. Graves     

Body size,demography and foraging in a socially plastic sweat bee: a common garden experiment

Phenotypic plasticity may evolve when conditions vary temporally or spatially on a small enough scale. Plasticity is thought to play a central role in the early stages of evolutionary transitions, including major transitions such as those between non-sociality and sociality. The sweat bee Halictus rubicundus is of special interest in this respect, because it is socially plastic in the British Isles: Nests are social or non-social depending on the environment. However, sociality comprises a complex suite of inter-related traits. To further investigate social plasticity in H. rubicundus, we measured traits that are potentially integral to social phenotype at a northern site, where nests are non-social, and a southern site where nests can be social. We found that foundresses at non-social sites were smaller, produced offspring of a size more similar to themselves, initiated nesting later, and took longer to produce their first female offspring. They began provisioning earlier in the day, finished earlier, and collected more pollen loads. Common garden experiments suggested that these differences represent mainly plasticity, as expected for traits involved in the overall plastic social phenotype, with only limited evidence for fixed genetic differences in foraging. Conditions during overwintering did not have major effects on a foundress' subsequent behaviour.     

Provisioning behavior and the estimation of investment ratios in a solitary bee,Calliopsis (Hypomacrotera) persimilis (Cockerell) (Hymenoptera: Andrenidae)

Summary One aspect of behavioral ecology that has received considerable attention, especially by students of social insects, is the relative amount of energy invested by parents in the rearing of male versus female offspring. Sexual selection theory makes predictions about how individuals should allocate their total investment in the sexes. To test these predictions we must accurately quantify the relative cost incurred by a parent in the production of an average individual of either sex. Body weight ratios are the most common estimate of cost ratio, but the correspondence between offspring body weight and energetic investment on the part of the parent has rarely been determined. Calliopsis (Hypomacrotera) persimilis is a solitary, ground-nesting bee whose natural history makes it particularly convenient for studies of investment patterns and foraging behavior. Each day females construct and provision from 1 to 6 cells in linear, closely-spaced series. Each cell is provisioned with pollen from Physalis Wrightii flowers, which is collected on two or three foraging trips. However, the temporal sequence in which two- and three-trip foraging bouts occur is not random. Females invariably begin each day provisioning cells with three trips worth of pollen and usually switch to provisioning the latter cells of the day with just two trips worth of pollen. The sex of the offspring within the same co-linear series of cells also varies non-randomly — female offspring predominate in the first cells of each series and male offspring in the latter cells. The correspondence between the number of foraging trips to provision a cell, the total time spent foraging, and offspring sex was determined for 36 cells. The data indicate a close, though not absolute, relationship between the number of foraging trips and the sex of the offspring: males usually received two trips of pollen, though some received three, whereas female offspring invariably received three trips worth of pollen. A number of potential estimates of the relative cost of female and male offspring production were calculated. Estimates of the cost ratio based on the amount of time spent foraging, adult dry body weight, and pollen ball dry weight all give similar values. Female offspring receive an energetic investment of from 1.3 to 1.5 times that of males. These results support the use of adult dry body weight ratios in the estimation of cost ratios.     

Slave addition increases sexual production of the facultative slave-making ant Formica subnuda

Formica subnuda is a facultative slave-making ant, and colonies without slaves are often found. We studied the effect of slave workers on sexual production of F. subnuda by experimentally increasing the proportion of slaves. We added c. 4000 worker pupae of the ant F. podzolica to 15 F. subnuda colonies and kept 15 colonies as controls. The following year we excavated all colonies, counted the proportion of slaves, the total number of workers (colony size) and the number of sexual offspring. The proportion of slaves was significantly higher in the slave-added colonies than in the control colonies. The total production of sexual offspring increased 57% in the treatment colonies in comparison to the controls. When colony size was adjusted to the number of sexual offspring, the treatment colonies produced significantly more sexual offspring than the controls. Slave addition did not alter sex ratios. We suggest that two alternative mechanisms, not mutually exclusive, caused the increase of sexual production in F. subnuda colonies: (1) Most of the added pupae were consumed and stored as fat body in workers at the end of the experimental year; the following spring the excess fat was metabolized and fed to the developing sexual larvae, or (2) a proportion of the added pupae hatched to become slaves; the following spring these slaves foraged actively for protein-rich food for the developing sexual offspring. Received: 27 April 1995/Accepted after revision: 23 October 1995     

Fitness benefits of multiple mating versus female mate choice in the cellar spider (Pholcus phalangioides)

Recent studies have demonstrated that mating with multiple males can be beneficial for females and her offspring even if males contribute nothing but sperm. This was mainly established for species in which sperm from several males mix in the reproductive tract of the female, thus allowing sperm competition and/or female sperm choice. However, in species with last male sperm precedence, female re-mating decides against the previous male by strongly limiting his reproductive success. We tested the effect of female re-mating behaviour using the cellar spider Pholcus phalangioides, which shows strong last males sperm precedence and moderate levels of polyandry under natural situations. We predicted that females prevented from remating even though they are receptive would show reduced reproductive success compared to females that accept two copulations and females that reject a second male, since the latter two treatments were allowed to behave according to their decisions. However, if the number of matings per se had an effect on oviposition or on offspring performance, double-mated females should perform better compared to both treatments of once-mated females. We measured female fecundity and fertility over a period of 140 days, comparable to the species' natural reproductive peak season. Two thousand one hundred and fifty-two offspring from 67 first egg sacs were reared under two feeding levels. We registered development time and survival, and measured offspring adult size and mass. We found a positive effect of double mating, as in this treatment, oviposition probability was higher compared to the other treatments. Interestingly, adult female offspring of the DM treatment that were raised under low food level had a higher condition index compared to those from FS and RM, but development time, size and mass at adulthood were not affected by mating treatment. Female choice only seemed to affect hatching latency of the offspring. Overall, the main predictor of female reproductive output and success was female body size.     

Enhancement of reproductive success through mate choice in a social rock-wallaby, Petrogale assimilis (Macropodidae) as revealed by microsatellite markers

The reason why a female who is socially paired to one particular male seeks extra-pair copulations (EPCs) with others has important implications in life history models and to the study of behaviour. The Allied rock-wallaby, Petrogale assimilis, lives in spatially isolated colonies in tropical north Queensland, Australia. Extensive observations of a colony at Black Rock showed that intense behavioural bonding occurs between pairs of adult males and females; about two-thirds of males paired with one female, the remainder paired with two females simultaneously. Single-locus microsatellite profiling determined the paternity of 63 offspring from 21 females for which long-term behavioural data were available. One-third of the young were fathered by males which were not paired socially with the mother. The mating system was heterogeneous: (1) all offspring of 11 females were fathered by the mother's partner, (2) all young of 5 females were fathered by extra-pair males, and (3) only some of the young of 5 females were fathered by their regular consort. Analysis of individual longitudinal demographic records showed that females whose young were always fathered by their consort had higher reproductive success than those whose young were always fathered as a result of (EPCs). However, females with some offspring fathered by their regular consort and others via EPCs had the highest probability of raising young to independence. These females were significantly more likely to have an offspring fathered as a result of an EPC if their previous young had failed to survive to pouch emergence. These results are consistent with the hypothesis that females choose mates for their genetic quality. Comparison of the males with which these females sought EPCs and the regular consorts suggested that arm length rather than body weight or testes size was used as the index of genetic quality. Results from a second colony of rock-wallabies in which the reproductive rate was accelerated were also consistent with the genetic-quality hypothesis. These results imply that by choosing better-quality fathers irrespective of social pairing, females are able to maximise their overall lifetime reproductive success, and presumably, those of their offspring. Received: 8 June 1997 / Accepted after revision: 28 February 1998     

Extra-pair paternity and intraspecific brood parasitism in wild zebra finches Taeniopygia guttata,revealed by DNA fingerprinting

Summary The frequency of extra-pair parentage in a wild population of zebra finches Taeniopygia guttata was examined by DNA fingerprinting. A total of 25 families, comprising 16 pairs of parents and 92 offspring (in broods of 1 to 6) were examined. Ten cases of extra-pair parentage, presumed to constitute intraspecific brood parasitism, were detected (10.9% of offspring or 36% of broods), including one possible instance of quasi-parasitism (parasitism by a female fertilized by the male nest owner). The average number of parasitic eggs per clutch detected by fingerprinting was 1.10±0.32 SD, very similar to the one egg difference in average clutch size between parasitised (6.0±0.82) and unparasitised nests (5.0±0.95). Two cases of extra-pair paternity (EPP) were detected among 82 offspring whose maternity was confirmed: 2.4% of offspring, or 8% of broods. In both cases EPP accounted for only a single offspring within a brood. Behavioural observations show that EPP occurs through extra-pair copulation rather than rapid mate switching. The results are discussed in the light of what is known about the fertile period and sperm precedence patterns in this species. Offprint requests to: T.R. Birkhead     

Asymmetric sibling rivalry and nestling growth in red-winged blackbirds (Agelaius phoeniceus)

Many birds hatch their offspring asynchronously, and the adaptive significance of this trait, if any, is controversial. David Lack suggested long ago that by facilitating brood reduction when resources are scarce, hatching asynchrony provides relief from the effects of overcrowding. Some field workers interpret this to mean that the growth and survival of survivors should rise following partial brood loss. Here we show in a 6-year study of red-winged blackbirds (Agelaius phoeniceus) that the presence or absence of marginal offspring in experimentally manipulated broods had virtually no effect upon the growth of core offspring, whereas alterations of the size of core brood had strong and significant effects. Nestling growth was, not surprisingly, slower in broods with partial brood loss. Intriguingly, marginal offspring showed significantly greater variation in mass. Core offspring are less sensitive to, but not exempt from, the inimical effects of resource shortfall than are marginal offspring. The phenotypic handicap appears to marginal offspring a caste of high-variance progeny whose fitness prospects rest upon levels of parental input (stochastic resources) and the size of the core brood (stochastic development). Received: 21 June 1999 / Revised: 5 June 2000 / Accepted: 25 June 2000     

Biased primary sex ratio in the bushcricket Poecilimon veluchianus,an insect with sex chromosomes

Offspring sex ratio at hatching was examined in the bushcricket Poecilimon veluchianus. Offspring sex ratios varied significantly between females (Fig. 1). Low mortality prior to sex determination established that this heterogeneity was already present in the primary offspring sex ratio. Sperm age and female age had no influence on offspring sex ratio (Fig. 2). Male age at copulation, however, correlated significantly with offspring sex ratio (Fig. 3). There were two types of males: one type produced predominantly daughters when young and an increasing proportion of sons with age. The other type produced, independent of age, 1:1 offspring sex ratios (Fig. 4). The two types of males seem to occur in approximately equal numbers. Sex ratio variation (1) may adaptively compensate for local sex ratio biases caused by sex-specific motility, or (2) it may be adaptive if there is a sex-differential effect of laying date on offspring fitness. Received: 14 March 1996/Accepted after revision: 24 June 1996     

Killer whale predation on marine mammals at Punta Norte,Argentina; food sharing,provisioning and foraging strategy

Summary The social dynamics of killer whales (Orcinus orca) that hunt marine mammals are apparently highly flexible, though strong individual associations do exist. The killer whales at Punta Norte offer an unusually detailed view of association patterns and foraging behaviour, and suggest a pattern of behaviour that optimizes hunting efficiency with exception only to strong associations between some individuals and the provisioning and training of offspring. The main points from this paper are as follows: First, hunting effort was concentrated where the capture rate was greatest. All pods selectively attacked the prey type for which they had the highest capture rate. The amount of southern sea lion prey captured was approximately equal to the estimated minimum energetic requirement for killer whales based on weight. Secondly, one whale in each pod did the majority of the hunting, and then provisioned the others in the pod. It was clear on numerous occasions that food was shared. A review of reported incidences of killer wales taking marine mammal prey suggests that it is common for a subset of the individuals in a pod to hunt. These results are discussed in the context of the evolution of foraging behaviour.Offprint requests to: A.R. Hoelzel at the first address     

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).     

Antipredator responses of California ground squirrels to rattlesnakes and rattling sounds: the roles of sex,reproductive parity,and offspring age in assessment and decision-making rules

California ground squirrels (Spermophilus beecheyi) and northern Pacific rattlesnakes (Crotalus viridus oreganus) have an adversarial relationship. Adults are partially protected by venom resistance and harass rattlesnakes in part to defend their more vulnerable offspring. Larger, warmer snakes are more dangerous than smaller colder snakes, and in escalated conflict squirrels could benefit from risk assessment strategies. Rattlesnakes often rattle at harassing squirrels and rattling sounds produce cues related to body size and temperature. In study 1 we played back rattling sounds from snakes that varied in dangerousness and evaluated the roles of sex and parity in squirrel risk assessment strategies. In general, squirrels tail flagged and stood bipedally more, and were slower to reapproach the playback speaker following playbacks of rattling sounds from more dangerous snakes. In comparison with males and nonmothers, mothers were most responsive to rattling sounds and more sensitive to variation in snake dangerousness. Mothers tail flagged more than males and nonmothers, and this behavior tracked variation in snake dangerousness most closely, perhaps reflecting the effects of snake size and temperature on pup vulnerability. These findings suggest that many aspects of squirrel antisnake behavior are governed by their effects on descendant kin. In study 2 we tested the effects of offspring age on mothers responses to live rattlesnakes and rattling sounds. According to the offspring value hypothesis, mothers should take more risks in defense of older offspring because they are more likely to survive to reproductive age. By contrast, under the offspring vulnerability hypothesis, older offspring are less vulnerable to predators and thus mothers should take fewer risks. Risk-taking, as measured by behaviors that bring the squirrel close to the snakes strike range, was either unaffected by or negatively correlated with offspring age. Thus, our findings suggest that whereas offspring value is unimportant in squirrel antisnake behavior, offspring vulnerability may affect maternal defense. We suggest that offspring vulnerability in mammals, in comparison with birds, may play a larger role in parental defense against predators.Communicated by P.A. BednekoffAn erratum to this article can be found at