The functional response of a hoarding seed predator to mast seeding

Mast seeding involves the episodic and synchronous production of large seed crops by perennial plants. The predator satiation hypothesis proposes that mast seeding maximizes seed escape because seed predators consume a decreasing proportion of available seeds with increasing seed production. However, the seed escape benefits of masting depend not only on whether predators are satiated at high levels of seed production, but also on the shape of their functional response (type II vs. type III), and the actual proportion of available seeds that they consume at different levels of seed production. North American red squirrels (Tamiasciurus hudsonicus) are the primary vertebrate predator of white spruce (Picea glauca) mast seed crops in many boreal regions because they hoard unopened cones in underground locations, preempting the normal sequence of cone opening, seed dispersal, and seed germination. We document the functional response of cone-hoarding by red squirrels across three non-mast years and one mast year by estimating the number of cones present in the territories of individual red squirrels and the proportion of these cones that they hoarded each autumn. Even though red squirrels are not constrained by the ingestive and on-body (fat reserves) energy reserve limitations experienced by animals that consume seeds directly, most squirrels hoarded < 10% of the cones present on their territories under mast conditions. Cone availability during non-mast years also reached levels that satiated the hoarding activity of red squirrels; however, this occurred only on the highest-quality territories. Squirrels switched to mushroom-hoarding when cone production was low and mushrooms were abundant. This resulted in type III functional response whereby the proportional harvest of cones was highest at levels of cone availability that were intermediate within non-mast years. Overall, more cones escaped squirrel cone-hoarding during a mast event than when cone production was low in non-mast years, which supports the predator satiation hypothesis. However, the highly variable seed escape in non-mast years may help to explain why all spruce cone production is not concentrated into fewer, larger, mast years.     

Rank-dependent hoarding effort in willow tits (<Emphasis Type="Italic">Parus montanus</Emphasis>): a test of theoretical predictions

Many birds and mammals store energy as hoarded food supplies. A supply of stored food is beneficial during periods when food is scarce, but building up and managing such a supply also entails costs. The optimal number of caches will be reached when the net benefit is at its maximum. If dominants can steal more stored food from subordinates than the other way around, the optimum will differ between these categories. A previous theoretical model of hoarding in groups with dominant and subordinate members produced three testable predictions: (1) hoarders should store more food as anticipated future conditions get worse; (2) subordinate flock members should store more food than dominants; and (3) dominants should increase hoarding relatively more than subordinates as conditions get worse. Here we present a field experiment on willow tits (Parus montanus) designed to test these predictions. We found support for all three. Hoarding increased as conditions got worse, subordinates stored at a higher rate than dominants, and dominants increased their hoarding effort relatively more than subordinates as conditions worsened. These results support the incorporation of information on dominance and food availability into models predicting food storage behaviour.Communciated by J. Dickinson     

The combined effects of seed perishability and seed size on hoarding decisions by Pére David’s rock squirrels

The food perishability hypothesis reasons that the perishability of nondormant acorns through rapid germination is the primary determinant of hoarding decisions (e.g., embryo removal in nondormant acorns in particular) by scatter-hoarding squirrels. However, we do not know whether seed size and its interactions with seed germination schedule affect squirrel’s hoarding decisions. By presenting pairs of acorns with contrasting germination/dormancy conditions and seed size, we investigated the relative importance of each target trait in determining the hoarding decisions of free-ranging Pére David’s rock squirrel (Sciurotamias davidianus) in Central China. Consistent with the food perishability hypothesis, the squirrels were highly sensitive to subtle differences of acorn germination status either within nondormant acorns or between nondormant and dormant acorns. Though there were no significant differences in seed hoarding and dispersal distance, the embryo-removal probability of nondormant acorns (especially those germinated) was much higher than that of dormant acorns prior to hoarding. Our results also support the seed size hypothesis. Large acorns were often hoarded more and moved farther than small acorns, and large nondormant acorns also had a higher probability of having their embryos removed. Moreover, the interactions between seed size and seed germination schedule had a large effect on whether a given acorn was hoarded or hoarded with its embryo removed. Our study indicates that the combined effects from seed germination schedule and seed size have the potential to determine hoarding decisions of scatter-hoarding squirrels.     

Interspecific competition in tree squirrels: do introduced grey squirrels (<Emphasis Type="Italic">Sciurus carolinensis</Emphasis>) deplete tree seeds hoarded by red squirrels (<Emphasis Type="Italic">S. vulgaris</Emphasis>)?

Red squirrels (Sciurus vulgaris) and introduced eastern grey squirrels (S. carolinensis) scatterhoard seeds of broadleaf trees. Scatterhoarded seeds are an essential resource in spring and their consumption increases red-squirrel fitness. We examined whether grey squirrels partly deplete the high-energy food resources cached by red squirrels, reducing their consumption, in two ways: (1) at the population level, comparing energy intake of feeding on cached seeds between a study site with red and grey squirrels and one with only red squirrels present; and (2) at the individual level, in the study site where species co-exist, relating hoard recovery of red squirrels to the amount of core-area overlap with grey squirrels. There were no significant site differences in the mean daily energy intake of red squirrels feeding on seeds recovered from caches. However, in the red-grey site, during spring, red squirrels that had a high percentage of their home-range core area overlapped by grey squirrels had a lower daily energy intake than low-overlap red squirrels. Body mass of red squirrels in spring was negatively correlated with the percentage of interspecific core-area overlap, but not with core-area overlap with other red squirrels. Our data suggest that interspecific competition for scatterhoarded seeds, with grey squirrels pilfering red squirrels' food caches, caused a reduced energy intake in red squirrels with a high degree of interspecific core-area overlap, and reduced body mass in spring. Therefore, cache pilfering is likely to reduce reproductive output in red squirrels, and thus play a role in the replacement of red by grey squirrels. Electronic Publication     

Interspecific competition between native Eurasian red squirrels and alien grey squirrels: does resource partitioning occur?

In heterogeneous environments, differential niche selection by two competing species will result in niche partitioning so that individuals of each species can maximise their fitness under different sets of environmental variables. Thus, niche partitioning is considered essential to allow co-existence of ecologically related species. To assess whether niche partitioning was occurring between native red squirrels and alien grey squirrels living together in a 13-ha high-quality mixed deciduous woodland in north Italy, we investigated temporal and spatial patterns in their activity and foraging behaviour between 1996 and 1998. We used live trapping and radio-tracking to study numbers, distribution and behaviour of squirrels. Daily and seasonal temporal activity patterns, and activity on the ground and in the trees, were similar in the two species. However, grey squirrels were more tree specialists and had a narrower tree-species niche width than red squirrels, in particular making greater use of oak. Other studies of red and grey squirrels in allopatry show that the two species differ in the extent they utilise oak. Overall, tree-species niche overlap was about 70%. Grey squirrels had larger home ranges than red squirrels. Home ranges and core areas of both species were larger in males than females. Also, intraspecific home range and core-area overlap patterns were similar to those found in allopatric populations of these species. Overall, there was no evidence that the use of space of one species was affected by the other. Our results show that there was no niche partitioning of activity or foraging behaviour in time or space during the study. This suggests that, at moderate grey-squirrel densities, red squirrels are unable to avoid competition with grey squirrels, and that competition for food and/or space will occur when these resources become limiting.     

Behavioral responses of territorial red squirrels to natural and experimental variation in population density

The relative scarcity of studies at the intersection of behavioral and population ecology is surprising given the presumed importance of behavior in density-dependent population regulation. Here we tested whether North American red squirrels (Tamiasciurus hudsonicus) adjust their behavior in response to local population density and whether they use rates of territorial vocalizations in their local neighborhood to assess density. We examined these relationships using 18?years of live trapping and 20?years of behavioral data that were collected across natural variation in local population density. To disentangle the effects of population density on behavior from those due to changes in per capita food abundance or changes in the frequency of antagonistic interactions with neighbors, we also experimentally manipulated population density with long-term food supplementation as well as perceived population density with long-term playbacks of territorial vocalizations. The frequency with which squirrels emitted territorial vocalizations was positively associated with local population density. In contrast, antagonistic physical interactions observed between squirrels and territorial intrusions were rare and the frequency of intrusions was weakly and negatively, not positively, associated with population density. Squirrels experiencing naturally and experimentally high density conditions spent less time in the nest and feeding but more time being vigilant. Similar density-dependent changes in behavior were observed in response to our manipulations of perceived population density, indicating that vocalization rates and not physical interactions or food abundance were the mechanism by which squirrels assessed and responded behaviorally to changes in local density.     

Seasonal variation in male-female competition,cooperation and selfish hoarding in a monogamous songbird

Both cooperation and conflict between the sexes are commonplace in monogamous mating systems. However, little is known about how cooperation and competition varies seasonally in monogamous species that maintain permanent territories. We presented territorial pairs of male and female New Zealand robins (Petroica australis) with a large supply of insect prey at monthly intervals for 2 years. Behavioural observations after food presentation were then made to quantify seasonal and sexual differences in aggressive interactions over prey, prey acquisition rates, mate provisioning, offspring provisioning, selfish food hoarding and cache retrieval. Data were used to evaluate sex-specific behavioural strategies of mediating competition for food. Results showed that males aggressively excluded females from experimental food sources year-round. Females only accessed food sources when males left them unattended. Consequently, females acquired fewer prey than males. After controlling for differences in prey acquisition, both sexes consumed similar amounts of prey in the non-breeding season. Even though males aggressively excluded females from accessing food sources directly, males fed large amounts of prey to females during the breeding season. Both sexes provisioned young at similar rates. Males cached less prey than females in the breeding season but more prey than females in the non-breeding season. Females showed similar caching intensities year-round. Although males tried to defend their hoards, females frequently retrieved male-made caches. Overall, results showed that although New Zealand robins cooperate to raise offspring during the breeding season, conflict between the sexes occurs year-round. Males and females display different behavioural strategies to gain access to experimental food sources, which appear to lessen male–female competition for food and evenly distribute food resources between the sexes.     

The association of female kin in the arctic ground squirrel Spermophilus parryii

Summary The association of kin in arctic ground squirrels (Spermophilus parryii) was studied near Haines Junction, Yukon Territory, during the summers of 1977, 1978, and 1979. Males dispersed in this population, thus only females were likely to live near relatives.Close female kin (sisters, mothers/daughters) had greater overlap of home ranges and interacted more amicably and less agonistically than did less closely related females. Closely related females clumped their young at emergence, whereas more distantly related females did not; there was little indication of clumping of pre-emerged young. The overlap of home ranges of distant relatives (known genetic relatives that had not associated in a natal burrow) was intermediate to that of close relatives and non-relatives. The types of interactions between distant relatives were more similar to those between non-relatives than between close relatives.I conclude that female arctic ground squirrels exhibit nepotism. Females may benefit from associations with relatives by sharing watching for predators once juveniles become conspicuous. Male arctic ground squirrels commit infanticide and several females may be more effective at protecting their young from infanticidal males than females living alone. I suggest that clumping of young by close relatives may provide a mechanism allowing distantly related females to identify each other.Address for offprint requests     

Seed predation and selection exerted by a seed predator influence subalpine tree densities

Strongly interacting species often have pronounced direct and indirect effects on other species. Here we focus of the effects of pine squirrels (Tamiasciurus spp.), which are a dominant pre-dispersal seed predator of many conifers including limber pines (Pinus flexilis) and whitebark pines (P. albicaulis). Pine squirrels depress seed abundance by harvesting most limber and whitebark pine cones on their territories. Pine squirrels further reduce seed availability for Clark's Nutcrackers (Nucifraga columbiana), the primary seed disperser of these pines, because selection exerted by pine squirrels has reduced the number of seeds per cone and causes seeds to be less accessible. We predicted that, if fewer seeds were available for dispersal by nutcrackers, pine recruitment should be suppressed in areas with pine squirrels. In support of this prediction, stand densities were about two times greater in areas where pine squirrels are absent than in areas where they are present. Alternative explanations that we considered do not account for these differences; however, precipitation may limit stand densities in the absence of seed limitation by pine squirrels. In sum, pine squirrels apparently depress limber and whitebark pine stand densities, with the potential for ecosystem impacts because these pines are foundation species within Western subalpine ecosystems.     

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     

Effects of relaxed predation pressure on visual predator recognition in the guppy

Many prey species have a genetic predisposition to recognise and respond to predators and can fine-tune their anti-predator behaviour following appropriate experience. Although the Trinidadian guppy ( Poecilia reticulata) has become a model species for the investigation of adaptive behaviour, the extent to which experience mediates predator recognition remains unclear. In this study, we examined the effects of relaxed predation pressure on patterns of anti-predator behaviour in populations differing in evolutionary history. The anti-predator behaviour of wild- and laboratory-born guppies from high- and low-predation localities in Trinidad were compared using three models resembling Crenicichla alta, a dangerous guppy predator, Aequidens pulcher, a less dangerous piscivore, and a snake. Snakes are not known to prey on guppies in Trinidad. Specifically, the following predictions were tested: (1) wild caught fish from the high-predation localities (where guppies co-occur with C. alta and A. pulcher) would respond to the three models according to their perceived level of threat, whereas guppies from the low-predation site would show a reduced response to all of the predator models; (2) high-predation laboratory-reared fish would display a reduced but qualitatively similar response to their wild counterparts; and (3) there would be no behavioural differences between wild- and laboratory-reared low-predation fish. In accordance with these predictions, the results revealed that wild fish originating from high-predation sites responded more strongly to the models than fish from low-predation sites. When reared in the laboratory, guppies from the high-predation population showed a reduced response compared to their wild-caught counterparts, but there was no difference in the behaviour of wild- and laboratory-reared low-predation fish. Model type affected predator inspection behaviour but not schooling tendency, and both wild- and laboratory-reared guppies were more wary of the fish models than the snake. These results suggest that early experience differentially mediates the anti-predator responses of fish from high-risk localities.     

Intensity of territorial defense in red squirrels: an experimental test of the asymmetric war of attrition

Summary Red squirrels defend exclusive, individual territories year round, 20% to 50% of females do not breed in any given year, and breeding females raise juveniles on their territories. Breeding is asynchronous, and the offspring of early-breeding females are more likely to hold an independently won territory than are late-born offspring. Based on the asymmetric war of attrition, we made the following predictions: (1) squirrels would respond more intensely to the calls of unfamiliar individuals than to the calls of neighbors; (2) breeding females would respond more intensely to unknown calls than would non-breeding females or males; (3) early-breeding females would respond more intensely than would late-breeding females to unknown calls; and (4) all classes of squirrels would respond similarly to the calls of neighbors. Playback experiments supported the predictions. Alternative hypotheses of kin selection, risk of infanticide, and seasonal difference in intruder pressure could not explain the results.     

Central place food caching: a field experiment with red-backed shrikes (Lanius collurio L.)

Summary Red-backed shrikes (Lanius collurio L.) stored experimentally presented mice (Mus musculus L) by reimpaling them on thorns of sloe bushes (Prunus spinosa L.) in the vicinity of the nest. Large mice (20 g) were stored further away from the nest than small mice (4 g), while the smallest mice (1 g) were transported directly to the nest and cut up there. Large prey required more round trips to deliver than smaller prey (4 g). Time to immobilize, load and deliver the prey and the proportion of undigestable tissue increased with prey size. Birds used stored prey as an alternative to hunting in other patches as expected from patch use models. The birds maintained a high rate of food delivery to the young by using stored mice during periods when their foraging success of natural prey was low. Several aspects of the shrike's food storing behaviour are in qualitative accord with suggestions derived from models assuming maximization of energy delivery rate.     

Interspecific associations of Cape ground squirrels with two mongoose species: benefit or cost?

Mixed-species associations have been described in many vertebrate species, but few behavioral studies have investigated associations between species from different mammalian orders. Cape ground squirrels (Xerus inauris) are highly social rodents that inhabit burrows with two species of mongoose, but the benefits of these interspecific associations to ground squirrels remain unresolved. We compared the behavior of squirrels while solitary, with conspecifics, and in the presence of suricates (Suricata suricatta) and yellow mongooses (Cynictis pencillatus). Squirrels spent less time alert and more time feeding when suricates were present, but increased vigilance in the presence of yellow mongooses. In a series of mobbing trials with a puff adder (Bitis arietans), a common predator of all three species, Cape ground squirrels were the most active in mobbing the snake. Our results suggest that Cape ground squirrels benefit from associating with suricates, but not necessarily with yellow mongooses. Both mongoose species benefit from the burrowing activities of the squirrels for thermoregulation and escape from predators, and a suite of other organisms may similarly benefit from the habitat modifications by Cape ground squirrels, suggesting they could be considered ecosystem engineers of the arid and semi-arid regions of southern Africa. Thus, the association between Cape ground squirrels and suricates appears mutually beneficial, whereas yellow mongooses may merely be commensals of the squirrels.     

Communal nesting in an ‘asocial’ mammal: social thermoregulation among spatially dispersed kin

Communal nesting can help defray the high cost of endothermic heat production in cold environments, but such social behavior is generally thought to be incompatible with the persistent defense of exclusive territories in typically ‘asocial’ animals. We examined the propensity for communal nesting in female red squirrels (Tamiasciurus hudsonicus), which maintain individual year-round territories, through intensive monitoring of litters over 22 years and by radio-tracking females during 3 years in late winter/early spring. Communal nesting was exceptionally rare during lactation: of 1,381 litters tracked to emergence, we observed a single instance in which two closely related (r?=?0.5) females pooled their litters into a single nest. In contrast, nest sharing between 2–3 females was relatively common in the late winter/early spring, prior to mating; at least 12 of 63 females (19 %) engaged in communal nesting during a year of systematic tracking of radio-collared females from late February to April. Communal nesting occurred more frequently when temperatures were colder, suggesting that such aggregations might function to reduce thermoregulatory costs. These social associations were typically, though not exclusively, between closely related individuals (r?≥?0.25 for seven of eight cases; mother–daughter dyads: four of eight), suggesting this cooperative behavior might evolve through kin selection and/or may reflect extended parental care. Our results demonstrate that female red squirrels engage in communal nesting, typically with closely related kin, despite a dispersed population structure that stems from the persistent defense of individual territories.     

Integrating archaeology and ancient DNA analysis to address invasive species colonization in the Gulf of Alaska

The intentional and unintentional movement of plants and animals by humans has transformed ecosystems and landscapes globally. Assessing when and how a species was introduced are central to managing these transformed landscapes, particularly in island environments. In the Gulf of Alaska, there is considerable interest in the history of mammal introductions and rehabilitating Gulf of Alaska island environments by eradicating mammals classified as invasive species. The Arctic ground squirrel (Urocitellus parryii) is of concern because it affects vegetation and seabirds on Gulf of Alaska islands. This animal is assumed to have been introduced by historic settlers; however, ground squirrel remains in the prehistoric archaeological record of Chirikof Island, Alaska, challenge this timeline and suggest they colonized the islands long ago. We used 3 lines of evidence to address this problem: direct radiocarbon dating of archaeological squirrel remains; evidence of prehistoric human use of squirrels; and ancient DNA analysis of dated squirrel remains. Chirikof squirrels dated to at least 2000 years ago, and cut marks on squirrel bones suggested prehistoric use by people. Ancient squirrels also shared a mitochondrial haplotype with modern Chirikof squirrels. These results suggest that squirrels have been on Chirikof longer than previously assumed and that the current population of squirrels is closely related to the ancient population. Thus, it appears ground squirrels are not a recent, human‐mediated introduction and may have colonized the island via a natural dispersal event or an ancient human translocation.     

Is the antipredatory response in behaviour reflected in stress measured in faecal corticosteroids in a small rodent?

Predation risk has been shown to alter various behaviours in prey. Risk alters activity, habitat use and foraging, and weight decrease might be a consequence of that. In mammals, studies on physiological measures affected by risk of predation, other than weight, are rare. We studied in two separate laboratory experiments foraging, hoarding behaviour and expression of stress measured non-invasively from the faeces in the bank vole (Clethrionomys glareolus), a common boreal rodent. Voles were exposed to predation risk using odours of the least weasels (Mustela nivalis nivalis). Distilled water served as control. In the first experiment, we found that foraging effort, measured as sunflower seeds taken from seed trays filled with sand, was significantly lower in trays scented with weasel odour. Both immediate consumption of seeds and hoarding were affected negatively by the weasel odour. Females hoarded significantly more than males in autumn. In the second experiment, the negative effect of weasel odour on foraging was consistent over a 3-day experiment, but the strongest effect was observed in the first night. Foraging increased over the time of the experiment, which might reflect either energetic compensation during a longer period of risk, predicted in the predation risk allocation hypothesis, or habituation to the odour-simulated risk. Despite decreased foraging under predation risk, stress measured as corticosteroid metabolite concentration in vole faeces was not affected by the weasel odour treatment. In conclusion, we were able to verify predation-risk-mediated changes in the foraging effort of bank voles but no physiological stress response was measured non-invasively, probably due to great individual variation in secretion of stress hormones.     

Strategies of homing behavior in the red squirrel,Tamiasciurus hudsonicus

Summary Seventeen red squirrels (Tamiasciurus hudsonicus) were displaced and released about 600 or 2000 m from their home site, and the routes they followed during several hours after release were recorded. The analysis of these routes shows that, in their first attempts to leave the release site area, the squirrels' orientation is non-random with respect to the home direction (Fig. 2a). However, after a few hundred meters in this initial direction, the animals tend to go back to the release site area, and then to perform a series of radial forays centered on the latter.The results suggest that, in the conditions of the experiment, the squirrels have some information on the direction, but not on the distance to the home site. Their behavior appears to be governed by two complementary (but occasionally conflicting) strategies, one being to use the directional information available, the other not to proceed further than a distance which corresponds to the distance normally covered in spontaneous exploratory trips. It is argued that this double strategy is appropriate for spontaneous homing trips consecutive to an exploratory foray.     

The effects of colony-level selection on the social organization of honey bee (Apis mellifera L.) colonies: colony-level components of pollen hoarding

Two-way selection for quantities of stored pollen resulted in the production of high and low pollen hoarding strains of honey bees (Apis mellifera L.). Strains differed in areas of stored pollen after a single generation of selection and, by the third generation, the high strain colonies stored an average 6 times more pollen than low strain colonies. Colony-level organizational components that potentially affect pollen stores were identified that varied genetically within and between these strains. Changes occurred in several of these components, in addition to changes in the selected trait. High strain colonies had a significantly higher proportion of foragers returning with loads of pollen, however, high and low strain colonies had equal total numbers of foragers Colony rates of intake of pollen and nectar were not independent. Selection resulted in an increase in the number of pollen collectors and a decrease in the number of nectar collectors in high strain colonies, while the reciprocal relationship occurred in the low strain. High and low strain colonies also demonstrated different diurnal foraging patterns as measured by the changing proportions of returning pollen foragers. High strain colonies of generation 3 contained significantly less brood than did low strain colonies, a consequence of a constraint on colony growth resulting from a fixed nest volume and large quantities of stored pollen. These components represent selectable colony-level traits on which natural selection can act and shape the social organization of honey bee coloniesCommunicated by R.F.A. Moritz     

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