Communal suckling in the cavy Galea musteloides

We quantified the extent of communal suckling in the cavy Galea musteloides. Six groups of animals were held in large indoor enclosures and suckling behavior was recorded over 113 h of observation. The groups contained 2–6 lactating females and 3–14 sucking pups. Due to the relative synchronization of births, 73% of the pups present in each group during lactation were non-offspring. Each of the 22 lactating females in the six groups suckled non-offspring in addition to her own offspring. On average, females suckled 86% of non-offspring present in their groups. Thus, 98% of all pups (n = 47) received milk from non-mothers. Although suckling frequencies were significantly higher for mothers with their own individual offspring than with non-offspring individuals, females invested more total time suckling all non-offspring than did suckling just their own; this was possible because for each mother many more non-offspring than offspring pups were present during lactation. Suckling bouts were significantly longer for mothers with their own individual offspring than with non-offspring individuals. The proportion of non-offspring suckling of mothers correlated negatively with the proportion of own young among the pups of a group. Non-offspring suckling did not affect future reproduction of females. Our observations demonstrate extensive practice of communal suckling in G. musteloides under laboratory conditions. Probably because all mothers of a group participated more or less equally in communal suckling behavior, the obvious cost of giving energetically expensive milk to non-offspring did not result in reduced (future) reproductive success. Potential benefits directly involved with communal suckling are unclear. More indirectly, communal suckling as well as birth synchrony might contribute to the formation of advantageous multi-litter kindergardens.     

Evidence of a maternal foraging cycle resembling that of otariid seals in a small phocid,the harbor seal

Summary Lactation strategies in the two largest families of seals have been characterized as a phylogenetic dichotomy, with sea lions and fur seals (Otariidae) exhibiting foraging cycles and true seals (Phocidae) a strategy of fasting. We show that a lactating phocid, the harbor seal, Phoca vitulina, has a foraging cycle similar to that of otariids. Time-depth recorders attached to lactating harbor seal mothers revealed that 9 of 11 females began bouts of diving, averaging 12–40 m, by mid-lactation (12 days). During the remainder of lactation, females made an average of seven diving trips, lasting about 7 h. They returned to the rookery during the interval between successive bouts to nurse their pups. Diving was more frequent during daylight than at night and diving bouts increased in duration as lactation progressed. The diving behavior of females that had weaned their pups and previously collected data from stomach lavage, suggest that the bouts of diving represent successful foraging. We propose that the lactation strategy of the harbor seal is intermediate to that of the otariids and other phocids studied. The harbor seal has a foraging cycle like the otariids, but typically resembles other phocids in length of lactation, rate of mass gain in pups, and in milk fat content. As harbor seals are among the smallest phocids, and only slightly larger than most otariids, it seems likely that maternal size constrains the amount of stored energy harbor seal females can bring to the rookery, forcing them to start feeding during the lactation period.Correspondence to: D.J. Boness     

Pup guarding by greater spear-nosed bats

Alloparental care poses an evolutionary dilemma because effort is expended on non-filial offspring. Thus, instances of alloparental care have been attributed to either mistaken identity, (i.e., recognition errors) or active cooperation. In greater spear-nosed bats (Phyllostomus hastatus), reproductive females roost together in stable long-term social groups in cave ceilings. Non-volant pups frequently fall from roost sites to the cave floor where they can die unless retrieved by an adult. In this study, we examined the function of adult female visits to non-filial young and tested whether visits were attributable to recognition errors or to cooperation. We found that females visited non-filial pups from their own social group more than expected. Females from different social groups attacked and sometimes killed pups, and male pups were attacked more frequently than female pups. Visits by group mates benefited fallen pups by reducing the likelihood of attack by females from other groups. In contrast to the mistaken identity hypothesis, we found that some females leave their own pups to approach and remain with group mates’ pups. We used microsatellite markers to estimate relatedness and test whether kinship could explain this alloparental care and found that females were unrelated to the pups they guard. We conclude that females who reside in highly stable social groups exhibit cooperative behavior that cannot be explained by kinship and is unlikely to be due to direct or generalized reciprocity. Instead, our data suggest that alloparental care likely involves a complex interplay between group membership and cooperative foraging.     

Sexual dimorphism in sea lion pups: differential maternal investment,or sex-specific differences in energy allocation?

Proximal mechanisms underlying a faster growth rate in male compared to female California sea lion pups were investigated. Males are significantly larger at birth than females. Specifically, we asked if differential maternal investment contributed to enhanced male growth via: (1) larger mothers having disproportionately more male pups, (2) more time and energy put into foraging by mothers of male pups, and (3) greater milk production in mothers of male pups. We also considered four aspects of differential energy utilization and acquisition by male and female pups: (1) male pups attempting to save energy for growth by changes in behavior, (2) longer suckling bouts with mother and more sneak suckling of non-filial females by male pups, (3) lower maintenance costs in males via a lowered resting metabolic rate, and (4) increased assimilation efficiency in males. Our study showed that there are no differences in the size of females or length of foraging trips for mothers of male and female pups. Male pups received more milk from their mothers, but the difference was no longer significant when the larger body size of males was considered. There were no differences in either the activity budgets or suckling behavior of male and female pups. Male pups, however, did have lower resting metabolic rates than females. We conclude that enhanced male perinatal growth is a consequence of a larger size at birth, proportionally more milk from mothers to support the greater demands of larger body size, and lower maintenance costs due to a lower resting metabolic rate. Received: 28 April 1995/Accepted after revision: 25 July 1995     

Reproductive success in a low skew, communal breeding mammal: the banded mongoose, Mungos mungo

In most cooperatively breeding species, reproduction is monopolised by a subset of group members. However, in some species most or all individuals breed. The factors that affect reproductive success in such species are vital to understanding why multiple females breed. A key issue is whether or not the presence of other breeders is costly to an individual’s reproductive success. This study examines the factors that affect the post-parturition component of reproductive success in groups of communal-breeding banded mongoose (Mungos mungo), where up to ten females breed together. Per-litter reproductive success was low (only 18% of pups survived from birth to independence). Whilst singular breeding was wholly unsuccessful, there were costs associated with breeding in the presence of increasing numbers of other females and in large groups. Synchronisation of parturition increased litter success, probably because it minimises the opportunity for infanticide or decreases competitive asymmetry between pups born to different females. There was no evidence of inbreeding depression, and reproductive success was generally higher in litters where females only had access to related males within their group. I conclude that communal breeding in female banded mongooses represents a compromise between the benefits of group-living and communal pup care on the one hand, and competition between females to maximise their personal reproductive success on the other. Such conflicts are likely to occur in most communal breeding species. Whilst communal breeding systems are generally considered egalitarian, negative effects of co-breeders on individual reproductive success is still an issue.     

Inter-litter competition and communal nursing among Norway rats: advantages of birth synchrony

Summary The effect of inter-litter competition on pup survival was investigated in pairs of female rats (Rattus norvegicus) living and breeding in the same environment. If a female gave birth when a 0- to 14-day-old litter was already present in the environment, her pups had a very high chance of surviving, similar to the situation in which no other litter was present. Moreover, the mother was likely to nurse communally with the mother of the 0- to 14-day-old litter. This communal nursing benefitted the newborn pups as evidenced by their being heavier at weaning than litters that were not nursed communally. In contrast, if a female gave birth when a 15-to 28-day-old litter was already present in the environment, her newborn pups were likely to die within 3 days postpartum, owing to the fact that they were often prevented from suckling at their mother's teats, resulting in milk deprivation, and were often beneath the older pups, resulting in physical trauma. These findings suggest that inter-litter competition is an important source of pup mortality when litters are born 15-28 days apart. The data are discussed in terms of the advantages of birth synchrony.     

Communal nursing in prairie dogs

Summary Within a natural population of blacktailed prairie dogs (Cynomys ludovicianus), 68% of sampled juveniles received milk from foster mothers via communal nursing (Table 1). In proximate terms, communal nursing may result because prairie dog mothers seem unable to discriminate between their own and others' offspring. In ultimate terms, both indirect selection and reduced predation on juveniles resulting from the formation of multilitter groupings have probably been important in the evolution of communal nursing.     

Offspring sex ratio in relation to female size in southern elephant seals,Mirounga leonina

Southern elephant seals Mirounga leonina display extreme sexual dimorphism. In addition females show great variation in size and stored resources at parturition. Therefore they present an excellent opportunity for examination of responses of sex ratio to resource availability. We studied the relationships between the size of southern elephant seal females at parturition and the size and sex of their pups at South Georgia over four breeding seasons. We found a large individual variation in maternal post-partum mass (range 296–977 kg, n=151). Larger mothers gave birth to larger pups, irrespective of the sex of their pup. Male pups were on average 14% larger than females at birth and consequently more costly to bring to parturition. Our results suggest that female southern elephant seals must weigh more than 300 kg if they are to breed at all, and more than 380 kg if they are to give birth to a male pup. Above this threshold the proportion of males among offspring rapidly increases with maternal mass, and stabilizes at a level not significantly different from parity. These results show that smaller females of southern elephant seals vary offspring sex ratio in a way that is consistent with theories on adaptive offspring sex ratio. A smaller mother with a male foetus may benefit from terminating her pregnancy and allocating the resources she saves to her own growth. She could then give birth to and raise a larger pup in the subsequent season.     

The social organization of the common vampire bat

Summary At a site in Costa Rica, three groups of 8–12 adult female vampire bats, Desmodus rotundus, utilize group-specific sets of hollow trees as day roosts. Long-term nonrandom associations between pairs of females, as measured by the proportion of time one bat spends roosting in the same tree with another bat over a 3 year period, occur even when preferences for particular trees are removed. Significant associations exist between both related and unrelated adult females. Adult male bats, however, show few associations with females or other males. By observing bats within trees and while foraging, and by monitoring feeding flights with radiotelemetry, the following potential benefits of association could be tested. Females roost together to (1) share a suitable microclimate, (2) avoid predators, (3) avoid ectoparasite infestations, (4) minimize travel to mobile prey animals, (5) respond to coercive males, (6) feed simultaneously from a bite, (7) remove ectoparasites by allogrooming, and (8) share food by regurgitating blood to other bats within roosts. The data do not indicate that any of the first five hypotheses provide significant benefits for long-term associations although predators and ectoparasite levels may cause occasional changes in roost sites. Simultaneous feeding was uncommon and apparently confined to females and their recent offspring. Allogrooming, although common, occurred independently of the presence of ectoparasites. Food sharing, however, occurred between both related and unrelated adult females with high levels of association and provides at least one selective advantage for maintaining cohesive female groups.     

Facultative sex-ratio adjustment in Norway rats: litters born asynchronously are female biased

Summary Previously we reported that inter-litter competition reduces the survival of pups born to pairs of female rats living and breeding in the same nesting environment. Inter-litter competition occurred when females gave birth asynchronously; specifically, when a female gave birth in the presence of 15 to 28-day-old pups, her newborn pups were likely to die as a result of nest intrusion by the older pups. In contrast, inter-litter competition occurred rarely when the two females gave birth synchronously. Because theories of facultative sex ratio adjustment predict that mothers giving birth in unfavorable circumstances should bias their offspring towards the more viable or less expensive sex, we predicted that litters born asynchronously would be female biased. Conversely, we also predicted that females giving birth under favorable conditions, i.e., synchronously, would bias their litters toward males. We found a female bias in asynchronous litters, but did not find a male bias in synchronous litters. Moreover, in contrast to other reports in the literature, the female bias in asynchronous litters was achieved without a reduction in litter size. Based on correlational data, we suggest several mechanisms that could produce this female bias: conditions at fertilization and implantation, time since the male last mated and number of pups suckling concurrently during gestation. Correspondence to: M.K. McClintock     

Synergism between sib-rearing and sex ratio in Hymenoptera

Summary In many bees and wasps, solitary females produce offspring without help from other females. The transition from lone mothers producing offspring to situations in which females often help to rear siblings is an important step in the origins of complex sociality and nonreproductive castes. Recent work on Hymenoptera has stressed the role of sex ratio variation in this transition; when a mother's brood is more female biased than average, older daughters are favored to help rear their younger siblings because they are more closely related to sisters than to their own offspring. Here the direction of causality is from biased sex ratios, which arise by some extrinsic mechanism, to the origins of sib-rearing (eusociality). We present a model in which there is a synergism between sib-rearing and female-biased sex ratios, which may either complement the sex ratio variation idea by increasing the rate at which helping spreads or be an alternative hypothesis about the origins of eusociality. The synergism in our model depends on three conditions. 1) Daughters that help cause more food to be provisioned per offspring, which in turn causes larger offspring. 2) Females gain more than males by being large, which favors mothers with helpers to produce a higher proportion of daughters. 3) A helper's inclusive fitness rises as her mother's brood becomes increasingly female biased because a female helper is more closely related to her sisters than to her brothers. A female helper may also be more closely related to her sisters than to her own offspring, but this particular sibling-offspring relatedness asymmetry is not required by the synergism model. These three conditions create a synergism which favors a rapid transition from solitary (subsocial) to eusocial. Demographic and ecological factors that facilitate the evolution of eusociality reduce the stringency of the relatedness asymmetry condition (3) required by our idea. The synergism model therefore complements factors other than relatedness that may have been important during the evolution of eusociality.     

Weaning in the guinea pig (<Emphasis Type="Italic">Cavia aperea</Emphasis> f. <Emphasis Type="Italic">porcellus</Emphasis>): Who decides and by what measure?

Offspring should be selected to influence maternal effort to maximize their own fitness, whereas mothers are selected to limit investment in present progeny. In mammals, this leads to a conflict over the amount of milk provided and the timing of weaning. The intensity and time course of such conflict has so far mostly been investigated experimentally in altricial rodents. However, it is expected that offspring options for conflict will depend on developmental state. We therefore investigated in the highly precocial domestic guinea pig (Cavia aperea f. porcellus) who decides over nursing performance and weaning and how pup state influences these decisions. Specifically, we tested whether a threshold mass of pups predicts weaning time. By exchanging older litters against neonates and vice versa, we produced a situation in which females differed in lactational stage from the cross-fostered pups. Our results indicate that females decide about the timing of weaning, as cross-fostered younger pups were weaned at a much younger age than controls and older pups benefited from continuing lactation of foster mothers. Growth rates did not differ in the treatment groups, and different weaning ages resulted in differing weaning mass refuting the hypothesis that weaning is based on a threshold mass of offspring. This constitutes clear evidence that in a precocial rodent, the guinea pig, decisions about maternal care are primarily determined by maternal state and little influenced by pup state despite the extreme precociality of offspring. We suggest that precocial pups show little resistance to early weaning when food is abundant, as they reach sufficient nutritional independence by the middle of lactation to enable independent survival.     

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.     

The effect of reproductive condition on the foraging behavior of female hoary bats,Lasiurus cinereus

Summary Female mammals experience larg changes in time and energy budgets associated with reproduction and these may influence the foraging strategies of individuals. I studied the changes in foraging behavior associated with reproduction in female hoary bats, Lasiurus cinereus. As lactation progressed, individuals departed to forage earlier in the evening and spent more time foraging per night and less time roosting with their young. Foraging time increased by at least 73% between early lactation and fledging and then declined as the young became independent. Females with two young foraged for longer than did those with one and females with pre- and postfledging young foraged in different habitats. The changes in foraging time suggest that foraging activity of female L. cinereus is constrained and individuals act as time minimizers, adjusting their foraging behavior to meet current energy demand. Predation risk is unlikely to constrain the behavior of these bats. However, maximizing energy intake throughout lactation may not be the optimal strategy because storing excess energy increases flight cost and may reduce foraging efficiency. The need to keep newborn young warm may also influence foraging time. Such constraints, causing changes in foraging activity, may alter the availability of habitats and prey and must be considered when modelling foraging strategies. In addition, changes in flight time may significantly alter the energy budgets of bats in different stages of reproduction.     

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).     

Nursing of own and foster offspring by Utah prairie dogs (<Emphasis Type="Italic">Cynomys parvidens</Emphasis>)

From 1995 through 2005, I studied nursing among Utah prairie dogs (Cynomys parvidens) living under natural conditions at Bryce Canyon National Park, UT, USA. I observed 850 aboveground nursings, which involved 122 mothers and 248 juveniles from 134 litters. Most of the mothers that nursed aboveground were middle-aged, and most nursing juveniles had been coming aboveground for 1–3 weeks. Most nursings involved a single juvenile, lasted 1–10 min, and occurred between 1800 hours and 2000 hours. Seventy-five percent of nursings (598/796) involved a mother suckling her own juvenile offspring; the other 24.9% (198/796) involved a mother suckling another mother’s offspring (i.e., communal nursing). Communal nursings involved juveniles of the home territory, and many communal nursings (74/198 = 37.4%) involved close kin such as half-siblings, grandoffspring, full-nieces, and full-nephews; other communal nursings (37/198 = 18.7%) involved more distant kin such as full-second cousins and full-third cousins. Of seven hypotheses that might explain the evolution of communal nursing, evidence supports the importance of two: elevated inclusive fitness via indirect selection and communal nursing as a cost of coloniality.     

Female foraging and male vigilance in white-tailed ptarmigan (Lagopus leucurus): opportunism or behavioural coordination?

For several species of birds, high rates of male vigilance are correlated with high rates of female foraging. This relationship is thought to ultimately result in higher reproductive success for females paired with highly vigilant males. However, previous research has not examined the behavioural mechanism that produces the correlation between male vigilance and rates of female foraging. Foraging females may take advantage of vigilance that males are using for other purposes. Alternatively, the purpose of male vigilance may be to increase females' ability to forage. We examined these alternatives by testing whether vigilance preceded or followed bouts of female foraging more often than would occur by chance alone, using simultaneous behaviour observations of pre-incubation pairs of white-tailed ptarmigan (Lagopus leucurus). Our results indicate that each member of a pair may influence the behaviour of the other. Females were more likely to initiate foraging bouts after males became vigilant than if their mate remained non-vigilant. Moreover, non-vigilant males were more likely to become vigilant if their mate was foraging than if she was engaged in some other activity. Despite the possibility that a sexual conflict exists as each member of a pair attempts to maximize its fitness, both sexes behave as though a major role of male vigilance is to enhance female foraging opportunities. Received: 3 May 1999 / Received in revised form: 14 June 1999 / Accepted: 16 June 1999     

Low roost-site fidelity in pallid bats: associated factors and effect on group stability

Despite potential costs of changing roost or densites, many animals frequently move between roosts or dens. Pallid bats (Antrozous pallidus) change diurnal roost sites frequently and also reportedly have a variety of cooperative social behaviors, many of which are associated with the care of developing offspring. Roost switching is likely to increase the costs of maintaining the group stability expected with social cooperation. Pallid bats roosting in rock crevices in central Oregon were studied with radiotelemetry to (1) examine characteristics of day roosts, (2) determine what ecological factors were correlated with low roost fidelity, and (3) examine the temporal stability of roosting groups of pregnant and lactating bats. Pallid bats changed roosts an average of once every 1.4 days throughout the summer. The bats exhibited seasonal shifts in roost use, occupying roosts behind thin slabs of rock in cool weather and roosts in deep rock crevices in warm weather. Roost switching was not correlated with daily variations in weather conditions or with structural characteristics of the diurnal roosts, although switching may have allowed bats to maintain familiarity with several roosts that vary in microclimate. Roost switching was positively correlated with ectoparasite load. High ectoparasite levels were correlated with lower body weights in lactating females (Fig. 3), suggesting that parasites may be costly to the bats. Roost switching may be a strategy to decrease ectoparasite loads by interrupting the reproductive cycles of those parasites that spend at least part of their life cycle on the walls of the roost. Both pregnant and lactating pallid bats frequently changed their diurnal roost location, but lactating bats tended to travel shorter distances between consecutive roosts. Lactating bats were more likely to continue to associate with particular roostmates despite changes in the location of the diurnal roost (Fig. 4) and were less likely to roost alone. Although the stability of groups of lactating bats was not absolute, evidence supported the prediction that such groups are more cohesive than are those of pregnant bats. Received: 20 June 1995/Accepted after revision: 13 July 1996     

Reproductive success of solitary and communal Philoponella oweni (Araneae: Uloboridae)

Summary Philoponella oweni (Uloboridae) is a facultatively communal orb-web spider found in arid regions in the southwestern United States. Within one habitat solitary adult females and communal groups of adult females coexist.This study compared the reproductive output (egg production) and reproductive success (production of live offspring) of solitary and communal females. There was no significant difference in the mean number of egg-cases produced by solitary and communal females, but communal females produced significantly more eggs per egg-case. However, communal females suffered significantly higher rates of egg-case parasitism from the pteromalid wasp Arachnopteromalus dasys. As a result, the mean number of liver offspring produced by solitary and communal females did not differ. Thus, both strategies led to roughly equal reproductive success. The advantages of aggregation of adults may be significant in the evolution of social behavior in spiders.     

The functional significance of infanticide in a population of California ground squirrels (Spermophilus beecheyi)

Naturally-occurring infanticide was observed in a population of California ground squirrels (Spermophilus beecheyi). In four seasons, 40 infanticides were observed. All victims were post-emergent pups. Of 37 killings in which the killer was sexed 36 were by females. All infanticidal females were mothers at the time they killed, but in no case was a mother seen to kill or harm her own young. The victim was cannibalized in 22 cases and taken immediately into the killer’s burrow in 16 others. In no case did killers gain access to the victimized mother’s burrow or territory and female pups were not killed preferentially over males. In light of evolutionary explanations, infanticide in this population may best fit the resource exploitation hypothesis, in which killers commit infanticide to gain a nutritional benefit. Resource competition is a possible auxiliary explanation, since any time a female kills unrelated young she is eliminating possible competitors to her own offspring. This behavior could confer a fitness advantage on killers or it could be an adaptively neutral, alternative feeding strategy. More data are necessary to distinguish between these hypotheses. Received: 26 January 1995/Accepted after revision: 9 September 1995