Infanticide by male house sparrows: gaining time or manipulating females?

The killing of genetically unrelated young by males has been viewed as a strategy that forces victimized females to advance the onset of their next fertile period, thus infanticidal males gain a time advantage that may be crucial to maximize reproductive success. Among females that may raise several broods in a year, a failure occurring relatively earlier in the time-course of the previous breeding attempt may result in an increased investment in the next breeding attempt. This female strategy may be exploited by males in their own interest, and may strongly select for male infanticidal behaviour. I demonstrate that, in the house sparrow, females mated with infanticidal males re-laid earlier, initiated more breeding attempts and fledged more offspring than females mated with non-infanticidal males. These results suggest that both the time saving and the manipulation of female investment are independent mechanisms conferring advantages that may have selected for male infanticide in the studied population.     

Some aspects of infanticide and intermale competition among langurs,Presbytis entellus,at Dharwar,India

Conspecific infanticides by male langurs depresses population growth in their densely populated habitats. An infanticidal model developed from demographic parameters obtained in the Dharwar study area predicts an almost stable population. On the other hand, a non-infanticidal model predicts a population growth by 2.6% per year. The effect of frequent troop usurpation and infanticide on the control of population growth must be strong if the natality rate is high. For subadult and juvenile surplus males, it must be difficult to survive and to mature in the all-male party in its poor habitat. However, from calculations for living adult males and the number of troops at Dharwar, most adult males are thought to be able to obtain a troop within five years of first challenge of usurpation. Dominant males do not always take over a troop containing more females than do subordinate males. These simple assessments require further intensive field studies to determine the precise differences between infanticidal and non-infanticidal populations and whether or not dominant males make a greater genetic contribution than subordinate males to subsequent generations.     

Sport Hunting,Predator Control and Conservation of Large Carnivores

Sport hunting has provided important economic incentives for conserving large predators since the early 1970''s, but wildlife managers also face substantial pressure to reduce depredation. Sport hunting is an inherently risky strategy for controlling predators as carnivore populations are difficult to monitor and some species show a propensity for infanticide that is exacerbated by removing adult males. Simulation models predict population declines from even moderate levels of hunting in infanticidal species, and harvest data suggest that African countries and U.S. states with the highest intensity of sport hunting have shown the steepest population declines in African lions and cougars over the past 25 yrs. Similar effects in African leopards may have been masked by mesopredator release owing to declines in sympatric lion populations, whereas there is no evidence of overhunting in non-infanticidal populations of American black bears. Effective conservation of these animals will require new harvest strategies and improved monitoring to counter demands for predator control by livestock producers and local communities.     

Infanticide among free-ranging langurs (Presbytis entellus) at jodhpur (Rajasthan/India): Recent observations and a reconsideration of hypotheses

Considerable disagreement characterizes the debate concerning frequency, causation, and function of infanticide in connection with adult male replacements in bisexual one-male troops of hanuman langurs (Presbytis entellus). Detailed observations are presented about two noninfanticidal and three infanticidal male changes including six eye-witness and five presumed cases of infanticide within three langur troops during a long-term study at Jodhpur, Rajasthan, India. The results do not support any explanatory hypotheses focussing on social crowding, regulation of population density, social stress, sexual frustration, incest avoidance, or social bonding, but are in general though not total agreement with the reproductive advantage hypothesis: mainly unrelated infants were killed (one possible exception), the infanticidal male generally sired the subsequent offspring (one exception), and the mean interbirth interval subsequent to infanticide is by 2.1 months shortened. Likewise, several cases of stress induced abortions occurred. It is demonstrated that postconception estrous behaviour is by no means a female counterstrategy to infanticide in order to confuse males concerning the issue of paternity, since an infanticidal male did not spare the subsequent offspring of mothers who copulated with him during pregnancy and pregnant females did not discriminate between fathers and non-fathers.     

Infanticide as an alternative male reproductive strategy in langurs: A mathematical model

This paper makes explicit some of the assumptions underlying the view that infanticide is one of two (or more) alternative reproductive strategies utilized by adult male langur monkeys (genus Presbytis). A mathematical model has been developed from these assumptions, and formulae derived which give the equilibrium proportion of infanticidal males expected in langur populations under any given set of reproductive and demographic conditions. Together with estimates of adult male reproductive success obtained from a previous analysis of langur infanticide, these formulae were then used to calculate the precise proportion of infanticidal males expected in natural populations of langurs characterized by specific average male tenures. Further, the number of generations required for such populations to reach their predicted equilibrium was estimated using a simple computer model of langur population dynamics. The present work has thus produced several quantitative predictions which are directly falsifiable with observational data obtainable from wild populations of langur monkeys. With only slight modification, the present model may also be applied to the numerous other primate and non-primate species whose mating systems include many of the same general features as that of langurs.     

Infanticide and effectiveness of pup protection in bank voles: does the mother recognise a killer?

Infanticide, the killing of conspecific young, has been documented in numerous species of mammals, especially rodents. In that infanticide is costly to the victim mother, natural selection should favour counter-strategies by females to protect their pups. We studied the frequency of infanticide by male and female bank voles (Clethrionomys glareolus) and the effectiveness of dams in recognising and deterring infanticide. In trials in which unprotected pups were exposed to voles of both sexes, one third of male and female intruders killed pups. When mothers were present at the nest site, not a single female and only 2 of 25 males were able to commit infanticide. Females acted aggressively towards all intruders and hence did not discern between infanticidal and non-infanticidal males and females. Aggression of dams against any intruder indicates that all strange individuals near the nest site of a territorial species form an equal threat to pups. However, the presence of the aggressive dam is sufficient to deter most intruders from harming offspring by keeping them at a distance from the nest site. Heat run before mating, in a species with postpartum oestrus and vulnerable pups in the nest, also might lure potential infanticidal males away from the nest site and obscure the nest location. Received in revised form: 17 October 2001 Electronic Publication     

New cases of inter-community infanticide by male chimpanzees at Ngogo,Kibale National Park,Uganda

Infanticide by males has been recorded in four chimpanzee populations, including that in Kibale National Park, Uganda. Some infanticidal attacks occur during inter-community aggression. The sexual selection hypothesis does not easily explain these attacks because they may not directly increase male mating opportunities. However, females in the attackers’ community may benefit by expanding their foraging ranges and thereby improving their reproductive success; thus infanticide may increase male reproductive success indirectly. We report two new cases of infanticide by male chimpanzees at Ngogo, Kibale National Park. Like two previous cases, these occurred during a boundary patrol and were almost certainly between-community infanticides. The patrolling males attacked despite the proximity of males from the victims’ presumed community. This probably explains why, unlike the earlier cases, they did not completely cannibalize their victims. Such attacks seem to be relatively common at Ngogo and infanticide may be an important source of infant mortality in neighboring communities. Our observations cannot resolve questions about the sexual selection hypothesis. However, they are consistent with the range expansion hypothesis: the infanticides occurred during a period of frequent encounters between communities associated with a mast fruiting event, and Ngogo community members greatly increased their use of areas near the attack site during another mast fruiting event one year later. Our observations contribute to growing evidence that lethal intergroup aggression is a common characteristic of wild chimpanzee populations.     

Infanticide in the evolution of reproductive synchrony: effects on reproductive success

Synchronous breeding in animals and plants has stimulated both a theoretical and empirical examination of the possible benefits of active synchronization. The selective pressures of predation and infanticide are the strongest candidates proposed to explain the evolution of reproductive synchrony. Alternatively, breeding asynchronously with conspecifics may ensure a greater availability of resources per breeder. However, the possible fitness benefits resulting from active asynchronization have not yet received attention in evolutionary ecology. Here we present a hypothesis, based on a graphical model, illustrating the costs and benefits of the two modes of reproduction. We tested the hypothesis empirically using a 2 x 2 full factorial study design, where reproductive synchrony and infanticide tactics were manipulated in enclosed populations of the bank vole. The results reveal a relationship between infanticide tactics and breeding synchrony as illustrated by our hypothesis. In general, female reproductive success (number and size of offspring surviving to weaning) was significantly lower in infanticidal populations. Moreover, an asynchronous breeding pattern proved to be advantageous in the noninfanticidal population but this advantage of asynchrony was lost as infanticide became common in the population. Our findings support the idea that synchronous reproduction could have evolved as a counterstrategy against infanticide.     

Is Mating Alone Enough to Inhibit Infanticide in Male Bank Voles?

Infanticide, the killing of conspecific young, is commonly recognized as an adaptive behavioural strategy enhancing the fitness of the perpetrator. Infanticide is supposed to be inhibited in several male rodent species after mating with a time lag to the time when perpetrators own offspring would be born. This is because males with no parental care do not recognize their own offspring. It has been suggested that copulation alone is enough to inhibit infanticidal behaviour in male rodents. Infanticidal behaviour occurs in more than 50% of male bank voles (Myodes glareolus), and offspring loss because of infanticide may have a great effect on breeding success and population recruitment. In a laboratory experiment, we studied whether infanticidal male bank voles after successful mating stop the killing of pups. Infanticidal males were paired with a female until successful copulation. After the young were born, the males’ infanticidal behaviour was studied from the time of expected birth of own pups until their post‐weaning age. We predicted that mated infanticidal males are inhibited from committing infanticide especially during the time period when pups are less than 10 d old. Against our prediction, 67% of the infanticidal males continued the killing of pups in the age of 3 d. Infanticidal behaviour remained stable, and half of the males were infanticidal still at the age of weaning of pups. Our results are contradictory to previous studies, as we observed no inhibition of infanticide during early life of pups nor increase in infanticide again when their own offspring would have reached the ‘safe’ age and size after weaning. We suggest that mating alone is not sufficient to inhibit infanticide. Thus, we suggest that other cues of the female with whom the male mated with or on her territory are needed for inhibition to occur.     

Infanticide and social flexibility in the genus <Emphasis Type="Italic">Gorilla</Emphasis>

Based on the cases of infanticide by male mountain gorillas reported from the Virunga volcanic region, the socioecological and life history features of gorillas satisfy the conditions for which infanticide may be expected. However, there are considerable variations in the occurrence of infanticide between habitats. We analyze the recent reports of infanticides that were directly observed or are suspected based on field evidence in two populations of eastern and western lowland gorillas (Kahuzi and Mbeli Bai, respectively) along with previous reports on mountain gorillas, and consider which social features are linked with and which factors influence the occurrence of infanticide in the gorilla populations. All victims were suckling infants and most of them were killed by males who seemed unrelated to them. Dependent infants are most vulnerable to infanticide when the protector male (its putative father in most cases) is absent, and so male protection ability seems to be important in determining female transfer decisions. Two cases observed in Kahuzi suggest that the infanticidal male may discriminate between infants to accept and those to kill according to his previous interactions with their mothers. Mating for a prolonged period prior to parturition is necessary for immigrant females to avoid infanticide by the new male of the group that they join. Infanticide was usually associated with female transfer, and the patterns of female association at transfer may shape variations in social structure between populations. Female mountain gorillas prefer large groups with multiple males and tend to transfer alone in order to seek more protection against infanticide in Virunga. By contrast, female eastern and western lowland gorillas tend to transfer with other females to small groups or solitary males, and maturing silverbacks take females to establish new groups through group fission in Kahuzi and Mbeli Bai. These differences may result in more multi-male and larger groups in the Virungas than in Kahuzi and Mbeli Bai. Rapid changes in density of gorilla social units and their relations following drastic environmental changes caused by recent human disturbances may also increase the probability of infanticide.     

Infanticide in <Emphasis Type="Italic">Cercopithecus mitis stuhlmanni</Emphasis> in the Kakamega Forest,Kenya: Variation in the Occurrence of an Adaptive Behavior

Infanticide by males is widespread across mammals and especially prevalent among primates. Considerable research has examined how fitness benefits can explain the occurrence of this behavior; less is known, however, about intrapopulation variation in its occurrence. We evaluated 10 infanticides by males in wild blue monkeys according to the sexual selection hypothesis. To explore intrapopulation variation in occurrence of infanticide, we compared these cases to 38 cases that were contextually similar but in which infanticide did not occur. We examined male reproductive benefit, infant age, maternal parity, postconception estrus, group defense, available mating partners, and context of takeover. We based comparisons on daily or near daily records of male presence in the study groups, infant birth dates, and male-female sexual interactions. Infanticides followed predictions of the sexual selection hypothesis: males were unlikely to kill their own offspring, the period for the mother’s return to conception was reduced by half, and males increased their chance of siring her next offspring. Difference in male reproductive benefit, costs, and motivation did not fully explain the observed variation in infanticide occurrence. Infants were more likely to be spared if they were older when a male first arrived, or if their mother had mated with the male in the second month after conception. The most important determinant of infant fate, however, was male identity, a finding consistent with 2 scenarios: 1) an infanticidal tendency may be influenced by a genetic polymorphism that is not fixed in this population or 2) infanticidal behavior may be a conditional male strategy. Further research on intrapopulation variation in infanticidal behavior should focus especially on characteristics of males.     

Infanticide and Maternal Offspring Defence in European Rabbits under Natural Breeding Conditions

Infanticide is an important source of mortality of dependent offspring in several mammal species, whereas female conspecifics are often the perpetrators. This has led to maternal counter‐strategies, such as the defence of the nests. However, cases of infanticide are hard to detect in the field, and studies on maternal offspring defence behaviour under natural breeding conditions are scarce. We conducted such a study on the European rabbit (Oryctolagus cuniculus), which is usually considered to show low maternal care. The study was carried out over 5 yr on a field enclosure population. We (1) studied infanticide rates and the impact of potential determinants: the group density and age structure of the females’ rank hierarchy within the groups; we used the latter as an estimator of social group stability. We (2) studied if mothers defend their breeding burrow against approaching, potentially infanticidal females. Overall, we recorded infanticide in 5% of all litters; infanticide was the cause in 12% of cases of litter mortality. The proportion of infanticide was 7% higher in groups where same‐age females occupied successive rank positions than in groups where the females’ rank hierarchy had a more heterogeneous and linear age structure. We hypothesize that social instability in the former groups was the reason for the increased infanticide risk. Infanticide rates were not correlated with group density and did not differ among mothers with different social ranks. Infanticide occurred exclusively during the first 10 d after parturition. During this time, mothers stayed closer to their breeding burrows than shortly before parturition or during later lactation. Moreover, mothers were more aggressive against other females in proximity to their breeding burrow than in more distant areas. We suggest that the pattern of spacing behaviour and intrasexual aggression of rabbit mothers are an adaptive response to the occurrence of female infanticidal behaviour.     

Infanticide in wedge-capped capuchin monkeys, Cebus olivaceus

Three cases of infanticide are reported from a well-studied population of wedge-capped capuchin monkeys, Cebus olivaceus, located in the Venezuelan llanos. For each case, we examined the social context and the consequences to the reproductive success of the infanticidal male. In 1 case, the male probably killed his own son. In 2 cases, the male killed an unrelated animal. In all 3 cases, the female conceived in the following breeding season, but in only 1 case did the infanticidal male possibly enhance his own reproductive success. These data are interpreted in the light of current hypotheses for infanticide.     

Infanticide in a chacma baboon troop

This report describes an infanticide and two attacks of an infant and a juvenile by a natal adult male in a troop of chacma baboons (Papio ursinus). The infanticidal male had become the dominant male in his troop five months before the infanticide suggesting that a trigger for infanticidal behavior is a rise in rank to dominant status.     

Presence of a conspecific increases superparasitism but not infanticide under self‐ and conspecific superparasitism in a semisolitary parasitoid,Echthrodelphax fairchildii (Hymenoptera: Dryinidae)

Female parasitoids are expected to avoid superparasitism (ovipositing in and/or on parasitized hosts) when unparasitized hosts are available. However, when the supply of unparasitized hosts is restricted, they are expected to self‐ as well as conspecifically superparasitize. One of the cues of a reduced availability of unparasitized hosts is the presence of a conspecific. Moreover, if the focal species can perform infanticide, after encountering a conspecific female, the females are expected to kill eggs existing in and/or on hosts when superparasitizing, because the eggs are more likely to be laid by others. In this study we investigated whether females of an infanticidal semisolitary parasitoid, Echthrodelphax fairchildii, increase their frequencies of superparasitism and infanticide after encountering a conspecific female. Echthrodelphax fairchildii females are capable of discriminating between self‐ and conspecific superparasitism until up to 0.75 h after the first egg was laid (self‐superparasitism frequency < conspecific superparasitism frequency). As expected, the female parasitoids were more likely to perform self‐ and conspecific superparasitism after they had encountered a conspecific. In particular, the self‐superparasitism frequency increased highly within a short period after the first oviposition, so that no difference between the self‐ and conspecific superparasitism frequencies was found. In contrast, the infanticidal‐probing frequency remained extremely low, irrespective of whether or not the female parasitoids had encountered a conspecific. Moreover, when superparasitizing, females usually laid female eggs. Possible causes for the low frequency of infanticidal probing and the female‐biased sex ratio are discussed.     

Propagation of electromagnetic radiation in mitochondria?

Infanticide by primate males was considered rare if groups contain more than one adult male because, owing to lower paternity certainty, a male should be less likely to benefit from infanticide. Guided by recent evidence for strong variation of infanticide in primate multi-male groups, we modelled the conditions for when infanticide should occur for a group with a resident and an immigrant male. Setting the parameters (e.g. infant mortality, reduction of interbirth interval, life-time reproductive success, genetic representation) to fit the conditions most commonly found in nature, we develop a game-theoretic model to explore the influence of age and dominance on the occurrence of infanticide and infant defence. Male age strongly impacts the likelihood of an attack which is modified by the father's defence. If the new male is dominant he is likely to attack under most circumstances whereas a subordinate male will only attack if the father does not defend. These model scenarios fit the conditions under which infanticide is known to occur in primate multi-male groups and offer an explanation why infanticide is common in some multi-male groups and rare in others. Overall, the benefits for infanticidal males are strongly governed by a reduced interbirth interval while advantages via improved genetic representation in the gene pool contribute but a minor fraction.     

DNA analyses support the hypothesis that infanticide is adaptive in langur monkeys.

Although the killing of dependent infants by adult males is a widespread phenomenon among primates, its causes and consequences still remain hotly debated. According to the sexual selection hypothesis, infanticidal males will gain a reproductive advantage provided that only unrelated infants are killed and that the males increase their chances of siring the next infants. Alternatively, the social pathology hypothesis interprets infanticide as a result of crowded living conditions and, thus, as not providing any advantage. Based on DNA analyses of wild Hanuman langurs (Presbytis entellus) we present the first evidence that male attackers were not related to their infant victims. Furthermore, in all cases the presumed killers were the likely fathers of the subsequent infants. Our data, therefore, strongly support the sexual selection hypothesis interpreting infanticide as an evolved, adaptive male reproductive tactic.     

Mating Strategies in Relation to Sexually Selected Infanticide in a Non-Social Carnivore: the Brown Bear

Based on the sexually selected infanticide (SSI) hypothesis, infanticide can be an adaptive mating strategy for males, but this is has rarely been documented in non‐social mammals. This phenomenon should not benefit females, so one would expect females to evolve mating counter strategies in order to protect their infants from infanticidal males. Cases of SSI are extremely difficult to document in the field, especially for non‐social species. Using field observations and genetic methods, we describe mating strategies employed by both sexes of brown bears (Ursus arctos) in relation to SSI. We present evidence for the first time suggesting that infanticide is an adaptive male mating strategy in this non‐social carnivore, as all requirements for SSI are fulfilled (1) infanticide shortens the time to the mother's next estrus, (2) the perpetrator is not the father of the killed infants, and (3) putative perpetrators sire the next litter. Moreover, all infanticide cases occurred during the mating season. We expected that primarily immigrant males were infanticidal, as in social species. However, we found that resident adult males commonly committed infanticide. Perhaps they recognize females they have mated with previously. Moreover, we used DNA‐based parentage testing to demonstrate a minimum of 14.5% of multiple paternities (up to 28% for litters with at least three young). Female promiscuity to confuse paternity may be an adaptive counter strategy to avoid infanticide.     

First direct evidence of infanticide and cannibalism in wild snub-nosed monkeys (Rhinopithecus bieti)

Although infanticide has been witnessed in many species of Colobinae, and a case was observed in a captive group of golden snub-nosed monkeys (Rhinopithecus roxellana), observed cases of infanticide in wild snub-nosed monkeys (Rhinopithecus spp.) have not previously been recorded. Here we describe the killing of a 1-month-old infant by a male in a black-and-white snub-nosed monkey (R. bieti) group at Xiaochangdu in Tibet. The infanticidal attack was witnessed as part of a long-term observational study of ecology and behavior that began in June 2003. The male was observed killing and eating the infant. The literature proposes three main explanations for infanticide: two adaptive hypotheses (sexual selection and resource competition), and one nonadaptive hypothesis (social pathology). Individual cases of infanticide, such as this one, are important for comparative purposes, but when examined on their own they are difficult to interpret in relation to established theoretical frameworks. The cases we describe here show some consistency with the sexual selection hypothesis, but the lack of critical information (i.e., as to paternity) makes it impossible to draw a firm conclusion. This is also the first described case of cannibalism in snub-nosed monkeys.     

The variable social organization of hanuman langurs(Presbytis entellus), infanticide,and the monopolization of females

Data from 24 wild populations of hanuman langurs (Presbytis entellus)in south Asia are used to test hypotheses seeking to explain variation in troop structure and the incidence of infanticide. The occurrence of infanticide is associated with a one-male troop structure and not with a high density. The density, predation, and economic-advantage hypotheses, as explanations for the occurrence of one-male and multimale troops, are not supported by the review. However, the monopolization hypothesis is not contradicted; the number of adult males per troop is significantly correlated with troop size and with the number of adult females per troop. Therefore it is suggested that a one-male troop structure will arise if a male is able to monopolize a group of females, a multimale troop if he cannot. One-male troops may predispose to infanticide because of high variance in male mating success and high intermale competition between groups rather than within troops. If female dispersion determines troop structure, it is speculated that females could manipulate males to form a multimale society if the advantages in terms of infant survival and intertroop conflict exceeded the costs in terms of not producing infanticidal “sexy sons.”