Land or lover? Territorial defence and mutual mate guarding in the crimson-breasted shrike

Crimson-breasted shrikes produce duets which are used in interactions with neighbours and intruders. We examined two major hypotheses explaining duetting, the territorial defence and mate guarding hypotheses, using playback experiments. The responses of 12 pairs towards solo male, solo female and stereo duet playback were analysed using principal component analysis and ANOVA. The measures of response intensity revealed in most cases highly coordinated responses of the pair partners. Although these joint responses could suggest cooperative territorial defence or mutual mate defence, responses to the treatments differed between males and females in terms of three variables. Females increased their answer rates significantly in response to unpaired female intruders, compared to unpaired male intruders. During duet playback, females directed their responses equally at both intruding pair members. In contrast, males were equally responsive towards unpaired male intruders and intruding pairs, but in the latter case directed their responses at the male pair member. In the presence of unpaired female intruders, males increased their solo song rates, but in the presence of intruding pairs, they increased overlapping rates and duetted with their mates. These divergent responses suggest that females use duets for mate guarding against unpaired females, whereas males use solo songs for mate guarding and both solos and duets in territorial defence.     

Dominions and site-fixed aggressive behavior in breeding male Leucorrhinia intacta (Odonata: Libellulidae)

Summary We report aggressive spacing behavior in male dragonflies, Leucorrhinia intacta, that is characterized by variations in the probability of chasing conspecific male intruders within a defined area around a male's perch. The chase probability depends on the total intruder pressure and the behavior and distance of the intruder from the perched male (Fig. 2). This nonexclusive, site-fixed area has been called a dominion.We also examined the distribution of intruders among the various behavior-distance categories. Chase rates (per 15 min) were correlated mostly with number of intruders that hovered (rather than flew) close to the territorial male. Finally, we examined the impact of the changing chase probabilities on the variation in aggressive interactions as a function of intruder pressure. We conclude that, to some extent, territorial males stabilized defense costs across a variety of intruder pressures by maintaining dominions.     

Reproductive benefits of infanticide in a biparental burying beetle Nicrophorus orbicollis

Summary Single females and pairs of Nicrophorus orbicollis Say were established as residents on carcasses to investigate interactions with intruders under natural conditions. When carcasses were exhumed 10–12 days later, one or more intruders were found on 19% of nests. Intruders expelled one or both of the residents and sometimes paired with a resident or another intruder of the opposite sex. An intruder tended to be larger than the same-sex adult it replaced on a carcass. A male intruder that joined a single female, on the other hand, was not necessarily larger than the resident female. There was indirect evident that intruders committed infanticide and then started their own reproductive attempt on the carcass. Follow-up experiments in the laboratory indicated that interactions between intruders and residents were agonistic and that infanticide regularly occurred as a consequence of a takeover. Use of a genetic marker demonstrated that male and female intruders obtained reproductive benefits from infanticide. However, intruders sometimes cared for mixed broods consisting of their own young and young of a prior resident.     

Asymmetric costs favor female desertion in the facultatively polyandrous northern flicker (Colaptes auratus): a removal experiment

Previous studies of biparental care in birds have focused on the males contribution and experimental removal of males. Woodpeckers, with a high level of paternal care including nocturnal incubation and brooding by males, offer a meaningful system in which to examine the importance of care by females. I studied the reproductive performance of 17 widowed male and 8 widowed female northern flickers (Colaptes auratus) relative to biparental (control) pairs. Of these single parents all widowed shortly after hatching, only one female abandoned its nest. Single parents boosted their provisioning rates to achieve 83% the provisioning rate of control broods, but reared significantly fewer young and young of poorer quality. However, single males, with 85% of the reproductive success of controls, were more successful than females with 43% the success of controls. Among widowed birds, a not significant lower survival was observed, but the chance of re-pairing with the same partner in a subsequent year was only 16% in the natural population, so long-term costs of desertion may be small. Although females seem to have the incentive and ability to desert, a lack of available males may constrain opportunities. This study demonstrates that when parental care roles are reversed in altricial birds, asymmetric benefits can favor female and not male desertion.     

Brood defense and filial cannibalism in the spottail darter (<Emphasis Type="Italic">Etheostoma squamiceps)</Emphasis>: the effects of parental status and prior experience

Male spottail darters (Etheostoma squamiceps) defend nest sites in which females deposit eggs over the course of several weeks. In laboratory experiments, I tested three predictions of the hypothesis that the presence of eggs increases the value of a nest site to male spottail darters: (1) males should preferentially defend nests with eggs over empty nests, and males that sired the eggs should (2) spend more time defending them and (3) consume less of their broods than should alloparent males. Parent males were tested using eggs they were guarding in the field; alloparents included males that were not defending eggs immediately prior to the experiment (inexperienced), and males that were defending eggs in the field when captured but were given eggs sired by a different male (experienced). In choice experiments in which males were offered nest sites differing only in the presence of eggs, males preferred to defend nest sites with eggs regardless of parental status. Parent males spent significantly more time defending eggs and consumed less of their broods than did inexperienced alloparent males. Experienced alloparent males were similar to parent males in brood defense but had levels of filial cannibalism similar to inexperienced males, suggesting that prior experience may influence brood defense but not egg consumption.     

Experimental evidence that bright coloration is not important for territory defense in purple martins

Summary I tested the hypothesis that bright breeding plumage in territorial males acts as a badge of fighting ability or aggressive motivation to intimidate intruders. Territorial male purple martins (Progne subis) whose iridescent blue plumage was lightened to mimic the appearance of subadult males did not suffer an increase in intruder pressure or loss of territory compared with control males. Bright plumage color itself did not deter intruders and was not important for successful territory defense. Furthermore, a bright coloration of owners was not associated with an increased level of aggression toward intruders. Results from parallel studies on this species suggest that bright coloration is important in territory acquisition. The effectiveness of badges of fighting ability and aggressive motivation in territory defense is limited by whether intruders benefit from assessing these traits in owners. Differences in signaling systems between species are due in part to differences in floater tactics and the mode of territory acquisition.     

Aggressive brood defense by females in the stomatopod Gonodactylus bredini

Summary Defense behavior by brooding Gonodactylus bredini females was studied in staged fights against conspecific challengers. Females brooding eggs were more successful at defending their dwelling cavities than were nonreproductive females. Females caring for larvae were no more successful at cavity defense than were controls.Brooding females do not defend cavities by using more aggressive acts or by engaging in longer fights than nonreproductive females. The increase in defense success seems to be due to a greater willingness to continue a contest after being struck by a challenger. Also, brooding females make more frequent use of threats, in particular the Meral Spread display, than controls.Females defend cavities as vigorously as if they still had eggs 4 or 5 days after their eggs have been removed. Within the first 24 h after oviposition, a female's success at defending a cavity is greatly reduced compared to that of a female just 1 day later.     

Female nest defense in a coral-reef fish, Dascyllus albisella, with uniparental male care

Intraspecific variation in the patterns of parental care has been observed in a variety of animals; however, the possibility of parental care by a non-caregiving parent of uniparental species has not been thoroughly explored. In the coral-reef damselfish, Dascyllus albisella, only males normally exhibit parental care. In this study, we examined the response of females of this species to egg predators after experimental male removal and an elevated level of egg predation, at two small patch reefs (reefs 1 and 2) in Hawaii. We tested theoretical expectations that a nest was defended only by females which had spawned in the nest, and that larger females had a higher likelihood of defense than smaller females. A nest was defended against egg predators more frequently by females that had spawned in that nest than would be expected by chance. Not all females that had spawned in a given nest participated in defense. There was a positive association between female body length and the likelihood of defense at reef 2, but not at reef 1. Within a set of females that had spawned in the same nest during the same nesting cycle, defending females had larger body lengths than non-defending females at reef 2 but not at reef 1. Lack of association between female size and likelihood of defense at reef 1 was unexpected, but may correlate with the smaller average female size and smaller size differences among females on that reef.     

Territory ownership and familiarity status affect how much male root voles (Microtus oeconomus) invest in territory defence

Neighbour–stranger discrimination occurs when individuals respond with more aggression to strangers than to territorial neighbours—a phenomenon termed the “dear enemy phenomenon” (DEP). We investigated the DEP with male and female root voles (Microtus oeconomus Pallas 1776) using field dyadic arena tests conducted in enclosures where we could test for the effects of familiarity (familiar versus stranger), ownership (resident versus intruder status) and resource-holding potential (body mass) on territorial behaviours. The results showed that males put more effort into territorial defence than females, and males could discriminate between neighbours and strangers. In males, aggressiveness was influenced by a significant two-way interaction between treatment and ownership. Male residents were more aggressive towards stranger intruders than towards neighbour intruders, while male intruders were less aggressive towards stranger residents than towards neighbour residents. In females, neither treatment nor ownership status had a significant effect on aggressiveness. Familiar males performed more social behaviours but less non-social behaviours than stranger males. Furthermore, there was a clear dominance hierarchy between residents and intruders in stranger dyads, with the male territory holders dominating the intruder in pairwise interactions. To our knowledge, these results demonstrate for the first time DEP in a small mammal with a known pedigree and present the first evidence for “prior resident advantage” in voles. We argue that both ownership status and familiarity status affect how much an individual invests in territory defence. The benefits of neighbour–stranger discrimination for male root voles and the absence of neighbour–stranger discrimination in female root voles are discussed.     

The effect of resource holding potential,nest size and information about resource quality on the outcome of intruder-owner conflicts in the sand goby

Summary Sand goby, Pomatoschistus minutus Pallas, males need a nest site for successful breeding. The value of the nest increases with size, as larger nests can hold more eggs. In the field using artificial nest sites, takeovers occurred in 29.0% of the cases. The winning intruders were on average 16.3% larger than the owners. These, in turn, were significantly smaller than the nonreplaced males. Replacement probability increased with nest size. Prior ownership was of no importance in experimental situations, but might have affected takeover probabilities in the field. Males assessed the size of a potential nest site on the basis of its external appearance. When intruders and owners were given opposite information about nest size the experiments resulted in takeover rates of around 50% in situations where takeover rates were expected to be extreme. This suggested that intruders might update their information about a site's true value during the fight. The continuous assessment, in turn, might affect their motivation to continue fighting. On average, larger males were more active in initiating displays. However, smaller opponents challenged their larger opponents more actively when they were defending a large nest than when they were defending a small nest. The greater willingness to defend might provide the intruder with more information about the real value of the nest.     

Parental investment, adult sex ratios, and sexual selection in a socially monogamous seabird

Although most birds are monogamous, theory predicts that greater female parental investment and female-biased adult sex ratios will lower the polygyny threshold. This should result in polygynous mating, unless obligate biparental care or the spatial and temporal distribution of fertilizable females constrains a male’s ability to take advantage of a lowered polygyny threshold. Here we present data on the extent of male sexually dimorphic plumage, adult sex ratios and breeding season synchrony in three populations of a socially monogamous seabird, the brown booby Sula leucogaster. For one of these populations, San Pedro Mártir Island, we also present data on differences in male and female parental investment, mortality and probability of pairing. The extent of plumage dimorphism varied among populations. Sex ratios were female biased in all populations. On San Pedro Mártir Island, parental investment was female biased, females failed more often than males to find a mate, but there was no polygyny. We suggest that on San Pedro Mártir: (1) a period of obligate biparental care coupled with a relatively synchronous breeding season constrained the ability of males to take advantage of a high environmental polygamy potential and (2) the resulting socially monogamous mating system, in combination with the female-biased adult sex ratio, caused females to be limited by the availability of males despite their greater parental investment. Received: 18 November 1999 / Accepted: 24 January 2000     

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

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

Interspecific aggression in Leucorrhinia dragonflies: a frequency-dependent discrimination threshold hypothesis

Summary Male Leucorrhinia dragonflies defend territories from conspecific and heterospecific intruders. Defense against heterospecifics is surprising, as mating and oviposition are the only activities that occur on the territories, and heterospecific males are not expected to pose a reproductive threat. L. frigida and L. intacta males respond aggressively with equal frequency and intensity against intrusions by conspecifics and heterospecifics. In contrast, L. proxima males respond more aggressively against conspecifics. The apparent lack of species discrimination shown by L. frigida and L. intacta males may result because territorial males that assess intruders (as do L. proxima) suffer a tactical disadvantage from hesitating when an intruder flies in. This assessment process may lead to reduced fighting success by the territorial male. Thus there is a tradeoff between assessment cost incurred when the intruder is a conspecific and benefit gained from avoiding conflict when the intruder is a heterospecific. Given this assessment cost, males of species that only rarely encounter conspecific intruders are more likely to evolve species discrimination than males of species that commonly encounter conspecific intruders.     

Why do female mice mate with multiple males?

Females often show multi-male mating (MMM), but the adaptive functions are unclear. We tested whether female house mice (Mus musculus musculus) show MMM when they can choose their mates without male coercion. We released 32 females into separate enclosures where they could choose to mate with two neighboring males that were restricted to their own territories. We also tested whether females increase MMM when the available males appeared unable to exclude intruders from their territories. To manipulate territorial intrusion, we introduced scent-marked tiles from the neighboring males into males' territories, or we rearranged tiles within males' own territories as a control. Each female was tested in treatment and control conditions and we conducted paternity analyses on the 57 litters produced. We found that 46 % of litters were multiply sired, indicating that multiple paternity is common when females can choose their mates. Intrusion did not increase multiple paternity, though multiple paternity was significantly greater in the first trial when the males were virgins compared to the second trial. Since virgin male mice are highly infanticidal, this finding is consistent with the infanticide avoidance hypothesis. We also found that multiple paternity was higher when competing males showed small differences in their amount of scent marking, suggesting that females reduce MMM when they can detect differences in males' quality. Finally, multiple paternity was associated with increased litter size but only in the intrusion treatment, which suggests that the effect of multiple paternity on offspring number is dependent on male–male interactions.     

Asymmetric sexual conflict over parental care in a biparental cichlid

Theoretical models predict that parents should adjust the amount of care both to their own and their partner’s body condition. In most biparental species, parental duties are switched repeatedly allowing for repeated mutual adjustment of the amount of care. In the mouthbrooding cichlid Eretmodus cyanostictus, terms are switched only once with females taking the first share. The timing of the shift of the clutch between mates strongly determines both partners’ brooding period and thereby their parental investment. Females signal their readiness to transfer the young several days before the male finally takes them, suggesting sexual conflict over the timing of the shift. In a lab experiment, we reduced the body condition of either the female or the male of a pair to test whether energy reserves affect the timing of the shift and whether female signalling behaviour depends on energetic state. Males with a lowered condition took the young later and incubated for a shorter period, which prolonged the incubation time of their female partners. When female condition was lowered, female and male incubation durations remained unchanged, although females signalled their readiness to shift more intensely. Our results suggest that males adjust their parental investment to own energy reserves but are unresponsive to their mate’s condition. Females appear to carry the entire costs for the male’s adjustment of care. We propose that intrinsic asymmetries in the scope for mutual adjustment of parental investment and the costs of negotiation crucially influence solutions of the conflict between sexes over care.     

Sex-biased costs in nest defence behaviours by lesser snow geese (Chen caerulescens): consequences of parental roles?

Nest defence behaviours, such as attacking predators and defending against predator attacks, expose birds to risk of injury and death. However, direct costs of such behaviours are poorly documented. To evaluate potential costs of nest defence behaviours in lesser snow geese (Chen caerulescens), we (1) estimated the proportion of interactions between arctic foxes (Alopex lagopus) and geese that resulted in physical contact and (2) examined how nest defence behaviours varied between male and female geese. We separated interactions into attacks initiated by foxes (attacks by foxes) and attacks initiated by geese (attacks by geese). Risks associated with attacks by geese were considerably lower than the risks associated with attacks by foxes; only 1 of 1,179 attacks by geese resulted in physical contact between foxes and geese, whereas 26 of 89 attacks by foxes involved such contact (two female geese were killed during these attacks). Attacks by geese were made almost exclusively by male geese (>97%), whereas female geese were involved in 75% of all attacks by foxes that resulted in physical contact with geese. There was, thus, a considerable difference in risks associated with male and female nest defence behaviours. We suggest that parental roles during nesting (i.e., females incubate and males guard) expose female geese to greater risk of injury and death. Male geese may, however, reduce the risk of injury or death to their mates with pre-emptive attacks on foraging foxes.     

How reliable are behavioral cues for assessment of male mating status in polyterritorial wood warblers,Phylloscopus sibilatrix?

It has been suggested that polyterritoriality in birds, i.e. mated males defending two spatially separated territories, deceives females into accepting polygyny against their best interest. Mated males singing on secondary territories should make it difficult for new arriving females to assess the mating status of unmated and mated males advertising to females. However, there have been objections to the deception hypothesis, suggesting that differences in territorial behavior between males of different mating status may make correct assessment possible. Polyterritorial males frequently leave their secondary territory to visit the female in the primary territory, thereby revealing their mating status to females in search of mates. The aim of this study is to investigate how reliable behavioral cues are in assessing the mating status of males in polyterritorial wood warblers. Our study shows that singing activity of mated males on secondary territories and unmated males was similar in the morning, while there was a significant difference in the afternoon. Unmated males spent, on average, more time on their territories than mated males spent on their secondary territories. However, there was considerable variation in both unmated and mated males in the frequency in which they left their territories. For instance, unmated males left their territories quite frequently early in the season compared to later. Hence, females must be able to interpret singing behavior and a male leaving his territory differently depending on time of the season and time of the day in order to correctly assess the mating status of advertising males. Estimating the increase in probability of finding an unmated male by using behavioral cues showed that females could increase the probability of finding an unmated male compared to random choice with respect to mating status. The uncertainty was, however, still considerable. The most important factor affecting the probability of finding an unmated male was the proportion of mated males on secondary territories compared to unmated males. Our study suggests that presence on a territory is not a reliable cue for the assessment of male mating status in polyterritorial wood warblers.     

Snake-directed mobbing by the Formosan squirrel Callosciurus erythraeus thaiwanensis

Summary The Formosan squirrel Callosciurus erythraeus thaiwanensis was often observed mobbing the snake Elaphe climacophora. A total of 36 natural and 21 experimentally induced mobbings were observed and analyzed. The number of attending squirrels per mobbing event varied from one to seven, and 84% of observed mobbings were performed by more than one individual. The duration of a mobbing bout increased directly with the number of assembled mobbers. In 68% of cases, one adult female was one of the mobbers; there was never more than one female per mob. The number of assembled males per mob varied from 0 to 5; 40% of the mobbings involved two or more males. This difference is correlated to a difference in spacing patterns of both sexes; female home ranges were distributed exclusively, whereas male home ranges overlapped each other. Females mated with multiple males whose home ranges overlapped theirs. Female's reproductive status (conception, lactation, and weaning) affected their intensity of response to the playback of mobbing calls, females in pup-rearing period being the most sensitive. One function of the mobbing appears to be defense of the young against snake predation. Females mobbed more intensively and longer than males. Males that had resided at the study site for at least 1 year tended to mob more frequently than intruders.     

Female-biased immigration and male peace-keeping in groups of the shell-dwelling cichlid fish Neolamprologus multifasciatus

The small cichlid fish Neolamprologus multifasciatus is endemic to Lake Tanganyika and is unique among fish because it lives in complex social groups with several reproductive males and females, all of which participate in defending their territory against neighbors and intruders. Individuals use empty snail shells for breeding and shelter. Previous parentage analysis using microsatellites suggested occasional exchange of individuals between groups. In field experiments, we found that females showed a higher tendency than males to migrate into territories already occupied by a resident pair. The phenomenon and causes of female-biased immigration were further investigated in aquarium experiments: Nine of 15 females, but only 2 of 15 equally sized males, settled in territories of established pairs. Territorial males exhibited more aggression toward strange males (potential reproductive competitors) than toward strange females (potential additional mates); their females were more aggressive toward strange females (probably competitors for shells) than toward strange males. Apparently, a conflict exists between the sexes regarding the immigration of additional females. This conflict seems to be the selective basis for observed active male interference in aggressive disputes between females in the territory. Interfemale tolerance in a group was greater in the male’s presence than in his absence. Received: 1 February 2000 / Received in revised form: 2 May 2000 / Accepted: 5 May 2000     

Evidence of equal maternal investment in the sexes in the polygynous and sexually dimorphic grey seal (Halichoerus grypus)

In polygynous and sexually dimorphic mammals, parents may be expected to bias their investment towards sons because variation in reproductive success is usually higher among males than among females. Moreover, male reproductive success often depends on adult body size, which, in turn, may depend on the level of parental investment. We therefore predicted that in the grey seal (Halichoerus grypus), a polygynous and sexually dimorphic phocid seal, females should invest more in individual sons than in individual daughters. We found that male pups were born heavier than female pups, but that the growth rates and suckling behaviour were similar for the two sexes. The growth rates and the birth weights were not correlated for the pups of either sex. Mothers did not behave differentially towards offspring of the two sexes, except that mothers of male pups spent more time in visual contact with their pups. Male and female pups had similar activity levels and begged at similar rates. We argue that reports of equal expenditure on the two sexes can be accepted as evidence of equal investment, provided that three assumptions are fulfilled. First, parental care must be costly to the parent. Second, energy expenditure must be the most important component of parental investment. Third, there must be no negative correlation between maternal body condition and the ratio of sons to daughters produced. We argue that these assumptions are met in our study, and that our results provide evidence of equal maternal investment in the sexes in grey seals.