Reproductive Consequences of Aggression in a Territorial Songbird

Territorial aggression can influence males’ ability to obtain high‐quality resources and access to mates; however, in many species, the reproductive consequences of variation in aggression are unknown. In this study, we investigated how individual variation in aggressive behavior relates to reproductive success in socially monogamous, genetically polygynous song sparrows (Melospiza melodia). Prior research in this species shows that male song sparrows differ in their willingness to engage in agonistic interactions with territorial intruders and that individual variation in aggression appears to have functional significance. Aggressive males have been shown to obtain territories where females produce larger clutch sizes, suggesting that individuals who display high levels of territorial aggression are defending high‐quality territories or females. Further, aggressive males are considered a greater threat to territory‐holding males than less aggressive males. In this study, we ask whether individual differences in aggression are linked to differences in extra‐pair reproductive success, annual reproductive success, and offspring quality. We did not uncover a relationship between aggression and annual reproductive success or patterns of extra‐pair paternity. However, we found that the nestlings of aggressive males grew at a faster rate than the nestlings of less aggressive males. Future studies should attempt to identify mechanisms to explain the relationship between offspring growth rate and male aggression and investigate whether faster offspring growth rates translate to greater survival and recruitment of offspring.     

Monogamy in a feeding generalist, <Emphasis Type="Italic">Chaetodon trichrous</Emphasis>, the endemic Tahitian Butterflyfish

Butterflyfishes have been well studied for their feeding ecology and mating systems. In particular, studies of corallivorous butterflyfishes have supported models of monogamy based on their predictable, low quality food; a patch of coral that is economically defensible by a pair. Moreover, pairs often exhibit trade-offs in territorial defense (greater by males) and feeding (greater by females) that improve their reproductive success. However, this model has not been well tested for more generalist feeders. In addition, recent hypotheses for monogamy in fish have emphasized parental care, but butterflyfishes do not provide parental care. This study tests five hypotheses for monogamy in the endemic Tahitian butterflyfish, Chaetodon trichrous: 1) uniform distribution of limiting resources, 2) joint defense of a territory, 3) low mate availability, 4) predator detection, and 5) benefits of cooperative behavior. Chaetodon trichrous was the most abundant butterflyfish in bays. Pairs jointly patrolled feeding territories. They preferentially fed over hard substrate other than live coral, however, this substrate was available outside of territories. They also ate plankton. Pairs were sorted positively for size, and all pairs were heterosexual. Males were larger than their partners, but females fed at higher rates. These results suggest that C. trichrous is monogamous, but reject the hypotheses that pairs form for joint defense of a territory (pairs swam together), that pairs remain together because of low mate availability (frequent interactions with neighbors), or that pairs form for predator detection (no homosexual pairs). Monogamy in C. trichrous is associated with the uniform distribution of hard substrate, although this resource is not limiting. Further, the higher feeding rate of females may represent a benefit provided by their monogamous mates.     

Intra- and Inter-seasonal Variation in the Socio-Spatial Behavior of Adult Male Collared Lizards, Crotaphytus collaris (Reptilia, Crotaphytidae)

When individuals maintain strong inter‐seasonal philopatry to the same territories, males may be able to re‐establish territory occupancy without intense intra‐sexual aggression, and instead spend more time courting females early in the reproductive season. Furthermore, when some males have prior experience defending the same territories, it may be necessary for young males to exhibit higher levels of aggression because they are establishing a territory for the first time. We tested these hypotheses by examining within‐season (1992 and 1997) temporal variation in the social behavior of adult male collared lizards of known age and prior territorial experience in a population where inter‐season philopatry to territories is high. Contrary to expectations, the frequency of aggression exhibited by males with and without prior territorial experience did not differ. The frequency of intra‐sexual aggression was higher in 1992 than in 1997, perhaps because male competitors were more abundant in 1992. Although there was an interactive effect of year, male display and patrol were low at the beginning of the reproductive season in Apr. and May, reached peaks during midseason in June, and then decreased as reproduction ended in July. The size of territories showed a similar pattern, with males defending larger areas in June. Our data support the philopatry hypothesis in that the establishment of territories occurred without high levels of aggression early in the season, perhaps because territory boundaries have been well defined by high rates of patrol and advertisement during the middle of the previous season. Inter‐sexual interactions were most frequent in June rather than at the beginning of the reproductive season. Adult females are producing their second clutches and yearling females are producing their first clutches in June. The high rate of inter‐sexual encounters in June supports the hypothesis that males allocate more time to courtship when females are receptive because there are more reproductively active females at this time. The temporal pattern of activities in adult Crotaphytus collaris appears to function as a compromise between competing intra‐ and inter‐sexual social demands on males, allowing males to maximize mating opportunities as well as maintain future access to productive territories.     

Experimental analysis of monogamy in the Caribbean cleaner goby, Gobiosoma evelynae

To investigate factors promoting monogamy, we studied the reproductive behaviour and ecology of the monogamous Caribbean cleaner goby at two sites near St Croix, U.S. Virgin Islands. We assessed mate fidelity and the amount of time required to acquire a new mate in experimentally widowed males and females. We also measured behavioural responses of pair members and single females to experimentally introduced conspecific intruders of both sexes. Finally, we evaluated the distribution of suitable habitat relative to existing goby territories. We found that pairs often separated due to males abandoning females and moving to a new territory, that most widowed fish soon acquired a new mate, and that there was strong aggression towards large same-sex intruders. We also found an abundance of suitable, unoccupied habitat. We conclude that pairs are maintained by intrasexual aggression that is related to both mate availability and other resources, such as high-quality, food-rich cleaning stations. Combined with observations of frequent pair separation, interterritory male movement, and relatively rapid remating by both sexes, these results suggest a complex mating system that is best classified as serial monogamy. Copyright 2003 Published by Elsevier Science Ltd on behalf of The Association for the Study of Animal Behaviour.      

Habitat selection and polygyny in breeding Corn Buntings Miliaria calandra

Unlike many other polygynous passerine species, female Corn Buntings Miliaria calandra apparently do not suffer costs by pairing polygynously, yet it is unclear whether this is because polygynous males hold the highest quality territories or because pairing with polygynous males is unimportant in determining female reproductive success. Male Corn Buntings on North Uist, Scotland, consistently defended territories which contained nesting habitat, and females often foraged outside male territories when provisioning nestlings. Females showed strong preferences for nesting in uncultivated land, and 80% of nests were under Hogweed Heracleum sphondylium, possibly because this provided cover against predation and the weather. When provisioning nestlings, females showed strong preferences for foraging in cereal crops, probably because this habitat provided better food resources and/or better cover from predators. Males were unpaired or paired with one to three females per breeding season, but variation in territory size or vegetation composition did not explain differences in the number of females paired with individual males. We suggest that when females neither gain benefits nor suffer costs by breeding polygynously, and males do not differ greatly in the areas of habitat selected, polygyny can arise through random female settlement within the nesting habitat.     

Caribbean damselfish with varying territory quality: correlated behaviors but not a syndrome

The behavioral syndrome hypothesis suggests that individualanimals within a population behave differently due to specificbehavioral types, and these should be consistent across behaviorsor in different contexts. In contrast, for animals that livewithin an environment in which territory quality can changeover time, natural selection should have favored behavioralflexibility and modulation of the cost of defense in relationto territory quality. This would require assessment of the territoryfollowed by displays of appropriate types and intensities ofbehavior. We examined the territorial behavior of male beaugregorydamselfish (Stegastes leucostictus) by enhancing territory qualityusing artificial breeding sites and comparing their behaviorto males on lower quality natural sites. When male fish weredefending high-quality artificial territories, they had higherlevels of aggression toward male conspecifics and courtshiptoward females than when on low-quality natural territories.We also found that aggression and courtship behaviors were correlatedon natural sites but not on artificial sites. Behaviors werenot correlated within individuals when males switched from naturalto artificial territories or from artificial to natural territories.These results indicate that males assess their current territoriesand adjust behaviors accordingly and that courtship and aggressivebehaviors are not linked within a permanent behavioral syndrome.     

The role of androgen receptors in regulating territorial aggression in male song sparrows

This paper examines the role that androgen receptors (ARs) play in modulating aggressive behavior in male song sparrows, Melospiza melodia morphna. Song sparrows are seasonally breeding, territorial birds that maintain year-round territories with male-female pair bonds formed during the spring breeding season. Plasma testosterone levels peak as territories are established and mates acquired. In late summer, testosterone levels fall and remain basal during the non-breeding season. We examined the role of ARs in regulating territorial aggression in captive song sparrows under short- and long-day conditions as well as just prior to, and at the start of the breading season in freely living birds using the nonsteroidal antiandrogen flutamide to block AR function. Birds were implanted with either empty or drug filled silastic implants for 18 to 42 days and then challenged with a novel male decoy to assess the individual birds level of male-male aggression. Freely living birds remained on their home territory and underwent a simulated territorial intrusion, whereas laboratory-held birds were assessed using a laboratory simulated territorial intrusion and remained in their home cage. Experimental treatment of male song sparrows decreased aggressive behavior during the pre-breeding life history substage (March-April) in freely living birds as well as in laboratory-held birds under long-day (16L:8D) conditions. During the early breeding substage (April-May) there was no measurable effect of flutamide treatment on aggressive behavior, nor was there a difference in behavior in the (8L:16D) laboratory birds. This demonstrates that ARs are an important component of the neuroendocrine control of aggressive behavior. Given that flutamide only affected aggression during the pre-breeding substage and in LD birds, the results suggest that AR dependent control of aggressive behavior changes as song sparrow life history states change.     

Possible relationships between morphology,territory quality,and skin color of American Kestrels1

ABSTRACT Carotenoid‐based coloration of skin and plumage has been found to be correlated with individual quality in many species of birds during the breeding season. However, less is known about the possible role of these signals during the nonbreeding season, particularly among nonpasserines that defend winter territories. American Kestrels (Falco sparverius) are sexually dimorphic raptors that defend winter territories and possess carotenoid‐based morphological features known to be correlated with individual quality. Much is known about winter territory use and habitat segregation by male and female kestrels, but possible relationships among morphological features, individual quality, and habitat quality have not been examined. Our objective was to examine possible relationships between morphology, territory quality, and skin color of American Kestrels. Male kestrels had brighter skin than females, and the skin color of male kestrels was positively correlated with size (wing chord and tail length) and territorial quality (hunting territories with less canopy and more grass cover). No such relationships were found for female kestrels. Skin color appears to be an honest indicator of quality for male American Kestrels and may serve both intersexual (territory acquisition) and intersexual (mate choice) functions during the breeding and nonbreeding seasons.     

Female Choice Predicts the Best Father in a Biparental Fish,the Midas Cichlid (Cichlasoma citrinellum)

The Midas cichlid is a monogamous, biparental species. It breeds in a highly competitive system where pairs have a low probability of raising fry to independence. Both parents must cooperate to retain the territory and protect the fry from predation. Previous experiments showed that females prefer large, aggressive and sexually experienced males as mates but males do not display any consistent preferences. Here I present the results of two experiments designed to see whether qualities preferred by females correlate with increased success in retaining territories and in providing parental care. Pairs with either large or aggressive males had an advantage in appropriating and holding a breeding territory; reproductive experience conferred no advantage in usurping a territory. Aggressive and reproductively experienced males had an advantage in defending the brood from predators of fry, but size had no effect. Thus, the qualities preferred by females confer advantages both in holding territories and in protecting fry. In contrast, males need not be selective because females, once in possession of a brood, defend it equally well regardless of size, aggressiveness, or reproductive experience. The system is one of mutually enforced monogamy based on female choice; females drive the system because they provide more investment than do males (combining gametes and time) and because this investment is a reliable resource to the male.     

Aggression of breeding eastern bluebirds (Sialia sialis) toward their mates and models of intra-and interspecific intruders

Hypotheses regarding the evolution of aggressive patterns during the breeding season in eastern bluebirds (Sialia sialis) were tested by placing models of eastern bluebirds and house sparrows (Passer domesticus) into territories of experimentally naive breeding pairs of eastern bluebirds during three stages of the nesting cycle. The responses of the resident males to models of male bluebirds decreased from the egg-laying to the nestling stage, and this response pattern can be interpreted as support for the protection-of-parentage hypothesis or the nest-site-defence hypothesis. The aggression of resident males to resident females during trials with models of male bluebirds weakly conforms to the pattern predicted by the protection-of-parentage hypothesis. The aggressive response to models of house sparrows also decreased from the egg-laying to the nestling phase in a manner consistent with the nest-site-defence explanation. No aggression toward mates by males or females was observed during trials with house sparrow models, an observation consistent with the predictions of the protection-of-parentage hypothesis.     

Disruption of Dear Enemy Recognition Among Neighboring Males by Female Leon Springs pupfish, Cyprinodon bovinus

Males defending territories often settle into adjacent areas, sharing a common border that is maintained by a reduced level of aggression known as dear enemy recognition. While social conditions may affect the dear enemy relationship among males, what role females play, if any, is unclear. In a field study of the highly promiscuous Leon Springs pupfish, Cyprinodon bovinus, we asked whether females influenced this relationship of neighbors to their advantage. We observed 16 territorial residents, mapping the precise location of each male's behaviors within its territory. Resident males engaged in less aggression against neighbors compared with intruders, and neighbors intruded less deeply into the residents’ territories than intruders. Despite this locality restriction, neighbors were responsible for as many spawning interruptions as intruders. Females did not spawn randomly in the males’ territories, nor did they spawn near territory centers where aggression was low. Rather, females were more likely to settle and spawn in the outer half of the territories where competition among males was highest. When a neighbor entered a resident's territory to interrupt a spawn, the female was more likely to leave the resident's territory for the neighbor's than to remain. These observations suggest that the female used the intrusion by the neighbor to engage the resident and interrupt the spawn as a measure of this male's quality and that, while neighboring males benefit from the presence of dear enemy recognition, females benefit from its disruption.     

Female mate choice based on territory quality in barn swallows

Female mate choice based on territory quality is difficult to study because territories often contain many resources, which are difficult to quantify. Here, using the barn swallow (Hirundo rustica gutturalis) breeding at an outdoor breeding site in Japan, where each male defends only a small territory containing old nests, we studied whether females choose social mates based on territory quality. Since the territories of this species contain few other resources, territory quality can easily be assessed by quantifying old nests in the territory. We made the following four observations: (1) male swallows displayed old nests in their territories to females; (2) the old nests used for the first clutch were less broken than the other old nests within the same territory; (3) territory quality, defined by the number of old nests weighted by the intactness of each old nest, predicted the productivity of the territory; and (4) males with better territories paired with females earlier, and hence bred earlier, than those with inferior territories. The relationships remained significant even after controlling for male morphological traits. Based on these results, we can infer that female swallows choose their mates based, in part, on territory quality.     

Nest and mate choice in the red bishop (Euplectes orix) : female settlement rules

We investigated female settlement in a colony of red bishops(Euplectes orix), a territorial and highly polygynous weaverbirdwidely distributed over sub-Saharan Africa. An earlier studyshowed that male reproductive success is mainly determinedby the number of nests a male builds in his territory, whichappeared to be a good indicator of male quality. Because malesprovide no parental care or food resources within the territory,females sharing a territory do not compete for material resourcesand might therefore be expected to settle preferentially interritories of males that build many nests to gain the possiblegenetic benefit of high-quality offspring. An analysis of femalesettlement, however, revealed that females did not show a preferencefor territories of males with many nests and that the distribution of female breeding attempts with regard to the number of vacantnests within a territory could be explained best by randomfemale settlement in 3 out of 4 years. Females settled moreoften than expected by chance (in 3 out of 4 years) in territoriesalready containing occupied nests, indicating that residentfemales did not discourage settlement of additional females.However, sharing a territory with other females might imposecosts in terms of an increased predation risk because nestsin territories that contained other nests with young sufferedfrom higher predation than nests in territories that did notcontain other nests with young. Females therefore might tradethe possible benefits of settling in territories of males withmany nests against the costs of sharing a territory with otherfemales. This might result in the mating pattern found withrandom female settlement and male reproductive success beingdirectly proportional to the number of nests built. We discuss possible implications of this mating pattern for sexual selectionon males.     

Prey availability affects territory size,but not territorial display behavior,in green anole lizards

The availability of food resources can affect the size and shape of territories, as well as the behaviors used to defend territories, in a variety of animal taxa. However, individuals within a population may respond differently to variation in food availability if the benefits of territoriality vary among those individuals. For example, benefits to territoriality may differ for animals of differing sizes, because larger individuals may require greater territory size to acquire required resources, or territorial behavior may differ between the sexes if males and females defend different resources in their territories. In this study, we tested whether arthropod abundance and biomass were associated with natural variation in territory size and defense in insectivorous green anole lizards, Anolis carolinensis. Our results showed that both male and female lizards had smaller territories in a habitat with greater prey biomass than lizards in habitats with less available prey, but the rates of aggressive behaviors used to defend territories did not differ among these habitats. Further, we did not find a relationship between body size and territory size, and the sexes did not differ in their relationships between food availability and territory size or behavioral defense. Together, these results suggest that differences in food availability influenced male and female territorial strategies similarly, and that territory size may be more strongly associated with variation in food resources than social display behavior. Thus, anole investment in the behavioral defense of a territory may not vary with territory quality.     

FOOD RESOURCES, TERRITORY DENSITY AND REPRODUCTIVE SUCCESS OF AN ISLAND SILVEREYE POPULATION ZOSTEROPS LATERALIS

We examined the relationships between food resources, territory density and some breeding parameters (individual reproductive success and parental age) of a Capricorn Silvereye population during two years' detailed study on Heron Island, Great Barrier Reef. The territorial behaviour of the Silvereyes included dawn song. aggression and distinctive vocalizations. During both years, about 85% of pairs occupied territories although the population sizes differed. The dispersion pattern of territories was not regular and was correlated with the density of figs and human food scraps. Nestlings were fed more figs as they grew, and the parents foraged in fig trees outside their territories. However, only about 60%, of territorial pairs fledged young. Pairs which fledged most young were older, fed more insects to their nestlings, and nested in areas containing more fruiting fig trees per territory. We conclude that food resources were in short supply, and that access to fig trees provided breeding birds with a quick energy source while they searched for insect protein for the young. The data are consistent with the predictions of the ‘sufficient resource‘ hypothesis of the adaptive significance of territoriality but not with its assumption that the principal benefit is the food within the territory.     

Winter territory prospecting is associated with life‐history stage but not activity in a passerine

Finding a high quality territory is essential for many animals to reproduce successfully. Despite its importance for fitness, we know little about the process of territory prospecting in wild birds, and whether individual traits and behaviours, such as personality, co‐vary with territory prospecting. Here, we use long‐term data from a wild, insular house sparrow Passer domesticus population to test three hypotheses about territory fidelity and prospecting: 1) house sparrows show high territory fidelity between years and also during winter. 2) Individuals will prospect for a breeding territory during their first winter whereas older, more experienced individuals will keep a territory from previous years and will, therefore, show no or reduced winter territory prospecting. 3) More active behavioural types will prospect more than less active behavioural types. We use data from four winters from automatically, daily recorded nest‐box visits of 188 birds of known age. The number of nest‐boxes that each individual visited within each winter was used as a proxy of winter territory prospecting. We show that house sparrows visit multiple nest‐boxes during their first winter, whereas older individuals keep territories year‐round and, potentially because of this, indeed show reduced winter territory prospecting. Activity was not associated with the number of nest‐boxes visited. Further research is needed to investigate whether time of territory and mate acquisition differs among individuals and the possible effect on lifetime fitness.     

Butterflyfish social behaviour,with special reference to the incidence of territoriality: a review

Synopsis Butterflyfishes (Chaetodontidae) are among the best studied of coral reef fishes. Feeding ecology and some aspects of behaviour have been firmly established. However, spacing behaviour remains controversial. Two major studies made in the 1970s concluded that the majority of species were not territorial. We suggest that these and other studies which have concluded that territories are not held have generally suffered from short observation periods, and have not mapped the ranges occupied by individuals. Further, low frequencies of agonistic behaviour have been interpreted as evidence of non-territoriality. By contrast, studies which have proven territoriality have had long observation periods and have mapped ranges. These have shown that territories are usually maintained with very little overt aggression. Spacing behaviour and feeding behaviour are clearly linked, with territoriality common among benthic-feeding species, especially obligate corallivores. Species with broad dietary flexibility tend to have flexible social systems, while plankton feeders are usually gregarious. The widespread occurrence of monogamy in butterflyfishes appears linked to territoriality, the majority of territorial species identified to date occurring predominantly as pairs. Data currently available suggest that this is because pair defence of the territory is more efficient than by individuals. However, several alternative hypotheses for the evolution of monogamy based on spawning constraints and predation risk cannot yet be ruled out.     

Why do territorial male Tengmalm's owls fail to obtain a mate?

Non-breeding may occur because non-breeders are immature or somehow physiologically incapable of breeding, or because of a lack of resources (e.g. food resources, mating partners) needed to breed. There is, however, a lack of experimental evidence on whether bachelor males possessing territories and nest-sites are able to breed when supplemented with extra food or provided with mating partners. In vole-eating Tengmalm's owl, Aegolius funereus, we provided supplementary food and transferred females in nest-boxes of non-breeding males. Bachelor males that we supplemented with food did not attract mates at a higher frequency than unfed control males, which suggests that a lack of food did not influence the ability to attract a mating partner. In contrast, bachelor males presented with a female seemed to breed more frequently than bachelor males in the control group without mate addition. This suggests that scarcity of females may be an important reason for the high proportion of non-breeding males in the population (c. 25%) and excludes the possibility that non-breeding males are physiologically unable to breed. The operational sex ratio of the owl population at the time of mating may be male-biased, and some males may thus remain unpaired. Habitat and nest-box quality also seemed to be lower among bachelors than among breeding males. Received: 22 September 1997 / Accepted: 29 December 1997     

鸟类婚配制度的生态学分类

在Ｅmlen和Ｏring鸟类婚配制度生态学分类系统的基础上,根据近年来鸟类行为生态学研究的成果,对鸟类的婚配制度进行了补充分类,并强调了应以进化稳定策略的观念来认识乌类的婚配制度。补充的鸟类婚配制度生态型包括：合作型一雄一雌制（cooperative monogamy)、临界型一雄一雌制（critical monogamy)、保卫雌性型一雄一雌制（female defense mognogamy)、从领域型一雄多雌制（poly-terri-tory polygyny)和社群繁殖制。合作型一雄一雌制的鸟 类雌雄个体爱力合作才保证繁殖的成功;临界型一悲欢离合一雌制鸟类雌雄个体都有多配倾向,但迫于生态压力必须共同抚育后代才能繁殖成功;保卫雌性型一雄一雌制的鸟 类通过保卫一个雌鸟不被其它雄鸟占有而保证繁殖成功,多领域型一雄多雌制的雄鸟通过占有多个领域而多个雌鸟交配;社群繁殖制的鸟 类由三个以上个体参与工部分参与繁殖,所有个体共同抚育一批后代,现有的鸟类婚配制度可以归为一雄一雌制（monogamy)、一雄多雌繁殖,所有个体共同抚育一批后代。现有的鸟类婚配制度可以归为一雄一雌制(polygyny)、一雌多雄制（polyandry)快速多窝型多配制（rapid-multiple-clutch polygamy)和社群繁殖制（social breeing systen）五大类型。     

Male song sparrows have elevated testosterone in response to neighbors versus strangers

Upon hearing a conspecific signal, animals must assess their relationship with the signaller and respond appropriately. Territorial animals usually respond more aggressively to strangers than neighbors in a phenomenon known as the “dear enemy effect”. This phenomenon likely evolved because strangers represent a threat to an animal's territory tenure and parentage, whereas neighbors only represent a threat to an animal's parentage because they already possess a territory (providing territory boundaries are established and stable). Although the dear enemy effect has been widely documented using behavioral response variables, little research has been conducted on the physiological responses of animals to neighbors versus strangers. We sought to investigate whether the dear enemy effect is observed physiologically by exposing territorial male song sparrows (Melospiza melodia) to playback simulating a neighbor or a stranger, and then collecting blood samples to measure plasma testosterone levels. We predicted that song sparrows would exhibit increased testosterone levels after exposure to stranger playback compared to neighbor playback, due to the role testosterone plays in regulating aggression. Contrary to our prediction, we found that song sparrows had higher testosterone levels after exposure to neighbor playback compared to stranger playback. We discuss several explanations for our result, notably that corticosterone may regulate the dear enemy effect in male song sparrows and this may inhibit plasma testosterone. Future studies will benefit from examining corticosterone in addition to testosterone, to better understand the hormonal underpinnings of the dear enemy effect.