Mating system structure and population density in a polygynous lizard, Sauromalus obesus (= ater)

Increases in population density often are associated with achange in mating system structure in numerous taxa. Typically,male interactions are minimal in extremely low density populations.As density increases, males exhibit territoriality but if densitybecomes too high, the energetic cost of defending a territorywill eventually outweigh the reproductive benefits associatedwith territoriality. Consequently, males in high density populationsmay abandon territoriality and adopt dominance polygyny, lekking behavior, or scramble competition. We investigated the relationshipbetween population density and mating system structure in threepopulations of the chuckwalla, Sauromalus obesus (= ater),near Phoenix, Arizona. Densities in the Phoenix Mountains (2.7chuckwallas/ha) were lower than any population previously studied.In the Santan Mountains (10.9 chuckwallas/ha), densities weresimilar to populations studied in the Mojave Desert, and inthe South Mountains (65 chuckwallas/ha), densities were the highest yet recorded. Male mating behavior was examined by determininghome range overlap and by making direct behavioral observations.Male home range size decreased with increasing population density.There was little overlap in home ranges among males in allthree populations, whereas home ranges of males and femalesconsistently overlapped, indicating that males were strictly territorial. This conclusion was supported by behavioral observationsof interactions among individuals in a natural setting. Thenumber of females wihin male territories was correlated withfood resources (plants) in all three populations. Female homerange size appeard to be related to food resources whereasmale home ranges appeared to be related to female distribution,population density, and geology. The retention of territoriality in spite of high population densities raises new questions aboutthe relationship between density and resource defense.     

Serial Monogamy as Polygyny or Polyandry?

Applications of sexual selection theory to humans lead us to expect that because of mammalian sex differences in obligate parental investment there will be gender differences in fitness variances, and males will benefit more than females from multiple mates. Recent theoretical work in behavioral ecology suggests reality is more complex. In this paper, focused on humans, predictions are derived from conventional parental investment theory regarding expected outcomes associated with serial monogamy and are tested with new data from a postreproductive cohort of men and women in a primarily horticultural population in western Tanzania (Pimbwe). Several predictions derived from the view that serial monogamy is a reproductive strategy from which males benefit are not supported. Furthermore, Pimbwe women are the primary beneficiaries of multiple marriages. The implications for applications of sexual selection theory to humans are discussed, in particular the fact that in some populations women lead sexual and reproductive lives that are very different from those derived from a simple Bateman-Trivers model.     

Territorial cooperation and social monogamy: factors affecting intersexual behaviours in pair-living snapping shrimp

Among the factors that may contribute to the evolution of social monogamy are selection for extended mate guarding of females and selection for territorial ‘cooperation’. Many socially monogamous taxa are also territorial, with ‘partners’ sharing a single territory, suggesting that one or both partners may benefit by sharing territorial maintenance. Snapping shrimp (genus Alpheus) are socially monogamous and territorial, living in excavated burrows or with host organisms, with females performing all parental care. The territorial cooperation hypothesis predicts that male and female partners share (1) territorial defence, resulting in a reduction in the risk of eviction from the burrow, (2) burrow construction duties, such that individuals in pairs spend less time in burrow construction relative to solitary individuals, and/or (3) foraging duties, by returning food to the burrow, where it is consumed by both partners. UsingA. angulatus as a model species, a territorial defence experiment revealed that females in pairs were significantly less likely than solitary females to be evicted by female intruders, but males in pairs were not significantly less likely than solitary males to be evicted by male intruders. A subsequent experiment revealed that paired males were significantly less likely to be evicted by an intruding male if paired with sexually receptive females than if paired with nonreceptive females. Another experiment revealed that (1) paired females spent significantly more time in burrow construction than paired males, and (2) both males and females consistently returned food items to the burrow, perhaps incidentally provisioning their mates. These data suggest that social monogamy may have been selected for in part because of the advantages of territorial cooperation, as both males and females are likely to benefit by dividing the labour of territorial defence and maintenance. These tests of the territorial cooperation hypothesis are synthesized with data from tests of the extended mate-guarding hypothesis to place snapping shrimp pairing behaviour into a larger construct incorporating both the influence of ecological pressures (territoriality) and mating interactions between the sexes. Copyright 2002 The Association for the Study of Animal Behaviour. Published by Elsevier Science Ltd. All rights reserved.     

Sex difference in the communicatory significance of localized defecation sites in Arabian gazelles (Gazella arabica)

Mammalian scent marking in localized defecation sites (latrines) has often been interpreted in the context of (male) territory defense. However, latrines could have different functions in males and females, especially where territorial males monopolize groups of females with stable social alliances and pronounced home range overlap. We investigated the communicatory significance of latrines in wild Arabian gazelles (Gazella arabica) and assessed the spatial distribution of latrines within home ranges. Latrine density and utilization was highest in the center of female group home ranges, and less frequent in peripheral home range sections, pointing towards communication within groups rather than towards territoriality. When considering male home ranges, latrine densities and utilization were higher in non-overlap zones, contradicting a territorial function. This pattern appears to be caused by more females than territorial males per given area establishing latrines. A subsequent survey of latrine utilization, based on camera trapping, suggests that males use latrines for territory defense: males visited latrines in overlap zones disproportionally more often than females, and successions of two males prevailed. Our study thus highlights that male territorial marking can be masked when males and females use the same marking system for different purposes.     

Increased vasopressin expression in the BNST accompanies paternally induced territoriality in male and female California mouse offspring

While developmental consequences of parental investment on species-typical social behaviors has been extensively characterized in same-sex parent-offspring interactions, the impact of opposite-sex relationships is less clear. In the bi-parental California mouse (Peromyscus californicus), paternal retrieval behavior induces territorial aggression and the expression of arginine vasopressin (AVP) in adult male offspring. Although similar patterns of territorially emerge among females, the sexually dimorphic AVP system has not been considered since it is generally thought to regulate male-typical behavior. However, we recently demonstrated that male and female P. californicus offspring experience increases in plasma testosterone following paternal retrieval. Since AVP expression is androgen-dependent during development, we postulate that increases in AVP expression may accompany territoriality in female, as well as male offspring. To explore this aim, adult P. californicus offspring that received either high or low levels of paternal care (retrievals) during early development were tested for territoriality and immunohistochemical analysis of AVP within the bed nucleus of the stria terminalis (BNST), paraventricular nucleus (PVN), and supraoptic nucleus (SON). Consistent with previous studies, high care offspring were more aggressive than low care offspring. Moreover, high care offspring had significantly more AVP immunoreactive (AVP-ir) cells within the BNST than low care offspring. This pattern was observed within female as well as male offspring, suggesting an equally salient role for paternal care on female offspring physiology. Regardless of early social experience, sex differences in AVP persisted in the BNST, with males having greater expression than females.     

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。     

Strong male/male competition allows for nonchoosy females: high levels of polygynandry in a territorial frog with paternal care

Our knowledge about genetic mating systems and the underlying causes for and consequences of variation in reproductive success has substantially improved in recent years. When linked to longitudinal population studies, cross-generational pedigrees across wild populations can help answer a wide suite of questions in ecology and evolutionary biology. We used microsatellite markers and exhaustive sampling of two successive adult generations to obtain population-wide estimates of individual reproductive output of males and females in a natural population of the Neotropical frog Allobates femoralis (Aromobatidae), a pan-Amazonian species that features prolonged iteroparous breeding, male territoriality and male parental care. Parentage analysis revealed a polygynandrous mating system in which high proportions of males (35.5%) and females (56.0%) produced progeny that survived until adulthood. Despite contrasting reproductive strategies, successfully reproducing males and females had similar numbers of mating partners that sired the adult progeny (both sexes: median 2; range 1-6); the numbers of their offspring that reached adulthood were also similar (both sexes: median 2; range 1-8). Measures of reproductive skew indicate selection on males only for their opportunity to breed. Reproductive success was significantly higher in territorial than in nonterritorial males, but unrelated to territory size in males or to body size in both sexes. We hypothesize that female polyandry in this species has evolved because of enhanced offspring survival when paternal care is allocated to multiple partners.     

Parental Investment and Territorial/Sexual Behavior in Male and Female Reed Warblers: Are they Mutually Exclusive?

The behavioral and endocrine changes from prenesting to feeding phases of reproduction were examined in free-living reed warblers. Apart from the onset of feeding behavior, these changes in males are particularly evident in song, territorial behavior and circulating levels of testosterone. The interactions between an individual's physiology and behavior during these behavioral transitions were tested first by correlating song and feeding rates with circulating levels of testosterone and secondly by implanting feeding males with steroids and documenting the resulting effects on song, territoriality and parental behavior. The physiological mechanisms associated with the transition from the sexual phase (nest-building and egg-laying) to the parental (incubation and feeding) in females were also examined. In this case, estrogens were implanted during the parental phase to enhance sexual behavior. The results demonstrate an antagonistic relationship between sexual/territorial behavior and parental care in males which is mediated by changes in circulating androgens and no such antagonism in females. The effects of parental manipulation on the feeding rates of the partners and the growth rates of offspring have been examined to shed light on the necessity for male parental input.     

Territorial behavior in Southeast Asian langurs: Resource defense or mate defense?

Between-group antagonism or territoriality in primates may serve two different but compatible functions: resource defense or mate defense. Females are expected to be involved more strongly in the first, males in the second. The resource defense hypothesis predicts that home range overlap should decrease as defensibility and population density increase, and that females should be involved in hostile between-group interactions. The mate defense hypothesis predicts that between-group relations should be hostile and that males should take the primary role in antagonistic encounters. In a comparative study of 12 populations of 6 Presbytis species in Southeast Asia, we found support for the mate defense hypothesis; only males produce loud calls, between-group antagonism is entirely a male affair, and neither defensibility nor population density determine spatial exclusivity or the level of antagonism. We discuss the differences between our findings and traditional interpretations of territorial behavior.     

Female preference for dominant males in the Mexican mojarra cichlid fish,Cichlasoma istlanum

It is widely assumed that female preference and male competition operate simultaneously during sexual selection. Dominance is likely an honest indicator of male quality, and females can identify and choose the dominant male to reproduce with individuals with greater competitive abilities, thus improving the quality and competitiveness of their offspring. In this context, few studies have investigated female preference in relation to male fighting ability. The Mexican mojarra, Cichlasoma istlanum, is a cichlid species native to the Balsas River basin. It is territorial during reproduction and provides parental care. Males commonly engage in territorial defence, whereas females care directly for offspring. This study examined whether females prefer dominant males that exhibit more aggressive behaviour. The authors conducted experiments using groups of two males and one female to test competitive ability in males and female preference. They also quantified the time during which the female associated with the dominant male and the subordinate male after observing the outcome of a fight between the two males. They found that Mexican mojarra females preferred dominant males and that the time females spent associating with males was positively related with their aggressive behaviours during competition. These results indicate that dominant males were more attractive than subordinate males to female Mexican mojarra. The relationship between female preference and male dominance in the Mexican mojarra demonstrates the importance of male competitive ability for future parental care in reproduction.     

The evolution of avian parental care

A stage model traces key behavioural tactics and life-history traits that are involved in the transition from promiscuity with no parental care, the mating system that typifies reptiles, to that typical of most birds, social monogamy with biparental care. In stage I, females assumed increasing parental investment in precocial young, female choice of mates increased, female-biased mating dispersal evolved and population sex ratios became male biased. In stage II, consortships between mating partners allowed males to attract rare social mates, provided a mechanism for paternity assessment and increased female ability to assess mate quality. In stage III, relative female scarcity enabled females to demand parental investment contributions from males having some paternity certainty. This innovation was facilitated by the nature of avian parental care; i.e. most care-giving activities can be adopted in small units. Moreover, the initial cost of care giving to males was small compared with its benefit to females. Males, however, tended to decline to assume non-partitionable, risky, or relatively costly parental activities. In stage IV, altriciality coevolved with increasing biparental care, resulting in social monogamy. Approaches for testing behavioural hypotheses are suggested.     

Compensatory investment in zebra finches: females lay larger eggs when paired to sexually unattractive males

The classical version of the differential allocation hypothesis states that, when females reproduce over their lifetime with partners that differ in their genetic quality, they should invest more in reproduction with high-quality males. However, in species with lifetime monogamy, such as the zebra finch, partner quality will typically remain the same. In this case, the compensatory investment (CI) hypothesis predicts higher investment for low-quality males, because low genetic quality offspring are more dependent on maternal resources. Here, we show that female zebra finches invested more resources, both in terms of egg volume and yolk carotenoid content, when paired to a low genetic quality male, as judged from his previous ability to obtain extra-pair paternity in aviary colonies. We also found that females deposited slightly larger amounts of testosterone into eggs when paired to a low parental quality male, as judging from his previous success in rearing offspring. This is, to our knowledge, the first experimental support for the CI hypothesis in a species with lifetime monogamy. We stress that in more promiscuous species, the benefits of classical differential allocation may partly be neutralized by the supposed benefits of CI.     

Pair territoriality in the longnose filefish,oxymonacanthus longirostris

The longnose filefish,Oxymonacanthus longirostris, usually lives in heterosexual pairs, the male and female swimming together and sharing the same territory. Pair territoriality in the species was examined in detail in relation to sexual differences in territorial defense activities. Rigorous pair territoriality was maintained only during the breeding season, although pairs used their home ranges exclusively to a certain extent, during the non-breeding season. The frequency of aggression against other conspecific pairs in the breeding season was higher than in the non-breeding season. Agonistic interactions appear to be over both mates and food resources, the strict pair territoriality in the breeding season possibly being due to mutual mate guarding. In intraspecific aggressive interactions, males usually led their partner females when attacking intruders. The feeding frequency of males was much lower than that of females in the breeding season. Mate removal experiments indicated that females could not defend their original territories solitarily and their feeding frequency decreased. Conversely, males could defend territories solitarily without a decrease in feeding frequency. These results suggest that males contribute most to the defense of the pair territory, with females benefiting from territorial pair-swimming with their partner males.     

Territoriality and male reproductive success in arctic ground squirrels

Although territorial defense is a common form of reproductivecompetition among male vertebrates, the exact reproductiveconsequences of this behavior are often poorly understood.To explore relationships between territoriality and reproductivesuccess in a nongroup-living mammal, we quantified patterns of space use, mating success, and fertilization success formales in a free-living population of arctic ground squirrels(Spermophilus parryii plesius). Because litters of this speciesare sired almost exclusively by a female's first mate, we predictedthat territory ownership would be associated with first accessto estrous females. During the 2-week period when mating occurred,each male in the study population attempted to defend a distinctportion of the habitat, although the success of this defensevaried among individuals. Twenty-six of 28 females monitoredmated with the male on whose territory they resided. However,the majority of females observed throughout estrus (65%; n= 20) also mated with at least one other male, indicating thatterritory ownership was not associated with exclusive accessto females. In contrast, territory ownership was significantly associated with first access to estrous females; 20 (71.4%)of 28 females mated first with the male on whose territorythey resided. In this regard, the behavior of S. parryii plesiusparallels that of socially monogamous birds in which territorialdefense by males functions to deter extrapair copulations byfemales. Although territorial defense represents an important component of male reproductive success in arctic ground squirrels,other aspects of male behavior (e.g., the ability to dominateagonistic interactions on the day of a female's estrus) arealso critical. We suggest that future studies of vertebratemating systems will benefit by viewing such defense as onlyone of multiple axes along which conspecific males compete foraccess to females.     

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.     

Mother's little helpers: Patterns of male care in mammals

Over the past 20 years, long-term studies have shown that mammalian fathers, brothers and even unrelated males may contribute substantially to the care of the young. Recently, more field studies of mammals with male care have further expanded interest in male care potential. Male mammals can never be sole care-givers because offspring depend on maternal milk, but assistance provided by males may allow females to produce more energetically costly litters. The need for such male care contributes to the development of obligate social monogamy or communal breeding.     

Impact of season and social challenge on testosterone and corticosterone levels in a year-round territorial bird

Plasma testosterone increases during breeding in many male vertebrates and has long been implicated in the promotion of aggressive behaviors relating to territory and mate defense. Males of some species also defend territories outside of the breeding period. For example, the European nuthatch (Sitta europaea) defends an all-purpose territory throughout the year. To contribute to the growing literature regarding the hormonal correlates of non-breeding territoriality, we investigated the seasonal testosterone and corticosterone profile of male (and female) nuthatches and determined how observed hormone patterns relate to expression of territorial aggression. Given that non-breeding territoriality in the nuthatch relates to the reproductive context (i.e., defense of a future breeding site), we predicted that males would exhibit surges in plasma testosterone throughout the year. However, we found that males showed elevated testosterone levels only during breeding. Thus, testosterone of gonadal origin does not appear to be involved in the expression of non-breeding territoriality. Interestingly, territorial behaviors of male nuthatches were stronger in spring than in autumn, suggesting that in year-round territorial species, breeding-related testosterone elevations may upregulate male-male aggression above non-breeding levels. In females, plasma testosterone was largely undetectable. We also examined effects of simulated territorial intrusions (STIs) on testosterone and corticosterone levels of breeding males. We found that STIs did not elicit a testosterone response, but caused a dramatic increase in plasma corticosterone. These data support the hypothesis that corticosterone rather than testosterone may play a role in the support of behavior and/or physiology during acute territorial encounters in single-brooded species.     

Genetic monogamy in Monteiro's hornbill, Tockus monteiri

Hornbills display a unique breeding habit in which the female seals herself into the nest cavity prior to egg laying and remains ensconced for most of the breeding cycle. This habit necessitates long-term sperm storage, which considerably lengthens the fertile period of the female. This in turn increases the amount of time available to females to engage in extrapair copulations. Because food delivered by the male is the primary source of energy for female maintenance, egg production, incubation, female moult and offspring provisioning, male parental investment (and thus cuckoldry) in these hornbills is extremely expensive. This combination of costly male parental care and female dependence potentially places hornbills at a critical endpoint in the spectrum of genetic promiscuity operating within pairs of socially monogamous birds. A paradigm of reproductive conflict suggests that these factors might allow males to coerce females into a state of genetic monogamy. Sperm storage, however, might allow female hornbills to thwart monopolization attempts by their mates. We used minisatellite multilocus DNA fingerprinting to analyse paternity in a Namibian population of Monteiro's hornbill, Tockus monteiri. Despite the ability of female hornbills to store sperm for long periods, we found no evidence of extrapair paternity (EPP). Although we cannot explicitly rule out a role for coercion in the evolution of the genetic monogamy we observed, the benefits of EPP appear to be devalued to the point that the reproductive interests of males and females overlap considerably. Copyright 2002 The Association for the Study of Animal Behaviour. Published by Elsevier Science Ltd. All rights reserved.     

Genetic analysis of male reproductive success in relation to density in the zebrafish, Danio rerio

Background

We used behavioural and genetic data to investigate the effects of density on male reproductive success in the zebrafish, Danio rerio. Based on previous measurements of aggression and courtship behaviour by territorial males, we predicted that they would sire more offspring than non-territorial males.

Results

Microsatellite analysis of paternity showed that at low densities territorial males had higher reproductive success than non-territorial males. However, at high density territorial males were no more successful than non-territorials and the sex difference in the opportunity for sexual selection, based on the parameter I _mates, was low.

Conclusion

Male zebrafish exhibit two distinct mating tactics; territoriality and active pursuit of females. Male reproductive success is density dependent and the opportunity for sexual selection appears to be weak in this species.     

Sex differences in biparental care as offspring develop: a field study of convict cichlids (<Emphasis Type="Italic">Amatitlania siquia</Emphasis>)

Parental investment theory states that parents should contribute more to older offspring. Differences between the sexes also influence how each parent contributes to offspring in biparental species. Here, we examined a naturally occurring population of biparental convict cichlids in Costa Rica to determine how each parent cared for offspring during two distinct offspring development stages. Consistent with the predictions of the reproductive value hypothesis, we hypothesized that the levels of parental contribution would be relative to the value that each parent places on a brood. We predicted that female parents would contribute more than male parents because female convict cichlids have lower future reproductive success than males. Additionally, we predicted that both parents should contribute more to older offspring, either due to the young’s increased susceptibility to predation (i.e., the vulnerability hypothesis) or because of the longer period of time parents have been interacting with older offspring (i.e., feedback hypotheses). This increase in investment by males should coincide with a change in the coordination of care between parents. Detailed observations of parental pairs in their natural habitat supported these predictions. Females contributed more to broods than males and were relatively unaffected by offspring age while males spent significantly more time with older, free-swimming fry. Additionally, males tended to leave younger offspring more than females did, and were more likely to do so consecutively with younger offspring. This suggests that the coordination of duties between parents changes as parental investment changes. Overall, these data support both the reproductive value and the vulnerability hypotheses, but not necessarily the feedback hypothesis.