Space,density and extra-pair matings have opposing impacts on male and female reproductive success

Many territorial species have a mating system characterized by males establishing home ranges in the breeding grounds prior to females, resulting in males competing for territories and females choosing a mate upon their arrival. It remains unknown, however, how the outcomes of decisions surrounding territory establishment and mate choice are influenced by the spatial configuration of the breeding grounds. We use a spatially explicit, individual-based model to investigate the sex-specific effects of these decisions on reproductive success. In our model, males that arrive earlier obtain higher quality territories and improve their chances for extra-pair copulations. Females can choose their mate to maximize the quality of the male or to attempt to minimize the density of other females near their nesting site to avoid competition. Females therefore face a tradeoff between high-density regions around high-quality males and low-quality males in areas of low competition. Our model predicts a negative correlation between male and female reproductive success under a wide range of conditions when the majority of the territories are on the margins of the breeding area. Most notably, this sexual conflict arises as an edge effect suggesting that fragmentation of breeding habitats could impact the consequences of mate choice in many species with territorial breeding habits.     

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.     

Genetic monogamy in single-male groups of acorn woodpeckers, Melanerpes formicivorus

Acorn woodpeckers have one of the most complex social systems of any bird species. Breeding units range in size from monogamous pairs to groups of 15 birds that include multiple breeding males and females as well as nonreproductive helpers-at-the-nest. Groups form when young remain at their natal nest to help their parents breed or when single-sex coalitions of siblings disperse to fill a reproductive vacancy on another territory. Plural breeding and helping behaviour are thought to be favoured through indirect fitness benefits for individuals that would otherwise be unable to breed due to a shortage of reproductive vacancies on territories with acorn stores. We report the results of multi-locus DNA fingerprinting of 51 offspring from 18 nests of 16 socially monogamous pairs of acorn woodpeckers. If socially monogamous females mate outside the pair-bond, indirect fitness benefits for cobreeders and helpers will be significantly reduced. Monogamous pairs accounted for all but one of the 51 offspring we tested; the single exception was apparently sired by the putative father, but the putative mother was excluded from maternity. Our results indicate that individuals remaining on their natal territories as helpers are generally the genetic offspring of the pair they help. They also suggest that single-sex coalitions offspring dispersing together from nests of socially monogamous pairs will be full-siblings.     

Mate choice by marsh wrens: the influence of male and territory quality

If females choose breeding situations to maximize their fitness, then those features of the male and/or territory that are important in mate selection by females should affect female fitness, be assessable prior to mating, and vary among males. The purpose of this study was to identify characteristics of male quality and territory quality that fit these criteria for marsh wrens Cistothorus palustris. The male's contribution to nest defence appeared not to affect female success or choice. However, females that received male assistance with feeding young produced more and heavier fledglings than females without assistance. Males that fed young did not attract more mates. Few males in this population fed young, and no relationship was found between feeding effort and physical or behavioural features of the male. Results of multivariate analyses suggest that, within sites, the measures of territory quality used here cannot explain the variance in the number of young fledged per female or male harem size. The examination of arbitrary territories showed that male pairing success, female settlement order and predation patterns were not significantly correlated between years. The results suggest that territory quality does not influence female success or choice. These results are discussed in the light of earlier attempts to relate features of males and territories to mate selection by female passerines.     

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.     

Multiple Ornaments in Male Northern Cardinals, Cardinalis cardinalis, as Indicators of Condition

Investigations of male ornaments in the context of sexual selection have tended to focus on single ornaments, although many species of birds possess multiple ornaments. Understanding the evolution of multiple ornaments requires knowledge of correlations among ornaments in the same individual and the extent to which ornament expression reflects individual condition and behavior. Variation in four male ornaments in socially monogamous, biparental northern cardinals (Cardinalis cardinalis) was related to body size, indices of condition, level of paternal care, and reproductive success. Redness of breast plumage positively predicted body size and negatively predicted nestling feeding rate. Bill color predicted current body condition, with birds with redder bills in better condition. Birds with smaller black face masks had greater reproductive success. These results are consistent with the hypothesis that different ornaments in male cardinals provide information on different aspects of condition and behavior.     

Costs and benefits of variable breeding plumage in the red-backed fairy-wren

The red-backed fairy-wren is a socially monogamous passerine bird which exhibits two distinct types of breeding male, bright males that breed in bright red and black plumage and dull males that breed in dull brown plumage. Most males spend their first potential breeding season in dull plumage and subsequent breeding seasons in bright plumage, but a relatively small proportion of males develop bright plumage in their first breeding season. This study quantifies morphology, behavior, and reproductive success of dull and bright males to assess the adaptive costs and benefits of bright plumage while controlling for age. Older bright males (two years of age or older) attempted to increase their reproductive success via copulations with extrapair females, whereas younger (one-year old) bright males and dull males did not. Thus, older bright males spent less time on their own territories, intruded on neighboring groups with fertile females more frequently, gave more courtship displays, and had larger sperm storage organs than did younger bright males and dull males. Microsatellite analyses of paternity indicate that the red-backed fairy-wren has extremely high levels of sexual promiscuity, and that older bright males had higher within-brood paternity than dull males or younger bright males. Regardless of age, bright males were more attractive to females in controlled mate choice trials than were dull males, and both age classes of bright males obtained higher quality mates earlier in the breeding season than did dull males, when nesting success was higher. In conclusion, although it appears that bright plumage increases access to higher quality mates, age also plays a central role in determining a male's overall reproductive success because of the high levels of sexual promiscuity exhibited by the red-backed fairy-wren.     

Mate acquisition tactics in polyandrous spotted sandpipers (Actitis macularia): the role of age and experience

We consider resource-defense polyandry and mate-access polyandryas female mating tactics in spotted sandpipers (Actitis macularia).These tactics can be distinguished by the resource females defend,female interclutch movement, expected reproductive success aftermoving, and male and female dispersion. We examine these characteristicsrelative to patterns observed in a 17-year study of spottedsandpipers, a species traditionally considered resource-defensepolyandrous. On average, 26% of spotted sandpiper females eachyear were monogamous. Older females were more likely to be polyandrous,and polyandrous females of each age employed different matingtactics. Yearlings were typically sequentially resourcedefensepolyandrous. Two-year-olds were primarily simultaneously polyandrous,exhibiting equivalent proportions of resource-defense and mate-accesspolyandry. Older females were primarily simultaneously resource-defensepolyandrous. Females tended to stay on territories where theyand/or their mates had greater breeding experience (i.e., manyclutches laid for females, many clutches diat hatched for males);females that moved went to territories where their mates hada history of breeding success. Location changes between clutchesby polyandrous females were better described by breeding experienceon a territory than by age.     

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.     

Reproductive success and survivorship of the Galapagos Hawk Buteo galapagoensis: potential costs and benefits of cooperative polyandry

The Galapagos Hawk Buteo galapagoensis is one of few species showing cooperative polyandry. As many as four males may mate with one female and share a territory and nest, Such groups, on average, produce more young than monogamous pairs, but on a per male basis (assuming that each male of a group has an equal chance to fertilize each young) polyandrous males produce fewer young per year than monogamous males. Territorial breeding birds show higher yearly survivorship (90%) than non-territorial, non-breeding birds (50% or less). All territorial birds seem to remain on their territories for life, and monogamous pairs actually constitute the remnants of polyandrous groups. Some potential factors producing such an unusual system are discussed. Limited breeding space and high mortality among non-territorial birds seem to work in favour of group breeding in this and other cooperative species. Polyandry may be favoured because of the reduced value of extra females in hawk breeding behaviour. Equal-status males may occur because of the lack of kinship among cooperating males and the high potential genetic cost of being a helper in a long-lived, low fecundity species. Other possible factors favouring group breeding are territory acquisition and the variable nature of the Galapagos climate.     

Territory quality drives intraspecific patterns of extrapair paternity

Despite the potential reproductive benefits of extrapair matings,extrapair paternity rates in many avian species often vary greatlyamong populations. Although ecological factors have been shownto influence intraspecific patterns of extrapair paternity insome species, for cooperatively breeding species living in familygroups, social/demographic factors may also play a role. Thisstudy examined how ecological factors related to territory quality(vegetation cover, insect abundance) and social/demographicfactors (group size, number of breeding pairs, genetic relatedness)influenced intraspecific patterns of extrapair paternity incooperatively breeding superb starlings, Lamprotornis superbus.Superb starlings inhabit spatiotemporally variable African savannaswhere high temporal variability drives reproductive decisions(adoption of breeding roles, offspring sex allocation) and whereterritories suitable for breeding are limited. Although extrapairpaternity rates were only 14% of offspring and 25% of nests,they varied greatly among groups, ranging from 4% to 32% ofoffspring and from 7% to 60% of nests. These among-group differencesin extrapair paternity were not related to social/demographicfactors but instead to territory quality; extrapair paternitywas higher on lower quality territories (lower vegetation coverand grasshopper abundance) than on higher quality territories(higher vegetation cover and grasshopper abundance). These resultssuggest that even in a heterogeneous landscape where suitablebreeding territories are limited, subtle differences in habitatquality can have profound effects on reproductive decisionsand patterns of extrapair paternity. Understanding the interactionbetween spatial (habitat heterogeneity) and temporal (temporalvariability) environmental variation will be important for determininghow environmental and social factors drive avian reproductiveand mating decisions.     

Male Seychelles warblers use territory budding to maximize lifetime fitness in a saturated environment

In cooperatively breeding species, helping at the nest and buddingoff part of the natal territory have been advanced as strategiesto increase fitness in an environment that is saturated withterritories. The importance of helping or territory buddingas a determinant of lifetime reproductive success (LRS) hasbeen debated because the potential benefits of both strategiescould not be separated. Here we test the causes and the immediateand future fitness consequences of single dispersal decisionstaken by male Seychelles warblers (Acrocephalus sechellensis).Males breeding in high-quality territories (high food abundance)have significantly higher LRS than similar-aged males buddingoff part of the parental territory. Initially, budders havea low reproductive success (because of limited food resourcesor absence of a breeding partner). However, they have a longlife span and inherit high-quality territories through sitedominance, by which they gain higher LRS than breeders on low-qualityterritories, helpers, or floaters. Experimental creation ofmale breeding territory vacancies showed that most young malesbecame budders because of intense competition for high-quality territories. The translocation of warblers to the previouslyunoccupied Aride Island shows that males behave according tothe expected fitness benefits of each dispersal strategy. Inthe absence of competition for territories on Aride, all youngmales bred in high-quality territories. However, after saturationof high-quality habitat with territories, most males becamebudders rather than breeders on low-quality habitat, helpers,or floaters.     

Monogamy in marine fishes

The formation of long-term pair bonds in marine fish has elicited much empirical study. However, the evolutionary mechanisms involved remain contested and previous theoretical frameworks developed to explain monogamy in birds and mammals are not applicable to many cases of monogamy in marine fish. In this review, we summarise all reported occurrences of social monogamy in marine fish, which has so far been observed in 18 fish families. We test quantitatively the role of ecological and behavioural traits previously suggested to be important for the evolution of monogamy and show that monogamous species occur primarily in the tropics and are associated with coral reef environments in which territory defence and site attachment is facilitated. However, there is little evidence that obligately monogamous species are smaller in body size than species that can adopt a polygynous mating system. We review the evidence pertaining to six hypotheses suggested for the evolution of monogamous pair bonds: (1) biparental care, (2) habitat limitation, (3) low population density/low mate availability/low mobility, (4) increased reproductive efficiency, (5) territory defence, and (6) net benefit of single mate sequestration. We outline predictions and associated empirical tests that can distinguish between these hypotheses, and assess how generally each hypothesis explains monogamy within and between breeding periods for species with different types of territories (i.e. feeding only or feeding and breeding). Hypotheses (1) and (2) have limited applicability to marine fishes, while hypotheses (3)-(5) have little empirical support beyond the species for which they were designed. However, the role of paternal care in promoting monogamous pair bonds is not explicit in these hypotheses, yet paternal care has been reported in more than 70 monogamous marine fish. We show that paternal care may act to increase the likelihood of monogamy in combination with each of the proposed hypotheses through decreased benefits to males from searching for additional mates or increased advantages to females from sequestering a single high-quality mate. Among species defending breeding and feeding territories, the benefits, both within and between reproductive periods, of sequestering a single high-quality mate (hypothesis 6) appear to be the best explanation for socially monogamous pairs. For species without parental care (i.e. holding only feeding territories), territory defence (hypothesis 5) in combination with the benefits of guarding a large mate (hypothesis 6) could potentially explain most instances of monogamy. Empirical studies of marine fishes over the past two decades are therefore slowly changing the view of monogamy from a mating system imposed upon species by environmental constraints to one with direct benefits to both sexes.     

Starting over: experimental effects of breeding delay on reproductive success in early‐arriving male American redstarts

Birds that arrive and breed early often have higher reproductive success than late individuals, either as a consequence of timing‐specific advantages (the timing hypothesis) or because these individuals and/or their resources are of higher quality (the quality hypothesis). In this study, we examined the potential influence of several factors affecting reproductive success by experimentally delaying breeding of early‐arriving male American redstarts Setophaga ruticilla, a species for which early male arrival is strongly related to increased reproductive success. Our manipulation involved the capture, holding, and release of males following pairing and territory establishment, resulting in the majority of subjects (67%) losing their initial mate (47%) or mate and territory (20%) and forcing them to start over approximately 12 d after their initial arrival. Males forced to start over (i.e. those losing their first territory and/or mate) did not experience any decrease in body condition, nor did their reproductive behaviour differ from that of early‐arriving control males. We found that naturally early‐arriving but experimentally manipulated males suffered reduced fledging success in comparison to early‐arriving males that bred early or late, but equivalent success in comparison to males that arrived and bred late. Based on our results, we propose that the relationship between early arrival and higher reproductive success in this species is mediated not simply by individual male quality or absolute arrival timing alone, but rather some other aspect of resource quality is likely important. We discuss and present evidence for two alternative explanations under the quality hypothesis: female quality and territory quality. To our knowledge, our study is the first to investigate the effects of experimentally delaying male breeding time, strengthening previous correlational evidence for resource quality as a potentially important selective agent driving early arrival in migratory birds.     

Relationship between arrival timing and breeding success of intra‐tropical migratory Fork‐tailed Flycatchers (Tyrannus savana)

For migratory birds, early arrival at breeding areas has many benefits, such as acquisition of better territories and mates. This strategy has been found in numerous species breeding at north‐temperate latitudes, but has not been yet reported for intra‐tropical migratory species. We evaluated the relationship between arrival date, initiation of breeding, and breeding success of Fork‐tailed Flycatchers (Tyrannus savana) breeding in southeastern Brazil and overwintering in northern South America. We color‐banded adult flycatchers during three breeding seasons and searched for them during the following breeding seasons. We also monitored nests from construction until either failure or fledging of young. We found that: (1) male Fork‐tailed Flycatchers arrived at the breeding site earlier than females, (2) males that arrived earlier had greater breeding success, and (3) nests where eggs were laid earlier in the breeding season were more likely to be successful than those where eggs were laid later. Male Fork‐tailed Flycatchers appeared to benefit from early arrival at a tropical breeding site, potentially mediated by their ability to acquire a high‐quality territory and mate as early as possible, and by the ability of their mate to begin breeding as early as possible. Breeding success for female Fork‐tailed Flycatchers may be determined primarily by a combination of the arrival date of their mate and how quickly they can begin breeding. Our results suggest that protandry occurs in an intra‐tropical migratory bird and that early arrival of males and early initiation of reproduction by females results in greater reproductive success. A better understanding of the underlying mechanisms that control the timing of migration and reproduction of this and other intra‐tropical migratory species is important for evaluating the challenges they face in light of current and future rapid environmental changes.     

Reproductive success in male Japanese minnows,Pseudorasbora parva: observations under experimental conditions

To evaluate the spawning success of male Japanese minnows,Pseudorasbora parva, and female mate choice, spawning behaviour was observed under both artificial and experimental conditions. Larger males had larger territories and greater reproductive success. The body weight of territorial males decreased during the maintenance of territories, while that of non-territorial males increased significantly. When the weight of non-territorial males exceeded that of territorial males, the former began to establish new territories on the substrate, suggesting a conditional strategy by non-territorial males to trade off immediate reproductive success with growth and hence improve future reproductive success. Females chose males with larger body size, probably based on dominance rank rather than the quality (or size) of territory. It was concluded that females choose males of higher dominance rank and that males compete for large territories, both of which play an important part in male reproductive success.     

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.     

EXTRAPAIR MATE CHOICE AND HONEST SIGNALING IN COOPERATIVELY BREEDING SUPERB FAIRY-WRENS

In many species of monogamous birds females copulate with males other than their social mates, resulting in extrapair fertilizations. Little is known about how females choose extrapair mates and whether the traits used to choose them are reliable indicators of male quality. Here we identify a novel male trait associated with extra-group mating success in the superb fairy-wren (Malurus cyaneus), a cooperatively breeding bird with one of the highest known frequencies of extra-group mating. Female fairy-wrens chose extra-group mates that molted earlier into breeding plumage. Males molted up to five months before the breeding season began, and only males that molted at least one month prior to its onset gained any extra-group fertilizations. This conclusion held after controlling statistically for the effect of age and social status on molt date. Once males acquired breeding plumage, they began courtship display to females on other territories. Thus, some males were displaying to females for several months before the breeding season began. This extraordinarily long period of advertisement by males may be facilitated by the long-term ownership of territories. We suggest that early acquisition of breeding plumage or the subsequent display behavior can be reliable cues for mate choice because they are costly to acquire or maintain.     

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.     

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.