Sexual selection in the monogamous barn swallow (Hirundo rustica). II. Mechanisms of sexual selection

Mechanisms of sexual selection in the monogamous, sexually dimorphic barn swallow (Hirundo rustica) were studied during a seven-year period. First, the sex ratio of reproducing adults was male-biased, and mated males had significantly longer tail ornaments than unmated males. Secondly, some of the unmated individuals later committed infanticide and became mated with the mother of the killed brood. Fathers of killed broods had significantly shorter tails than other males, and there was a tendency for infanticidal males to have longer tail ornaments than other unmated males. Thirdly, long-tailed male barn swallows were more successful in acquiring extra-pair copulations than other males, and females involved in extra-pair copulations, as compared to females not involved in such copulations, had mates with shorter tail ornaments. Fourthly, male barn swallows having long tails as compared to short-tailed males acquired mates in better body condition. Females mated to long-tailed males reproduced earlier, laid more eggs and were more likely to have two clutches than were females mated to short-tailed males. Finally, females mated to long-tailed males put more effort into reproduction than did other females, as evidenced by their relatively larger contribution to feeding of offspring. Thus, at least five different components of sexual selection affected male reproductive success. Selection arising from differential success during extra-pair copulations, differential reproductive success and differential male reproductive effort thus accounted for most of the selection on tail ornaments in male barn swallows.     

Do female great tits (Parus major) assess males by eavesdropping? A field study using interactive song playback

Male singing behaviour correlates with extra-pair success in several passerine birds. Singing interactions during territorial contests provide relative information on the males involved. Such information may be important in female extra-pair behaviour and eavesdropping on singing interactions among males may allow females to make such relative assessments. We used interactive playback to instigate singing contests with male great tits during the peak fertile period of their mate in an attempt to alter females'' assessment of mates'' quality relative to neighbours (potential extra-pair partners). We escalated a contest to one male (by overlapping his songs) and then subsequently de-escalated a contest (by alternating) to a neighbour. Intrusions onto neighbouring territories by females mated to either treatment male were then monitored. Females mated to escalation treatment males were more likely to intrude following playbacks than females mated to de-escalation treatment males. Although the absolute song output of males did not differ between treatments, males produced more song relative to playback in de-escalation treatments and relative song output was positively correlated with female intrusions. Therefore, female great tits eavesdrop on singing interactions and change their visitation rates to neighbouring territories according to their mate''s singing performance relative to neighbours.     

Intraspecific differences in benefits from feeding in mixed-species flocks

The Madagascar Paradise Flycatcher Terpsiphone mutata and Common Newtonia Newtonia brunneicauda frequently form two-species flocks in the deciduous dry forest of western Madagascar. In T. mutata , some males have long tails, while other males and females have short tails. When foraging in mixed flocks, each type of bird captured prey more rapidly than otherwise, but the degree of increase in feeding rate was smaller in long-tailed males. When in mixed flocks, all T. mutata caught prey on leaves in the canopy where N. brunneicauda foraged. Long-tailed males changed feeding habits from sallying when not in mixed flocks, whereas short-tailed birds showed no change of feeding habit. The elongated tails of long-tailed males may have made their foraging less efficient owing to decreased agility in the canopy. N. brunneicauda is monomorphic and often formed groups of three to five individuals. In monospecific flocks, subordinates fed at low rates on branches owing to frequent hostile encounters. When foraging in mixed flocks, however, subordinates foraged among leaves, and their feeding rates increased because the frequency of intraspecific interference decreased greatly. Dominants did not show any difference in feeding pattern with social situation. Thus, heterospecific flocking was more advantageous for subordinates.     

保加利亚北部地区匈牙利小家鼠的空间组织结构

本文研究了保加利亚北部地区的匈牙利小家鼠在繁殖期的空间关系和分布。这种动物最普遍的空间关系是一雄一雌,也观察到两个或三个雄鼠与一个雌鼠、一个雄鼠与两个或三个雌鼠的组织形式。很少观察到雌性的社群小组。雌鼠之间的巢区没有发现重叠。一般雄鼠的巢区大于雌鼠。有一半雄鼠的巢区与一个雌鼠的巢区相重叠,其他的雄鼠与两个成年雌鼠的巢区相重叠。这些结果表明,匈牙利小家鼠具有多变的社群繁殖婚配体制,即单配制和多配制共存。比较普遍的一雄一雌的配对形式和雌性的独居特性,以及雌性之间巢区的相对独立性,表明单配制是该物种的主要婚配体制。     

Breaking the rules: sex roles and genetic mating system of the pheasant coucal

Generally in birds, the classic sex roles of male competition and female choice result in females providing most offspring care while males face uncertain parentage. In less than 5% of species, however, reversed courtship sex roles lead to predominantly male care and low extra-pair paternity. These role-reversed species usually have reversed sexual size dimorphism and polyandry, confirming that sexual selection acts most strongly on the sex with the smaller parental investment and accordingly higher potential reproductive rate. We used parentage analyses and observations from three field seasons to establish the social and genetic mating system of pheasant coucals, Centropus phasianinus, a tropical nesting cuckoo, where males are much smaller than females and provide most parental care. Pheasant coucals are socially monogamous and in this study males produced about 80% of calls in the dawn chorus, implying greater male sexual competition. Despite the substantial male investments, extra-pair paternity was unusually high for a socially monogamous, duetting species. Using two or more mismatches to determine extra-pair parentage, we found that 11 of 59 young (18.6%) in 10 of 21 broods (47.6%) were not sired by their putative father. Male incubation, starting early in the laying sequence, may give the female opportunity and reason to seek these extra-pair copulations. Monogamy, rather than the polyandry and sex-role reversal typical of its congener, C. grillii, may be the result of the large territory size, which could prevent females from monopolising multiple males. The pheasant coucal’s exceptional combination of classic sex-roles and male-biased care for extra-pair young is hard to reconcile with current sexual selection theory, but may represent an intermediate stage in the evolution of polyandry or an evolutionary remnant of polyandry.     

The adaptive significance of monogamy in the golden-reumped elephant-shrew

Golden-rumped elephant-shrews Rhynchocyon chrysopygus are primarily monogamous, males and females defending joint territories for long periods and probably for life. However, males may occasionally solicit extra-pair copulations from neighbouring females and annex their territories if the resident male disappears for any reason. Males provide no direct paternal care, and the adaptive significance of monogamy in this species is unclear. Although it is possible that males provide some indirect benefits to the female and her offspring, these are unlikely to be substantial. Males and females spend little time together, so males are unlikely to provide additional protection from predators, and there is no evidence that the presence of a territorial male reduces the costs of territorial defence for the female. Females were able to breed successfully when mated bigamously, demonstrating that the undivided assistance of the male is not essential for successful rearing of offspring. The fact that male elephant-shrews occasionally attempt to defend more than one female suggests that polygyny would be beneficial for males. However, defending two territories is costly, resulting in increased activity and weight loss, and higher rates of intrusion by neighbouring males. Female elephant-shrews do not occupy particularly large home ranges for their body size. The results suggest that the high costs of defending a territory large enough to encompass the ranges of more than one female, combined with the greater probability of being cuckolded, make defending more than one female a poor option for male elephant-shrews.     

Social Organization of Lepilemur ruficaudatus

We characterize the spatial organization of red-tailed sportive lemurs (Lepilemur ruficaudatus) as a key aspect of their social organization and social system. Sportive lemurs are small (<1000 g), nocturnal and folivorous primates endemic to Madagascar. We studied a population of 57 individually-marked individuals in Kirindy Forest, western Madagascar, between 1995 and 2001. We radio-tracked 20 males and 26 females of the marked population to obtain detailed information on the size and location of their home ranges. Census data and morphometric measurements provided complementary data sets. Males and females occupied small (<1 ha) home-ranges. Long-term records from 9 individuals revealed home-range stability over several years. In 4 cases home ranges overlapped extensively with that of one member of the opposite sex; in 2 cases, a spatial association of 1 male and 2 females occurred. However, home ranges overlapped very little with neighboring individuals of both sexes. During the study period, spatially associated individuals used on average 5.6 sleeping trees within 117 days, but they spent on average only about every fourth night together. The data suggest that home ranges in red-tailed sportive lemurs are exclusively used by pairs or trios and that the modal social organization of red-tailed sportive lemurs is pair-living.     

The effect of breeding density and male quality on paternity-assurance behaviours in the house sparrow, <Emphasis Type="Italic">Passer domesticus</Emphasis>

Several factors can influence the risk of cuckoldry through extra-pair paternity for male birds. The number of neighbouring males is thought to affect the chance of females engaging in extra-pair copulations, and species which breed both socially (colonially) and solitarily provide an ideal opportunity to test the effect of close proximity on extra-pair behaviour and paternity guards. In this study, the extent to which male house sparrows, Passer domesticus, used two alternative strategies, namely frequent copulation and mate-guarding, to ensure paternity was investigated. We also examined how males vary the two paternity guards according to their breeding sociality. Pairs at the dense colony started to copulate at a higher rate at the beginning of the fertile period than those of the medium-sized colony and solitary breeding pairs. Male house sparrows appear to fine-tune their strategies according to the breeding density. Both strategies are alternatively used in the weak fertile period but are simultaneously used in the peak fertile period. Our results suggest that males modify their strategy according to their individual abilities: mate-guarding intensity was positively correlated with the black breast badge size.     

Off-territory movement of male American Redstarts (Setophaga ruticilla) in a fragmented agricultural landscape is related to song rate, mating status and access to females

Male songbirds often move off-territory to pursue extra-pair fertilizations. This movement represents a trade-off between paternity gain and loss and can be influenced by male quality and access to fertile females. Access to females could be reduced in fragmented landscapes that have small patches and low connectedness. We studied movement and extra-pair fertilization success of radio-tracked male American Redstarts (Setophaga ruticilla) in forest patches in an agricultural landscape in Alberta, Canada, over 2 years. Males spent an average of 18% of their time off-territory, mostly intruding onto adjacent territories and rarely moving between patches. They averaged 0.8 trips/h, with mean trip duration of 17 min and mean trip distance of 104 m. Less time was spent off-territory when their mate was nest-building and males intruded most often onto territories with nest-building females. Males with higher song rates and more nearby females intruded most onto other territories. Monogamous males in better condition with higher song rates spent the most time off-territory. However, males with more nearby females and higher local breeding synchrony spent the least time off-territory, suggesting these males face a trade-off between seeking extra-pair fertilizations and protecting against cuckoldry. Forest cover was not an important predictor of movement. Investment in off-territory movement did not predict extra-pair fertilization success or probability of cuckoldry. However, few tracked males achieved extra-pair fertilizations (1/22 tracked males vs 18/57 non-tracked males), possibly an artefact of low sample size or the effect of radio transmitters on female choice.     

Spacing pattern in a social group of stray cats: effects on male reproductive success

Reproductive consequences of male spacing patterns have received relatively little attention in nonterritorial mammals, in particular in group-living species, where most studies have focused on the relation between social rank and reproductive success. We investigated the effects of spacing pattern on male reproductive success within a social, nonterritorial, promiscuous population of stray cats, Felis catus. Male home ranges overlapped home ranges of many females, consistent with a promiscuous mating system. Furthermore, males with the largest home ranges included the most female home ranges; they successfully reproduced with these females and had the highest reproductive success. Home range size predicted male reproductive success even when controlling for the effect of social rank. However, males also reproduced with females whose home range did not overlap their home range, suggesting that males can make quick excursions outside their home range to find new mating opportunities. We conclude that, in group-living situations, a male's ability to maintain a large home range may be one of the principal causes of variation in mating success in the stray cat.     

Intra-group home ranges in a female-biased group of sex-changing fish

Simultaneous loss of multiple males from large, multi-male social groups of the coral reef fish, Anthias squamipinnis, induces an equivalent number of females to change sex. One-to-one replacement could be produced in large groups if the group was organized according to either of two hypotheses. The group could consist of an aggregation of single-male, multi-female units with the home ranges of particular females falling within the home ranges of each male, and with sex change controlled separately within each unit. Alternatively, females might range freely throughout the group, with sex change controlled by differential interaction rates among males and females. In the latter case, female home ranges would overlap substantially with the home ranges of several males. To choose between these hypotheses, the three-dimensional, intra-group home ranges of five males and five adult females were measured in a group containing five males and 32 adult females. All 10 fish had definable home ranges. Home ranges of females were larger than home ranges of males. All females ranged much farther outside of the home range of individual males than inside, and most females spent most of the observation time outside of the home range of any one male. These results effectively disproved the hypothesis of spatially-defined, single-male, multi-female units. Rather, large groups consist of females ranging freely over the discrete home ranges of several males.     

Space Use and Social Mating System of the Hantavirus Host, <Emphasis Type="Italic">Oligoryzomys longicaudatus</Emphasis>

The long-tailed mouse, Oligoryzomys longicaudatus (Cricetidae: Sigmodontinae), is the major host of Andes hantavirus, the etiological agent of hantavirus pulmonary syndrome in the south of Argentina and Chile. Studying the ecology of this species is necessary to understand how Andes hantavirus is maintained in nature. In this study, we examine the home range size and intra- and intersexual overlap degree of male and female O. longicaudatus in order to elucidate the mating system of this species. To our knowledge, this research provides the first documentation, obtained from a specific design, of spacing and mating systems in this species in Argentina. The study was conducted seasonally from April (autumn) 2012 to October (spring) 2013 in a shrubland habitat of Cholila, Andean region, Argentina. We studied spacing patterns using 59 and 51 home ranges established by adult males and females, respectively, in two 3.24 ha capture-marked and recapture grids. Significant differences between sexes in home range size and overlap degree were found. Male home ranges were always larger than those of females. We observed exclusive space use both among males and females (13.15 ± 18.67, and 3.60 ± 3.43%, respectively). Considering only those males that get access to receptive females (40%), average intersexual overlap value was about 30.82 ± 19.73%. Sexual differences in home range sizes and the spatial avoidance between breeding males, that would reflect intrasexual competition for receptive females, allows us to propose a polygynous mating system for O. longicaudatus.     

The Round-Eared Sengi and the Evolution of Social Monogamy: Factors that Constrain Males to Live with a Single Female

Animal dispersion in space and time results from environmental pressures, and affects the outcome of a species' social organization. When females are solitary, males may either roam or be pair-living. We studied possible environmental influences affecting the social organization of the round-eared sengi ( Macroscelides proboscideus ) in a semi-desert in South Africa, using trapping and radio-tracking across 2.5 yr. Adult sex ratios did not deviate from 1:1 and we found no indication of sexual dimorphism in body mass. Females maintained exclusive areas, which had little overlap (<4%) with neighbouring females (NF), and males overlapped predominately only with the home range of single females. Generally, inter- and intra-sexual overlap with neighbouring individuals was low (3–6%) for both sexes, indicating territoriality and pair-living. Pairs were perennial and territories were maintained year-round. However, males generally maintained much larger areas than females, which were sensitive to population density. Male space use appeared to be primarily limited by the presence of neighbouring males. Female home ranges were smaller-sized despite changes in population density, possibly for energetic efficiency. Some paired males attempted to take over widowed females, but shifted back to their original home range following the intrusion of an un-paired male. We conclude that social monogamy is the predominant social organization in round-eared sengis in a semi-desert that may have resulted from females living solitarily in small exclusive territories, balanced sex ratios, and from a low variation of body mass between males.     

Explicit experimental evidence for the effectiveness of proximity as mate-guarding behaviour in reducing extra-pair fertilization in the Seychelles warbler

Extra-pair copulations (EPCs; copulations outside the pair bond) are widespread in birds and may result in extra-pair fertilizations (EPFs). To increase reproductive success, males should not only seek to gain EPFs, but also prevent their own females from gaining EPFs. Although males could reduce the number of EPCs by their mates, this does not necessarily mean that they reduce the number of EPFs; indeed several studies have found no association between EPCs and EPFs. Male Seychelles warblers (Acrocephalus sechellensis) follow their partner closely during the period when the pair female is most receptive (fertile period). We show that males that guarded their mates more closely were less likely to have extra-pair young in their nest. This study on the Seychelles warbler is the first to provide explicit experimental evidence that mate guarding is effective in reducing EPFs. First, in territories where free-living males were induced to stop mate guarding during the pair female's fertile period, extra-pair parentage was higher than in the control group. Second, in the experimental group, the probability of having an extra-pair nestling in the nest was positively associated with the number of days during the fertile period for which mate guarding was artificially stopped. Thus, male mate guarding was effective in reducing the risk of cuckoldry.     

Spatial Dispersion and Nonmigratory Spawning in the Bluehead Wrasse (Thalassoma bifasciatum)

Earlier studies of the behavior of the bluehead wrasse Thalassoma bifasciatum have shown it to be a migratory spawner with large terminal-phase males defending temporary spawning territories. We describe a variant social structure where fish occupy permanent home ranges, spawn within or near these home ranges, and are not territorial. Movements of identified terminal-phase and intermediate males and females were mapped in the backreef areas of three coral reefs in Puerto Rico. Locations of spawns were then compared with these home-range maps. All fish used most or all of their morning home range during the afternoon spawning period. Both terminal-phase males and females spawned within or near their home ranges. Males were relatively tolerant of other males during the afternoon spawning period: only one in five encounters between large males resulted in aggressive chases. The location of chases bore no relation to spawning sites, areas of morning foraging, or to home-range borders. A Monte Carlo computer simulation was used to measure home-range dispersion of terminal-phase and intermediate males at our main site. If males were territorial, we should expect their home ranges to be significantly overdispersed within the cumulative area they occupied. During the first year male home ranges were overdispersed but during the second year they were randomly dispersed, even though the same number of terminal-phase males occupied the same study site both years. Differences in social behavior between backreef areas and other areas reported in the literature, and found in other portions of our reefs, may be related to differences in feeding ecology between populations and between different portions of the same reef.     

Male extraterritorial forays, age and paternity in the socially monogamous reed bunting (Emberiza schoeniclus)

Genetic studies have shown that extra-pair paternity is widespread among socially monogamous bird species. Yet, the role of males and females and their behavior leading to this mixed reproductive strategy is poorly understood. Here, we analyze paternity in relation to male age and mating behavior in the socially monogamous reed bunting (Emberiza schoeniclus). We report a positive relation between male extraterritorial forays and success in obtaining extra-pair fertilizations. Extraterritorial forays tended to increase in frequency with male age and older males sired a larger number of extra-pair offspring than young males. Identified extra-pair sires were old in nine out of ten cases. The likelihood of being cuckolded was not affected by male age. Although based on correlative data, our results highlight age-dependent explorative male behavior as a key determinant for the understanding of extra-pair mating in the reed bunting. We do, however, emphasize the need for further studies to reveal the role of females in extra-pair copulations and fertilizations.     

Paternal care declines with increased opportunity for extra-pair matings in fairy martins

Following the discovery that extra-pair fertilizations are common in many birds, it has been predicted that male participation in parental care may be influenced by their opportunity for extra-pair copulations. However, such a trade-off between male contribution to parental care and the availability of fertile females has not been confirmed. Here we use a novel remote monitoring technique to show that participation in incubation by male fairy martins, Hirundo ariel, declines with the increasing availability of fertile females in the breeding colonies. Furthermore, male contribution to incubation is most responsive to change in the availability of fertile females in the early morning, when most copulations occur, and also if their clutches are smaller than average. Both of these patterns support the presence of a trade-off between parental and extra-pair copulation effort. We suggest that this trade-off may be widespread among the 90% of bird species where males contribute to parental care.     

Social System of <Emphasis Type="Italic">Microcebus berthae</Emphasis>, the World’s Smallest Primate

Our goal was to provide a first characterization of the social system of pygmy mouse lemurs (Microcebus berthae), the worlds smallest primate species. During a 4-mo field study of 12 females and 27 males, we combined capture-recapture and morphometric data with detailed behavioral observations of individually marked subjects and genetic paternity analyses of a population in Kirindy Forest, western Madagascar. We describe the social organization of Microcebus berthae as a solitary forager living in an individualized neighborhood system characterized by extensive intra- and intersexual home range overlap of adult individuals within a male-biased population. Male and female pygmy mouse lemurs inhabited home ranges (males: 4.9 ha; females: 2.5 ha) that are more than twice as large as those of sympatric Microcebus murinus. On average, pygmy mouse lemurs of both sexes spent about half of the days sleeping alone. Preliminary analysis of genetic population structure suggests female philopatry and male dispersal. Sleeping associations of variable composition that consisted not preferentially of close relatives and proximity during part of the nightly activity contributed together with regular social interactions to the maintenance of a social network. The spatial distribution pattern of adult males and females, the absence of sexual size dimorphism, relatively large male testicular volume and moderate female estrous synchrony suggest a promiscuous mating system with a high potential for scramble competition. In general, there are many similarities between the social system of Microcebus berthae and those of other Microcebus spp. However, striking differences with sympatric gray mouse lemurs (Microcebus murinus) in female home range size, dispersion and sleeping behavior indicate the existence of species-specific socioecological adaptations in closely related species occupying very similar ecological niches.     

Movements and home range of ferrets (Mustela furo) at Pukepuke Lagoon,New Zealand

Abstract

Ferrets at Pukepuke Lagoon were live-trapped during the periods November 1970 to August 1973 and January 1976 to July 1977. Although the sex ratio of the 75 trapped ferrets was equal, there was a strong bias towards male captures in the earlier period (58.4%) and towards female captures in the later period (68.7%). Habitat modification and changes in the trapping programme for the later period probably resulted in the increased abundance and capture rate of females. Between consecutive captures 58.2% of male movements exceeded 250 m, as against on1y 39.7% of female movements; the mean distance between consecutive captures was 435 m for males and 285 m for females. Mean monthly movements varied from 700 m in January to 315 m in June for males, and from 360 m in March to 150 m in September for females. Juveniles of both sexes moved similar, intermediate distances. Dispersing juveniles were trapped from January until April; males tended to appear in traps earlier in the year than females. Only 4 of 14 juvenile males setNed as residents, hut 15 of 28 juvenile females did so. Male and female home ranges overlapped extensively, but within each sex individual ranges were usually separate. Resident males maintained a well defined spacing pattern, adjusting their boundaries if a juvenile settled or a neighbouring range became vacant. In 1976 six females had overlapping ranges in an area of albundant food. Female ranges (12.4 ha) were on average smaller than male ranges (31.3 ha); home ranges of both sexes were smaller during the breeding season (August-February) than at other times.     

Extra-pair matings and mate guarding in the common murre Uria aalge

Forced extra-pair copulations (FEPC) are frequent and mate guarding well developed among common murres. Male murres were at the colony almost continuously through the pre-laying period, but females were present only infrequently, and the frequency of FEPCs was significantly correlated with the relative number of males present. Males vigorously defended their partners from other males and females attempted to resist extra-pair matings. Females whose mates were absent were particularly vulnerable to FEPCs. The number of extra-pair copulations performed by males was estimated to vary between 0–32/season. The reproductive successes of males not performing FEPC, and those performing at the mean and maximum rate are estimated. Males most active in FEPC may substantially increase their reproductive success compared with males not performing FEPC. Ecological conditions in the common murre probably favour prolonged sperm storage, which in turn provides opportunities for sperm competition and favours both effective mate guarding and extra-pair copulations.