Intraspecific competition and the maintenance of monogamy in tree swallows

Intraspecific competition for access to breeding resources maylimit male mating success typically monogamous birds. We examinedthe potential for intraspecific competition to limit polygynyin tree swallows at Beaverhill Lake, Alberta, Canada. In thispopulation, polygynous males raisedmore fledglings than monogamousmales, and there was little or no cost to females from nestingpolygynously. Under these conditions one might expect polygynyto be more common than that observed(8% of males). We foundthat females were most aggressive toward conspecific intrudersearly in the breeding season. This aggression was associatedwith (1) females settling farther apart than expected underrandom settlement, (2) later settlement by secondary than bymonogamous females, and (3) no relationship between female settlementdate and male territory size instead of the negative correlationexpected if females settled randomly without competition. Earlyin the season, males also settled farther apart than expectedif they had settled randomly, and among males with two or morenest boxes on their territory, males with widely separated nestboxes were more likely to be polygynous. Monogamy is probablythe most common pairing association in this population becauseintraspecific competition for nest sites prevents most malesfrom gaining a territory with nest sites far enough apart topermit two females to breed without one female excluding theother. Females appeared to be defending an area surroundingtheir nest box to limit nest usurpation or intraspecific broodparasitism, rather than to limit any loss of male parental carefrom polygyny.     

Effects of site fidelity and breeding performance on mate retention in a short‐lived passerine,the tree swallow Thachycineta bicolor

We studied the relationship among re‐mating, site fidelity and breeding performance in the tree swallow Tachycineta bicolor using 16 y of data on reproductive biology in a population breeding in nest boxes near Ithaca, New York. Of 217 pairs for which both members survived the non‐breeding season, 76% mated with a new partner and 24% reunited with their previous mate. Pairs did not increase their breeding success by breeding together for more than one breeding season. Males produced fewer fledglings after breeding with a new partner, but females neither increased nor decreased their success when breeding with a new mate. Females who bred with a new partner were younger than females that reunited with their previous mates, and they were more likely to move to a different nest box. Males that bred with a new mate were of similar age to males that reunited, and they did not move more often. The probability of breeding with a new partner was better predicted by female age than by previous breeding success, suggesting that re‐mating was not strongly affected by past breeding performance. Because younger females change breeding sites more frequently than do older females and females that mated with a new partner were younger than females that reunited with their previous mates, we suggest that the tendency of tree swallows to change partners between years is a by‐product of lower site fidelity of younger females rather than a strategy for increasing breeding success.     

Large males have a mating advantage in a species of darter with smaller, allopaternal males Etheostoma olmstedi

Theory suggests that males that are larger than their competitors may have increased mating success, due to both greater competitive ability and increased attractiveness to females. We examined how male mating success varies with male size in the tessellated darter Etheostoma olmstedi. Previous work has shown that large males tend to move around and breed in vacant breeding sites, and consequently provide less care for their eggs, while smaller individuals can be allopaternal, caring for the eggs of other males as well as for their own. We studied female egg deposition in a natural breeding population using artificial breeding sites and in the laboratory, where female choice of spawning site was restricted to two breeding sites tended by two males of different sizes. In both the field and the laboratory, nests tended by larger males were more likely to receive new eggs. Additionally, the mean size of males associated with a nest was positively correlated with both the maximum coverage of eggs at the nest and the number of times new eggs were deposited. We discuss how the increased mating success of larger males, despite their decreased parental care, may help explain allopaternal care in this species [Current Zoology 56 (1): 1-5, 2010].     

Size-mediated dominance and aggressive behavior of male Japanese fluvial sculpin Cottus pollux (Pisces: Cottidae) reduce nest-site abundance and mating success of conspecific rivals

Despite growing evidence for plasticity in the mating patterns of nest-holding animals in relation to the changes in nest abundance, the effects of aggressive interaction by dominant males on nest availability for conspecific rivals remains unclear. To quantify the effects of male–male competition on nest-site choice and mating success of the male Japanese fluvial sculpin Cottus pollux, we conducted experiments on 5 males from different 5 size classes under both sufficient and shortage nest-abundance conditions. Nest-choice experiments showed that both male size class and nest-abundance condition had significant effects on the nesting rates of males. Following the nest-choice experiments, 10 gravid females were added in the experimental tanks. Mating experiments revealed that male size, nesting rate before addition of females, and the number of courtship attempts on females were valid variables of male mating success, regardless of nest-abundance conditions. After achieving initial mating success, the largest nesting male exhibited more frequent aggressive interaction with other conspecific males than he did before obtaining eggs in his nest. Our results suggest that size-mediated dominance and aggressive behavior of males may disrupt nest acquisition of other conspecific males, and may consequently result in extreme variation in mating success among males even under sufficient nest-abundance conditions.     

Is egg carrying attractive? Mate choice in the golden egg bug (Coreidae,Heteroptera)

In several species of fish, females select males that are already guarding eggs in their nests. It is a matter of debate as to whether a female selects a good nest site for her offspring (natural selection) or a male for his attractiveness (sexual selection). The golden egg bug, Phyllomorpha laciniata Vill, resembles fish in the sense that mating males carry more eggs than single males, but in the bugs, female mate choice is decoupled from egg site choice. The sexual selection hypothesis predicts that if females select males using male egg load as a cue for male quality, they should not mate with a male when eggs are removed, regardless of his mating attempts. When individual females were enclosed with an egg-loaded male and an unloaded male, they mated equally often with both males, although the loaded males courted more. In addition, when only successful males were used, females mated equally often with the loaded male and the unloaded male irrespective of sex ratio. Male choice rather than female choice affected mating frequency when sex ratio was equal. Therefore, females do not select the male by the eggs he carries, but successful males may receive many eggs due to egg dumping by alien females while they mate or as a consequence of mate guarding.     

Nonlinear effects of female mate choice in wild threespine sticklebacks

Sexual selection by mate choice represents a very important selective pressure in many animal species and might have evolutionary impacts beyond exaggeration of secondary sexual traits. Describing the shape and strength of the relationships linking mating success and nonsexual traits in natural conditions represents a challenging step in our understanding of adaptive evolution. We studied the effect of behavioral (nest site choice), immunological (trematode level of infection), genetic diversity (measured by mean d2) and morphological (standard length and pectoral fin size) traits on male mating success in a natural population of threespine sticklebacks Gasterosteaus aculeatus. Male mating success was measured by microsatellite genotyping of embryos used to infer female genotypes. First, we analyzed all territorial males (full analysis) but also considered independently only males with a nonzero mating success (reduced analysis) because some of the males with no eggs could have been part of a later breeding cycle. Multiple linear regressions identified a significant negative effect of parasite load in the full analysis, whereas no linear effect was found in the reduced analysis. The quadratic analyses revealed that nest location and parasite load were significantly related to mating success by positive (concave selection) and negative (convex selection) quadratic coefficients respectively, resulting in a saddle-shaped fitness surface. Moreover, there were significant interactions between nest location, mean d2 and parasite load in the reduced analysis. The subsequent canonical rotation of the matrix of quadratic and cross-product terms identified two major axes of the response surface: a vector representing mostly nest site choice and a vector representing parasite load. These results imply that there exists more than one way for a male threespine stickleback to maximize its mating success and that such nonlinear relationships between male mating success induced by female mate choice and male characteristics might have been overlooked in many studies.     

Effects of nestboxes on the breeding biology of Southern House Wrens Troglodytes aedon bonariae in the southern temperate zone

Nestboxes are known to increase clutch size, enhance breeding success and affect the social mating system of several cavity nesters. Although in recent years various cavity nesters have been studied in nestboxes in South America, the effects of boxes on the biology of the study species are unknown. We evaluated the effects of nestboxes on the breeding biology and social mating system of Southern House Wrens Troglodytes aedon bonariae by comparing birds breeding in nestboxes and tree cavities in two cattle ranches in Buenos Aires Province, Argentina. Southern House Wrens nesting in boxes had higher breeding success but, contrary to studies on the temperate zone, we did not find differences in clutch size between Wrens breeding in nestboxes and tree cavities. The main causes of nest failure in tree cavities were nest predation and flooding of the cavity (70 and 23% of the failures, respectively) while in nestboxes predation and desertion were the most important causes of failure (38 and 34% of the failures, respectively). The social mating system of Southern House Wrens is monogamy with biparental care, and neither was affected by the boxes. Males did not attract secondary females to additional nestboxes; however, nestboxes are safer breeding sites than tree cavities, and females seemed to prefer males with nestboxes on their territory. These results suggest that nest quality alone might be not enough for secondary females to accept polygyny.     

Male mating success in the Azorean rock-pool blenny: the effects of body size, male behaviour and nest characteristics

Larger male Azorean rock-pool blennies Parablennius sanguinolentus parvicornis received more female visits, were courted more often by females and received more spawnings. Larger males also received a higher number of male intrusions, attacked more conspecifics and defended larger territories. Larger males showed more nest cleaning behaviour and a marginally non-significant trend for higher egg fanning rate. Male courtship, male attack rate against conspecifics and parental behaviour were all correlated with the frequency of female spawnings received by each male even when controlling for male size, suggesting that these behaviour patterns influence male mating success. On the other hand, a positive partial correlation was found between female courtship and the frequency of female spawnings, controlling for the number of female visits, which suggests a role for female mate choice on male mating success. Finally, males nesting in chambers in the bottom of pools received more spawnings than males nesting either in crevices or under boulders. However, nest opening area was associated significantly negatively with male mating success, when controlling for male size. Thus, the present data suggest strongly that male characteristics overrule nest characteristics in determining male mating success in the Azorean rock-pool blenny.     

Determinants of male mating success in a natural population of a stream goby of the genus Rhinogobius

The mating success of individually identified males of a stream goby, Rhinogobius sp. CB (cross‐band type), was recorded over one breeding season. One to 3 year‐old males were active in mating, but 3 year‐old males, which accounted for 36% of the population, amounted to c . 70% of the total males guarding eggs in the nest. Three quarters of the breeding males had only one brooding cycle, but the others had two or three. All the latter males changed their nest sites between cycles within a riffle. In 1 and 2 year‐old males, the number of brooding cycles contributed more to the mating success than egg mass size in one brooding cycle. For mating success of 3 year‐old males, the egg mass size in one brooding cycle, which can be enhanced by spawning with a large female or multiple females, was as important as the number of brooding cycles. These male reproductive tactics could be attributed to the age‐related ability of nest construction and mate acquisition.     

Breeding ecology and male spawning success in two hill-stream populations of the freshwater goby,Padogobius martensi

Synopsis This investigation was carried out at two study sites to examine the influence of shelter availability, nest site quality and male body size on spawning success of malePadogobius martensi. At Stirone Stream (first study site) individuals were not randomly distributed with respect to usable stones; larger males and females were consistently found to occupy larger stones. Laboratory experiments showed that the bottom surface area of the shelter was the relevant cue for the choice. The number of nesting males settled in a given area increased with increasing availability of local stone cover; inter-nest distance was directly correlated with male size. At Rivarolo Stream (second study site), which was characterized by lower nest density and a greater shortage of large stones, the pattern of shelter occupation was similar. These results suggest the presence of competition among individuals for the possession of larger-sized shelter sites. At both sites, the number of eggs in the nest increased as stone size increased; however, at the first study site, male body size played a major role in male spawning success as it concerns both the numbers of females mated and the number of eggs, while at the second study site, male body size was not shown to influence male spawning success. Factors responsible for between-site differences in male spawning success are discussed.     

Female mate choice across mating stages and between sequential mates in flour beetles

Few studies have examined how female premating choice correlates with the outcome of copulatory and post-copulatory processes. It has been shown that polyandrous Tribolium castaneum females discriminate among males before mating based on olfactory cues, and also exert cryptic choice during mating through several mechanisms. This study tested whether a male's relative attractiveness predicted his insemination success during copulation. Bioassays with male olfactory cues were used to rank two males as more and less attractive to females; each female was then mated to either her more attractive male followed by less attractive male, or vice versa. Dissections immediately after second copulations revealed a significantly higher percent of successful inseminations for females that remated with more attractive males compared with those that remated with less attractive males. These results indicate that cryptic female choice during copulation reinforces precopulatory female choice in T. castaneum, and suggest that females could use cryptic choice to trade up to more attractive males, possibly gaining better phenotypic or genetic quality of sires.     

Successive nest building and polygyny of Fan-tailed Warblers Cisticola juncidis

The polygynous mating system of the Fan-tailed Warbler Cisticola juqcidis was investigated between 1978 and 1981. The male warbler builds many nests unaided; however, he has no more than one active vacant nest for courting at any time. Nest building lasted from April until August. After completing the outer fabric of a nest, the male advertised it and led a female to the nest site by a unique invitation flight. On average a male built 6.5 nests during one breeding season and three of them were accepted by females. The most successful male completed 18 nests, and mated with 11 females. Out of a total of 111 males which established a territory, 30 had no mate, 14 were monogamous, and the rest were polygynous. About 50 to 70% males were polygynous over the four years. The sex ratio varied from 1.41:1 to 2.17:1 (females: male) in the breeding population. It was partly caused by the presence of 'floating males'. After the completion of the outer fabric of the nest, the male warbler did not take any further role in nesting and caring for young.
The polygynous mating system of the Fan-tailed Warbler is characterized by successive nest building. Its extreme development results from the long breeding period and the male having no role in parental care.     

Nest box use by woodland dormice (Graphiurus murinus): the influence of life cycle and nest box placement

The use of nest boxes by the woodland dormouse, Graphiurus murinus, was investigated over a 13-month period in a riverine forest of the Great Fish River Reserve, South Africa. We predicted that some characteristics of nest box placement would affect nest box use and that the seasonal pattern of nest box use would be linked to the species' life cycle and physiological and socioecological characteristics. Generalized linear models indicated that the time since nest box installation and nest box height above ground positively affected the frequency and intensity of nest box use. Male and female dormice, as well as adults and juveniles, did not differ in the number of nest boxes used and equally occupied individual nest boxes. The percentage of nest boxes used peaked during spring and summer (breeding period) and dropped during winter (hibernation). However, whereas significantly more males were caught during the mating season (spring), the number of females occupying nest boxes was constant during the year. As female dormice successfully bred in the nest boxes, the observed sexual patterns suggest that (artificial) nest sites represent an important resource for females, whereas females seem to constitute the main resource for males, as predicted by the socioecological model.     

The influence of nest predation on mating strategies under polygyny

The mating status of an advertising male is one important factorfor female assessment. In species where already-mated malestry to attract additional females at some distance from theirmate, mating status may be difficult to assess. Limitationsin the time available to breed and a decrease in reproductivesuccess over the breeding season could make the costs outweighthe benefits of a time-consuming mate assessment process, whichautomatically postpones the onset of breeding. Time and energyconstraints will then limit the behavioral options for femalesin search of breeding opportunities, and females would suffera cost when assessing male mating status. Female choice maythen be better understood by analyzing the behavioral optionsfor females; the success of a strategy in which costly assessmentsare avoided and females choose mates indiscriminately, disregardingmating status ("fast" strategy), could be compared to the successof a strategy in which females carefully try to assess matingstatus ("coy" strategy). Nest predation should be included insuch an analysis because it influences the amount of male assistancefemales of polygynous males receive, as nest predation for onlyone of a polygynous male's two females leads to more male assistancefor the other female. In this study, I compare the outcome ofa "fast" strategy with that of a "coy" strategy under differentrates of nest predation. Although high probabilities of becominga mate of a polygynous male associated with a fitness reductiondecrease the relative success of a "fast" strategy, simulationsshow that high rates of nest predation considerably lower thedifference in the outcome of the two strategies. Therefore,the cost of assessing mating status must be relatively smallto be economical.     

Lifetime mating success in the damselfly Coenagrion puella

Lifetime mating success of male azure damselflies (Coenagrion puella) was measured in a natural population. The major determinant of mating success is the number of days a male spends at the breeding site, which is mostly determined by a male's adult lifespan. Long-lived males have a higher mating rate than short-lived males, and daily mating rate increases with age up to 6 days, then falls. Large males live longer, but have a lower daily mating rate than small males. These effects of size are very weak, accounting for no more than 2% of the daily variance in mating success. The only overall effect of size on lifetime mating success is that males at both extremes of the size distribution are more likely to fail to mate. Chance differences in the number of females encountered are sufficient to account for the remaining variance in mating success. The weather is also shown to have a major effect on mating success. We draw attention to the ways in which it may be misleading to draw conclusions about the action of sexual selection from studies of daily, rather than lifetime, reproductive success. We provide evidence to support the view that variance in male reproductive success is neither evidence for sexual selection, nor a measure of its intensity.     

Does part reproductive history predict competitive interactions and male mating success in pupfish?

Two experiments were performed to determine the effect of past territoriality and mating success on subsequent dominance and ability to attract females. First, in 14 staged, pair-wise encounters, the development of nuptial coloration and agonistic behaviour was examined in two types of males: previously territorial (PT) and previously non-territorial (PNT) males. Previously territorial males developed a more intense nuptial coloration and won all contests. Second, to test the males' ability to attract females, other factors were controlled that could have affected female mate choice, such as male size, quality of breeding substrate and male-male interactions. In 40 trials, PT males were more active, developed a more intense nuptial coloration, courted females more vigorously, spawned sooner and had a higher mating success than PNT males. Thus, in pupfish, Cyprinodon pecosensis, previous reproductive history is a good indicator of both dominance and mating success. Only males in good physical condition achieve high mating success, and both intra- and inter-sexual selection are important in maintaining the close correlation between expression of nuptial coloration, a second sexual trait, and other male attributes, such as physical condition and vigour, that allow a male to secure and defend a territory against rivals and to attract and spawn with females.     

Mating behaviour and mating success in woodfrogs Rana sylvatica

Mating behaviour of woodfrogs was investigated in a woodland pond in southeastern Michigan. Males became sexually mature one year before females and outnumbered females by 5.6 to 1 at the breeding site. Males employed searching behaviour to obtain mates. Yearly male mating success varied from 0 to 2 matings, and larger males had a greater probability of mating than smaller males. The mating season lasted less than 10 days. Most egg masses were deposited in 1 m² of a 256-m² pond. Experimental introduction of egg masses to the breeding site indicated that presence of eggs was sufficient to stimulate egg deposition. The influence of selection for synchronous sexual receptivity in females, communal egg deposition, and delayed female sexual maturation on male mating behaviour and variation in male mating success is discussed.     

Distribution and reproductive success of the Pied Flycatcher Ficedula hypoleuca in relation to forest patch size and vegetation characteristics; the effect of scale

We studied habitat choice, distribution and reproductive success in the Pied Flycatcher Ficedula hypoleuca in relation to the spatial structure of a fragmented forest area. Two hierarchical levels of forest habitat structure were used: (1) forest patch size (macrohabitat) and (2) vegetative structure within forest patches (microhabitat). In spring, both males and females settled preferentially in large and medium-sized forest stands (>1 ha) where breeding density was also higher than in small stands (<1 ha). Stands <5 ha were occupied later, and the proportion of unpaired males was higher there than in large stands (>5 ha). The known age distributions of breeding birds and breeding success were independent of forest patch size. Nest predation rate was not associated with stand size or nest distance from the forest edge. The preference of breeding birds for large forest patches was presumably related to the higher amount of resources (nest sites, mates, food) a large patch can offer for reproduction compared with a small patch. At the microhabitat level, territories of old males were characterized by relatively more deciduous trees that contained more invertebrate food than coniferous tree-dominated territories of yearling males. The reproductive success of old males, as measured by the number of fledged young per male, was higher than that of yearling males. This suggests that the larger body size and blacker plumage of old males possibly contributed to male dominance and that old males excluded younger males from preferred habitats through territorial behaviour. Our results suggest that habitat choice of the Pied Flycatcher was affected by both micro- and macrohabitat. On the basis of settlement pattern, density and reproductive success, the distribution of Pied Flycatchers across forest stands of different sizes followed the ideal-free distribution model, whereas at the microhabitat level, age-related unequal distribution of males followed the ideal-despotic model of Fretwell and Lucas.     

Use of novel nest boxes by carmine bee‐eaters (Merops nubicus) in captivity

Carmine bee‐eaters make attractive additions to zoo aviaries but breeding programs have had challenges and limited success. The objectives of this study were to document nesting behavior of Carmine bee‐eaters in a captive setting and compare reproductive success between a novel nest box (plastic, 17 × 30 × 22 cm) and a PVC pipe model used previously (30 cm long, 8 cm in diameter). Three bee‐eater pairs were given access to seven nest chambers (six novel boxes, one PVC model). Behavioral observations occurred during a 15‐min period in the morning or afternoon before egg production and continued until chicks fledged for a total of 87 observation periods (21.75 hr). All occurrences by an individual bird entering or exiting a nest tunnel, food provision, and the time (min) spent inside a nest cavity were documented. Additionally, daily temperature within each nest chamber was recorded. Before eggs were produced the average daily temperature (23.02°C) within the nest chambers did not differ, suggesting that nest cavity choice was not influenced by temperature. No differences were detected among pairs in percent of observed time spent inside their nest cavities or number of times a nest tunnel was entered during the incubation or fledging periods. During incubation females spent a greater percent of observed time inside the nest cavity than males (P=0.02). During the fledging period food provision did not differ between the pairs, however males entered their nest tunnels more often per hour than females (P=0.03), and males tended to provide food more often than females (P=0.053). Two pairs nested in novel nest boxes and successfully fledged one chick each. The pair that nested in the PVC model did not fledge a chick. A nest box that aids in keeping eggs intact is essential for breeding bee‐eaters in captivity, and maintaining captive populations will provide opportunities for zoo visitors to enjoy these birds and will reduce the need to remove birds from the wild. Zoo Biol 0:1–13, 2007. © 2007 Wiley‐Liss, Inc.     

SEXUAL SELECTION AND FITNESS VARIATION IN A POPULATION OF SMALLMOUTH BASS,MICROPTERUS DOLOMIEUI (PISCES: CENTRARCHIDAE)

Monogamy is often presumed to constrain mating variance and restrict the action of sexual selection. We examined the reproductive patterns of a monogamous population of smallmouth bass (Micropterus dolomieui), and attempted to identify sources of within-season fitness variation among females and known-age males. Many males did not acquire a nest site, and many territorial males were unsuccessful in acquiring a mate. The likelihood that territorial males mated depended on several aspects of nest sites. Mated males of age three were larger than the average size of age-three males in the population. The mean sizes of age-four and age-five mated males were not different from the average of same-age males in the population. Thus, selection resulting from the acquisition of a mate favored large size among only age-three males. Timing of nest construction and breeding among territorial males was negatively related to male size and did not depend on male age after taking male size into account. Indirect evidence (numbers of eggs deposited in nests) suggests that the timing of spawning among females was also negatively related to female size. Fertility selection favored early reproduction within the season by males of all ages, but large male size was favored among only age-four males. The combined early breeding of fecund females and female mate choice of large males may explain the positive correlation between the size of age-four males and the number of eggs acquired. Despite large differences of female fecundity, however, the variance of relative mate number contributed about two times more than the variance of relative fertility among females to the total variance of relative fitness within each sex.