Thomas W. P. Friedl

Sexual selection in the lek-breeding european treefrog : Body size, chorus attendance, random mating and good genes

Variation in male reproductive performance and success was studied over 2 years in a population of individually marked European treefrogs, Hyla arborea , a hylid frog with prolonged breeding season and a lek mating system. Reproductive success in males was not related to body size, age or growth rate, and there was no evidence for size- or age-assortative mating. Male mating success was significantly correlated with chorus attendance, that is, the number of nights spent calling at the breeding site. There were no significant differences with regard to body size, age or growth rate between males surviving to the next breeding season and those not surviving. However, surviving males showed a significantly higher chorus attendance than nonsurviving males, which is a novel finding for lekking anurans. Furthermore, males that survived from the previous breeding season spent significantly more nights calling at the breeding site than males that were observed for the first time. Since calling in frogs is costly, these results suggest that chorus attendance reflects male quality. The distribution of male mating success did not differ from a Poisson distribution, indicating a random mating pattern. Since male quality is related to chorus attendance, females that mate randomly are likely to mate with high-quality males and thereby gain indirect genetic benefits without incurring costs of extended mate searching and mate assessment. We suggest that a similar mating pattern is found in many lek-breeding hylid frogs in which male mating success is mainly determined by chorus attendance.

Determinants of male mating success in the red bishop (Euplectes orix)

Abstract We studied sexual selection in the red bishop, Euplectes orix, a colonial, polygynous weaverbird widely distributed over sub-Saharan Africa. Male reproductive success measured in terms of the number of nests accepted by females and the number of eggs and nestlings in all the nests on a males territory varied considerably. The standardized variance (variance/mean2) in male reproductive success ranged from 0.505 to 1.737 in different years, indicating a high potential for sexual selection in this species. An analysis of genetic parentage for 432 nestlings by non-radioactive, multilocus DNA fingerprinting confirmed that male reproductive success (number of young sired on the territory) in this species can be reliably estimated by the measures introduced above. In all 4 study years there was a strong positive correlation between male mating success and the total number of nests that males built in their territories. The number of nests built can be partitioned into the number of weeks a male held a territory and his nest-building performance. Both factors exert a significant positive effect on male mating success and in combination explained between 53.3 and 86.3% of the variation in male reproductive success. Male morphological characters were found to be of no importance. Males that established a territory in the following season built more nests and held their territories for longer than males that did not establish a territory in the following season, suggesting that these measures might be indicators of male condition and quality. Male nest-building performance (number of nests built per week) seems to be unrelated to male condition or quality.

Temporal changes in demand for and supply of nests in red bishops (Euplectes orix): dynamics of a biological market

The red bishop (Euplectes orix) is a highly polygynous and colonial weaverbird. Males construct several nests within their territories to which they try to attract females, and females are solely responsible for incubation and raising offspring. In this paper, we describe the characteristics of the red bishop’s mating system as a biological market and investigate the role of nests built by males as a traded commodity in a mating market. As timing of breeding in red bishops in arid and semi-arid zones depends on rainfall patterns which are often unpredictable, there are temporal changes in demand for and supply of nests within a breeding season, with breeding activities of males and females being highly synchronised. We found that males increased their nest-building speed with increased female breeding activity independently of rainfall, indicating that supply follows demand in this mating market. The supply of nests was always larger than the demand for nests. Construction costs for nests increased with demand for nests as indicated by shorter nest-building duration and shorter building delays between two consecutively built nests at times of high breeding activity. Males as a trading class are chosen according to the age of their nests offered, with young nests having a higher probability of being accepted by females. Furthermore, female choosiness with regard to nest age decreased when their own market value decreased, as predicted by biological market theory. The temporal changes of breeding activity together with the female preference for young and fresh nests require that males quickly adjust nest-building activity to varying female demand for new nests. However, males with a better adjustment of building speed to female breeding activity did not gain higher mating success.

EXTRAPAIR FERTILIZATIONS IN RED BISHOPS (EUPLECTES ORIX): DO FEMALES FOLLOW CONDITIONAL EXTRAPAIR STRATEGIES?

Abstract Although many studies have revealed differences in patterns of extrapair paternity between species and between populations of the same species, possible differences in female extrapair behavior within populations have received far less attention. Here, we propose and test the hypothesis that females of the highly polygynous Red Bishop (Euplectes orix) follow two different extrapair strategies, with benefits of extrapair copulations depending on the quality of their social mate (”conditional-extrapair-strategy hypothesis”). In particular, we propose that females mated to low-quality males enhance the genetic quality of their offspring by performing extrapair copulations with males of higher quality than their social mate (in accordance with the good-genes hypothesis), whereas females mated to high-quality males perform extrapair copulations as insurance against temporary infertility of their social mate caused by sperm depletion due to frequent copulations (in accordance with the fertility-insurance hypothesis). Several predictions derived from our hypothesis, regarding differences in frequency and distribution of both unhatched eggs and extrapair young between high-quality and low-quality males, are tested and shown to be supported. Most importantly, hatching failures occurred more often in territories of high-quality males than in those of low-quality males, and the proportion of unhatched eggs within the territory was positively correlated to breeding synchrony for high-quality males only. Those results suggest that sperm depletion is a risk for females mated to high-quality males with many mating opportunities. The fact that broods of high-quality males without unhatched eggs had a significantly higher proportion of extrapair young than broods of high-quality males with unhatched eggs is consistent with the hypothesis that females mated to high-quality males were successful in performing EPCs as insurance against temporary infertility of their social mates. Furthermore, genetic fathers of extrapair young hatched in territories of low-quality males were found to be of higher quality than the corresponding social fathers, whereas no difference in quality was found between genetic and social fathers of extrapair young hatched in territories of high-quality males. Although we cannot exclude the possibility that the observed pattern was a result of females mating randomly from within the available pool of extrapair males, it is in accordance with our prediction that females mated to low-quality males perform extrapair copulations with males of higher quality than their social mate to enhance offspring fitness. Overall, the data presented here suggest that females within our study colony adjusted their extrapair behavior to the quality of their social mate, even if we cannot entirely rule out alternative explanations for some of the results obtained. Thus, the study provides support for the conditional-extrapair-strategy hypothesis, which states that within populations, females may follow different extrapair strategies to gain different benefits. Fertilizaciones Extra-pareja en Euplectes orix: ¿Siguen las Hembras Estrategias Extra-pareja Condicionales?

Effects of behavioural time budgets and nest-building efficiency on male reproductive performance in red bishops ( Euplectes orix )

In this study we have investigated the effect of nest-building behaviour, courtship behaviour, and male–male interactions on male reproductive performance of the red bishop (Euplectes orix), a highly polygynous, colonially breeding weaverbird species. Previous studies on red bishops have revealed that male reproductive success is mainly determined by the number of nests built in a territory, and that reproductive success and the number of nests both vary substantially among males. Here we test whether the high variation observed in reproductive performance is related to male behavioural patterns (as assessed by time-budget analysis) and/or nest-building efficiency (as assessed by the number of fibres incorporated in the nest in unit time). We collected data on male time budgets and analysed behavioural patterns for three breeding seasons. A greater number of nests built was positively related to both the amount of time allocated to nest-building behaviour and the efficiency of nest building. Neither the amount of time spent in courtship behaviour nor the amount of time spent in male–male interactions was related to reproductive success. Male reproductive success, irrespective of the number of nests built, was partly determined by the number of fibres incorporated, suggesting some importance of nest quality in terms of nest chamber density to male reproductive success. There were no trade-offs with regard to time allocated to different behaviour. Instead, males differed with regard to their territory attendance and activity levels, which might be because of differences in energy resources and may thus reflect inherent differences in male body condition and, ultimately, male genetic quality.