Genetic monogamy in the absence of paternity guards: the Capricorn silvereye, Zosterops lateralis chlorocephalus, on Heron Island

We investigated the genetic mating system of a socially monogamous passerine bird, the Capricorn silvereye Zosterops lateralis chlorocephalus, on an island of the Great Barrier Reef. There were no cases of extrapair paternity (EPP) among 122 offspring from 53 broods detectable by minisatellite or microsatellite DNA fingerprinting. Behavioral observations of paired birds showed that this was not a consequence of efficacious paternity guards and that females did not engage in extrapair copulation (EPC). Frequency of intrapair copulations was also low, with only 14 cases observed during 199 hours of observations of the 11 focal pairs in the fertile periods of females, and this was consistent with anatomical features of the cloacal protuberance in males. In this population, young birds form life-time pair bonds soon after gaining independence but females are obviously not attempting EPC possibly to redress this early mate choice. This is despite the fact that they breed in high density with a synchronous start and asynchronous spread of laying in a protracted season and males do not positively exhibit mate guarding behavior when females are fertile. Our results support high fidelity of socially monogamous birds on islands and are consistent with the hypothesis that sexual selection is reduced where genetic variation in fitness is limited.

Extra-pair paternity and mate-guarding behaviour in the brown thornbill

We used multilocus DNA fingerprinting to assess parentage in the brown thornbill, Acanthiza pusilla, a socially monogamous Australian passerine. Extra-pair paternity was uncommon (6.2% of 178 offspring; 11.9% of 67 broods) and there was no evidence of intra-specific brood parasitism. Extra-pair paternity was limited because pairs spent more time together when females were fertile and males were able to evict intruding males before they could approach the female. Males were responsible for the close proximity of partners during the fertile period. Mate guarding therefore appears to be a male tactic aimed at preventing female infidelity rather than a cooperative behaviour of the pair aimed at preventing extra-pair copulations and/or female harassment. Females did not attempt to escape male guarding and were rarely observed to solicit copulations from intruding males. Nevertheless, females paired to smaller and younger males were more likely to cuckold their mates than females paired to larger and older males. This suggests that females may be more likely to seek or accept extra-pair matings when paired to small, young males or that old, large males are better at preventing their mates from engaging in extra-pair copulations. We found that male age but not male size influences mate-guarding behaviour. Older males tended to respond more aggressively to intruders. We therefore speculate that the relationship between male size/age and extra-pair paternity in brown thornbills may arise because female thornbills prefer large males as mates but are unable to express this preference as easily when paired to older males.

Behaviour of the black flying fox Pteropus alecto: 2. Territoriality and courtship

Flying foxes are commonly thought of as highly social mammals, yet little is known about the dynamics of their social interactions at a day roost. The aim of the present study was to examine the nature of the seasonal activities of territoriality and courtship amongst wild flying foxes in Australia. Focal observations were conducted at two permanent roosts of black flying foxes Pteropus alecto during periods of peak social interaction in the summers of 1999 and 2000 in urban Brisbane, Queensland. Observations of male territoriality were conducted at dawn and began eight weeks prior to the commencement of mating. The majority of defense bouts (87%) consisted of ritualised pursuit, while 13% of bouts involved physical contact expressed as either wrestling or hooking. One male with an unusually large territory took significantly longer to defend it than other males with less territory to defend. Observations of courtship revealed repetitive courtship sequences, including pre-copulatory approaches by the males, copulation attempts and grooming/resting periods. Thirty-four complete courtship sequences incorporating 135 copulation attempts were recorded over two seasons. Females actively resisted courtship approaches by males, forcing males to display a continuous determination to mate over time where determination can be considered an indicator of 'fitness'. The courtship bout length of females with suckling young was significantly longer ((x) over bar +/- SE; 230.9 +/- 22.16 s) than that of females unencumbered by large pups (158.5 +/- 9.69 s), although the length of copulations within those courtships was not (45.6 +/- 5.19 versus 36.2 +/- 3.43 s).

Gender-role alternation in the simultaneously hermaphroditic freshwater snail Physa acuta: not with the same partner

In simultaneous hermaphrodites, gender conflicts that arise from two potential mates sharing the same gender preference may be solved through conditional reciprocity (or gamete trading). Conditional reciprocity had initially been considered widespread, but recent studies suggest that its real occurrence may have been overestimated, possibly because most mating observations have been performed on isolated pairs of individuals. Some resulting patterns (e. g., non-random alternation of sexual roles) were indeed compatible with conditional reciprocity but could also have stemmed from the two partners independently executing their own mating strategy and being experimentally enforced to do so with the same partner. Non-random alternation of gender roles was recently documented in the simultaneously hermaphroditic freshwater snail Physa acuta. To distinguish between conditional and unconditional gender alternations, we observed copulations of individually marked snails reared at three contrasted densities. We showed that density affected the overall frequency of copulations during the first 2 days of the experiment with high-density boxes showing more copulations than low density boxes, but it did not affect gender alternation patterns. A change in gender role was observed more often than expected by chance over two successive copulations by the same individual, confirming previous studies. However, gender switches did not preferentially occur with the same partner. We conclude that gender alternation is not due to conditional reciprocity in P. acuta. It may rather stem from each individual having a preference for gender alternation. We finally discuss the mechanisms and the potential extent of this unconditional reciprocity.

Condition-dependent genetic benefits of extrapair fertilization in female blue tits Cyanistes caeruleus.

In many socially monogamous birds, both partners perform extrapair copulations (EPC). As this behaviour potentially inflicts direct costs on females, they are currently hypothesized to search for genetic benefits for descendants, either as 'good' or 'complementary' genes. Although these hypotheses have found some support, several studies failed to find any beneficial consequence of EPC, and whether this behaviour is adaptive to females is subject to discussion. Here, we test these two hypotheses in a natural population of blue tits by accounting for the effect of most parameters known to potentially affect extrapair fertilization. Results suggest that female body mass affected the type of extrapair genetic benefits obtained. Heavy females obtained extrapair fertilizations when their social male was of low quality (as reflected by sexual display) and produced larger extrapair than within-pair chicks. Lean females obtained extrapair fertilizations when their social mate was genetically similar, thereby producing more heterozygous extrapair chicks. Our results suggest that mating patterns may be condition-dependent.

Effect of mating history on gender preference in the hermaphroditic snail physa acuta

Several internally fertilizing hermaphroditic animals can only perform one sexual role at a time. In such species, two individuals that engage in a copulation may have different interests in acting as male or female. A gender choice must be made which, if both individuals have the same preference, may give rise to a severe sexual conflict. Here we tested the hypothesis that gender choice could be influenced by mating history, using the freshwater snail, Physa acuta. We recorded the copulatory behaviour of 240 pairs composed of a focal individual and a partner, each either short- or long-isolated. We found that the time to the first copulation was unaffected by isolation status, suggesting that first contacts in this species are random processes. In contrast, the duration of copulations and the frequency of rejection behaviours suggested that individual gender preference switches from male biased to female biased as isolation increases. In addition, snails rejected copulations more frequently when presented to a partner with the same isolation status. Reciprocity, measured as the rate of gender swapping between the first and second copulations, was high irrespective of gender status. We suggest possible evolutionary causes for this gender preference switch and discuss its potential importance in natural population as well as its consequences for the maintenance of hermaphroditism

Age and multiple mating effects on reproductive success of Grapholita molesta (Busck) (Lepidoptera, Tortricidae)

The reproductive success of the oriental peach moth was evaluated in four experiments: 1) assessment of the mating duration, fecundity, fertility and longevity of females paired with virgin and immediately mated males; 2) mating duration, spermatophore size, fecundity, fertility and longevity in females paired with virgin and up to four times mated males; 3) receptivity of females to additional copulations after mating with virgin or mated males, and the effects of this behavior in female fecundity, fertility and longevity; 4) influence of insects age in the reproductive output. Males (33%) could copulate immediately after a previous copula. They were fertile until the fourth mating, but only in the first copula they transferred the longest (1.43 ± 0.10 mm) and widest (0.83 ± 0.11 mm) spermatophore, presenting the fastest mating duration (34.8 ± 2.62 min). A high proportion of females copulated by non-virgin males (84%) was receptive to other copulas, in comparison to those copulated by virgin males (32.4%). However, the fecundity, fertility, and longevity were similar among females that copulate once or more. The age was the most important factor affecting reproductive variables, where one and three day old insects had a significant higher fecundity, fertility and presented a shorter mating duration in comparison with older individuals. Results pointed out that the reproductive capacity of Grapholita molesta changes a little with respect to the analyzed factors, highlighting the elevated biotic potential of the species.

Biotic potential and reproductive parameters of Spodoptera eridania (Stoll) (Lepidoptera, Noctuidae) in the laboratory

Biotic potential and reprodutcive parameters of Spodoptera eridania (Stoll) (Lepidoptera, Noctuidae) in the laboratory: This study aimed to evaluate the biotic potential and reproductive parameters of Spodoptera eridania (Stoll, 1782) under controlled conditions (25 ± 1ºC, 70 ± 10% RH and 14 hour photophase). The longevity, pre-, post- and oviposition periods, fecundity and fertility of 15 couples was evaluated. The longevity of females (10.80 days) was not significantly higher than those of males (9.27 days). The mean durations of the pre, post and oviposition periods were 2.067, 0.600 and 8.133 days, respectively. The mean fecundity per female was 1,398 eggs and the mean fertility was 1,367.50 larvae. On average, females copulated 1.133 times. A strong positive correlation was observed between the number of mating and fecundity (r = 0.881, P <0.001). However a strong negative correlation was observed between the number of copulations and the duration of the pre-oviposition period (r = -0.826, P = 0.002) and longevity (r = -0.823, P = 0.001). The biotic potential of S. eridania was estimated at 1.894 x 10(25) individuals/female/year. The net reproductive rate (Ro) was 560.531 times per generation and the mean generation time (T) was 35.807 days. The intrinsic rate of increase (rm) was 0.177, with a finite rate of increase (l) of 1.193, per week

Extra-pair paternity, testes size and testosterone level in relation to colour polymorphism in the barn owl Tyto alba

In many bird populations, individuals display one of several genetically inherited colour morphs. Colour polymorphism can be maintained by several mechanisms one of which being frequency-dependent selection with colour morphs signalling alternative mating strategies. One morph may be dominant and territorial, and another one adopt a sneaky behaviour to gain access to fertile females. We tested this hypothesis in the barn owl Tyto alba in which coloration varies from reddish-brown to white. This trait is heritable and neither sensitive to the environment in which individuals live nor to body condition. In Switzerland, reddish-brown males were observed to feed their brood at a higher rate and to produce more offspring than white males. This observation lead us to hypothesize that white males may equalise fitness by investing more effort in extra-pair copulations. This hypothesis predicts that lighter Coloured males produce more extra-pair young, have larger testes and higher levels of circulating testosterone. However, our results are not consistent with these three predictions. First, paternity analyses of 54 broods with a total of 211 offspring revealed that only one young was not sired by the male that was feeding it. Second, testes size was not correlated with male plumage coloration suggesting that white males are not sexually more active. Finally, in nestlings at the time of feather growth testosterone level was not related to plumage coloration suggesting that this androgen is not required for the expression of this plumage trait. Our study therefore indicates that in the barn owl colour polymorphism plays no role in the probability of producing extra-pair young.

Blood corticosterone levels and intersexual selection games: best-of-bad-job strategies of male common lizards

Glucocorticoids affect physiology and behaviour, reproduction and potentially sexual selection as well. Shortterm and moderate glucocorticoid elevations are suggested to be adaptive, and prolonged and high elevations may be extremely harmful. This suggests that optimal reproductive strategies, and thus sexual selection, may be dose dependent. Here, we investigate effects of moderate and high elevations of blood corticosterone levels on intra- and intersexual behaviour and mating success of male common lizards Lacerta vivipara. Females showed less interest and more aggressive behaviour towards high corticosterone males and blood corticosterone levels affected male reproductive strategy. Males of moderate and high corticosterone elevations, compared with Control males, showed increased interest (i.e., higher number of chases, tongue extrusions, and approaches) towards females and high corticosterone males initiated more copulation attempts. However, neither increased male interest nor increased copulation attempts resulted in more copulations. This provides evidence for a best-of-a-bad-job strategy, where males with higher corticosterone levels compensated for reduced female interest and increased aggressive female behaviour directed towards them, by showing higher interest and by conducting more copulation attempts. Blood corticosterone levels affected intrasexual selection as well since moderate corticosterone levels positively affected male dominance, but dominance did not affect mating success. These findings underline the importance of female mate choice and are in line with adaptive compensatory behaviours of males. They further show that glucocorticoid effects on behaviour are dose dependent and that they have important implications for sexual selection and social interactions, and might potentially affect Darwinian fitness.

The relative fitness of Drosophila melanogaster (Diptera, Drosophilidae) that have successfully defended themselves against the parasitoid Asobara tabida (Hymenoptera, Braconidae)

Drosophila melanogaster larvae defend themselves against parasitoid attack via the process of encapsulation. However, flies that successfully defend them selves have reduced fitness as adults. Adults which carry an encapsulated parasitoid egg are smaller and females produce significantly fewer eggs than controls. Capsule-bearing males allowed repeated copulations with females do not show a reduction in their number of offspring, but those allowed to copulate only once did. No differences were found in time to first oviposition in females, or in time to first copulation in males. We interpret the results as arising from a trade-off between investing resources in factors promoting fecundity and mating success, and in defence against parasitism. The outcome of this investment decision influences the strength of selection for defence against parasitism.

Behavioral and spatial analysis of extraterritorial movements in red foxes (Vulpes vulpes)

Abstract: Movements away from the natal or home territory are important to many ecological processes, including gene flow, population regulation, and disease epidemiology, yet quantitative data on these behaviors are lacking. Red foxes exhibit 2 periods of extraterritorial movements: when an individual disperses and when males search neighboring territories for extrapair copulations during the breeding season. Using radiotracking data collected at 5-min interfix intervals, we compared movement parameters, including distance moved, speed of movement, and turning angles, of dispersal and reproductive movements to those made during normal territorial movements; the instantaneous separation distances of dispersing and extraterritorial movements to the movements of resident adults; and the frequency of locations of 95%, 60%, and 30% harmonic mean isopleths of adult fox home territories to randomly generated fox movements. Foxes making reproductive movements traveled farther than when undertaking other types of movement, and dispersal movements were straighter. Reproductive and dispersal movements were faster than territorial movements and also differed in intensity of search and thoroughness. Foxes making dispersal movements avoided direct contact with territorial adults and moved through peripheral areas of territories. The converse was true for reproductive movements. Although similar in some basic characteristics, dispersal and reproductive movements are fundamentally different both behaviorally and spatially and are likely to have different ultimate purposes and contrasting effects on spatial processes such as disease transmission

Mating system and exclusive postzygotic paternal care in a Neotropical harvestman (Arachnida: Opiliones)

This study tests predictions of the hypothesis of evolution of paternal care via sexual selection by using the Neotropical harvestman Pseudopucrolia sp. as the model organism. Females use natural cavities in roadside banks as nesting sites, which are defended by males against other males. Females leave the nests after oviposition, and all postzygotic parental care is accomplished by males, which protect the eggs and nymphs from predators. We provided artificial mud nests to individuals in the laboratory and conducted observations on the reproduction of the species. Male reproductive success was directly related to nest ownership time: the longer a male held a nest, the higher his chances of obtaining copulations. All males that succeeded in mating and obtaining one clutch eventually mated with additional females that added eggs to the clutch. Thus, desirable males were not limited to monogamy by paternal care. Experimental manipulations demonstrated that guarding males were more attractive to females than were nonguarding males and also that males guarded unrelated eggs. Finally, we found that females and nonguarding males spent more time foraging than guarding males. We use our data to contrast hypotheses on the origin and maintenance of paternal care and to provide a critical assessment of the hypothesis of the evolution of paternal care via sexual selection. (C) 2009 The Association for the Study of Animal Behaviour. Published by Elsevier Ltd. All rights reserved.

Distribuição diária do comportamento da mosca da fruta Anastrepha zenildae Zucchi, (Diptera: Tephritidae) em laboratório.

The knowledge of Anastrepha zenildae behavioral aspects combined with the biology of Tephritidae may contribute to monitoring and control programs of this fruit fly that is considered as economically important to the Rio Grande do Norte state. In order to characterize the daily activity profile of this species, we studied the behaviors of resting, locomotion, feeding, cleaning, courtship, copulation and oviposition of animals submitted to an artificial 12:12h light-dark cycle (750:1lux) with controlled temperature (26±2 °C). The observations were made with groups of 16 males and 16 females during 3 consecutive days each generation from parental to F5. Resting, locomotion, feeding and cleaning data were recorded as frequency and time of occurrence by scanning technique in 15 minutes windows per hour, with a record each minute. Courtship, copulation and oviposition were recorded as frequency, time of occurrence and duration by al occurrences technique. Resting was the most frequent behavior with males resting more than females. Locomotion was more evident in the first half of the ligh phase with higher values in females. Cleaning and feeding behaviors were more frequent in the second half of the light phase for both sexes with females eating more frequently than males. During the courtship, males were grouped in lek formations showing wings vibration and pheromone liberation. Courtship occurred more frequently 4 to 7 h after lights on (81,9%) with copulations being more frequent 6 h after lights on with a mean duration of 58,1±40,4 min. Copulation attempts were observed in males inside and outside the lek with aggressive behavior being observed only between males in the lek. Oviposition behavior was similar to that described for other species of the genus with a peak of this activity 2-3 h after the lights on, mean duration of 43, 7±34, 8 sec and 2 to 5 eggs by event. According to the results, sexual behavior of A. zenildae is temporally different of other sympatric species of the genus, being favorable to the reproductive isolation as well as the use of resources as oviposition substrate.

Ecologia comportamental reprodutiva de Diastatops obscura Fabricius (Insecta, Odonata)

In this thesis I discuss the reproductive behaviour and ecology of the libellulid Diastatops obscura Fabricius, 1775, (Insecta: Odonata) in natural conditions. Populations of this species were studied on the middle stretch of the Pitimbu River, Parnamirim municipality, Rio Grande do Norte, Brazil, during four discontinuous periods between 2002 and 2004. The objectives include the description of strategies and behaviors of both sexes, with especial interest in the intra-male competition for territories and females, the mate selection by females and the importance of male body size and other secondary characters on their reproductive success; from an adaptationist point of view. It was observed that the behavior of males and females in the reproductive areas are interrelated : the males came earlier to compete for the best territories and the females waited the result of that competition to be fertilized by dominants males, which preferably occupied areas near the river margin. The reproductive success of males with territories on the margin, estimated by number of copulations, ovipositions and days acting as territorial, was better than obtained by more separated territorial males and by satellite males. The body size of males is an important factor for the copulation and oviposition taxes and for the number of territorial days, favoring the biggest individuals. I also discuss the apparently importance of wing brilliance and wing integrity on male reproductive success. On inter-sexual relationships, I proved that females of D. obscura participate in mate selection, rejecting non-territorial males or substituting their sperm for other of higher status