Nest concealment and parental behaviour interact in affecting nest survival in the blackcap (Sylvia atricapilla): an experimental evaluation of the parental compensation hypothesis

Nest concealment varies strongly within populations of many species. Although some studies have revealed the beneficial effects of concealment in mitigating predation pressure on nests, other studies were unable to find similar effects. One potential reason for the mixed results is that parental behaviour may compensate for the effects of nest cover, and specifically designed experimental studies are needed to reveal this compensation. I studied the effects of concealment on the probability of nest survival in the blackcap (Sylvia atricapilla), by experimentally manipulating the degree of nest-foliage cover. There was a significant effect of the treatment depending on nest type and the phase of nesting. Whereas there was no effect of concealment on nest survival in natural nests, there was a positive effect in real nests baited with plasticine clutches (i.e. without parental activity). Parents probably behaviourally compensated for poor concealment in natural nests (nest guarding, defence). In line with this, there was no effect of concealment on nest survival during incubation, whereas there was probably a positive effect in the nestling phase. Parents spent more time on the nest during incubation (80%) than during the care of nestlings (40%) and, consequently, had more opportunities to compensate for poor cover. In general, we cannot use single measures of behaviours or states (nest concealment) as an indication of predation risk because of the capacity for compensation in other behaviours.Communicated by C. Brown     

Mating patterns and a test for female mate choice in Etheostoma spectabile (Pisces,Percidae)

Neither size nor breeding color correlated with spawning success of male orangethroat darters, Etheostoma spectabile (Pisces, Percidae), under natural field conditions. When females were presented experimentally with a simultaneous choice they spent no more time in proximity to large than small males, and were subsequently no more likely to spawn with large than with small males. Females also displayed no preference for bright versus dull males. Males and females did not differ significantly in size. Etheostoma spectabile may lack sexual size dimorphism as a result of the lack of female choice for size and the ineffectiveness of male attempts to monopolize females, or selection may be for increased size of females. Males are not dwarfs because of sperm competition. Contest competition among males appeared to be important in initiating spawnings but many males obtained spawnings by participating in ongoing spawning events. Etheostoma spectabile is an example of a sexually dimorphic species with no evident female preference for male size or color.     

Higher rate of nest loss among primary than secondary females: infanticide in the great reed warbler?

An individually marked population of great reed warblers Acrocephalus arundinaceus was studied between 1984 and 1991 in South Central Sweden. The fates of 279 nests were analysed for the 25 days following clutch initiation. The average frequency of nest loss (43%) did not differ significantly between years or between four periods of the breeding season. After a breeding failure, 70% of the females laid a replacement clutch, most of them together with the same male. On polygnous territories, males assisted the female who first hatched young regardless of her initial mating status (Fig. 2). According to the acutal status of the females attending the nests, for each day nests were classified as being of either monogamous (M), primary (P) or secondary (S) status. From egg-laying to fledging the rate of nest loss decreased among nests of primary status whereas it increased among nests of monogamous and secondary status. During the egg-laying period, the rate of nest loss was 3 times higher among nests of primary than among nests of monogamous and secondary status (Fig. 3). Thus, the high loss level among nests of primary status during the laying period was closely associated with the presence of a female with a less advanced nest on the territory. All nests were situated in reed beds above deep water and most of them at a height at which possible inter-specific nest predators would have caused disturbance to the nest itself. The suspicion that secondary females committed sexually selected infanticide was supported by an experiment with dummy eggs that revealed bill peck markings identical to those obtained from great reed warblers (Fig. 4).     

Female yellowhammers (Emberiza citrinella) prefer yellower males: a laboratory experiment

The importance of male plumage coloration as a signal of male dominance and a cue for female choice in the monogamous yellowhammer, Emberiza citrinella, was tested in two sets of experiments in an indoor aviary. Dominance was tested by introducing two individuals, with no previous experience of each other, in an aviary with food and water. Aggression occurred more often between two males than between a male and a female and more frequently between pairs of males including at least one old male than between two young males. Dominance was not related to male colour in trials between males of the same age class, but it was in trials between an old and a young male, often differing in colour. Thus, age may be a more important determinant of dominance than colour. Female preference for more colourful males was tested by allowing hormone-induced females to choose between a more and a less colourful dummy male. Females spent more time in front of more colourful than drabber males and also more often perched beside colourful males than duller individuals. Although male colour cannot be ruled out as a dominance signal, the results suggest that male colour is primarily used as a signal in mate choice. Female choice may hence be responsible for maintenance of bright plumage in the male yellowhammer.     

Black beak tip coloration as a signal of phenotypic quality in a migratory seabird

Many different seabirds including terns have black color at the tip of their beak, but a yellow, orange, or red color of the rest of the beak. The tip of the beak of Arctic terns Sterna paradisaea is black, while the rest of the beak is deep red during the breeding season. We hypothesized that the black coloration of the tip of the beak is a reliable signal of phenotypic quality maintained by the costs of deposition of melanin. In this paper, we tested a number of predictions arising from this hypothesis. Consistent with the hypothesis, we found that (1) Arctic terns mated assortatively with respect to beak color score; (2) early breeding adults had larger beak color scores than late breeders; (3) clutch size was larger in adults with large beak color scores; (4) adults with a small beak color scores were more likely to die in the immediate future than individuals with large scores; and (5) beak color score was related to environmental conditions at the breeding grounds, as reflected by the North Atlantic Oscillation and the amount of fertilizer used in agriculture, and hence, leaking to coastal waters and environmental conditions at the Antarctic wintering areas as reflected by the Southern Oscillation. These findings suggest that beak color score provide reliable information about individual phenotypic quality.     

Courtship in fast water currents by a male stream goby ( Rhinogobius brunneus) communicates the parental quality honestly

The stream goby Rhinogobius sp. DA (dark color type) shows exclusive paternal care of the eggs. Males court females in the stream current and previous field observations suggest that females favor males that perform their courtship display in faster water currents, and that such males may have high parental ability because of good physical condition. To validate these observations we examined female choice under controlled laboratory conditions. Mate choice experiments indicated clearly that females preferred males that courted in the faster currents, whereas neither sexually different morphological traits, such as body size, nor the water current alone were important. The experiments with food supply treatments indicated that only males of high physical condition are able to court in the fast currents. Furthermore, males that courted in the fast currents achieved good egg survival, whereas males that were unable to court in the currents did not, due to cannibalism of their egg clutch. Thus, the maximum speed of the water current in which a male courts should be indicative of his quality and of the subsequent survival of eggs under his care. We conclude that Rhinogobius sp. DA females utilize the male courtship display in the water current as an honest indicator of parental quality.Communicated by K. LindströmAn erratum to this article can be found at     

Food site residence time and female competitive relationships in wild gray-cheeked mangabeys (<Emphasis Type="Italic">Lophocebus albigena</Emphasis>)

Authors of socioecological models propose that food distribution affects female social relationships in that clumped food resources, such as fruit, result in strong dominance hierarchies and favor coalition formation with female relatives. A number of Old World monkey species have been used to test predictions of the socioecological models. However, arboreal forest-living Old World monkeys have been understudied in this regard, and it is legitimate to ask whether predominantly arboreal primates living in tropical forests exhibit similar or different patterns of behavior. Therefore, the goal of our study was to investigate female dominance relationships in relation to food in gray-cheeked mangabeys (Lophocebus albigena). Since gray-cheeked mangabeys are largely frugivorous, we predicted that females would have linear dominance hierarchies and form coalitions. In addition, recent studies suggest that long food site residence time is another important factor in eliciting competitive interactions. Therefore, we also predicted that when foods had long site residence times, higher-ranking females would be able to spend longer at the resource than lower-ranking females. Analyses showed that coalitions were rare relative to some other Old World primate species, but females had linear dominance hierarchies. We found that, contrary to expectation, fruit was not associated with more agonism and did not involve long site residence times. However, bark, a food with a long site residence time and potentially high resource value, was associated with more agonism, and higher-ranking females were able to spend more time feeding on it than lower-ranking females. These results suggest that higher-ranking females may benefit from higher food and energy intake rates when food site residence times are long. These findings also add to accumulating evidence that food site residence time is a behavioral contributor to female dominance hierarchies in group-living species.     

Selected polyandry: female choice and inter-sexual conflict in a small nocturnal solitary primate (<Emphasis Type="Italic">Microcebus murinus</Emphasis>)

Sex-specific interests over the maximization of reproductive success lead to an inter-sexual conflict over the optimal mating system in a species. Traditionally, the outcome of this inter-sexual conflict has been studied from the male perspective but it also depends on female mating strategies, such as manipulating the temporal distribution of sexual activity, advertisement, and mate choice. We used a small nocturnal primate, the gray mouse lemur (Microcebus murinus) to determine the relative importance of female mating strategies on the outcome of this conflict in a species where females are solitary during their activity period. We studied their mating behavior over three consecutive annual mating seasons and determined the genetic relationships among more than 300 study animals to quantify individual reproductive success. We found that most females were receptive asynchronously. Females did not exhibit any obvious direct mate choice, probably due to a highly male-biased operational sex ratio and the corresponding costs of choosiness. However, females exercised indirect choice for multiple matings. They mated with 1–7 males up to 11 times during their single night of receptivity. As a result, mixed paternity was common but heavier males sired more offspring, meaning that indirect female choice for superior males cannot be excluded. Females exhibited a mixed mating strategy, avoiding costly direct mate choice but still counteracting male efforts to monopolize mating, successfully increasing genetic variability among offspring. Thus, females had a major influence on the outcome of the inter-sexual conflict despite male monopolization attempts.Communicated by J. Setchell     

Intermittent breeding as a cost of site fidelity

Intermittent breeding (skipping a breeding season) can be the result of an adaptive decision by a focal individual, trading current reproductive success in favour of future reproductive success (residual reproductive value hypothesis). In contrast, an individual can also be forced by conspecifics to abandon the familiar breeding site and refrain from breeding due to lack of suitable alternative breeding sites or mates (competition hypothesis). I studied intermittent breeding in the territorial and site-faithful Eurasian oystercatcher Haematopus ostralegus, using a dataset covering 20 years. Intermittent breeding (in total 86 cases) occurred among breeders that formerly bred in high- as well as low-quality territories. The main factor associated with intermittent breeding in high-quality sites was death of a mate, while in low quality sites divorce was the most prominent factor. In 93% of the cases birds were forced to cease breeding due to pressure from conspecifics consistent with the competition hypothesis. There was no association between intermittent breeding and promotion to a territory of better quality. Instead, oystercatchers returned to breeding habitat of similar quality and at a very close distance (median distance 128 m) from the previous breeding location. Breeding absences lasted on average 2.4 years, with a maximum of 9 years, and the quality of the territory obtained after the absence varied with the duration of it. Birds who re-bred in a high-quality territory acquired this on average faster than those that re-bred in a low-quality territory, indicating that birds in high-quality sites are better competitors.     

Experimental and natural changes in the peacock's (Pavo cristatus) train can affect mating success

Petrie et al. (1991) demonstrated a correlation between the degree of elaboration of peacocks' trains and their mating success, and also showed that this correlation occurred because females preferred to mate with the male that had the most elaborate train of those sampled on the lek. Although these data suggest that female choice is responsible for non-random mating in this species, they do not conclusively show that train morphology is the cue that females respond to, because they do not rule out the possible influence of another unidentified variable which is correlated with train elaboration. This paper presents an experimental test of the importance of the peacock's train in determining male mating success. If the number or arrangement of eye-spots in the peacock's train influences mating success, then changing the number of eye-spots should change mating success. This prediction was tested in an experiment where the trains of male peafowl (Pavo cristatus) were manipulated by removing a number of eye-spots between mating seasons. Peacocks with eye-spots removed showed a significant decline in mating success between seasons compared with a control group. This result, together with the observational data, supports the hypothesis that the peacock's train has evolved, at least in part, as a result of female choice.     

Multiple mate choice criteria and the importance of age for male mating success in the microhylid frog, Cophixalus ornatus

We examined multiple mate choice criteria in Cophixalus ornatus, a terrestrial breeding, microhylid frog. Mate choice consisted of three stages: mate attraction (male calling), courtship (male behavior between the call site and the nest), and nest site selection by the female. For male C. ornatus, the possession of a call with low dominant frequency relative to calling neighbors increased the probability that they would attract females. Dominant frequency was negatively correlated with age independent of male mass and snout vent length. When escorting the female from the call site to their nest, males traveled along more convoluted paths than when returning to the nest alone. The convolution of the path was, therefore, considered an aspect of courtship. Females released eggs into nests with structural characteristics typical of nests constructed by older males. Thus, females increased their chances of locating an acceptable nest by preferentially approaching males with lower dominant frequencies. This study is the first to demonstrate that age, independent of mass or snout-vent length, can influence call characteristics in anurans, and it is also the first to demonstrate the importance of male age to female mate choice in an amphibian.     

Asexuals looking for sex: conflict between species and mate-quality recognition in sailfin mollies (Poecilia latipinna)

When two closely related species are sympatric the process of species recognition (identifying conspecifics) and mate-quality recognition (increased fitness benefits) can yield a conflict when heterospecifics resemble high-quality conspecifics. Conflict in species versus mate-quality recognition may serve as a possible mechanism for the persistence of unisexual, gynogenetic Amazon mollies (Poecilia formosa). Amazon mollies require sperm from closely related species (e.g., sailfin mollies, P. latipinna) to start embryogenesis but inheritance is strictly maternal. When choosing mates, male sailfin mollies from populations sympatric with Amazon mollies may rely on traits indicating species identity rather than those indicating mate quality. Conversely, males from allopatric populations may rely more on traits indicating mate quality. Previous work has found that male sailfin mollies in sympatry exhibit a significantly greater mating preference for female sailfin mollies over Amazon mollies compared to males in allopatry. In addition, male sailfin mollies prefer to associate with and produce more sperm in the presence of larger conspecific females, which are more fecund. We hypothesized that male sailfin mollies experience a conflict in species recognition and mate-quality recognition in the presence of Amazon mollies that are relatively larger than female sailfin mollies. To test this hypothesis, we paired males from sympatric and allopatric populations with a larger Amazon molly and a smaller female sailfin molly. We scored the number of mating attempts that males directed to conspecific and heterospecific females. Males in most sympatric and allopatric populations demonstrate no clear preference for conspecifics. In addition, we found some evidence for a difference in mating preference between allopatric and sympatric populations with males from allopatry showing a greater heterospecific mate preference. These results indicate a conflict between species and mate-quality recognition. In sympatry this conflict may contribute to the persistence of gynogenetic Amazon mollies.     

Odour-based species recognition in two sympatric species of sac-winged bats (<Emphasis Type="Italic">Saccopteryx bilineata,S. leptura)</Emphasis>: combining chemical analyses,behavioural observations and odour preference tests

Combining chemical analysis and odour preference tests, we asked whether two closely related sympatric species of sac-winged bats use odour for species recognition. Males of the two sister species Saccopteryx bilineata and Saccopteryx leptura have pouches containing an odoriferous liquid in their antebrachial wing membrane, which is used in S. bilineata during courtship displays. Although both species occasionally share the same daytime roosts and are morphologically similar, there is no evidence for interbreeding. We compared the production and composition of the wing sac odorant in male S. leptura and S. bilineata and performed odour preference tests with female S. bilineata. Similar to male S. bilineata, male S. leptura cleansed and refilled their wing sacs with secretions, but they spent more time each day in doing so than male S. bilineata. Chemical analysis by gas chromatography and mass spectrometry revealed that male Saccopteryx carried species-specific scents in their wing sacs. Binary choice tests confirmed that female S. bilineata preferred the wing sac scents of male S. bilineata to those of the sister species, suggesting that the species specificity of male wing sac scents maintain the pre-mating isolation barrier between these closely related species.     

Exogenous testosterone increases female aggression in the European starling (<Emphasis Type="Italic">Sturnus vulgaris</Emphasis>)

In the European starling, Sturnus vulgaris, optimal mating systems differ between males and females. Males gain from polygyny, whereas monogamy increases female fitness. The cost of polygyny to females lead to intense female–female competition, and it has previously been shown that the intensity of female aggression during the pre-breeding period can predict the realised mating system. The physiological regulation of such female aggression in starlings is not yet known. This study examines the role of testosterone in mediating aggressive behaviours involved in intra-specific reproductive competition in female starlings. Testosterone levels were experimentally elevated with testosterone implants in females during the pre-laying period. To simulate a situation in which an additional female tried to mate with the focal female’s mate, a caged female was presented close to a nest-site to which the male could attract a secondary female. Testosterone was significantly related to several behaviours involved in female–female interactions. Females with testosterone implants spent significantly more time close to the caged female and produced more song bouts than control females. In contrast, male behaviour was unrelated to the experimental status of the mate. Females mated to males that attracted a secondary female were less aggressive towards the caged female than those that remained monogamously mated. The effect of exogenous testosterone in this study indicates that androgens may mediate social behaviours in female starlings during the breeding season.     

Changes in singing behavior of male black-capped chickadees (Parus atricapillus) following mate removal

We removed the mates of ten male black-capped chickadees (Pares atricapillus) during the nest-building period to determine the effect of female presence on dawn singing. During the first dawn chorus following mate removal, males sang significantly longer, increased movement within their territory, and increased the percentage of their territory covered while singing. After the female was returned, these parameters returned to the pre-removal values. Males did not alter the frequency range or modal frequency of their songs when the mate was removed, nor did they change the degree of frequency shifting in the fee-bee song. We conclude that dawn singing in the black-capped chickadee acts, in part, as an intersexual signal, and that the behavior of frequency shifting in the song may be directed more toward rival males than females. Correspondence to: K. Otter     

Dynamic female preference for multiple signals in <Emphasis Type="Italic">Rhinogobius brunneus</Emphasis>

Many species base their choice of mates on multiple signals which provide them with different kinds of information. Choosers may assess the signals together to evaluate the overall quality of potential mates, but individuals often pay attention to different signals in different contexts. In Rhinogobius brunneus, a fish displaying exclusive male parental care, females generally prefer males showing larger first dorsal fins (FDF) and more active courtship displays as mates. Females choosing a mate usually initially assess the FDF and later utilize courtship for the final decision. In our experiments, females with different hunger states used different signals when selecting mates. Females in both hunger states preferred males with larger FDF in the first stage. In the second stage, well-fed females showed highly repeatable choice, whereas poorly fed females responded only to variation in the courtship activity of males. The males preferred by poorly fed females exhibited significantly higher offspring survival than nonpreferred males. Under conditions of food shortage, males allocate more energy to future reproduction at the expense of the present brood, and females may prioritize signals predictive of offspring survivorship over signals reflecting other aspects in male quality to minimize the losses in direct benefits. We conclude that R. brunneus females may employ information from both signals but dynamically adjust their prioritization of each signal to current conditions to ensure the choice that is currently most adaptive.     

污染场地河道底泥浸出液对斑马鱼胚胎的毒性效应

为了评价某污染场地河道底泥对水生生物的毒性效应及其潜在生态风险,以斑马鱼(Brachydanio rerio)为受试生物,进行了河道底泥浸出液对斑马鱼胚胎的发育毒性实验。结果表明,5种底泥浸出液(A1、A2、A3、A4、A5)对斑马鱼胚胎的发育均有毒性效应,浸出液对斑马鱼胚胎的死亡率存在一定的剂量-效应关系,随着浸出液体积分数的增加,死亡率显著上升。浸出液对胚胎孵化抑制作用明显,随着浸出液体积分数的增加,孵化率显著下降。且能引起斑马鱼胚胎多种中毒症状,如水肿,鱼体弯曲,发育不全等异常现象,表明该段河道底泥存在潜在的生态风险。因此,在后期的污染场地河道整治中,需对河道底泥进行挖掘清理和无害化处理。     

Use of trace element analysis of feathers as a tool to track fine-scale dispersal in birds

Tracking dispersal and migratory movements of animals over small and large spatial scales is a challenge. In birds, a promising yet underutilized tool is the trace elemental composition of feathers. The elemental profile of a feather may reveal information about the geographic origin of a bird provided that molting occurs on the breeding grounds and that elemental differences exist between breeding areas. Here, we explore the use of trace elemental composition in body feathers of the Puget Sound white-crowned sparrow Zonotrichia leucophrys pugetensis as a tool (1) to discriminate among birds collected in four different song dialect populations along a 400-km stretch of the Pacific Northwest coast and (2) to assign males singing nonlocal dialects in one population to potential natal populations. Inductively coupled plasma mass spectrometry detected 34 trace elements in sampled feathers and in a discriminant function analysis seven of these elements differed among the four source populations. Half of the six nonlocal dialect singers, who were likely to have immigrated into the focal population, were assigned to a population that matched their song dialect. Our study suggests that feather microchemistry is a promising tool for identifying geographic origins of dispersing birds over small geographic scales and in combination with other markers, such as song, may give insight into ecological and evolutionary processes.     

Indirect genetic benefits of polyandry in a spider with direct costs of mating

The search for the evolutionary explanation of polyandry is increasingly focused on direct and indirect selection on female resistance. In a polyandrous spider Stegodyphus lineatus, males do not provide material benefits and females are resistant to remating. Nevertheless, polyandrous females may obtain indirect genetic benefits that offset the costs associated with multiple mating. We manipulated the opportunity for females to select between different partners and examined the effect of female mating history (mated once, mated twice, or rejected the second male) on offspring body mass, size, condition, and survival under high- and low-food rearing regimens. We found that multiple mating, not female choice, results in increased female offspring body mass and condition. However, these effects were present only in low-food regimen. We did not find any effects of female mating history on male offspring variables. Thus, the benefits of polyandry depend not only on sex, but also on offspring environment. Furthermore, the observed patterns suggest that indirect genetic benefits cannot explain the evolution of female resistance in this system.     

Regulation of worker reproduction in bumblebees (Bombus terrestris): workers eavesdrop on a queen signal

In the annual bumblebee Bombus terrestris, the onset of queen-worker conflict over male production is seasonally and socially constrained. Workers will do better if they start to reproduce (the so-called competition phase) only after ascertaining that larvae are committed to gyne development but before the season ends because they gain more by rearing sister-gynes than their own sons. Here, we tested two nonmutually exclusive hypotheses as to what triggers the onset of worker reproduction: Workers can directly monitor larval development and/or workers eavesdrop on the queen signal that directs gyne development. Exposing workers to gyne larvae through a double mesh did not advance the competition phase compared to control colonies. However, when workers, but not the queen, were allowed contact with gyne larvae, both the competition phase and gyne production were advanced. Thus, while larvae do not emit a volatile pheromone that discloses their developmental route, the physical contact of workers with such larvae triggers early competition phase. However, workers exclusively exposed to worker larvae (colonies prevented from producing gyne larvae) started to reproduce at the same time as control colonies. Replacing the resident queen with an older queen (from gyne-rearing colonies) advanced the competition phase, irrespective of worker age. The results are consistent with the hypothesis that workers eavesdrop on the queen pheromones. This is adaptive because it allows workers a broader time-window for reproduction and thus to gain fitness from rearing both sister-gynes and sons before the season ends without affecting colony development.