Coping with Shifting Nest Predation Refuges by European Reed Warblers Acrocephalus scirpaceus

Predation, the most important source of nest mortality in altricial birds, has been a subject of numerous studies during past decades. However, the temporal dynamics between changing predation pressures and parental responses remain poorly understood. We analysed characteristics of 524 nests of European reed warblers monitored during six consecutive breeding seasons in the same area, and found some support for the shifting nest predation refuge hypothesis. Nest site characteristics were correlated with nest fate, but a nest with the same nest-site attributes could be relatively safe in one season and vulnerable to predation in another. Thus nest predation refuges were ephemeral and there was no between-season consistency in nest predation patterns. Reed warblers that lost their first nests in a given season did not disperse farther for the subsequent reproductive attempt, compared to successful individuals, but they introduced more changes to their second nest sites. In subsequent nests, predation risk remained constant for birds that changed nest-site characteristics, but increased for those that did not. At the between-season temporal scale, individual birds did not perform better with age in terms of reducing nest predation risk. We conclude that the experience acquired in previous years may not be useful, given that nest predation refuges are not stable.     

Responses of breeding Reed Warblers Acrocephalus scirpaceus to mounts of Sparrowhawk Accipiter nisus, Cuckoo Cuculus canorus and Jay Garrulus glandarius

Mounts of a Cuckoo Cuculus canorus , a Sparrowhawk Accipiter nisus and a Jay Garrulus glandarius were presented at nests of Reed Warblers Acrocephalus scirpaceus at all breeding stages. Response strength could best be classified according to approach distance (to the mount) and vocalizations used. Warblers reacted much more aggressively to a mount on the nest than to one 3 m adjacent; they approached much more closely, were less likely to sing but often gave a frenzied-sounding rasp (which was never given to an adjacent mount). Reaction was more aggressive to a Cuckoo, which was often attacked, than to a Sparrowhawk, of which they were wary; response to a Jay was intermediate. Reaction was more aggressive after the clutch had been completed than before. After the brood had fledged, parents still responded strongly to the Jay and Sparrowhawk, but responded to the Cuckoo only briefly before ignoring it. These results are consistent with the Reproductive Value-Stimulus Value hypothesis. They also demonstrate that Reed Warblers can tell Cuckoos from Sparrowhawks, as can many non-host species. The close resemblance of these two species is often suggested to be mimicry of the Sparrowhawk by the Cuckoo but, if this is so, it is not successful. Alternatively, the similarity could be mimicry to reduce predation on the Cuckoos themselves, a chance resemblance, or due to similar selective forces acting on both species, as each would presumably benefit from reduced conspicuousness.     

Effects of mate removal on the behaviour and reproductive success of Reed Warblers Acrocephalus scirpaceus

The Reed Warbler Acrocephalus scirpaceus is a largely monogamous insectivorous passerine in which males and females provide equal care by day to eggs and chicks. Polygyny occurs occasionally, with males leaving one female unaided. When females were temporarily removed from three recently-completed clutches their males deserted and resumed the high song levels typical of unmated males. Males may desert either because they are physically incapable of incubation or because the energy expenditure needed for a male to return to an equivalent stage in the breeding cycle is much lower than for a female to do so. Widowed females ( n = 7 ), however, continued the breeding attempt alone, with similar incubation levels but higher provisioning rates than those of control females. In three out of four mid-season broods raised by lone females all fertile eggs were reared to healthy fledglings (in the fourth brood the female died), while only one of four late-season nests produced any fledglings (which were underweight). Late-season control nests were as successful as earlier ones. Loss of male help led to starvation of chicks, but caused no adverse effects during incubation. This explains the small changes in widows' sitting levels during incubation, but much greater effects after hatching. Females may need male help to rear late broods (but not early broods) as days are shorter and food is scarcer. Males may normally help at nests, even those in the mid-season, because in stressful spells (even for a few days) such help is vital for successful breeding but in good periods it costs the male little.     

Parental Investment and Territorial/Sexual Behavior in Male and Female Reed Warblers: Are they Mutually Exclusive?

The behavioral and endocrine changes from prenesting to feeding phases of reproduction were examined in free-living reed warblers. Apart from the onset of feeding behavior, these changes in males are particularly evident in song, territorial behavior and circulating levels of testosterone. The interactions between an individual's physiology and behavior during these behavioral transitions were tested first by correlating song and feeding rates with circulating levels of testosterone and secondly by implanting feeding males with steroids and documenting the resulting effects on song, territoriality and parental behavior. The physiological mechanisms associated with the transition from the sexual phase (nest-building and egg-laying) to the parental (incubation and feeding) in females were also examined. In this case, estrogens were implanted during the parental phase to enhance sexual behavior. The results demonstrate an antagonistic relationship between sexual/territorial behavior and parental care in males which is mediated by changes in circulating androgens and no such antagonism in females. The effects of parental manipulation on the feeding rates of the partners and the growth rates of offspring have been examined to shed light on the necessity for male parental input.     

Sexual Differences in the Responses of Captive Great Reed Warblers (Acrocephalus arundinaceus) to Variation in Song Structure and Repertoire Size

The function and evolution of repertoires in songbirds is still not clearly understood. This study involves the great reed warbler Acrocephalus arundinaceus and playback of recorded song to both captive males and females under controlled laboratory conditions. Females were implanted with oestradiol and the copulation solicitation display used as an index of sexual response. Males were not implanted and crest-raising used as an index of aggressive response. Two types of songs were used (short and long) and two sizes of repertoire (high and low). Females only responded to long songs, and responded significantly more to high than low repertoires. It was concluded that long songs have a sexual function and that high repertoires are the result of intersexual selection through female choice. Males responded to short and long songs, but did not discriminate between high and low repertoires. Adding a live male to the experiment made no difference to female responses but males responded significantly more. When field studies are also taken into consideration the overall conclusion is that song quality and repertoire size is particularly important in female choice, and that short songs and a male presence are more important in aggressive behaviour. Integration of both field and laboratory studies on males and females should lead to more rigorous testing of hypotheses concerning the function and evolution of bird songs.     

Stable isotope ratios in winter‐grown feathers of Great Reed Warblers Acrocephalus arundinaceus,Clamorous Reed Warblers A. stentoreus and their hybrids in a sympatric breeding population in Kazakhstan

Analyses of the stable isotope composition of feathers can provide significant insight into the spatial structure of bird migration. We collected feathers from Great Reed Warblers Acrocephalus arundinaceus, Clamorous Reed Warblers A. stentoreus and a small sample of their hybrids in a sympatric breeding population in Kazakhstan to assess natural variation in stable isotope signatures and delineate wintering sites. The Great Reed Warbler is a long‐distance migrant that overwinters in sub‐Saharan Africa, whereas the Clamorous Reed Warbler performs a short‐distance migration to the Indian sub‐continent. Carbon (δ¹³C), nitrogen (δ¹⁵N) and deuterium (δD) isotope signatures were obtained from winter‐grown feathers of adult birds. There were highly significant differences in δD and less significant differences in δ¹³C between Great and Clamorous Reed Warblers. Thus, our results show that the stable isotope technique, and in particular the deuterium (δD) signal, resolves continental variation in winter distribution between these closely related Acrocephalus species with sympatric natal origin. The isotope signatures of hybrid Great × Clamorous Reed Warblers clustered with those of the Great Reed Warblers. Hence, a parsimonious suggestion is that the hybrids undergo moult in Afrotropical wintering grounds, as do the Great Reed Warblers. The observed δD values fell within the range of expected values based on available precipitation data collected at precipitation stations across the wintering continents of each species. However, the power to predict the winter origin of birds in our study system using these data was weak as the expected values ranged widely at this broad continental scale.     

Nest defence,enemy recognition and nest inspection behaviour of experimentally parasitized Reed Warblers Acrocephalus scirpaceus

Capsule Reed Warblers in a regularly parasitized population do not recognize Cuckoo Cuculus canorus as a special enemy and do not change their behaviour at nest immediately after being parasitized.

Aims To assess if an intruder near the nest influences the behaviour of the Cuckoo host.

Methods Host responses to Cuckoo, control Pigeon dummies and human intruder were observed. Host behaviour at 71 nests was video-recorded for 30 minutes at four experimental groups of nests: Cuckoo dummy, Cuckoo dummy + Cuckoo egg, Pigeon dummy, human intruder.

Results Reed Warblers did not respond differently to the Cuckoo and the control species. The experimental procedure had no significant effect on the behaviour of hosts during the study period. We were unable to find any differences in the time spent at the nest, clutch inspection behaviour and nest defence behaviour between morning and afternoon experimental groups. Our results do not support the hypothesis that afternoon laying by the Cuckoo is maintained by a selection pressure from the host. We observed no ejection or egg-pecking during the 30-min period after the experimental parasitism.

Conclusions Low aggression and non-specificity of host responses in our study area are in line with the fact that the Reed Warbler is an intermediate rejecter of Cuckoo eggs as expected from the spatial habitat structure hypothesis.     

Enemy Recognition of Reed Warblers (Acrocephalus scirpaceus): Threats and Reproductive Value Act Independently in Nest Defence Modulation

Organisms should respond more aggressively towards species perceived as a danger to their offspring, but intensity of defence may be gauged by the value of current offspring weighed against the value of future reproductive opportunities. We tested whether defensive responses of nesting reed warblers (Acrocephalus scirpaceus) are the result of an interaction effect between the type of stimulus confronted and the value of the warbler’s nesting attempt. We quantified the ability of reed warblers to discriminate among brood parasites, nestling predators and non‐threatening species at different stages of the breeding cycle. We also determined whether variables that influence the value of offspring, such as time of season, size and age of clutch or brood, and time of day and number of visits to the nest, explain variation in the intensity of defence recorded during the egg and nestling stages. Responses to the three stimuli differed significantly, as reed warblers consistently directed their mobbing calls and attacks towards parasites, whereas they were less conspicuous when confronted with models of predators. Reed warblers modulated their responses towards each stimulus in accordance with the threat each posed at a specific nesting stage, whereas they were not affected by other variables relative to their reproductive potential. The churr call, however, was uttered independently of the stimulus, as it was triggered by the mere presence of nestlings in the nest.     

Early settling the following spring: a long-term benefit of mate desertion by male Great Reed Warblers Acrocephalus arundinaceus

In a population of the Great Reed Warbler Acrocephalus arundinaceus in central Japan 33 cases of mate desertion by males were recorded during eight breeding seasons. Thirty of these occurred at the end of the breeding season. The absence of the male parent during the nestling period affected neither the frequency of nestling starvation nor the return of females. Eight deserters were confirmed to have begun moulting prior to the fledging of their young. Males which had abandoned their territory relatively early in the summer tended to settle earlier and to be polygynous the following spring. As earlier settling is an important factor influencing polygyny in males, an increased chance of early settling the following season can be regarded as a long-term benefit of mate desertion.     

Longer is fatter: body mass changes of migrant Reed Warblers (Acrocephalus scirpaceus) staging at Eilat,Israel

Ecological barriers are the riskiest phases of the annual migrations for migratory birds. Comparatively little field data exists pertaining to the ability of migratory birds to prepare for the challenges of crossing ecological barriers, or their ability to recuperate afterward. Migrating Reed Warblers (Acrocephalus scirpaceus) were captured in Eilat, Israel, during their spring and autumn migrations. Data on spring and autumn body masses, their inter-annual variation, and the pattern of body mass increase were analysed. The birds show a significant inter-annual variation in their body mass and body condition index in both seasons, which is consistent with the data from other sites and for other passerine species. During stopovers, mass gain occurred in both seasons. Birds in poor initial condition, and those that stop over for a longer period of time, gained more body mass faster. In spring, but not in autumn, the progress of the season was also an important factor; late-arriving birds gained more fuel faster. The average rate of fuel gain was 0,157g·day^?1 ± 0.018 SE.     

Catabolic enzyme activities in the pectoralis muscle of premigratory and migratory juvenile Reed Warblers Acrocephalus scirpaceus (Herm.)

Summary Differences in catabolic capacity in the pectoralis major muscle of premigratory and migratory Reed Warblers were examined. The oxidative capacity was greater in the migratory birds, probably reflecting a training effect caused by the increased locomotion activity just prior to migration.Fatty acid oxidation was already high in the premigratory birds, possibly reflecting a saving of carbohydrate for anabolic purposes (feather growth) during the moult. Glycolytic capacity was slightly increased in the migratory birds, suggesting an increased carbohydrate oxidation during migration in spite of the great contribution of fatty acid oxidation.     

Molecular phylogeny of Palearctic-African Acrocephalus and Hippolais Warblers (Aves: Sylviidae)

Phylogenetic relationships of the reed warbler group (genera Acrocephalus, Hippolais, Chloropeta; Aves: Passeriformes) and their potential relatives were studied using nucleotide sequences (1 kb) of the mitochondrial cytochrome b gene. This species-rich but morphologically poorly differentiated group of insectivorous passerines is distributed in Eurasia, Africa, and Australasia. Intergeneric relationships were poorly resolved, but monophyly of the reed warbler group (including Chloropeta) versus other Sylviidae was strongly supported. A basal polytomy within the reed warbler group consists of seven branches and may indicate a rapid early radiation. In the genus Acrocephalus three major clades were identified, which corresponded to phenotypic groups characterized by body size and plumage patterns. However, current delimitation of some subgenera (Acrocephalus, Lusciniola, Bebrornis) is at variance with our phylogeny estimate, and appropriate revisions are proposed. The genus Hippolais, which may or may not be monophyletic, consisted of two well-supported clades of four species each. Some Acrocephalus taxa whose species status had been doubted (griseldis, tangorum, orientalis, australis) proved to be highly distinct genetically. Genetic distances between members of two pairs of allopatric Hippolais taxa (caligata/rama; opaca/elaeica) were as large or larger than between other closely related warbler species. Overall, cytochrome b sequences resolved phylogenetically young relationships quite well, whereas more ancient nodes remained poorly resolved.     

Responses of Male and Female Black Grouse to Male Vocal Display

The evolution of leks may be explained by several hypotheses. The ‘female preference’ hypothesis, which states that females favour males that have aggregated, has recently gained some empirical support. Low-quality, unattractive males may, however, settle near attractive males, as predicted by the ‘hotshot’ hypothesis. We tested whether black grouse Tetrao tetrix females use auditory cues to find the preferred leks, and whether males respond to vocal display emitted on leks. We conducted a playback experiment with male vocal display (rookooing) on leks, where the visiting females and displaying males were counted. The number of males tended to increase more on playback leks. Specifically, the number of 1-year-old males was greater on playback sites than on control sites. Also, the number of females, in relation to the lek size before the start of the experiment tended to increase. In addition, we used aviary playback trials to test whether females distinguish between single-male and multi-male displays. In a choice test, females showed greater preference for the ‘multi-male’ tape. The tendency for increased male numbers on playback leks resulted from increased visits of young, mobile males which were attracted to leks that they perceived to be large. This suggests a ‘hotshot’-type mechanism in the settlement of young males. Because females also responded to the supplemented auditory advertisement, or directly to the increased number of males, the ‘female preference’ hypothesis is also supported. Females may, at least in part, base their decision of which lek to visit on auditory cues, but visual contact to males may be also needed.     

Effect of Post-Weaning Individual Housing on Autonomic Responses in Male Rats to Sexually Receptive Female Rats

Post-weaning individual housing induces significant alterations in the reward system of adult male rats presented with sexually receptive female rats. In this study, we examined the effects of post-weaning individual housing on autonomic nervous activity in adult male rats during encounters with sexually receptive female rats to assess whether different affective states depending on post-weaning housing conditions are produced. Changes in heart rate and spectral parameters of heart rate variability indicated that in post-weaning individually housed male rats, both sympathetic and parasympathetic activity increased with no change in the sympathovagal balance, while in post-weaning socially housed male rats, both sympathetic and parasympathetic activity decreased with a predominance of parasympathetic activity. These two patterns of shifts in sympathovagal balances closely resembled changes in autonomic nervous activity with regard to classical appetitive conditioning in male rats. The autonomic changes in male rats housed individually after weaning corresponded to changes associated with the reward-expecting state evoked by the conditioned stimulus, and the autonomic changes observed in male rats housed socially after weaning corresponded to changes associated with the reward-receiving state evoked by the unconditioned stimulus. These results suggest that different affective states were induced in adult male rats during sexual encounters depending on male–male social interactions after weaning. The remarkable change caused by post-weaning individual housing may be ascribed to alteration of the reward system during sexual encounters induced by deficiency of intermale social communication after weaning.     

A Novel Function for the Hox Gene Abd-B in the Male Accessory Gland Regulates the Long-Term Female Post-Mating Response in Drosophila

In insects, products of the male reproductive tract are essential for initiating and maintaining the female post-mating response (PMR). The PMR includes changes in egg laying, receptivity to courting males, and sperm storage. In Drosophila, previous studies have determined that the main cells of the male accessory gland produce some of the products required for these processes. However, nothing was known about the contribution of the gland''s other secretory cell type, the secondary cells. In the course of investigating the late functions of the homeotic gene, Abdominal-B (Abd-B), we discovered that Abd-B is specifically expressed in the secondary cells of the Drosophila male accessory gland. Using an Abd-B BAC reporter coupled with a collection of genetic deletions, we discovered an enhancer from the iab-6 regulatory domain that is responsible for Abd-B expression in these cells and that apparently works independently from the segmentally regulated chromatin domains of the bithorax complex. Removal of this enhancer results in visible morphological defects in the secondary cells. We determined that mates of iab-6 mutant males show defects in long-term egg laying and suppression of receptivity, and that products of the secondary cells are influential during sperm competition. Many of these phenotypes seem to be caused by a defect in the storage and gradual release of sex peptide in female mates of iab-6 mutant males. We also found that Abd-B expression in the secondary cells contributes to glycosylation of at least three accessory gland proteins: ovulin (Acp26Aa), CG1656, and CG1652. Our results demonstrate that long-term post-mating changes observed in mated females are not solely induced by main cell secretions, as previously believed, but that secondary cells also play an important role in male fertility by extending the female PMR. Overall, these discoveries provide new insights into how these two cell types cooperate to produce and maintain a robust female PMR.     

Satellite Tagging and Biopsy Sampling of Killer Whales at Subantarctic Marion Island: Effectiveness,Immediate Reactions and Long-Term Responses

Remote tissue biopsy sampling and satellite tagging are becoming widely used in large marine vertebrate studies because they allow the collection of a diverse suite of otherwise difficult-to-obtain data which are critical in understanding the ecology of these species and to their conservation and management. Researchers must carefully consider their methods not only from an animal welfare perspective, but also to ensure the scientific rigour and validity of their results. We report methods for shore-based, remote biopsy sampling and satellite tagging of killer whales Orcinus orca at Subantarctic Marion Island. The performance of these methods is critically assessed using 1) the attachment duration of low-impact minimally percutaneous satellite tags; 2) the immediate behavioural reactions of animals to biopsy sampling and satellite tagging; 3) the effect of researcher experience on biopsy sampling and satellite tagging; and 4) the mid- (1 month) and long- (24 month) term behavioural consequences. To study mid- and long-term behavioural changes we used multievent capture-recapture models that accommodate imperfect detection and individual heterogeneity. We made 72 biopsy sampling attempts (resulting in 32 tissue samples) and 37 satellite tagging attempts (deploying 19 tags). Biopsy sampling success rates were low (43%), but tagging rates were high with improved tag designs (86%). The improved tags remained attached for 26±14 days (mean ± SD). Individuals most often showed no reaction when attempts missed (66%) and a slight reaction–defined as a slight flinch, slight shake, short acceleration, or immediate dive–when hit (54%). Severe immediate reactions were never observed. Hit or miss and age-sex class were important predictors of the reaction, but the method (tag or biopsy) was unimportant. Multievent trap-dependence modelling revealed considerable variation in individual sighting patterns; however, there were no significant mid- or long-term changes following biopsy sampling or tagging.     

Genotype-by-Environment Interactions for Female Mate Choice of Male Cuticular Hydrocarbons in Drosophila simulans

Recent research has highlighted the potential importance of environmental and genotype-by-environment (G×E) variation in sexual selection, but most studies have focussed on the expression of male sexual traits. Consequently, our understanding of genetic variation for plasticity in female mate choice is extremely poor. In this study we examine the genetics of female mate choice in Drosophila simulans using isolines reared across two post-eclosion temperatures. There was evidence for G×Es in female choosiness and preference, which suggests that the evolution of female mate choice behaviour could differ across environments. However, the ranked order of preferred males was consistent across females and environments, so the same males are favoured by mate choice in spite of G×Es. Our study highlights the importance of taking cross-environment perspectives in order to gain a more comprehensive understanding of the operation of sexual selection.