Detrimental impacts of radiotransmitters on juvenile Louisiana Waterthrushes

ABSTRACT.   The Louisiana Waterthrush (waterthrush; Seiurus motacilla ) is a forest-dwelling, Nearctic-Neotropical migratory passerine that nests along streams. We attached radiotransmitters (0.6–0.8 g) to 12 nestling waterthrushes using snug, elastic loops. At three nests, adult waterthrushes were videotaped removing radio-tagged young from the nest. In addition, we recovered nine radio-backpacks (with two still attached to the carcasses of nestlings) near nests within a few days after attaching transmitters. Only one of 12 radio-tagged young was relocated more than 24 h after attaching the transmitter. Thus, the method of transmitter attachment we used was not effective. Using snug, nonelastic loops (e.g., nylon) for the harness may reduce the loss of transmitters, but may injure the skin as fledglings grow. Other possible alternatives include (1) gluing the transmitter to skin on the back of nestlings, (2) capturing fledglings in mist nets and attaching transmitters a week or more after fledging by which time contour feathers have grown and the likelihood of a parent removing the transmitter may be reduced, or (3) attempting to monitor fledglings without attaching transmitters. The success of the latter two alternatives would likely be enhanced by attaching transmitters to adults and then tracking them to locate their still-dependent fledglings.     

Effects of ectoparasites of young on parents' behaviour in a Mediterranean population of Blue Tits

In altricial birds, adults are expected to alleviate the impact of parasitism on their young by increasing parental care. We experimentally investigated the effects of blowfly parasitism on adult behaviour in the nest cavity in a population of Blue Tits in which nestlings are heavily parasitized by blowfly larvae. The behaviour of males and females was different, since only the females carried out nest sanitation activities related to the presence of parasites. The time females devoted to nest sanitation was higher in parasitized nests than in experimentally deparasitized ones, suggesting that chicks and adults might share the costs of blowfly parasitism.     

Females compensate for moult‐associated male nest desertion in Hooded Warblers

Uniparental offspring desertion occurs in a wide variety of avian taxa and usually reflects sexual conflict over parental care. In many species, desertion yields immediate reproductive benefits for deserters if they can re‐mate and breed again during the same nesting season; in such cases desertion may be selectively advantageous even if it significantly reduces the fitness of the current brood. However, in many other species, parents desert late‐season offspring when opportunities to re‐nest are absent. In these cases, any reproductive benefits of desertion are delayed, and desertion is unlikely to be advantageous unless the deserted parent can compensate for the loss of its partner and minimize costs to the current brood. We tested this parental compensation hypothesis in Hooded Warblers Setophaga citrina, a species in which males regularly desert late‐season nestlings and fledglings during moult. Females from deserted nests effectively doubled their provisioning efforts, and nestlings from deserted nests received just as much food, gained mass at the same rate, and were no more likely to die from either complete nest predation or brood reduction as young from biparental nests. The female provisioning response, however, was significantly related to nestling age; females undercompensated for male desertion when the nestlings were young, but overcompensated as nestlings approached fledging age, probably because of time constraints that brooding imposed on females with young nestlings. Overall, our results indicate that female Hooded Warblers completely compensate for male moult‐associated nest desertion, and that deserting males pay no reproductive cost for desertion, at least up to the point of fledging. Along with other studies, our findings support the general conclusion that late‐season offspring desertion is likely to evolve only when parental compensation by the deserted partner can minimize costs to the current brood.     

The presence of kleptoparasitic fledglings is associated with a reduced breeding success in the host family in the barn owl

Fledgling birds sometimes abandon their own nest and move to neighboring nests where they are fed by host parents. This behaviour, referred to as ‘nest‐switching’, is well known in precocial birds that are mobile soon after hatching and can easily reach foster nests. In contrast, due to the difficulty of observing nest‐switching in territorial altricial birds, the causes and consequences of moving to others’ nests are poorly known in this group of birds. Nest‐switchers can be adopted by the foster parents or they can steal food from the host parents meant for their offspring, a form of kleptoparasitism, which may result in reduced breeding success of the host nest. In Israel, 12 barn owl fledglings left their natal nests and were found in 9 host nests out of 111 monitored nests (8.1%). Nest‐switchers that fledged earlier in the breeding season flew shorter distances to reach host nests probably because the density of nests with younger nestlings is higher early in the season. The number of host nestlings fledged and the percentage of nestlings fledged was lower in host nests than in nests without switchers. The occasional nest‐switchers were always older than host nestlings (respectively 80 and 50 days of age, on average) and host parents fledged fewer young when nest‐switchers occupied host nests with younger nestlings. This suggests that nest‐switchers are kleptoparasites because the presence of the older alien fledglings is associated with a lower breeding success of the host parents.     

Nest as an extended phenotype signal of female quality in the great reed warbler

Extended phenotypes with signalling function are mostly restricted to animal taxa that use construction behaviour during courtship displays. However, they can be used also as post‐mating signals of mate quality, allowing individuals to obtain reliable information about their partners. Nest size may have such a signalling function and a lot of indirect evidence supports this view. However, direct evidence based on an experimental approach is still widely missing. Here we test the role of nest size in post‐mating signalling of mate quality in the great reed warbler Acrocephalus arundinaceus, a passerine with female‐restricted nest‐building behaviour. Based on observational data, clutch size, nestling weight, brood size and fledglings’ propensity to return to their natal site positively correlated with nest size. Moreover, we experimentally enlarged great reed warbler nests to investigate whether this manipulation affects male investment in feeding. We found that males fed their nestlings significantly more intensively on enlarged nests than those on control nests. This suggests that nest size in this species serves as a signal of female quality or willingness to invest in reproduction and that it pays males to enhance their feeding effort according to this signal. Thus, we provide convincing evidence that animal communication takes place through the extended phenotypes and that post‐mating signalling of quality is not restricted only to males, but may function equally well in females.     

Sex-differences in nest defence in Fieldfares Turdus pilaris in relation to their size and physical condition

The nest defence of solitary-breeding Fieldfares Turdus pilaris against a human intruder, a corvid and a hawk was studied in a mountain birch forest in central Norway in 1990–2002. Females defended their nests against a human intruder more vigorously than did males before hatching, and vice versa after hatching. Although both sexes increased their nest defence, only males showed a significant increase and showed a greater defence than females at the chick stage. Females with 1-week-old nestlings had longer wings, were heavier and in a better physical condition than those that failed to get nestlings to that stage. Females, but not males, lost condition during nesting. Birds in better condition were more aggressive. The pattern of nest defence by males and females against the corvid was similar to that shown against human intruders, whereas males took greater risks than females towards the raptor. There was weak evidence that birds that were more successful showed greater levels of nest defence. The sex-role reversal in nest defence may be connected to differences in costs in relation to the renesting potential and to the physical condition of the sexes.     

Polygyny in the pied flycatcher,Ficedula hypoleuca: a test of the deception hypothesis

The deception hypothesis has been proposed as an explanation for polygyny in the pied flycatcher. According to this hypothesis, already-mated males hide their mating status with polyterritorial behaviour and thereby increase their chances of obtaining a second mate. In a study area at Oslo, Norway, secondary females raised 84% as many fledglings as did concurrent monogamous and primary females. The unmated males sang most of the time near their nest site, whereas the already-mated males frequently disrupted singing for longer periods in their secondary territories to visit their primary nest; such visits to the primary nest occurred both before and after the time of their second mating. The behaviour of the males suggests that deception of females is not an evolutionarily stable strategy, as an observant female would soon discern the male's status. Another difficulty with the deception hypothesis is that secondary females laid larger clutches than primary females. The number of young fledged from secondary nests was not dependent on the distance to the primary nest. The deception hypothesis was not supported by the data, and the reduced reproductive success of secondary females may be explained by the cost of searching for a mate.     

COOPERATIVE BREEDING OF THE NORTHWESTERN CROW CORVUS CAURINUS IN BRITISH COLUMBIA

The behaviour of helpers at nests of Northwestern Crows was studied on Mandarte Island and Mitlenatch Island, British Columbia. Not all nests had a helper and there was only one helper per nest. Helpers participated in varying degrees in the defence of the territory and nest, feeding of the nestlings and fledglings and they cached food on the territory. Adult males fed helpers, and helpers obtained most of their food on the adults' territory. Adults with helpers laid larger clutches and produced more fledglings per nest than adults without helpers. It is suggested that cooperative breeding in the Northwestern Crow is of recent origin.     

Does Social Mating System Influence Nest Defence Behaviour in Great Reed Warbler (Acrocephalus arundinaceus) Males?

In birds with biparental care, males and females often conflict over how much care to provide to their offspring and it may be substantially influenced by increased level of polygamy. In accordance with sexual conflict theory, males of socially polygynous bird species provide much less care to their nestlings than do males of most socially monogamous species. Most of previous studies, however, have used feeding behaviour as an index for variations in male parental care only. However, this may be skewed if polygynous males compensate for lower feeding assistance through the provision of other parental care such as protection of nests from predators. In this paper, we examine nest defence behaviour in the facultatively polygynous great reed warbler with respect to sex and type of social mating system. We recorded latency to the first arrival, distance from the predator and defensive reaction of each parent towards a human intruder. Socially polygynous males with two simultaneously active nests defended primary females’ nests less vigorously than socially monogamous males, whereas no differences were found between monogamous and primary females. Generally, however, they took a bigger role in nest defence than males in all cases. Our results support an idea that sexual conflict is driven by polygamy and that type of social mating system can influence nest defence behaviour of facultatively polygynous birds. This finding should be taken into consideration when studying nest defence parental care in polygynous mating systems.     

Breeding success of the Great Bustard Otis tarda in Zamora Province, Spain, in 1984

Great Bustards are still fairly abundant (5-2 nests/1000 ha) in Zamora. Overall fecundity was high (92% of adult females attempt to nest; 2–47 eggs/clutch; 89% egg fertility), but pre- and post-hatching mortality were also high (50% of eggs laid; 57% of nestlings hatched), due mainly to agricultural machinery but also to natural predators. Calculated annual yields per 100 adult females were: 227 eggs, 102 hatchlings, 44 fledglings. Production of 44 fledglings/y could sustain a stable adult population (mortality approx. 8%/y) if mortality of immatures is not more than 18–22%/y.     

Brood parasitism increases provisioning rate, and reduces offspring recruitment and adult return rates, in a cowbird host

Interspecific brood parasitism in birds presents a special problem for the host because the parasitic offspring exploit their foster parents, causing them to invest more energy in their current reproductive effort. Nestling brown-headed cowbirds (Molothrus ater) are a burden to relatively small hosts and may reduce fledgling quality and adult survival. We documented food-provisioning rates of one small host, the prothonotary warbler (Protonotaria citrea), at broods that were similar in age (containing nestlings 8–9 days old), but that varied in composition (number of warbler and cowbird nestlings) and mass, and measured the effect of brood parasitism on offspring recruitment and adult returns in the host. The rate of food provisioning increased with brood mass, and males and females contributed equally to feeding nestlings. Controlling for brood mass, the provisioning rate was higher for nests with cowbirds than those without. Recruitment of warbler fledglings from unparasitized nests was 1.6 and 3.7 times higher than that of fledglings from nests containing one or two cowbirds, respectively. Returns of double-brooded adult male and female warblers decreased with an increase in the number of cowbirds raised, but the decrease was more pronounced in males. Reduced returns of warbler adults and recruitment of warbler fledglings with increased cowbird parasitism was likely a result of reduced survival. Cowbird parasitism increased the warblers’ investment in current reproductive effort, while exerting additional costs to current reproduction and residual reproductive value. Our study provides the strongest evidence to date for negative effects of cowbird parasitism on recruitment of host fledglings and survival of host adults.     

Intruders in Nests of the Spotless Starling: Prospecting for Public Information or for Immediate Nesting Resources?

The prospective behaviour for nests by conspecific intruders may be a strategy to gather public information for future reproduction or to secure resources for immediate reproduction. Although the relationship between the sex, age and breeding experience of intruders and the sex and age of residents may be crucial for understanding the significance of nest‐prospecting behaviour, a precise determination of these traits has rarely been addressed in field studies. In a free‐ranging population of spotless starlings ( Sturnus unicolor), inexperienced birds were the predominant intruders. The ratio of male‐to‐female intruders was higher among birds without previous breeding experience, although our results did not allow us to determine whether more inexperienced males or females intruded nests. The average annual rate of intrusion of either sex was strongly correlated with the number of fledglings produced in the study colony the previous year, which seems to indicate that the inexperienced intruders were, in general, yearlings. The proportion of the sexes among the intruders in each study year was correlated with the average age of male owners but not with the age of female owners. This result suggests that the presence of intruding males in the nests was affected by an increased opportunity to find vacancies in nesting territories as resident males became older. This finding supports the hypothesis that, in most cases, birds visited nests to obtain personal information about nesting resources. However, the intrusion rates were higher when there were nestlings than during the incubation period, which suggests that collecting public information was also involved in the intruding behaviour. Both sexes regularly attacked intruders, which implies that the intruders inflicted some kind of cost on the owners. Female floaters were more frequently chased, probably because they regularly lay parasitically in conspecific nests.     

Patterns of affiliation among immature rhesus monkeys (Macaca mulatta)

The affiliative interaction patterns of the immature members of a group of rhesus monkeys at the Yerkes Regional Primate Research Center reflected a strong bias toward matrilineal kin, although this effect was modified by age and sex variables. Association with kin decreased with age, particularly for males. Juvenile males showed less of a kin bias in their behavior than did juvenile females, especially for grooming. Juvenile males also exhibited a preference for interaction with other males. The diminished association with kin and the same-sex bias may be reinforced in adolescence as adult males begin to aggressively target adolescent males involved in agonistic encounters with females or immatures. Adolescent males did not decrease their levels of social interaction relative to those of adolescent females; however, these males preferentially associated with other males (predominantly their own age-sex class) and specifically avoided females and younger animals, both kin and nonkin. Avoidance may diminish conflicts with females or immatures which could result from association, thereby decreasing the potential for selective aggressive interference by adult males. Juvenile and adolescent females maintained strong ties with their kin and preferentially associated with other females and immatures. The breadth of interaction of females with other females may facilitate the establishment of dominance relationships as females mature. Familiarity and predictability may also decrease the necessity of more severe agonistic interaction.     

Brood provisioning rates and fledgling behavior of Cordilleran Flycatchers in southwestern Colorado

The behavior of young songbirds after fledging is one of the least understood phases of the breeding cycle, although parental provisioning rates and movement of fledglings are key to understanding life history evolution. We studied Cordilleran Flycatchers (Empidonax occidentalis) at two sites in southwestern Colorado, USA, from 2012 to 2017. We banded and sexed breeding adults to determine the relative contributions of males and females to nestling and fledgling care, and attached radio‐transmitters to nestlings to facilitate observations of brood behavior after fledging. Females made 60% and 78% of total observed feedings of nestlings and fledglings, respectively. Parental provisioning rates increased with nestling age, and per‐nestling provisioning rates increased with brood size. Parental provisioning rates declined just before fledging, then increased just after fledging. Fledging times of individuals in broods were asynchronous and concentrated during the late afternoon and early evening. Males stopped caring for fledglings before females even though this species is single‐brooded, with some late‐season broods being abandoned by males. Broods spent the first three weeks after fledging within 400 m of nests, after which they began to disperse. Most aspects of the breeding biology of Cordilleran Flycatchers in our study, including the duration of nestling and fledging periods, female‐dominated provisioning, and movement patterns of fledglings, were similar to those of other Empidonax species. However, the times when young fledged were not concentrated in the morning as reported in most other songbirds, and this result warrants additional study of the timing of fledging in ecologically and taxonomically similar species. The increased per‐nestling provisioning rate with increasing brood size was unexpected, and additional study is needed to determine if this increase results from a trade‐off between adult annual survival and productivity favoring increased provisioning of young in larger broods, or from the existence of high‐quality individuals where larger clutches and higher provisioning rates are linked.     

Aggression in the early phases of the reproductive cycle of the male three-spined stickleback (Gasterosteus aculeatus)

Male sticklebacks with nests were more aggressive towards other males than were males which had yet to build nests.Males with nests were more aggressive towards other males than towards non-gravid females. This was not true of fish that had yet to build nests.The increased aggression shown to other males by males with nests was reduced after gonadectomy to the level characteristic of fish that had yet to build nests. This reduction occurred in the first week after the operation. Gonadectomy had no immediate effect on the aggression shown to non-gravid females.The aggression of males that had still to build nests was not reduced by gonadectomy.Gonadectomy resulted in a complete absence of nest building and nest-directed behaviour.     

Plastic sex-roles in the common goby – the effect of nest availability

One of the most important factors shaping animal sex-roles is the operational sex ratio, since a skew of this ratio promotes mating competition in the more abundant sex. In this study, we wanted to see if a change in sex-roles, related to an environmental induced change in operational sex ratio, could be demonstrated in a field situation. Common goby ( Pomatoschistus microps ) males build nests under bivalve shells and provide exclusive parental care. Therefore, available nest sites are crucial for successful breeding in this species and should also affect the operational sex ratio. We addressed the question of whether the availability of nest sites, through an effect on the operational sex ratio, affects courtship and mating competition in the common goby. We increased nest site availability in the field by adding potential nest material (bivalve shells) to one location. We then compared the reproductive behaviour of the gobies at this and another location with lower nest site abundance. In general, the reproductive behaviour differed as predicted. Under nest shortage, males occupied new nests and received eggs faster than under nest excess. Behavioural observations of nests revealed that males initiated courtship more often than females where nest sites were abundant, whereas female courtship dominated where nests were scarce. Males more often rejected females attempting to enter their nests under nest shortage, male sneaking attempts were also more common in this bay. Male-male aggression was frequent at both locations, whereas agonistic interactions between females only occurred in the nest shortage bay. Thus, males should be subject to sexual selection in both bays even though courtship roles differed. These results suggest that nest site availability affects mating competition and courtship roles in the common goby.     

Effects of long-acting testosterone treatment on freeliving pied flycatchers, Ficedula hypoleuca, during the breeding period

Breeding behaviour and spermatogenesis of all males in a population of pied flycatchers, Ficedula hypoleuca, were experimentally prolonged by long-acting testosterone. The behaviour of these birds was compared with that of birds from an untreated population and a population where the males were given the vehicle only. The testosterone treatment prolonged territorial behaviour, and a higher percentage of nests without a feeding male during the nestling period was found in the experimental area. The proportion of polygamous males did not however differ between the experimental and control areas. The feeding activity of those testosterone-treated males returning to their home territories was markedly depressed. The female of such a pair did not compensate for the male's low feeding activity, but fed the nestlings as she would have done with a normal untreated male at the nest. On the other hand, females without a male present during the nestling period compensated for their mates' absence by increasing their own feeding frequency. It was also shown that the song of the male pied flycatcher is dependent upon androgens. The testosterone-treated males produced fewer fledglings than the control males.     

European starlings: nestling condition, parasites and green nest material during the breeding season

Male European starlings (Sturnus vulgaris) intermingle fresh herbs, preferably species rich in volatile compounds, into their dry nest material. In a field study, we investigated whether these herbs affect the mite and bacteria load of the nests and the condition of the nestlings either directly or via parasite control. We examined the amount of herbs and the number of plant species males carried into their nests, the variation of volatile compounds in the headspace air of the nest boxes and mite/bacteria load of the nests throughout the season. The amount of herb material and the number of plant species, the number of substances emanated by these plants and the infestation of the nests with bacteria and mites (Dermanyssus gallinae) increased with season. In a field experiment, we exchanged natural starling nests with experimental nests with or without herbs. We found that the herbs had no effect on the mites but fewer bacteria were sampled in nests with herbs than in nests without herbs. The body mass of the fledging was not related to the season or the mite/bacteria load of the nests. However, nestlings from nests with herbs fledged with higher body mass than nestlings from nests without herbs. Both bacteria and mite load were related to nestling mortality. In nests containing no herbs, the numbers of fledglings declined significantly with the increasing mite load while the mites had no effect on the number of fledglings in nests with herbs. Thus, the nest herbs counteracted the effect of the mites. In conclusion, it seems that volatile herbs can reduce bacterial but not mite infestation of the starling nests. The positive influence of herbs on nestling growth indicates that herbs either directly (perhaps as immunostimulants) improve the condition of the nestlings and help them cope with the harmful effects of mites, or they provide a nest environment beneficial for the nestlings‘ development by the reduction of germs.     

The function of aggressive chases by breeding Black and Red Kites Milvus migrans and M. milvus during the post-fledging dependence period

Black and Red Kites Milvus migrans and M. milvus chase other raptors approaching their nests. The study of this behaviour during the post-fledging period suggested that it reflects mainly, but not only, anti-predator behaviour. The frequency of vigilance and aggressive chases decreased through the post-fledging period as predicted by theoretical models of nest defence. Although predation risks were similar, Black Kites invested more time chasing intruders than did Red Kites. Black Kites, unlike Red Kites, chased away intruding juveniles, which may be interpreted as a behaviour to avoid investment in unrelated fledglings. Black Kites usually nest in loose colonies where the risk of, and selection pressures against, accidental investment in unrelated fledglings is likely to be greater than for Red Kites. Differences in aggressive chases by Black and Red Kites are better related to this than to different predation risks.