Territorial behavior of Western Sandpipers on their nonbreeding grounds: effect of sex and foraging interference

ABSTRACT Nonbreeding shorebirds often alternate social structure between anonymous flocks and territorial behavior in response to different environmental factors. To evaluate specific drivers for one species, we studied the spacing behavior of wintering Western Sandpipers (Calidris mauri) at Bahía Santa María, northwestern Mexico, using behavioral observations. The density, population structure, and territorial behavior of Western Sandpipers differed among three habitat types. Cattail marshes supported 110 birds per ha, 49% males, and no territorial birds. Mangrove flats supported 288 birds per ha, 58% males, and 5% territorial birds. Brackish flats supported 365 birds per ha, 76% males, and 7% territorial birds. Territories consisted of rectangular strips (5–9 m long, N= 77). Territory length was not related to either bird density or number of territorial birds by plot, but was positively related to nearest bird distance. Aggression rate was inversely related to territory length, suggesting that territory length is set by the costs of defense. Foraging rate was independent of territory length, and prey densities in territories did not differ from those in areas used by nonterritorial birds. Males were more likely to be territorial and had a higher aggression rate than females, suggesting that males, which forage more on surface prey, were more affected by foraging interference. Our results suggest that the territorial behavior of Western Sandpipers in our study was an opportunistic strategy to reduce foraging interference. The variation in spacing behavior we documented provides evidence that interference competition affects the social structure of Western Sandpipers during the nonbreeding season.     

Should I Stay or Should I Go Now: Late Establishment and Low Site Fidelity as Alternative Territorial Behaviors

In species in which males defend territories for breeding, males may differ in territorial behavior; alternative behaviors among territorial males are not well understood. In our long‐term study of partially‐migratory song sparrows, we have observed that most territorial males establish territories before females begin nesting and remain site‐faithful both within and between breeding seasons; however, some males establish territories later in the season (late establishers) and/or change territory locations either within or between seasons (movers). Whether late establishment or moving are equally successful strategies for territory defense, or best‐of‐bad‐job options, is not known. Here, we compare the frequencies of these behaviors to demographic variables over a 9‐yr period and compare lifetime tenure and early season nesting success for males who differ in site fidelity and timing of territory establishment. Across years, late establishing was negatively correlated with the return rate of previously territorial males; moving was positively correlated with the number of occupied territories at the start of the breeding season (territory density). While moving was independent of number of years on territory, late territory establishment only occurred in a male’s first year as a territory holder. Of 88 males, 25% established their first territory late, primarily in undefended space; 31% moved. Late and early establishers did not differ in lifetime tenure; movers, however, had longer lifetime tenure than site‐faithful males. Among early establishers, movers and non‐movers did not differ in the number of successful early nests/year or number of young fledged/year; among late establishers, however, movers had significantly higher early nesting success by both measures. Late establishers who moved had higher early season nesting success and higher early season nesting success/year than site‐faithful early establishers. Thus, individual variation in the timing of territory establishment and site fidelity may be facultative alternative territorial strategies.     

Autumnal territorial aggression is independent of plasma testosterone in mockingbirds

Mockingbirds (Mimus polyglottos) show intense territorial activity in the autumn as newcomers attempt to establish space within resident populations. Examination of autumnal territorial behavior showed that unmated males sing more and engage in more territorial fights than mated males. Newcomers that have just acquired space also sing more and show more territorial fights than birds resident to the population for at least one prior season. Among established residents, the average number of territorial fights was greater in birds that shared more territory boundaries with new residents. Radioimmunoassay of plasma samples taken from males during the molt and following the onset of territorial defense showed that during both periods plasma concentrations of testosterone (T), dihydrotestosterone (DHT), and estradiol were basal or below the sensitivity of the assay system. Moreover, groups of males that differed in song and territorial aggression did not differ in plasma concentrations of T, DHT, or luteinizing hormone (LH). Hormone analyses confirm measurements on several other avian species suggesting that sex steroid concentrations are low in the fall and winter and that variations in aggressive behavior at this time of year may be unrelated to LH and androgen levels. Our observations contribute to a growing body of work in temperate passerines indicating that the role of androgens in mediating aggressive challenge may be restricted to the breeding season. The possible hormonal basis (if any) of song and territorial aggression in mockingbirds outside the breeding season remains obscure.     

Territory establishment,song learning strategies and survival in song sparrows

In most songbirds, the processes of song learning and territory establishment overlap in the early life and a young bird usually winds up with songs matching those of his territorial neighbors in his first breeding season. In the present study, we examined the relationships among the timing of territory establishment, the pattern of song learning and territorial success in a sedentary population of song sparrows (Melospiza melodia). Males in this population tend to learn their songs from their neighbors and consequently they show high song sharing with neighbors and use these shared songs preferentially in interactions with them. Males also show significant variation in the timing of territory establishment, ranging from their natal summer to the next spring. Using a three-year dataset, we found that the timing of territory establishment did not systematically affect the composition of the song repertoire of the tutee: early establishers and late establishers learned equally as much from their primary tutors and had a similar number of tutors and similar repertoire sizes, nor did timing of territory establishment affect subsequent survival on territory. Therefore, the song-learning program of song sparrows seems versatile enough to lead to high song sharing even for birds that establish territories relatively late.     

Social factors affect the singing rates of female northern cardinals Cardinalis cardinalis

Social factors, such as the duration of territorial occupancy or of time paired with a mate can affect the rate at which individual birds sing. This study examined the influence of duration of pair‐bond and territory occupancy as well as date on the rate of singing by male and female northern cardinals Cardinalis cardinalis. When differences in breeding status were controlled, female cardinals sang at higher rates earlier in the season. Females in newly formed pairs sang at significantly higher rates than those in pairs that had previously bred together. In contrast, male cardinals did not show significant variation in the rate of singing throughout the season. The song rates of males in newly formed and established pairs did not differ significantly. Song rates for males and females in mated pairs were not significantly correlated. This study suggests that social factors have a strong effect on the rate at which female cardinals sing. It is possible that increased intra‐sexual aggression by females when they are establishing a new territory with a new mate leads to this higher level of song output. Once females have established a territory and acquired a mate, dear‐enemy effects might diminish the need for acoustic territorial defence. Differences in the constraints of nesting or the attraction of extra‐pair mates might explain the sexual differences in male and female singing behaviour.     

Social status influences microhabitat selection: breeder and floater Eagle Owls Bubo bubo use different post sites

Social status can be reflected in many aspects of an individual’s behaviour and ecology, including habitat use and conspecific interactions. In territorial species where at least two social groups – breeding birds and non‐territorial floaters – are recognized, the diverse tasks associated with territorial ownership can lead territory holders to behave differently from the non‐territorial part of the population. Territory holders defend their breeding area and reproduce, whereas floating individuals are dispersing and lead a more transient life, during which they do not show any territorial behaviour even when settling in a more or less fixed area (known as the stop phase). As social interactions are based on visual and vocal cues, the use of specific sites for sending and/or receiving signals can be a crucial choice in an animal’s life. By analysing the post‐site selection of Eagle Owl Bubo bubo breeders and floaters during their nocturnal activity, we found that: (1) territory holders selected more visible and dominant posts than non‐territorial floaters; (2) the choice of posts made by floating individuals did not differ between the wandering and stop phases of dispersal; and (3) floating females intruded more frequently than floating males within a breeder’s home‐range. These findings highlight the fact that two social strategies are possible within the same species, depending on an individual’s social status and its related tasks. Breeders could take advantage of visible locations to declare their status as territory holders, whereas floaters could benefit from a more secretive life to wander unnoticed among occupied territories. This secretive life would help floaters to reduce the risks associated with conspecific aggression. Finally, the greater occurrence of floating females within breeders’ home‐ranges can be explained by the fact that female incursions in a breeder’s home‐range are less risky than male intrusions.     

Why Do Males of Parnassius imperator Fight for Bare Rocks but Not the Nectar Flower During Mate Selection?

Territoriality is an important male behavioral trait used in mate selection of butterflies, but the factors determining the outcome of territorial contests remain controversial. We studied the foraging and territorial behaviors of males in the Parnassius imperator butterfly. Adult males were selective about nectar sources, but they did not defend nectar flowers. By contrast, they fought over areas of bare rocks because bare rocks were used as lek locations to attract and court receptive females. The ownership of a lek, rather than sex pheromones or wing pigments, played the key role in attracting females. In an owner replacement experiment, the owner of a territory always defeated intruders, indicating that perching order, not relative body size, determined the outcome of territorial contests. Territories are small areas of bare rocks that lack food resources, so intruders do not challenge the ownership of a resident in a location where they can easily find vacant territories. Males that first perch in a territory have an advantage compared to intruders in that they can forage on flowers in the vicinity of the territory. This may help explain why satiated owners always defeated fatigued intruders. Our findings suggest that early perching on bare rock allows male butterflies of P. imperator to occupy a territory, attract receptive females and exclude later‐arriving competitors.     

What do foraging hummingbirds maximize?

Summary Hainsworth and Wolf (1976) reported that under certain conditions hummingbirds made food choices which did not maximize their net rate of energy intake while foraging. They concluded that the birds were not foraging optimally. We show here that their birds probably maximized a different utility function, the net energy per unit volume consumed (NEVC), which appears to be an optimal choice on a time scale longer than that of a foraging bout. Our own experiments with Archilochus colubris support the conclusion that hummingbirds make foraging decisions that maximize NEVC. A simulation model shows that, in nature, NEVC maximization would require fewer foraging trips and visits to fewer flowers per day to balance daily energy budgets. For territorial birds this can lead to smaller territory sizes and reduced costs of territorial defense. Plants that evolutionarily increase corolla length to enhance pollinator specificity need only increase nectar concentration slightly to maintain the same net energy per unit volume consumed (NEVC) by a given hummingbird pollinator.     

Differences in venom composition between orb-weaving and wandering Hawaiian Tetragnatha (Araneae)

Spider venoms are complex mixtures of toxins that are used primarily for immobilizing prey. There is evidence of chemical variation in spider venoms among close relatives, yet few studies have analysed their evolution within an ecological and phylogenetic framework. On the Hawaiian archipelago, Tetragnatha, a cosmopolitan orb-weaving genus, has undergone a radiation in which a monophyletic lineage has abandoned web-building and become obligately wandering foragers. This study compares venom composition and details of feeding behaviour between orb-weaving and wandering Hawaiian Tetragnatha. Protein gel electrophoresis patterns indicated that relative to orb-weavers, wandering species had a reduced concentration of low molecular weight (<14kDa) components. Both orb-weaving and wandering Tetragnatha captured flying prey (adult lepidopterans, dipterans), but wandering spiders also captured non-flying prey (insect larvae, spiders). There were no distinct differences between orb-weavers and wanderers in prey capture and immobilization sequences, or in the paralytic effects of bites on prey. However, prey bitten by wanderers took longer to be permanently immobilized than prey bitten by orb-weavers. Contrary to predictions, there was no indication that web-loss in this group was associated with an increase in venom potency.     

Food supply controls the body condition of a migrant bird wintering in the tropics

The hypothesis that migratory bird populations are limited during the non-breeding season is increasingly supported by empirical studies that also suggest consequences that carry-over into subsequent seasons. Although variation in food supply is the best supported explanation for non-breeding season limitation, the ecological mechanisms and physiological consequences are not well understood. We both supplemented and reduced Ovenbird (Seiurus aurocapilla) food availability on replicate plots in Jamaica in each of 3 years to determine the direct role of food in limiting the physical condition of Ovenbirds. Annual variation in rainfall and food supply created a natural experiment in parallel with manipulations. Sex and age-classes of Ovenbirds did not respond differently in terms of body condition to either food manipulation or natural variation in environmental conditions, suggesting that this population is not structured by strong dominance relationships. Ovenbird body mass, fat, and pectoralis muscle shape were positively and predictably related to manipulated food availability. Feather regrowth rate also responded positively to food supplementation and negatively to food reduction in the drier of 2 years. Prior to manipulation, annual variation in body mass corresponded to annual variation in food supply and rainfall, providing additional, correlational evidence of food limitation. Since multiple intercorrelated body condition indices of Ovenbirds responded directly to food supply, and since food supply influenced body condition independently of other habitat features, we argue that food is a primary driver of non-breeding season population limitation. Moreover, since these effects were observed during the late non-breeding period, when individuals are preparing to migrate, we infer that food availability likely initiates carry-over effects.     

Body weight, gonad development and moult in the Tawny owl (Strix alum)

Gonad development, moult and seasonal changes in body weight and composition in the Tawny owl Strix aluco were studied by examining the carcasses of 369 owls (mostly road casualties) supplemented by 112 weights of live birds. In breeding females laying was preceded by the accumulation of fat and to a lesser extent protein which meant that they weighed more at this time (February/March) than at any other. Females declined in weight after laying but were still heavy during incubation. In contrast, males and non-breeding females did not increase in weight before the start of the breeding season. Juveniles reached or even exceeded adult weight well before independence due to the deposition of fat. Even after the exclusion of diseased or contaminated individuals, 9·4% of the birds examined were identified as starving; most of these were in the autumn and were probably newly-independent young wandering in search of territories. In both sexes gonad maturation was of brief duration coinciding with the period (mid-March to mid-April) in which eggs are normally laid. Ovarian growth was biphasic. In the three months prior to the breeding season ovarian condition in different birds was positively correlated with body weight and it appeared that the largest ovarian follicles of females in poor condition failed to attain the size from which rapid growth to final ovulation occurs. in males testis size in the breeding season was correlated with pectoral muscle weight (an index to protein condition) but not body weight. The majority of adults commenced wing moult in June. The average duration of primary moult was estimated to be 77 days. Healthy birds replaced the primaries of both wings at the same rate but most diseased birds moulted asymmetrically and/or out of season. First-year birds renewed their body feathers between September and November. In the Tawny owl territory establishment, breeding and moult are temporally separated.     

EXPERIMENTS ON THE LIMITATION OF BIRD NUMBERS BY TERRITORIAL BEHAVIOUR

(1) This paper examines removal and other experimental studies on the role of territorial behaviour in the limitation of bird densities. Experimental design is discussed, as are the types of conclusions that can be drawn. (2) Experiments have been conducted on more than 40 species from a wide range of taxonomic groups. Most provided evidence for density limitation, probably mediated by territorial behaviour, and for the existence of a non-territorial sector able to take territories when territory owners were removed or when additional habitat was made available. Experiments in the breeding season indicated that some surplus birds, although younger on average then territorial birds, were sexually mature and able to breed when given the chance. In some species, replacement birds bred less well than other territorial birds, but in other species no difference was apparent between the two groups. (3) Other experiments indicated that, while density was limited by territorial behaviour in good habitats, this was not the case in poor habitats, and that some individuals would move from poor to good habitat when territory owners were removed from good habitat. (4) In some species, non-territorial birds of breeding age were present in the population despite the existence of vacant territorial sites, while in others replacements were observed on good territories but not on poor ones. The implications were that site quality influenced whether settlement, defence and breeding occurred, and that some individuals had more stringent site requirements than others. In some seabirds' sites in the centre of a colony were more attractive than sites on the edges. (5) Many land-bird species that have been studied show two peaks of territorial activity each year, in autumn and in spring, and a limitation on breeding density can occur at either or both seasons, depending on conditions. (6) The fact that densities of many bird species fluctuate greatly from year to year is not inconsistent with the limitation of density by territorial behaviour. Several mechanisms are apparent through which density might be limited by territorialism at different levels in different years, so that surplus non-territorial birds are present in years of low territorial density, as well as in years of high territorial density. (7) Experiments have shown that density is limited by the presence of territorial birds, in some species at different levels in different areas or years. The next step is to find whether density is regulated by the presence of territorial birds over a period of years in a density-dependent manner. For this, observational data are required to find whether the proportion of birds that is excluded by territorial behaviour each year varies with the total number available for settlement. Such studies can be made only on species in which non-territorial birds can be counted accurately, as well as territorial ones.     

Patterns of territorial space use by Shining Sunbeams (Aglaeactis cupripennis), tropical montane hummingbirds

For many territorial hummingbirds, habitat use is influenced primarily by the interaction between resource acquisition and non-foraging behaviors such as territory advertisement and defense. Previous research has highlighted the importance of foraging-associated habitat features like resource density and distribution in determining the space-use patterns of hummingbirds. Less is known, however, about how habitat selection associated with non-foraging behaviors influences space use by territorial species. We used radio telemetry to examine patterns of territorial space use by Shining Sunbeams (Aglaeactis cupripennis) in high Andean montane forests near Manu National Park, Peru, and Bosque Comunal “El Carmen” near Chordeleg, Ecuador. We quantified within-territory habitat characteristics related to resource acquisition and non-foraging behaviors such as territory advertisement and defense. We found that Shining Sunbeams showed high use of core areas in territories where foraging effort was relatively low. We found no relationship, however, between the position of core areas and habitat characteristics associated with territory defense, predator avoidance, or other non-foraging behaviors. We also found no relationship between use of non-core areas and habitat use based on resource acquisition. Thus, patterns of territorial space use by Shining Sunbeams may be characterized by core areas not determined by foraging behavior. Further studies examining territorial behaviors and the influence of intrusion pressure will help identify the underlying determinants of territory space use by this and other species of Andean hummingbirds.     

The significance of seed food in chick development re‐evaluated by tracking day‐to‐day dietary variation in the nestlings of a granivorous passerine

The dietary adjustment of nestlings of granivorous birds to a seed diet and the different morphological characteristics of ingested food have rarely been examined in natural conditions. It has been suggested that the provision of cereal grains to nestlings of some seed‐eating bird species in modern agroecosytems is the result of poor food conditions after agricultural intensification. We analysed the abundance of invertebrate prey in the main foraging habitat of parent birds, daily changes (from hatching to fledging) in the efficiency of cereal seed digestion, and the dietary characteristics, diet composition and prey type delivered to nestlings of the Yellowhammer Emberiza citrinella. Analysis of faecal sacs from nests located in breeding habitat with an abundant invertebrate fauna revealed no relationship between the proportion of cereal seeds in the diet of nestlings and the food supply in the main foraging sites of the parents. Neonate nestlings (1 day old) exclusively received weakly chitinized invertebrate prey (arachnids and flies), but from the second day of life the nestlings were fed a variety of highly chitinized invertebrate prey, the percentage biomass of which did not change for the remainder of the nesting period. Cereal grains started to be delivered to 3‐day‐old nestlings and were already efficiently digested, and the percentage biomass of this food type increased progressively with nestling age. We suggest that the provisioning of cereal grains to nestlings is not forced by external factors, such as modern agricultural intensification; rather, it is an intentional behaviour of parent birds aimed at achieving physiological adjustment to seed food in the early stages of ontogeny.     

Mixed patterns of morphological adaptation to insularity in an aerial displaying bird,the Common Snipe Gallinago gallinago

On islands, colonizing birds may evolve behavioural and morphological adaptations to the new environment, often resulting in changes in body size and reduction or even total loss of flight. These island populations have therefore been used to test hypotheses related to adaptations for flight. However, in certain species in which flight is used not only in foraging and migration but also in mating displays, disentangling the effects of natural and social selection is difficult. Thus, sedentary populations of species that perform aerial displays (such as the Common Snipe Gallinago gallinago that breed in the Azores archipelago) may offer an opportunity to separate the effects of natural and social selection on morphology. If insular Common Snipe respond to the characteristic ecological context of oceanic islands, we expect them to differ from migratory conspecifics in body size and by having relatively smaller and more rounded wings. On the other hand, if social selection exerts a more powerful force over the morphology of this species, we expect that sedentary and migratory birds will not differ in flight‐related characters. We tested these hypotheses by comparing morphological characters measured on live Common Snipe captured in the Azores during the breeding season with those measured on migratory specimens hunted during autumn/winter in mainland Portugal. Sedentary Azorean birds were smaller and had relatively shorter tails but did not show the tendency for insular birds to possess more rounded wings as described in other taxa, including in the Azores. Bergman's rule might explain the difference in body size and shorter tails may be responsible for behavioural differences between populations. The lack of difference in wing shape might be explained by the need of the Common Snipe to perform aerial displays during courtship, suggesting an effect of social selection on the migratory strategy of this species.     

Early Comers Become Owners: Effect of Residency Experience on Territorial Contest Dynamics in a Lycaenid Butterfly

A positive relationship between ownership and contest success is widespread among territorial butterflies. We tested the hypothesis that owners win simply because they have a longer experience occupying a given territory than do intruders in territorial butterflies. Using the lycaenid butterfly Chrysozephyrus smaragdinus , we staged contests between two males that had experienced different residency periods at the same contested territory, similar to owners and intruders in natural contests. We found that males that had occupied the contested territory for a longer period (a day) always defeated the shorter occupancy males (15 min). Therefore, residency period plays a key role in settling territorial contests. As residency confers advantages in a time-dependent manner, the mechanism of owner dominance could be an increase in motivation to fight as local familiarity increases. Moreover, this result predicts a population-level territorial dynamics. If residency period per se confers advantages in each contest, males arriving at a territory early in their adult season should acquire territorial dominance because they can remain there for a certain period with only a few rivals; late-appearing males do not have this advantage. As predicted, we found that males appearing at territory sites early in the season were more likely to hold their territories. This finding provides new evidence for the evolution of protandry in butterflies.     

The natural plasma testosterone profile of male blue tits during the breeding season and its relation to song output

In male birds, the gonadal hormone testosterone (T) is known to influence territorial and mating behaviour. Plasma levels of T show seasonal fluctuations which vary in relation to mating system and social instability. First, we determined the natural T profile of male blue tits Parus caeruleus during the breeding season. We found that plasma levels of T increased at the onset of nest building. Thus, the increase in circulating T was not associated with territory establishment, nor with the fertile period of the males' mates. In most individuals, T levels dropped to values close to zero during the period of chick feeding. Second, we investigated the relationship between plasma levels of T and male age, size, and singing behaviour. During the mating period, T levels did not differ between 1 yr old and older males and did not correlate with body size or condition. However, song output during the dawn chorus tended to be positively correlated with T levels. Therefore, if high T levels are costly, song output might be an honest indicator of male quality in blue tits. Finally, we show that plasma levels of T are significantly higher during the night than during the day. This pattern has also been observed in captive non-passerine birds, but its functional significance remains unknown.     

Territorial aggression does not feed back on testosterone in a multiple-brooded songbird species with breeding and non-breeding season territoriality,the European stonechat

Testosterone mediates reproductive behaviours in male vertebrates. For example, breeding season territoriality depends on testosterone in many species of birds and in some, territorial interactions feed back on testosterone concentrations. However, the degree to which territorial behaviour and testosterone are associated differs even between species with seemingly similar life histories, especially between species that also defend territories outside the breeding season. Here, we investigate the link between territorial behaviour and testosterone in European stonechats. Previous studies found that territorial aggression in stonechats depends on testosterone in a breeding, but not in a non-breeding context. We investigated whether stonechats show a rise in testosterone during simulated territorial intrusions (STI) during the breeding season. Post-capture testosterone concentrations of males caught after an STI were not higher than those of males caught in a control situation regardless of breeding stage. However, most of the males would have been able to mount a testosterone response because the same individuals that did not increase testosterone during the STI showed a substantial increase in testosterone after injections of gonadotropin-releasing hormone (GnRH). GnRH-induced and post-capture testosterone concentrations were positively correlated and both decreased with successive breeding stages. Further, territory owners with a short latency to attack the decoy expressed higher post-capture testosterone concentrations than males with a longer latency to attack the decoy. Thus, there is no evidence for behavioural feedback on testosterone concentrations during male-male interactions in stonechats. In combination with previous studies our data suggest that testosterone functions as an on/off switch of high intensity territorial aggression during the breeding season in stonechats. The among-species variation in the androgen control of territorial behaviour may be only partly a result of environmental differences. Instead, potential differences in how territoriality evolved in different species may have influenced whether and how a reproductive hormone such as testosterone was co-opted into the mechanistic control of territorial behaviour.     

Breeding limits foraging time: evidence of interrupted foraging response from body mass variation in a tropical environment

Birds should store body reserves if starvation risk is anticipated; this is known as an ‘interrupted foraging response’. If foraging remains unrestricted, however, body mass should remain low to limit the predation risk that gaining and carrying body reserves entails. In temperate environments mass gain in female birds during breeding is often attributed to egg formation and mass loss after incubation to flight adaptation or the effect of reproductive workload, rather than as a result of an adaptive interrupted foraging response to the limited foraging time or unpredictable foraging conditions that breeding demands. In tropical environments, foraging conditions vary more within the breeding season than in temperate environments, and so studies in tropical environments are more suited to decouple the potentially confounded effects of increase in body reserves versus egg formation on the body mass of breeding birds. In this study, we test whether breeding results in an interrupted foraging response in a tropical savannah system using body mass data collected over a 15‐year period from female common bulbuls Pycnonotus barbatus. This species breeds both in the wet and dry season, despite fewer resources being available in the dry season. Breeding stage predicted female body mass: body mass peaked abruptly during incubation, but was not closely associated with the egg‐laying stage, and declined during brood rearing. Breeding females were heavier in the dry season than in the wet season. In the dry season, heavier birds were more likely to incubate eggs or brood chicks. These observations suggest that increased body reserves are required to buffer the consequence of limited foraging time or impoverished foraging conditions, which may be most pronounced during incubation and in the dry season, respectively. Such mass increases are consistent with an interrupted foraging response, which may apply to temperate zone birds experiencing foraging restrictions during breeding.     

Social factors affect the singing rates of female northern cardinals Cardinalis cardinalis

Social factors, such as the duration of territorial occupancy or of time paired with a mate can affect the rate at which individual birds sing. This study examined the influence of duration of pair-bond and territory occupancy as well as date on the rate of singing by male and female northern cardinals Cardinalis cardinalis . When differences in breeding status were controlled, female cardinals sang at higher rates earlier in the season. Females in newly formed pairs sang at significantly higher rates than those in pairs that had previously bred together. In contrast, male cardinals did not show significant variation in the rate of singing throughout the season. The song rates of males in newly formed and established pairs did not differ significantly. Song rates for males and females in mated pairs were not significantly correlated. This study suggests that social factors have a strong effect on the rate at which female cardinals sing. It is possible that increased intra-sexual aggression by females when they are establishing a new territory with a new mate leads to this higher level of song output. Once females have established a territory and acquired a mate, dear-enemy effects might diminish the need for acoustic territorial defence. Differences in the constraints of nesting or the attraction of extra-pair mates might explain the sexual differences in male and female singing behaviour.