Aggression of breeding eastern bluebirds (Sialia sialis) toward their mates and models of intra-and interspecific intruders

Hypotheses regarding the evolution of aggressive patterns during the breeding season in eastern bluebirds (Sialia sialis) were tested by placing models of eastern bluebirds and house sparrows (Passer domesticus) into territories of experimentally naive breeding pairs of eastern bluebirds during three stages of the nesting cycle. The responses of the resident males to models of male bluebirds decreased from the egg-laying to the nestling stage, and this response pattern can be interpreted as support for the protection-of-parentage hypothesis or the nest-site-defence hypothesis. The aggression of resident males to resident females during trials with models of male bluebirds weakly conforms to the pattern predicted by the protection-of-parentage hypothesis. The aggressive response to models of house sparrows also decreased from the egg-laying to the nestling phase in a manner consistent with the nest-site-defence explanation. No aggression toward mates by males or females was observed during trials with house sparrow models, an observation consistent with the predictions of the protection-of-parentage hypothesis.     

Intra- and interspecific agonistic behaviour in sympatric harriers during the breeding season

We analysed the temporal and sexual patterns of intra- and interspecific aggression in sympatric harriers during the breeding season, to determine the main resource defended (food, nest sites, mates) and how factors such as body size or breeding system (territorial versus colonial) influence aggressive behaviour. We predicted that if aggression is (at least partly) related to competition for food, the hen harrier, Circus cyaneus, because of its large size and territorial system, should be more aggressive (both intra- and interspecifically) than the smaller, colonial Montagu's harrier, Circus pygargus. The intraspecific aggression rate of both harriers peaked early in the season, was mainly intrasexual and increased with the number of neighbours. These patterns support the mate competition hypothesis to explain intraspecific aggression in both species. Montagu's harriers were more aggressive towards conspecifics than hen harriers. Their aggression rates towards heterospecifics were high at the start of the season then decreased, supporting the hypothesis that interspecific aggression serves primarily for nest site defence. Hen harriers showed lower interspecific aggression rates in the prelaying period and a gradual increase throughout the breeding cycle, particularly by females, who hunt around the nest for food for the nestlings. These patterns correspond to food competition, as food resources around the nest are probably more important and interspecific intrusion more costly for territorial hen harriers than for colonial Montagu's harriers. Copyright 2002 The Association for the Study of Animal Behaviour. Published by Elsevier Science Ltd. All rights reserved.     

Brief report: Outbreak of severe aggression in captive Macaca mulatta

An outbreak of severe aggression occurred among females in a rhesus macaque breeding group at the California Primate Research Center four years after the group was established. During the breeding season in which this occurred, the incidence of injured females in other breeding groups at the Primate Center was significantly higher than in the previous year. This breeding season was the first in which a large number of females reached sexual maturity. The group in which the most severe aggression occurred contained the largest number and proportion of maturing females. Evidence suggests that the simultaneous maturation of a large cohort of adolescent females may be associated with increased levels of aggression, and that this aggression may be intensified by certain aspects of captivity.     

Reproductive tactics of the ringed plover Charadrius hiaticula

Reproductive tactics of ringed plovers Charadrius hiaticula were studied at three localities in SW Sweden during five seasons. The usual clutch size is four, but removal experiments showed that females can produce five eggs in succession, with similar intervals between all eggs. High predation made mean breeding success per clutch low, 6.3% of eggs resulting in fledged young. Replacement clutches were common, and many pairs laid again after rearing their first brood to fledging. Egg laying spanned three months, much longer than for other waders in this region. Between years, reproductive success varied unpredictably with time of the season, but averaged over several years, the expected success was low and similar for the different parts of the breeding season. Chicks from late clutches had similar survival and recruitment as others. Because of the long breeding season and high rate of nest failure a female may produce up to five clutches of four eggs per season, containing in total about 3.7 times her own mass. Yearly local survival of adult females and males was estimated to 84.6 and 88.6%, respectively. Ability to produce many clutches with similar expected success throughout the season favours a long reproductive period, sometimes leading to double-brooding. Possible life-history trade-offs are discussed.     

Seasonal variation in reproductive traits of the oriental shrimp Palaemon macrodactylus (Crustacea: Caridea: Palaemonidae) in a non-native population

The magnitude of variations in reproductive traits of Palaemon macrodactylus females throughout a breeding season was studied in a non-native population at Mar del Plata harbor, Argentina. Fecundity, egg size, reproductive output, weight and elemental composition of eggs, and larvae were analyzed in females collected at the beginning, in the mid point, and near the end of a reproductive season and designated as early, middle season, and late females. The highest reproductive output was observed in early females, while the highest fecundity and egg volume occurred in middle season females. Eggs and larvae showed larger body mass in early than in late females. Embryos from early females contained and consumed more carbon during development than embryos from late females, and they also used part of the available nitrogen. Differences in reproduction were observed among the three groups of females. On the one hand, late females matured early but had a poor first reproduction, with few embryos and high egg loss; however, they had longer reproductive life and an enhanced reproductive output in the following season when they became early females. On the other hand, females collected at the midpoint in the reproductive season matured later and had the highest fecundity and egg volume. In addition, larvae with different characteristics resulted from each type of female and were presumably well adapted to the conditions prevailing at the moment they hatched. The extended reproductive period and the diversity of embryos and larvae produced may favor the invading ability of the species.     

Seasonal sex role changes in the blenniid Petroscirtes breviceps, a nest brooder with paternal care

The plasticity of the sex roles in the blenniid fish Petroscirtes breviceps , a nest brooder with exclusive paternal care, was studied throughout an 8 month breeding season. Males performed most courtships early and late in the breeding season, whereas females performed most in the middle of the season. These results indicated that the sex of individuals initiating courtship changed seasonally, with courtship role reversal in the middle of the season. Intrasexual aggression in both sexes occurred much more frequently in mid-season than in the early and late seasons. Males frequently fought when available nest sites were limited, regardless of the presence of females, suggesting that males competed for nests in order to qualify to mate (resource competition). In contrast, courting females fought only in mid-season, when females' relative success in entering nests decreased, indicating that females competed for limited mating opportunities (mating competition). The reversed courtship roles and female mating competition in mid-season suggested that the sex roles in P. breviceps changed seasonally from the conventional roles to reversed roles and back again during one breeding season. This study provides the first empirical evidence of multiple changes in the sex roles of animals within a breeding season.     

The effect of heated effluent on egg production in the estuarine prawn Upogebia Africana (Ortmann)

Egg production of a population of Upogebia africana (Ortmann) in a pond containing water heated by a thermal power station at Knysna, South Africa was compared with that of the population in the adjacent estuary and in two other estuaries. The breeding cycle, proportions of ovigerous females, incubation time for eggs at different temperatures, number of eggs produced, sex ratios, and size-frequency of males and females were all investigated. The number of eggs produced/100 prawns was found to be nearly twice as high in the population exposed to heated effluent as compared with the population in the adjacent estuary. The increase was due to a combination of a change in sex ratio resulting in more females in the population and a greater proportion of larger females in the pond. Since the number of eggs produced/female is positively related to the size of the prawn, large females in warmer water produced more eggs than smaller females in the adjacent estuary. Exposure to heated effluent did not alter the initiation or duration of the breeding season, nor cause precocious maturity; nor did it alter the number of eggs carried per female at various sizes.     

Temporal patterns of circulating LH and steroid hormones in a brood parasite, the Brown-headed cowbird, Molothrus afer.

The breeding behaviour of female Brown-headed cowbirds, Molothrus ater , is remarkably distinct from that of females of most other passerine species. Cowbirds are brood parasites that do not build nests, incubate eggs or feed young. Instead, they have an extended egg-laying stage, and deposit eggs in the nests of hosts within large breeding territories that are defended from other female cowbirds. In this communication, we report the changes in plasma levels of circulating hormones and in morphological characteristics that occur in free-living female Brown-headed cowbirds over a breeding season.
Plasma LH in female cowbirds increases at the onset of breeding and remains elevated for approximately eight weeks, at which time levels decline to basal values. This pattern is similar to that of follicle development, and is consistent with the prolific reproductive output by individuals of this species. Circulating E, levels are quite variable and do not demonstrate any discernible pattern, while plasma B is elevated during the egg-laying period and declines subsequently.
Testosterone titre (T) is highest early in the breeding season, when territorial interactions are generally most frequent. T levels decline slowly thereafter, reaching their lowest values at the season's end. Plasma DHT levels are slightly higher than those for T; DHT exhibits little change until late in the breeding season, when it declines. Fat scores are highest prior to egg laying, and fat reserves diminish as the breeding season progresses. In contrast, body weight increases during the reproductive season, probably as a result of developing eggs in the ovary. These results are consistent with the hypothesis that temporal patterns of hormone secretion are correlated with the expression of social systems.     

Costs and Benefits of Competitive Traits in Females: Aggression,Maternal Care and Reproductive Success

Recent research has shown that female expression of competitive traits can be advantageous, providing greater access to limited reproductive resources. In males increased competitive trait expression often comes at a cost, e.g. trading off with parental effort. However, it is currently unclear whether, and to what extent, females also face such tradeoffs, whether the costs associated with that tradeoff overwhelm the potential benefits of resource acquisition, and how environmental factors might alter those relationships. To address this gap, we examine the relationships between aggression, maternal effort, offspring quality and reproductive success in a common songbird, the dark-eyed junco (Junco hyemalis), over two breeding seasons. We found that compared to less aggressive females, more aggressive females spent less time brooding nestlings, but fed nestlings more frequently. In the year with better breeding conditions, more aggressive females produced smaller eggs and lighter hatchlings, but in the year with poorer breeding conditions they produced larger eggs and achieved greater nest success. There was no relationship between aggression and nestling mass after hatch day in either year. These findings suggest that though females appear to tradeoff competitive ability with some forms of maternal care, the costs may be less than previously thought. Further, the observed year effects suggest that costs and benefits vary according to environmental variables, which may help to account for variation in the level of trait expression.     

Factors influencing the onset and duration of receptivity of female purple rock crabs, Hemigrapsus sexdentatus (H. Milne Edwards, 1837) (Brachyura: Grapsidae)

The effects of males, field, and laboratory conditions on the receptivity of females were tested in the New Zealand purple rock crab Hemigrapsus sexdentatus. Onset and duration of female receptivity is of interest because it influences the time available for mating and therefore the operational sex ratio (OSR), male-male competition, and the extent of sperm competition. Females were receptive once a year for a short time prior to oviposition. The breeding season was highly synchronised and lasted for about 3 weeks (from the end of March to mid-April; southern autumn), after which, almost all females carried eggs. We found few receptive females in the field (0% to 4.9%) during the breeding season despite a large number of crabs examined (935 in 1999 and 555 in 2000), suggesting that females are receptive for less than a day. The onset of the breeding season was the same for the wild crabs and those held in field cages, but the duration of receptivity increased to several days for caged females. The onset of the breeding season of females in the laboratory was earlier compared to females in the field and had, overall, a longer breeding season. Females isolated from males stayed receptive significantly longer (5.5 days) than females caged with males (3.3 days), suggesting that the duration of female receptivity is adjusted according to the presence or absence of males. Our results suggest that females have some control over their receptivity in relation to male presence, and this could influence the outcome of sexual selection.     

Song repertoires and seasonal song in the yellowhammer, Emberiza citrinella

Songs were recorded from male yellowhammers, Emberiza citrinella, in England and West Germany. Sonagraphic analysis revealed three major types of song structure in both populations. Most males had a repertoire of two distinct song types, and sharing between individuals was extremely rare. Most song is produced late in the breeding season, and playback experiments confirmed that it is correlated with an increase in territorial aggression at this time. Some speculations are advanced concerning the origins and functions of song repertoires in the yellowhammer.     

Androgen‐metabolizing enzymes show region‐specific changes across the breeding season in the brain of a wild songbird

The Lapland longspur (Calcarius lapponicus) is an arctic‐breeding songbird that shows rapid behavioral changes during a short breeding season. Changes in plasma testosterone (T) in the spring are correlated with singing but not territorial aggression in males. Also, T treatment increases song but not aggression in this species. In contrast, in temperate‐zone breeders, song and aggression are highly correlated, and both increase after T treatment. We asked whether regional or temporal differences in androgen‐metabolizing enzymes in the longspur brain explain hormone‐behavior patterns in this species. We measured the activities of aromatase, 5α‐reductase and 5β‐reductase in free‐living longspur males. Aromatase and 5α‐reductase convert T into the active steroids 17β‐estradiol (E₂) and 5α‐dihydrotestosterone (5α‐DHT), respectively. 5β‐Reductase deactivates T via conversion to 5β‐DHT, an inactive steroid. We examined seven brain regions at three stages in the breeding season. Overall, aromatase activity was high in the hypothalamus, hippocampus, and ventromedial telencephalon (containing nucleus taeniae, the avian homologue to the amygdala). 5β‐Reductase activity was high throughout the telencephalon. Activities of all three enzymes changed over time in a region‐specific manner. In particular, aromatase activity in the rostral hypothalamus was decreased late in the breeding season, which may explain why T treatment at this time does not increase aggression. Changes in 5β‐reductase do not explain the effects of plasma T on aggressive behavior. © 1999 John Wiley & Sons, Inc. J Neurobiol 41: 176–188, 1999     

Female brown-headed cowbirds', Molothrus ater, organization and behaviour reflects male social dynamics

In four large aviaries, we studied social assortment and reproductive behaviour of female brown-headed cowbirds housed with males differing in age class and in corresponding levels of intrasexual interaction. Juvenile and adult females resided with either (1) adult males, (2) juvenile males, (3) adult and juvenile males, or (4) no males. We observed social behaviour of males and females from September 1999 through to the 2000 breeding season. Throughout the year, males in the different conditions showed different levels of social interaction, with adult males being most engaged in male-male interactions and juvenile males being least engaged in male-male interactions. Females in the four conditions differed in their associations with males and with other females. In conditions with adult males, females spent more time near males, approached males more often, and associated with females less. During the breeding season, females in conditions with more adult males copulated more and produced more fertile and viable eggs. In the condition where females were housed with juvenile males, they copulated less, laid fewer eggs and destroyed more of their eggs. Results indicate that throughout the year, females are sensitive to male age and behaviour in their social group, and that this sensitivity can have reproductive consequences. Copyright 2002 The Association for the Study of Animal Behaviour. Published by Elsevier Science Ltd. All rights reserved.     

Sexual dimorphism, female reproduction and egg incubation in the oriental leaf-toed gecko (Hemidactylus bowringii) from southern China

We studied sexual dimorphism, female reproduction and egg incubation of the oriental leaf-toed gecko (Hemidactylus bowringii) from a population in southern China. The largest male and female in our sample were 60 and 57 mm snout-vent length (SVL), respectively. Males are the larger sex; sexual dimorphism in head size and tail length (TL) is evident in juveniles and adults, with males having larger heads as well as longer tails than females. Oviposition occurred between late May and late July. Females switched from laying two eggs early in the breeding season to 1-2 eggs later in the season. Clutch mass and egg mass were both independent of female SVL, whereas relative clutch mass was negatively correlated with female SVL. The previous conclusion that female H. bowringii lay a single clutch of eggs per breeding season is unlikely to be true. Thermal environments experienced by H. bowringii eggs affect incubation length as well as morphological and locomotor phenotypes of hatchlings. Hatchlings from eggs incubated at 30 degrees C were larger (SVL, tail length and body mass) and performed better in the racetrack than their counterparts from eggs incubated at 24 degrees C. Temperatures suitable for embryonic development are relatively high in H. bowringii, primarily as a consequence of the adaptive response to warm environments in southern China.     

Factors affecting aggression among females in captive groups of rhesus macaques (Macaca mulatta)

Captive groups of primates often exhibit higher rates of aggression than wild, free-ranging groups. It is important to determine which factors influence aggression in captivity because aggression, particularly intense aggression, can be harmful to animal health and well-being. In this study, we investigated the effect of ground substrate as well as season, rank, age, and group size on rates of agonistic interactions per female in seven captive groups of rhesus macaques (n = 70 females, 1,723 focal samples) at the California National Primate Research Center. Agonistic interactions were divided into three categories: displacements, mild aggression, and intense aggression. Females living in enclosures with gravel substrate were 1.7 times more likely to be involved in intense aggression (e.g. chases and physical contact) than females living in enclosures with grass (Poisson regression model: P < 0.001). High-ranking females were at least 1.3 times more likely to be involved in mild (e.g. threats and lunges) aggression than lower-ranking females (low rank: P = 0.03; mid rank: P = 0.001). Females of all ranks were 1.5-1.9 times more likely to be involved in both intense and mild aggression during the breeding season than other seasons. Age and group size did not affect rates of mild or intense aggression. These findings indicate that although some aggression appears to be natural and unavoidable, i.e. aggression during the breeding season, the well-being of captive macaques can be improved by developing grass substrate in outdoor enclosures.     

Androgen-metabolizing enzymes show region-specific changes across the breeding season in the brain of a wild songbird.

The Lapland longspur (Calcarius lapponicus) is an arctic-breeding songbird that shows rapid behavioral changes during a short breeding season. Changes in plasma testosterone (T) in the spring are correlated with singing but not territorial aggression in males. Also, T treatment increases song but not aggression in this species. In contrast, in temperate-zone breeders, song and aggression are highly correlated, and both increase after T treatment. We asked whether regional or temporal differences in androgen-metabolizing enzymes in the longspur brain explain hormone-behavior patterns in this species. We measured the activities of aromatase, 5alpha-reductase and 5beta-reductase in free-living longspur males. Aromatase and 5alpha-reductase convert T into the active steroids 17beta-estradiol (E(2)) and 5alpha-dihydrotestosterone (5alpha-DHT), respectively. 5beta-Reductase deactivates T via conversion to 5beta-DHT, an inactive steroid. We examined seven brain regions at three stages in the breeding season. Overall, aromatase activity was high in the hypothalamus, hippocampus, and ventromedial telencephalon (containing nucleus taeniae, the avian homologue to the amygdala). 5beta-Reductase activity was high throughout the telencephalon. Activities of all three enzymes changed over time in a region-specific manner. In particular, aromatase activity in the rostral hypothalamus was decreased late in the breeding season, which may explain why T treatment at this time does not increase aggression. Changes in 5beta-reductase do not explain the effects of plasma T on aggressive behavior.     

Consequences of a female-biased sex-ratio in a socially monogamous bird: female-female pairs in the Roseate Tern Sterna dougallii

In the socially monogamous gulls and terns, female-biased sex ratios are sometimes revealed by the occurrence of ‘supernormal clutches’, which are usually attended by female-female pairs or other multi-female associations. We studied these phenomena in the endangered Roseate Tern Sterna dougallii at Bird Island, USA, from 1970 to 1995. DNA-techniques were used to sex breeding adults in 1992–94. Supernormal clutches (with three or four eggs) have comprised 1–7% of all Roseate Tern clutches at Bird Island since at least 1970, probably increasing in frequency since 1980. Supernormal clutches were spatially clustered; most were laid late in the peak period of nesting during each season. More than 80% of supernormal clutches and at least 7% of normal clutches were attended by multi-female associations; most of these were female-female pairs, with a few trios (male + two females, or three females) and one quartet (four females). More than half of the multi-female associations attended normal clutches. Some female-female pairs were maintained for up to five years. The age-distribution of females mated to females did not differ significantly from that of females mated to males. Females mated together usually laid eggs synchronously (±2 days). Such females laid fewer eggs than females mated to males (means 1.20 versus 1.73), and had lower fertility and hatching success (about 46% versus 98%); they were less successful in raising young from eggs that did hatch (means 58% versus 73%), but this difference was not significant. Their overall breeding success was much lower (about 0.34 fledglings per female versus 1.35). The sex-ratio of breeders was about 127 females to 100 males; about 20% of breeding females did not have male mates. Female Roseate Terns that do not obtain male mates appear to be of low phenotypic ‘quality’ - based on late laying, small clutches and small eggs. Our data support the hypothesis that such females have a higher fitness if they mate with each other and raise a few young than if they do not breed at all.     

Sex, age, experience and condition as factors affecting arrival date in prospecting common terns, Sterna hirundo

Several studies have shown that seabird colonies consist to a large extent of young nonbreeders (prospectors). These individuals appear at the colony later in the season than established breeders. The reasons for this late arrival have remained unclear in most cases, mainly because of technical difficulties in collecting sufficient data from nonbreeding individuals. We used a novel transponder system to identify remotely the members of a common tern colony, including nonbreeders, during eight breeding seasons and we combined the system with automatic balances. Ninety-two per cent of prospectors returned for the first time when 2 years old and 88.9% of recruits to the breeding population had spent at least one previous season at the colony as prospectors. In both sexes, most individuals prospected for one season, but more males than females prospected for more than one season, although a higher proportion of females started breeding without a previous prospecting phase. Terns arrived earlier in the season the older they were and the more experience of the colony they had, but experience proved to be more important than age. Prospectors gained about 3 weeks with a previous prospecting season whereas an additional year of age allowed birds to arrive only about 6 days earlier. Prospectors returning later in the season arrived with lower body masses. Males on average arrived earlier at the colony than females. Copyright 2003 Published by Elsevier Science Ltd on behalf of The Association for the Study of Animal Behaviour.      

Context-Dependent Plastic Response during Egg-Laying in a Widespread Newt Species

Previous research on predator-induced phenotypic plasticity mostly focused on responses in morphology, developmental time and/or behaviour during early life stages, but the potential significance of anticipatory parental responses has been investigated less often. In this study I examined behavioural and maternal responses of gravid female smooth newts, Lissotriton vulgaris, in the presence of chemical cues originating from invertebrate predators, Acilius sulcatus water beetles and Aeshna cyanea dragonfly larvae. More specifically, I tested the extent of oviposition preference, plasticity in egg-wrapping behaviour and plasticity in egg size when females had the possibility to lay eggs at oviposition sites with and without predator cues during overnight trials. I found that individuals did not avoid laying eggs in the environment with predator cues; however, individuals that deposited eggs into both environments adjusted the size of the laid eggs to the perceived environment. Females deposited larger eggs earlier in the season but egg size decreased with time in the absence of predator cues, whereas individuals laid eggs of average size throughout the investigated reproductive period when such cues were present. Also, egg size was found to be positively related to hatching success. Individuals did not adjust their wrapping behaviour to the presence of predator cues, but females differed in the extent of egg-wrapping between ponds. Females’ body mass and tail depth were also different between ponds, whereas their body size was positively associated with egg size. According to these results, female smooth newts have the potential to exhibit activational plasticity and invest differently into eggs depending on temporal and environmental factors. Such an anticipatory response may contribute to the success of this caudate species under a wide range of predator regimes at its natural breeding habitats.     

The reliance on distant resources for egg formation in high Arctic breeding barnacle geese Branta leucopsis

Breeding in the high Arctic is time constrained and animals should therefore start with their annual reproduction as early as possible. To allow for such early reproduction in migratory birds, females arrive at the breeding grounds either with body stores or they try to rapidly develop their eggs after arrival using local resources. Svalbard breeding barnacle geese Branta leucopsis have to fly non‐stop for about 1100 km from their last continental staging site to the archipelago making the transport of body stores costly. However, environmental conditions at the breeding grounds are highly unpredictable favouring residual body stores allowing for egg production after arrival on the breeding grounds. We estimated the reliance on southern continental resources, i.e. body stores for egg formation, in barnacle geese using stable isotope ratios in the geese's forage along the flyway and in their eggs. Females adopted mixed breeding strategies by using southern resources as well as local resources to varying extents for egg formation. Southern capital in lipid‐free yolk averaged 41% (range: 23–65%), early laid eggs containing more southern capital than eggs laid late in the season. Yolk lipids and albumen did not vary over time and averaged a southern capital proportion of 54% (range: 32–73%) and 47% (range: 25–88%), respectively. Our findings indicate that female geese vary the use of southern resources when synthesising their eggs and this allocation also varies among egg tissues. Their mixed and flexible use of distant and local resources potentially allows for adaptive adjustments to environmental conditions encountered at the archipelago just before breeding.