High levels of extra-group paternity in a population of Australian magpies Gymnorhina tibicen: evidence from microsatellite analysis

Breeding systems vary widely in birds, from monogamous pairs through to complex group systems where subordinates assist breeding individuals to rear young each season. The Australian magpie varies geographically both in plumage patterns and social organization. Some populations of both eastern and western plumage forms are plural breeders with group size varying from three to over 15 mature individuals. This study used variation at microsatellite loci to determine the level of extra-group paternity in a population of the western form near Perth in Western Australia. Extra-group paternity was the highest recorded for any bird species to date (82%) and indicates that few offspring within a territory are sired by the social partner of the female. In addition, the data indicated that nearly 10% of juveniles were not the genetic offspring of any female within their territory, suggesting some intraspecific brood parasitism. Taken together, these findings are remarkable considering the highly territorial nature of the species and the extent of territorial defence practised by all members of the group towards extra-group conspecifics during daylight hours.     

Territory Characteristics among Three Neighboring Chimpanzee Communities in the Taï National Park,Côte d'Ivoire

We studied territory characteristics among three neighboring chimpanzee communities in the Taï National Park, Côte d'Ivoire, and compared them with other chimpanzee populations. We characterized territories and ranging patterns by analyzing six variables:, (1) territory size, (2) overlap zone, (3) territory utilization, (4) core area, (5) territory shift, and (6) travel distance. Data collection covered a period of 10 mo, during which we simultaneously sampled the local positions of mostly large parties, including males in each community, in 30-min intervals. In Taï, chimpanzees used territories in a clumped way, with small central core areas being used preferentially over large peripheral areas. Although overlap zones between study communities mainly represented infrequently visited peripheral areas, overlap zones with all neighboring communities also included intensively used central areas. Territory utilization was not strongly seasonal, with no major shift of activity center or shift of areas used over consecutive months. However, we observed shorter daily travel distances in times of low food availability. Territory sizes of Taï chimpanzees tended to be larger than territories in other chimpanzee communities, presumably because high food availability allows for economical defense of territorial borders and time investment in territorial activities. Therefore we suggest, that use of territory in Taï chimpanzees is strongly influenced by intercommunity relations. To understand differences in territory characteristics between various populations, it is of major importance to consider not only the intracommunity but also the intercommunity context.     

Scent‐marking behavior by subordinate Eurasian beavers

Chemical communication by scent‐marking is widespread among mammals and can serve different purposes, such as territory demarcation, mate attraction and self‐advertisement. In this study, we examined scent‐marking behavior by subordinate Eurasian beavers (Castor fiber) in a population in southeast Norway. We conducted scent experiments in 18 family groups, simulating a territory intruder using experimental scent mounds (ESM). In total, we recorded 196 territorial reactions (destroying and/or overmarking of scent mounds). Subordinates contributed 14% of all territorial reactions and first overmarked 12% of the ESM, and the number of subordinates in the family group did not increase the total number of territorial reactions. This suggests that the contribution by subordinates plays a minor role in territorial defense. The number of territorial reactions by subordinates was positively related to the age of their parents, suggesting that subordinates might take over territorial duties with increasing parental senescence, likely because they have increased chances of inheriting the territory. Increased experience in territorial activities possibly helps subordinates to successfully gain and defend a territory of their own.     

Flock, pair and group living lifestyles without cooperative breeding by Australian Magpies Gymnorhina tibicen

The social behaviour of Australian Magpies Gymnorhina tibicen was studied at Linton in the Manawatu district, New Zealand, from June 1978 to November 1982. Territorial magpies defended all-purpose areas averaging 5 ha in size, and there was no correlation between territory size and the number of occupants. The mean adult survival rate was 0.85, and average annual productivity was 0.96 juveniles per breeding female. Nomadic flocks formed in the spring and foraged on open, treeless pasture. They mainly comprised 1–3-year-old magpies which did not breed. Territories were defended by monogamous pairs, and by groups ranging from three to six individuals. Some territorial groups probably originated in the flock, and may therefore have consisted of unrelated birds. Other territorial groups formed when juveniles did not disperse away from the natal territory, and were called family groups. Non-breeders did not help at the nest in any group, and there was no difference in the production of young by females breeding in group-defended territories compared with those in pair-defended territories. The opportunity to disperse into non-territorial flocks may have liberated Australian Magpies from cooperation.     

Social organization in the pheasant (Phasianus colchicus): harem formation, mate selection and the role of mate guarding

Harem formation and mate selection were studied in the pheasant in order to determine the advantages of territorial harem defence polygyny to the two sexes. We investigated the factors affecting harem size and the advantage to a female in remaining with one territorial male during breeding.
Female group size declined during late March and early April as females moved from large overlapping ranges into smaller, more widely dispersed breeding ranges. The proportion of female groups accompanied by males increased during this period.
Some males had a disproportionate share of females. Settled females were monogamous but, because a female's nest was generally outside the male's territory, her home range was larger than his territory.
Harem members were usually from the same winter group. Harem size was not related to territory quality in terms of food supply or nesting cover. Females were loyal to one male in more than one year even if his territory position changed. Older, territory-owning males had more females, both adult and immature, than males with newly-established territories. Harem size was not correlated with territory size.
We conclude that the mating system of the pheasant is based on mate guarding which protects females not only from the risk of predation or injury, but also from excessive energy expenditure incurred through being chased by other males. When escorted by a territorial male, females spent three times as much time feeding, one-fifth as much time running, and one-tenth as much time alert, as they did when not guarded.     

Is Territory Defence related to Plumage Ornaments in the King Penguin Aptenodytes patagonicus?

Colourful ornaments in monogamous birds may be directed at potential mates or other conspecifics to signal individual condition, reproductive status or fighting ability, especially in monogamous and territorial species. We investigated whether the size of the orange auricular patch may be an indicator of aggressiveness in the king penguin Aptenodytes patagonicus , a monogamous and territorial seabird. The relationship between auricular patch size and defence behaviour was explored relative to territory location (centre vs. periphery of the colony), period of reproduction (early vs. late), state of reproduction (incubation vs. brooding) and sex. The proportion of time spent in territorial defence and the rate of aggressive behaviours were positively correlated with auricular patch size, mainly because central birds were more aggressive than peripheral birds and also had larger patch sizes. The period of reproduction, state of reproduction and sex did not interact with patch size to affect aggressiveness. Our results suggest that the size of the auricular patch in king penguins may be a reliable signal allowing individuals to evaluate the quality of mates or competitors in terms of aggressiveness. Whether aggressiveness is directly linked to patch size or indirectly through body condition, however, remains to be determined. In any event, birds with larger patches seem to gain central territories in the colony, thereby increasing their reproductive success. Finally, our study adds to the growing evidence that the evolution of sexually monomorphic ornaments may stem from mutual sexual selection.     

Territoriality and male reproductive success in arctic ground squirrels

Although territorial defense is a common form of reproductivecompetition among male vertebrates, the exact reproductiveconsequences of this behavior are often poorly understood.To explore relationships between territoriality and reproductivesuccess in a nongroup-living mammal, we quantified patterns of space use, mating success, and fertilization success formales in a free-living population of arctic ground squirrels(Spermophilus parryii plesius). Because litters of this speciesare sired almost exclusively by a female's first mate, we predictedthat territory ownership would be associated with first accessto estrous females. During the 2-week period when mating occurred,each male in the study population attempted to defend a distinctportion of the habitat, although the success of this defensevaried among individuals. Twenty-six of 28 females monitoredmated with the male on whose territory they resided. However,the majority of females observed throughout estrus (65%; n= 20) also mated with at least one other male, indicating thatterritory ownership was not associated with exclusive accessto females. In contrast, territory ownership was significantly associated with first access to estrous females; 20 (71.4%)of 28 females mated first with the male on whose territorythey resided. In this regard, the behavior of S. parryii plesiusparallels that of socially monogamous birds in which territorialdefense by males functions to deter extrapair copulations byfemales. Although territorial defense represents an important component of male reproductive success in arctic ground squirrels,other aspects of male behavior (e.g., the ability to dominateagonistic interactions on the day of a female's estrus) arealso critical. We suggest that future studies of vertebratemating systems will benefit by viewing such defense as onlyone of multiple axes along which conspecific males compete foraccess to females.     

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.     

Hormonal responses to male-male social challenge in the blue tit Cyanistes caeruleus: single-broodedness as an explanatory variable

The "challenge hypothesis" posits that when established social orders are challenged, plasma testosterone (T) in socially monogamous breeding male birds will temporarily increase to facilitate aggressive responses. However, not all birds conform to predictions. To expand upon past findings, we examined effects of direct territorial challenge on T levels in the blue tit (Cyanistes caeruleus). We found that simulated territorial intrusions caused a decline in plasma T during both territory establishment and laying/incubation. Conversely, corticosterone (CORT) levels dramatically increased. We also examined challenged blue tit males for levels of corticosterone-binding globulin (CBG), a carrier molecule that displays affinity for both CORT and T in birds. Although the CBG showed increased occupation by CORT during challenge, effects were not accompanied by a significant increase in the unbound T fraction. Thus, competitive hormone interactions on the CBG do not seem sufficient to explain changes in circulating T levels. To place our results within the context of past findings, we compared all socially monogamous birds tested to date for plasma levels of T during situational territorial intrusion experiments. We found that birds raising only one brood per season (e.g., the blue tit) consistently show no increase in plasma T but instead show elevations in circulating CORT. Thus, we suggest that single-broodedness plays an important role in determining patterns of hormone change and should be considered in future discussions of hormone-behavior interactions.     

Mating systems and territory in Lapwings Vanellus vanellus

In a population of Lapwings Vanellus vanellus studied for 5 years near Bergen, southwester Norway, 23–41% of the males were polygynous, having two and sometimes three mates, whereas 50–77% were monogamous and 0–21% remained unmated. Bachelors held territories in or immediately adjacent to the study area each of the years. As predicted from the Polygyny Threshold model, primary females generally laid eggs earlier than monogamous females, but the overlap in laying dates was substantial between these two classes. The number of females and start of breeding on the territories were analysed in relation to nine variables pertaining to safety from predation and to food. Of these, territory size correlated most consistently with number of females (positively) and the laying of first egg (negatively). Numbers of females observed were not significantly different from numbers expected from territory size in 4 out of 5 years; however, earlier start of breeding on large territories indicated that females did not settle according to the Neutral Mate Choice model. The Female-biased Sex Ratio hypothesis was refuted, unmated territorial males being available throughout the breeding seasons.     

Pair formation in the Mongolian gerbil

This report describes the initial phase of social organization in Mongolian gerbils, Meriones unguiculatus, in large outdoor enclosures. These conditions offer a comparatively free choice for the establishment of bonds and social structure. In five replicate 20-day studies, the only social unit that formed was the male-female pair. The order of events was related to four sequential 5-day periods: (I) males established territories by male-male competition (1st to 2nd period); (II) females visited more than one male initially (1st and 2nd period), but developed a preference for one male or territory (3rd to 4th period); (III) as the females developed a preference, they also challenged other females having a preference for the same male or territory (approximately the 3rd period); (IV) as one female became locally dominant, some territorial males started chasing all but the dominant female. Males remained more active in territorial defence throughout the study. The defence became more intense as the females became strongly attached to one male and/or territory only. However females, after having established themselves in one particular male territory, spent significantly less time in the preferred area during the day of oestrus compared with the days before and after. Copulatory behaviour patterns, where observed, did not occur exclusively in interactions between territorial mates.     

Shadow competition in wild juvenile sea-trout

Shadow competition occurs in a group of sit-and-wait predators when those closer to a source of mobile prey reduce the feeding success of those further from the prey source. It was examined in territorial juvenile sea trout Salmo trutta in a small stream. The fry formed groups of two to six fish with adjacent territories and a social hierarchy within each group. It was hypothesized that: (i) as group size increased, the mean number of prey eaten per fish within a group decreased and the variability in prey consumption between fish increased; (ii) prey consumption by individual fish decreased with increasing distance from the food source; (iii) group size increased as the mean water velocity immediately upstream from a group, and hence potential drifting food, increased. Five groups of fry were fed on small shrimps released upstream from each group at a rate of one every 15 s over a 10 min period, this procedure being repeated over 5 days to provide five replicates per group. Experiments were performed three times in 1967, 1969 and 1974 to provide information on 45 groups of fry. The first and third hypotheses were supported, but the second was only partially supported. In 1967 and 1969, territory size and shrimp consumption by individual fry decreased with increasing distance from the food source. This also occurred in 1974, except during a critical period for survival when fry density was exceptionally high with large numbers of sea trout lacking territories. This resulted in sea trout fry with the largest territories eating fewer shrimps than those with medium-sized territories because they spent more time defending their territories against sea trout lacking territories. This study is the first to demonstrate shadow competition in a vertebrate species, but has also shown that territorial defence may modify the consequences of shadow competition when densities are high and there is strong competition for the acquisition of a territory.     

Reproductive success and survivorship of the Galapagos Hawk Buteo galapagoensis: potential costs and benefits of cooperative polyandry

The Galapagos Hawk Buteo galapagoensis is one of few species showing cooperative polyandry. As many as four males may mate with one female and share a territory and nest, Such groups, on average, produce more young than monogamous pairs, but on a per male basis (assuming that each male of a group has an equal chance to fertilize each young) polyandrous males produce fewer young per year than monogamous males. Territorial breeding birds show higher yearly survivorship (90%) than non-territorial, non-breeding birds (50% or less). All territorial birds seem to remain on their territories for life, and monogamous pairs actually constitute the remnants of polyandrous groups. Some potential factors producing such an unusual system are discussed. Limited breeding space and high mortality among non-territorial birds seem to work in favour of group breeding in this and other cooperative species. Polyandry may be favoured because of the reduced value of extra females in hawk breeding behaviour. Equal-status males may occur because of the lack of kinship among cooperating males and the high potential genetic cost of being a helper in a long-lived, low fecundity species. Other possible factors favouring group breeding are territory acquisition and the variable nature of the Galapagos climate.     

Polygyny and the breeding success of the great reed warblerAcrocephalus arundinaceus

Settlement date, mating status, and breeding success of individually marked great reed warblers, Acrocephalus arundinaceus, were studied during the 1980–84 breeding seasons in Kahokugata, Ishikawa Prefecture, Japan. Twenty-five per cent of the territorial males were polygynous, of which the majority were bigamous. The settling periods of both sexes were long, extending for 65 days in males and 49 days in females. About 80% of males and females settled in the first half of the settling period, and the settlement date of 28–54% males overlapped with that of females. Many of the late settlers were bachelors and the males which mated earlier tended to be polygynous. The timing of a male's settlement is important in acquiring mates. Fifty-five per cent of eggs laid were lost before fledging, mainly due to predation. The mean number of fledglings was 3.19 per primary female, 2.41 per secondary female, and 2.80 per monogamous female. Comparison of the number of fledglings of females which mated during the same period showed that the presence of another female in the same territory did not adversely affect the breeding success of either of the polygynous females. Polygynous males have the advantage of decreasing the risk of breeding failure under high predation pressure.     

Seasonal Changes of Vocal Rates and Their Relation to Territorial Status in Male Galápagos Sea Lions (Zalophus wollebaeki)

Male vocal displays play an important role in sexual selection through both male–male competition and female choice. While this is supported by numerous studies in birds, much less attention has been paid to the role of such displays in mammals. We investigated the function of vocal displays in a polygynous mammal, the Galápagos sea lion (Zalophus wollebaeki). In our study population a large proportion of mature males are unable to establish a territory, providing the opportunity to compare the vocal behaviour between territorial and non‐territorial males. We examined how seasonal patterns of vocalizations differed between territorial and non‐territorial males and how the number of females present in a territory influenced behaviour of territorial males. We found that territorial males vocalized at higher rates than non‐territorial males, and territorial males vocalized more at the onset than towards the end of territory tenure. During the onset of territory‐tenure vocalizations of territorial males were directed more often towards other males than females. Furthermore, we found that vocalizations of territorial males were not only given in male–male interactions, but were also actively directed towards females. Territorial males vocalized at higher rates when more females were present in their territory. Our results suggest that vocalizations are important in male–male interactions, are relevant in territorial disputes and are used as a proxy for females to assess a male’s quality.     

The Social System of a Communal Breeder,the Bay-winged Cowbird Molothrus badius

Molothrus badius (bay-winged cowbird), an icterine blackbird with cooperative breeding, shares behavioural and ecological characteristics with other communal nesters: it is sedentary, has a high annual survival rate (76.2%) and a strong nest-site tenacity (mean breeding dispersal of 41.9 and 89.4 m for males and females). Behavioural data, including collective agonistic displays, suggest group territoriality. Before egg hatching most breeders occurred as single pairs showing territorial behaviour (82% of nests), and nesting was usually solitary (distances to nearest nests of 25–103 m). Most breeders were apparently monogamous, with a 2.5% incidence of extrapair copulations in the territory during clutch formation. During the nestling stage one to four helpers occurred at 95% of M. badius nests. Most helpers were 1–2 years old, but older breeding adults (mostly males) that failed to rear their own offspring helped at the end of the season. The number of helpers increased (up to 4) with nestling age. Helpers were also recruited during the postfledging period, and group size reached up to 10 adults at this stage. Helpers mobbed predators and brood parasites, and provided 35% of the nestling food. Provisioning rate was positively and significantly correlated with number of helpers, although age of nestlings was the best predictor of overall food delivery rate. The helping system was almost obligate and productivity comparisons between nests with/without helpers are not possible. Data suggest that helpers increased the breeding success per nest. The correlation between the provisioning rates of parents and helpers was negative but non-significant. In 18% of nests 3 to 4 individuals were present before the nestling period, including cases of apparently polyandrous trios and one case of joint nesting by two pairs. Within Brown 's (1987) categories of social organization M. badius is mainly group territorial with plural nesting. Habitat requirements of M. badius are wide and nest sites do not appear to limit breeding. Kinship plays a role in the social system, as 9 of 12 helpers marked as nestlings helped their parents.     

European robins (Erithacus rubecula) lack an increase in testosterone during simulated territorial intrusions

The challenge hypothesis (Wingfield et al. in Am Nat 136:829–846, 1990) predicts that circulating testosterone increases when socially monogamous male birds are challenged during breeding. Although the challenge hypothesis has been confirmed in large-scale interspecific comparisons of seasonal hormone profiles, experimental tests of the challenge hypothesis are still uncommon and the results equivocal. We tested one of the predictions of the challenge hypothesis by investigating the behavioural and hormonal response of free-living European robins during simulated territorial intrusions (STIs) in the breeding season. We conducted STIs by placing a stuffed decoy in a territory and playing robin song. After the behaviour of the focal male had been recorded for at least 10 min, it was captured and a blood sample was taken immediately. Controls were caught within 10 min of the first response of the territory owner. Hormone concentrations were measured by radio-immunoassay. Although previous studies have shown that testosterone has an impact on aggression, European robins do not respond to STIs by increasing circulating levels of testosterone.     

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.     

Couch potatoes do better: Delayed dispersal and territory size affect the duration of territory occupancy in a monogamous mammal

In territorial, socially monogamous species, the establishment and defense of a territory are an important strategy to maximize individual fitness, but the factors responsible for the duration of territory occupancy are rarely studied, especially in long‐lived mammals. A long‐term monitoring program in southeast Norway spanning over 18 years allowed us to follow the individual life histories of Eurasian beavers (Castor fiber) from adolescence in their natal family group to dispersal and territory establishment until the end of territory occupancy. We investigated whether territory size, resource availability, population density, and dispersal age could explain the duration of territory occupancy, which ranged from 1 to 11 years. The duration of territory occupancy was positively related to dispersal age, suggesting that individuals that delayed dispersal had a competitive advantage due to a larger body mass. This is in support with the maturation hypothesis, which states that an animal should await its physical and behavioral maturation before the acquisition of a territory. Further, we found that individuals that established in medium‐sized territories occupied them longer as compared to individuals in small or large territories. This suggests that large territories are more costly to defend due to an increased patrolling effort, and small territories might not have sufficient resources. The lifetime reproductive success ranged from zero to six kits and generally increased with an increasing duration of territory occupancy. Our findings show the importance of holding a territory and demonstrate that dispersal decisions and territory selection have important consequences for the fitness of an individual.     

Behaviour and breeding biology of the cooperatively breeding Grey-backed Fiscal Shrike Lanius excubitorius in Kenya

Grey-backed Fiscal Shrikes Lanius excubitorius were studied over a 2j year period near Lake Naivasha, Kenya. Grey-backs are cooperative breeders, with group sizes ranging from two to II. Only one pair breeds per group, with all other group members aiding in the rearing of young. The study population ranged from 64 to 79 individuals that occurred in from 13 to 16 groups. Non-breeding helpers made up to 66% of the population, with male helpers being more numerous overall than females. The annual survival rate was 65%, with no differences detected between the survival of males and females, or of breeders and helpers. Only male helpers were observed to acquire breeding status within the natal territory. Some female helpers acquired breeding positions in territories adjacent to their natal territories. Group territorial displays occurred throughout the year but were most pronounced prior to breeding during rainy periods. Reproductive success was very low, with only 14.5% of the recorded breeding attempts leading to fledged young. Large groups (four or more birds) had greater reproductive success than small groups, but many factors other than, or in addition to, group size may have influenced this pattern. The breeding male contributed the most food to the incubating female and to the nestlings. Male helpers and the breeding female contributed more to nestlings than did female helpers. Observations on the post-fledging period indicate that socialization and establishment of dominance may be of importance in group-living species living in a restricted ecological and social setting.