Territory structure of the Robin Erithacus rubecula outside the breeding season

On the basis of radio-tracking data from 51 individuals, the space use of Robins Erithacus rubecula outside the breeding season was non-uniform and multinuclear. On average, territorial Robins spent 78% of their time in one or more resource patches represented by bushes, which accounted for only 1% of a convex polygon encompassing each range. A suite of range polygons was developed to quantify range structure. One of these polygons showed similar size to, and high overlap with, defended-area polygons and so was a good estimate of territory. In addition, the area of bushes in each polygon suggested Robins occupied constant resource territories. There was undefended neutral ground between territories, which, although seldom visited, was crossed to achieve intrusions. Over the short term (days), Robins showed high fidelity to the resource patches they defended. Over the longer term (months), most individuals showed some degree of range shift. It is concluded that if the information concerning territories is to be maximized, empirically derived sampling protocols should be used when measuring territory parameters.     

Breeding of the Seychelles Magpie Robin Copsychus sechellarum and implications for its conservation

The total population of the Seychelles Magpie Robin Copsychus sechellarum declined from 38–41 birds in 12–13 territories in 1977–1978 to 17–21 birds in eight to nine territories in 1988–1990 and was entirely confined to Fregate Island (210 ha) in the Seychelles. After a successful cat eradication program in 1981–1982, recruitment improved, although the abandonment of agriculture had caused a reduction in the amount of feeding habitat and hence in the carrying capacity of the island. The population declined because of the failure of recruitment to compensate for the annual adult mortality. Foraging activity of the Magpie Robin was greater in high-quality territories (measured by soil invertebrates available), leading to increased reproductive success. Through supplementary feeding, five times as many recruits were produced. Of the 11.5 potential annual breeding recruits, 5.3 are required to compensate for adult mortality, and the other 6.2 recruits can be regarded as “surplus” contributing to an increase. Magpie Robins prefer to breed in rotten trees, which are a scarce resource. The greater the distance between the nest site and feeding areas, the less time was spent in incubation and nest guarding, resulting in greater egg loss. Because of lack of suitable areas for establishing territories, many young Magpie Robins became “floaters”. Nest disturbance, both by these floaters and by the introduced Indian Mynah Acridotheres tristis, had adverse effects on the breeding success of robins. A recovery plan has been designed to save the Magpie Robin. Territories have been improved for feeding (by tree planting) and for breeding (by providing nestboxes and reducing nest disturbances). Given the vulnerability of one small island, the presence of surplus birds (supported mainly by supplementary food) and the suitability of neighbouring Aride Island (68 ha), successful translocations to this island took place in 1992 and 1994. Given the presence now of 47 individuals on two islands, it is hoped that the species will pull back from the brink of extinction.     

Age‐related sexual plumage dimorphism and badge framing in the European Robin Erithacus rubecula

Male and female European Robins Erithacus rubecula display their red breasts in year‐round territorial contests. Despite the clear signalling role of the red breast, little is known about its sexual dimorphism or trends in size when Robins age. We studied these patterns in resident and migrant Robins in a Mediterranean population. Both male and female first‐year Robins had smaller red breasts than second‐year individuals. Females, but not males, showed a reduction in badge area after the second year of life, leading to sexual dimorphism in red breast area in the oldest Robins. The grey fringe around the red breast showed a steady increase in width in males when ageing, but not in females, also leading to sexual dimorphism in this trait among oldest Robins. The contrast between the red breast and its surrounding grey fringe was higher than that between the breast and the back at both high and low environmental light intensities. This suggests that the grey fringe could function as a frame to highlight the perimeter of the red breast, and that the Robin’s plumage is equally suited to display in open areas and forest understoreys. Our study suggests that the extent of the red breast and its grey frame could confer information about the age and sex of the birds, and thus be used as a signal in territorial contests and mating decisions.     

Breeding ecology of the Clay-coloured Robin Turdus grayi in lowland Panama

The breeding ecology of Clay-coloured Robins Turdus grayi was studied in 1979 in the former Canal Zone of lowland Panama. Two study areas were chosen close together, Summit Gardens (27-3 ha), an ecological island of habitat suitable for Clay-coloured Robin surrounded by forest, and Morgan's Gardens (3–6 ha), a similar island surrounded by deforested areas and settlements. Eighty-three active nests were found and checked. The breeding season lasted from 2 March to 21 June. In Morgan's Gardens the first broods were raised in the dry season and second broods in the rainy season; in Summit Gardens most broods started at the beginning of the rainy season. In comparison to the closely related Blackbird T. merula in Poland, Clay-coloured Robins build nests on isolated trees or bushes, usually in more conspicuous sites (interpreted as an anti-predator adaptation) and on flexible, horizontal branches off the main trunk. Over 45% of broods were destroyed by predators but other factors causing brood loss were negligible. Nests built on palms were considerably safer. In the dry season the predation rate was low (5%) and increased during the course of the rainy season. Unlike Blackbirds, Clay-coloured Robin nestlings were given a lot of fruit; their diet was diversified. Starvation occurred in 60% of nests although usually only the youngest nestling died. Breeding density in Morgan's Gardens at 50 pairs per 10 ha was much higher than in Summit Gardens at 15>8 pairs per 10 ha. Breeding losses were lower, the nestlings were fed more fruit, the average nestling weight was lower and the production of fledglings per breeding pair was twice as high in Morgan's Gardens. It was concluded that a strategy of settling in a rather overpopulated place with hard foraging conditions but lower predation was better than settling in a area with a better food supply but higher predation.     

Survival of the Orange-tufted Sunbird,Nectarinia osea

Cape Sugarbirds are southern African endemics, found in fynbos vegetation of the Western and Eastern Cape Provinces of South Africa, and they feed almost exclusively on the nectar of Protea inflorescences. Breeding male Sugarbirds are highly territorial and defend resources for themselves, their mates and their offspring. Sugarbirds, however, often leave their breeding territories in search of food during the dry season and return to the same breeding sites each year, thus experiencing an annual cycle of movement from one food source to another. Male territory size as well as breeding success, were determined over two breeding seasons for a population in the Helderberg Nature Reserve in the Western Cape Province. Sugarbird fledgling success was significantly greater for males defending large territories. Males moved or increased the size of their territories between seasons, possibly to improve breeding success, but no changes in territory size were observed during the breeding season.     

Territorial behaviour of the Robin Erithacus rubecula: the importance of vegetation density

Robin Erithacus rubecula territories were mapped in an isolated copse of mixed deciduous woodland. An index of vegetation density, measured at numerous randomly chosen sites within the copse, was found to be higher in areas occupied by Robins. Both vegetation density and territory area were compared with measures of breeding success. In all cases vegetation density was correlated with breeding success and territory area was not. Possible evolutionary advantages to defending areas with dense vegetation are discussed.     

Seasonal change in Red-backed Shrike Lanius collurio territory quality - the role of nest predation

This paper examines the relationships between territory preference, reproductive performance and nest predation in a Red-backed Shrike Lanius collurio population breeding in rapidly changing farmland. Territory preference was measured by (a) the order of laying dates in territories within years and (b) territory occupation frequency over a five-year period. Seven environmental variables were tested but only the density of Sloe (Blackthorn) Prunus spinosa in territories was correlated positively with territory occupation frequency. Within years, Red-backed Shrikes started to breed earlier in territories with more Sloe, but reproductive performance was not related consistently to territory differences. A high proportion of Sloe-rich territories was vacant each year. Thus, the observed negative association between settlement order and amount of Sloe may reflect a seasonal change in habitat preference. Early breeders prefer Sloe-rich territories whereas late birds prefer Sloe-poor territories. Supporting evidence is that the proportion of occupied territories that was Sloe-rich was higher for early than for average or late first breeding attempts. Moreover, in contrast to early in the season, the probability of a replacement clutch following nest failure was not associated with the amount of Sloe, but increased with increasing densities of Juniper Juniperus communis. Late in the nesting season, pairs breeding in Sloe-rich territories suffered higher nest predation than those breeding in Sloe-poor territories. In early nesting attempts, however, the relationship was reversed. Nest predation is probably an important selective agent on habitat selection since it was the principal factor explaining variation in the numbers of fledglings produced among territories. Thus, the Red-backed Shrike may shift territorial preferences adaptively as the season progresses to match within-season changes in predation pressure in different types of territories.     

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.     

Possible relationships between morphology,territory quality,and skin color of American Kestrels1

ABSTRACT Carotenoid‐based coloration of skin and plumage has been found to be correlated with individual quality in many species of birds during the breeding season. However, less is known about the possible role of these signals during the nonbreeding season, particularly among nonpasserines that defend winter territories. American Kestrels (Falco sparverius) are sexually dimorphic raptors that defend winter territories and possess carotenoid‐based morphological features known to be correlated with individual quality. Much is known about winter territory use and habitat segregation by male and female kestrels, but possible relationships among morphological features, individual quality, and habitat quality have not been examined. Our objective was to examine possible relationships between morphology, territory quality, and skin color of American Kestrels. Male kestrels had brighter skin than females, and the skin color of male kestrels was positively correlated with size (wing chord and tail length) and territorial quality (hunting territories with less canopy and more grass cover). No such relationships were found for female kestrels. Skin color appears to be an honest indicator of quality for male American Kestrels and may serve both intersexual (territory acquisition) and intersexual (mate choice) functions during the breeding and nonbreeding seasons.     

Norfolk Island Robins are a distinct endangered species: ancient DNA unlocks surprising relationships and phenotypic discordance within the Australo-Pacific Robins

Uncertain taxonomy hinders the effective prioritization of taxa for conservation. This problem is acute for understudied island populations in the southwest Pacific Ocean, which are increasingly threatened by habitat loss, predation and climate change. Here, we offer the first test of taxonomic limits and phylogenetic affinities of the iconic Pacific Robin radiation (Petroica multicolor) in order to prioritize the conservation of its nominotypical subspecies, the endangered Norfolk Island Robin (P. m. multicolor). We integrate phylogenetic analyses of ancient DNA and quantitative measures of plumage and morphometric variation to show that the Norfolk Island Robin should be recognized as a distinct species. Phenotypic and genetic datasets contradict the longstanding treatment of Pacific Robins (including Norfolk Island Robins) and Scarlet Robins (P. boodang) as a single species. Instead, we show that Norfolk Island Robins are deeply divergent from Scarlet Robins and have more genetic similarity to Red-capped Robins (P. goodenovii) than to other Pacific Robins. This finding is unrepresentative of the current taxonomic and conservation status of the Norfolk Island Robin, which we propose should be recognised as an endemic endangered species. Our study clearly shows that in the absence of contemporary tissues, ancient DNA approaches using historical museum specimens can address taxonomic questions that morphological traits are unable to resolve. Further, it highlights the need for similar studies of other threatened Norfolk fauna with uncertain taxonomic status in order to ensure appropriate conservation prioritization.     

Territorial song and song neighbourhoods in the Scarlet Rosefinch Carpodacus erythrinus

Throughout the range of the Scarlet Rosefinch, its territorial song consists of 3–9 (usually 4–5) elements, of which there are 5 different types. The differences lie in the way the pitch of the element changes in time (frequency "slope") and the width of the frequency band. Within a given type of song, the various elements can be present in almost any combination. Therefore, so many song types can be formed that the songs in even small parts of the species' area are clearly distinct from one another. Despite this capacity for variation, however, by chance identical songs may be sung in widely separated parts of the area, in some cases by different subspecies.
The species has not developed large-scale dialects or regiolects based on a song tradition acquired during an early imprinting phase. Scarlet Rosefinches tend to breed in small colonies, groups of up to about 15 pairs characterized by the same type of song (song neighbourhoods, formed by the development of a microlect).
Microlects develop by a founder effect. When males, near one-year old or older, join one another to form isolated colonies after arrival in the breeding region, they adopt ("learn") the song type that will eventually characterize the colony from the first male to arrive at the site. After the colony has been founded, in most cases each male uses only one type of song during a breeding season, with practically no variation of the temporal and frequency parameters.
Singing the same type of song, the members of a colony accept one another sufficiently to allow the breeding territories to be closely packed. It appears that a long-lasting capacity for acoustic learning, in combination with colony-like breeding and great ecological flexibility, has allowed the Scarlet Rosefinch to become the most successful species of the genus Carpodacus .     

Pair territoriality in the longnose filefish,oxymonacanthus longirostris

The longnose filefish,Oxymonacanthus longirostris, usually lives in heterosexual pairs, the male and female swimming together and sharing the same territory. Pair territoriality in the species was examined in detail in relation to sexual differences in territorial defense activities. Rigorous pair territoriality was maintained only during the breeding season, although pairs used their home ranges exclusively to a certain extent, during the non-breeding season. The frequency of aggression against other conspecific pairs in the breeding season was higher than in the non-breeding season. Agonistic interactions appear to be over both mates and food resources, the strict pair territoriality in the breeding season possibly being due to mutual mate guarding. In intraspecific aggressive interactions, males usually led their partner females when attacking intruders. The feeding frequency of males was much lower than that of females in the breeding season. Mate removal experiments indicated that females could not defend their original territories solitarily and their feeding frequency decreased. Conversely, males could defend territories solitarily without a decrease in feeding frequency. These results suggest that males contribute most to the defense of the pair territory, with females benefiting from territorial pair-swimming with their partner males.     

Family-based territoriality vs flocking in wintering common cranes Grus grus

We describe winter territoriality in common cranes, Grus grus , a long-lived migrant species with long-lasting pair bonds and parental care extending throughout the winter. Cranes are territorial in the breeding season, and usually gregarious during migration and wintering. Only 2% of the families present in our study area were territorial, all other families foraged in flocks with immatures and adult pairs. Territorial pairs defended the same winter territory year after year, but only when they had offspring. They were gregarious otherwise. The average breeding success measured throughout several years was higher in territorial pairs. Winter territories were small (0.7 km² on average), but included a higher diversity of habitats than the areas visited by gregarious birds (11.7 km² on average). Adults of territorial families showed longer vigilance times, and lower food intake rates than did adults in flocks, which were compensated with a longer time spent foraging per day. The accumulated daily food intake did not differ between adults in flocks and in families. We suggest that winter territoriality is a facultative strategy, conditioned by parental experience and habitat availability.     

EXTRAPAIR MATE CHOICE AND HONEST SIGNALING IN COOPERATIVELY BREEDING SUPERB FAIRY-WRENS

In many species of monogamous birds females copulate with males other than their social mates, resulting in extrapair fertilizations. Little is known about how females choose extrapair mates and whether the traits used to choose them are reliable indicators of male quality. Here we identify a novel male trait associated with extra-group mating success in the superb fairy-wren (Malurus cyaneus), a cooperatively breeding bird with one of the highest known frequencies of extra-group mating. Female fairy-wrens chose extra-group mates that molted earlier into breeding plumage. Males molted up to five months before the breeding season began, and only males that molted at least one month prior to its onset gained any extra-group fertilizations. This conclusion held after controlling statistically for the effect of age and social status on molt date. Once males acquired breeding plumage, they began courtship display to females on other territories. Thus, some males were displaying to females for several months before the breeding season began. This extraordinarily long period of advertisement by males may be facilitated by the long-term ownership of territories. We suggest that early acquisition of breeding plumage or the subsequent display behavior can be reliable cues for mate choice because they are costly to acquire or maintain.     

Mate choice in Darwin's Finches

Female Geospiza conirostris on Isla Genovesa, Galapagos, pair preferentially with males who have had previous breeding experience. They choose mates on the basis of courtship behaviour and black adult plumage. By mating with experienced black males, they gain a fitness advantage in terms of fledgling production and recruitment of young into the breeding population. Behavioural signs of past breeding experience and black plumage are reliable age- and condition-dependent traits. We suggest that females use conspicuous black plumage to identify old males at a distance, then interactions through courtship to modify initial assessments. Females paired with inferior males may increase the genetic quality of their offspring by extra-pair copulations; results of heritability analysis of morphology are consistent with this suggestion. Females change mates at a frequency of 12–27% per breeding season. They re-pair with males who are generally old, experienced, and hold territories adjacent to the deserted male. Females that re-pair gain a benefit, whereas males who are deserted within a breeding season incur a cost of more than 50% of their future potential production for that season. We conclude that females in choosing males seek reliable indicators of potential parental care, and in addition they may seek indicators of genetic quality.     

Costs of reproduction in male lizards, Sceloporus virgatus

Models of life history evolution typically assume a balance between the benefits of current reproductive activity and the costs to future reproductive success or survivorship, but empirical studies often find positive correlations between such components of fitness in undisturbed animal populations. I examined possible survivorship costs of reproduction in free-ranging male lizards, Sceloporus virgatus , and found that males with low levels of mating success were less likely to survive to the following breeding season. I also investigated two possible indicators of reproductive effort, increase in ectoparasite load and decrease in body weight during the breeding season. Levels of parasitism with trombiculid mites at the end of the breeding season were not associated with any measure of fitness or body condition (mating success, survivorship to the following year, relative weight loss). Yearling males (which have low levels of mating success) usually gained weight during the breeding season, while older males generally lost weight during this period. This suggests that young males may have postponed reproduction in favor of body growth and that seasonal weight loss of older males might reflect reproductive effort. Within the group of older males, individuals with the highest levels of mating success did not have high levels of either weight loss or mortality. Mate guarding behavior, an alternative to the aggressive territorial behavior typical of many lizard species, may allow certain males to obtain mates without expending large amounts of energy or exposing themselves to great mortality risks.     

Effects of breeding success, mate fidelity and senescence on breeding dispersal of male and female blue-footed boobies

1. Understanding the effects of individual and population factors on variation in breeding dispersal (the movement of individuals between successive breeding sites) is key to identifying the strategies behind breeders' movements. Dispersal is often influenced by multiple factors and these can be confounded with each other. We used 13 years of data on the locations, mates, breeding success and ages of individuals to tease apart the factors influencing breeding dispersal in a colonially breeding long-lived seabird, the blue-footed booby Sula nebouxii. 2. Breeding dispersal varied among and within years. Males dispersed further in years of higher population density, and late breeding males and females dispersed further than early breeders. This temporal variation related to changes in competition for territory was taken into account in all tests of individual factors influencing breeding dispersal. 3. Individuals that retained their mates from the previous year dispersed shorter distances than those that changed their mates. 4. The effect of previous breeding success depended on mate fidelity. Unsuccessful breeding induced greater dispersal in birds that changed their mates but not in birds that retained their mates, indicating that breeders who change mates may take their own previous breeding experience into account during habitat selection. Faithful individuals may have to stay close to their previous sites to encounter their mates. 5. Male divorcees dispersed over shorter distances than their former mates, possibly because males contribute more than females to establishing territories. 6. Dispersal of males and females declined with increasing age over the first 10-11 years of life, then increased in old age, possibly due to senescent decay in the ability to compete for mates and territories.     

5. GENERAL NOTES

Olive Thrushes Turdus olivaceus olivaceus in Grahamstown, South Africa, were present in their territories throughout the year. Mist-net captures showed no seasonal fluctuation in the population density of adults. The breeding season had a main peak from August to November and a minor peak in April. Song output was greatest at the beginning of the breeding periods, when territorial boundaries were most keenly contested. There was a positive correlation between seasonal variation in breeding intensity and song output. The adult survival rate was estimated at 80%, and 56% of the fledglings were alive in their natal territories at the age of independence. In this study area, only females built nests and brooded nestlings. Only males sang. Both males and females defended their territories.     

The role of androgen receptors in regulating territorial aggression in male song sparrows

This paper examines the role that androgen receptors (ARs) play in modulating aggressive behavior in male song sparrows, Melospiza melodia morphna. Song sparrows are seasonally breeding, territorial birds that maintain year-round territories with male-female pair bonds formed during the spring breeding season. Plasma testosterone levels peak as territories are established and mates acquired. In late summer, testosterone levels fall and remain basal during the non-breeding season. We examined the role of ARs in regulating territorial aggression in captive song sparrows under short- and long-day conditions as well as just prior to, and at the start of the breading season in freely living birds using the nonsteroidal antiandrogen flutamide to block AR function. Birds were implanted with either empty or drug filled silastic implants for 18 to 42 days and then challenged with a novel male decoy to assess the individual birds level of male-male aggression. Freely living birds remained on their home territory and underwent a simulated territorial intrusion, whereas laboratory-held birds were assessed using a laboratory simulated territorial intrusion and remained in their home cage. Experimental treatment of male song sparrows decreased aggressive behavior during the pre-breeding life history substage (March-April) in freely living birds as well as in laboratory-held birds under long-day (16L:8D) conditions. During the early breeding substage (April-May) there was no measurable effect of flutamide treatment on aggressive behavior, nor was there a difference in behavior in the (8L:16D) laboratory birds. This demonstrates that ARs are an important component of the neuroendocrine control of aggressive behavior. Given that flutamide only affected aggression during the pre-breeding substage and in LD birds, the results suggest that AR dependent control of aggressive behavior changes as song sparrow life history states change.     

The relative costs and benefits of territorial defense and the two conditional male mating tactics in the Comanche Springs pupfish (Cyprinodon elegans)

We examined the aggressive costs and reproductive benefits of territorial defense and its alternatives in a population of the Comanche Springs pupfish (Cyprinodon elegans). The breeding system was characterized by three different male mating tactics: territorial defense, satellite positioning, and sneak spawning. The mating tactic adopted by males reflected the males' sizes. Territorial residents were the largest, satellites were medium-sized, and sneakers were the smallest adult males observed. Consistent with the hypothesis that primary mating tactics are relatively high-cost, high-benefit strategies, we found that territorial males engaged in a number of aggressive encounters but had higher reproductive successes than any other males. However, our observations did not support the premise that conditionally breeding males engage in relatively low-benefit, low-cost tactics. Medium satellites and small sneakers acquired fewer spawns than did territorial males, but both satellites and sneakers were involved in as much aggression as territorials. That is, the data supported the prediction that satellite or sneaker males with the inability to compete for territories would attempt to accrue some reproductive opportunities in the presence of territorial males despite the high costs of spending time on the breeding grounds. Adopting conditional tactics appeared to allow satellites and sneakers to make the best of a bad situation. During a subsequent breeding season, large males were absent from the population, and medium-sized males established territories. The aggressive behavior exhibited by medium residents was similar to the previous year, but these males did not acquire higher reproductive successes than medium satellites had. The implications of switching tactics and the influence of operational sex ratios on the costs and benefits of the male tactics are briefly addressed. Electronic Publication