Isolation and characterization of polymorphic microsatellite loci in the plural cooperatively breeding superb starling,Lamprotornis superbus

Superb starlings (Lamprotornis superbus) are separate nesting plural cooperative breeders endemic to East Africa that show high levels of reproductive conflict and low reproductive skew. I isolated 31 polymorphic microsatellite loci from the superb starling genome and characterized them in approximately 31 individuals. The numbers of alleles per locus ranged from two to 16 and observed heterozygosities ranged from 0.03 to 0.88. Many of these loci amplified in other passerine species including the European starling, Sturnus vulgaris, and a variety of other starlings and mockingbirds.     

Temporal but not spatial environmental variation drives adaptive offspring sex allocation in a plural cooperative breeder

Cooperatively breeding birds have been used frequently to study sex allocation because the adaptive value of the sexes partly depends upon the costs and benefits for parents of receiving help. I examined patterns of directional sex allocation in relation to maternal condition (Trivers-Willard hypothesis), territory quality (helper competition hypothesis), and the number of available helpers (helper repayment hypothesis) in the superb starling, Lamprotornis superbus, a plural cooperative breeder with helpers of both sexes. Superb starlings biased their offspring sex ratio in relation to prebreeding rainfall, which was correlated with maternal condition. Mothers produced relatively more female offspring in wetter years, when they were in better condition, and more male offspring in drier years, when they were in poorer condition. There was no relationship between offspring sex ratio and territory quality or the number of available helpers. Although helping was male biased, females had a greater variance in reproductive success than males. These results are consistent with the Trivers-Willard hypothesis and suggest that although females in most cooperatively breeding species make sex allocation decisions to increase their future direct reproductive success, female superb starlings appear to base this decision on their current body condition to increase their own inclusive fitness.     

Influence of habitat and season on foraging flock composition in the European Starling (Sturnus vulgaris)

Summary This study examines the relative importance of habitat and season in determining size and composition of foraging flocks of European Starlings (Sturnus vulgaris). We hypothesized that if season was the principal determinant of flock size, variation across seasons (within habitat) would be greater than variation across habitats (within season). The calendar year was divided into 6 seasons corresponding to major stages in the annual cycle of starlings. After measuring the availibility of 15 foraging habitats occurring within the study area, we drove a 184 km route weekly (over a 17 month period) to census starling flock size, species composition, habitat use, and activity.Although flock size was influenced by both season and habitat, it varied relatively less within seasons (across habitats). Only five of nine seasons showed significant habitat effects, and two (of the five) were significant mainly because of large aggregations in habitats with unusually abundant food (e.g. feedlots, cornfields). In contrast, eight of nine habitats showed significant seasonal effects. The only exception (hay) was little used (<1% of all birds observed). For starlings the activities associated with seasonal events of the annual cycle were a major influence on patterns of social foraging. Habitat factors were less important because of constraints imposed by seasonal factors, but under some conditions (e.g. drought, clumped food supply) they became dominant.     

Starling foraging success in relation to agricultural land-use

Changes in agricultural land-use have been suggested to contribute to the decline of several bird species through negative effects on their food supply during breeding. One important change in land-use has been loss of pastures, especially permanent pastures. In this study we investigated how different forms of agricultural land-use affected foraging success of a declining bird species, the European starling Sturnus vulgaris . We let caged starlings forage in different forms of agricultural fields and determined time spent foraging and foraging success. The starlings' activity level (time spent actively foraging) as well as the number of prey caught per time unit was strongly related to the abundance of prey in soil samples. Also the body mass change during the experiment was positively related to activity level and prey capture rate. We found consistent differences in foraging variables between habitats. In spring sown grain starlings were least active and found fewer prey items at a lower rate than in any other habitat. The other three habitats differed less, but in general mowed hay fields appeared slightly more valuable than the cultivated and natural pastures. We did not find any differences between natural and cultivated pastures in foraging variables. Thus, starling foraging success is higher in grass-covered fields than in cultivated fields, but the management of the grass-covered fields mattered less. The results are consistent with starlings having higher population densities and breeding success in areas with higher availability of pasture. We suggest that the physical structure of the habitat (sward height) and moisture may be additional variables that need to be taken into account to explain starling breeding density and success in the agricultural landscape.     

Association of drongos with myna flocks: Are drongos benefitted?

Common myna(Acridotherus tristis) and jungle myna(Acridotherus fuscus) forage in pure and mixed flocks of various sizes in fallow lands. These flocks were often found associated with drongos that forage individually on the insects herded out by the movements of the flocking myna. We report here the benefits and costs of such association to drongos and mynas. Drongos had a tendency to associate with larger (> 21) than smaller (<20) flocks irrespective of the species composition of the flocks. Drongos associated with larger flocks showed increased foraging trips and harvested more insects in a given time than those that were either isolated or were associated with small flocks. The food range of drongos and mynas differed significantly indicating that they do not compete with each other. Thus our results indicate that drongos are benefitted by this association; however this association neither benefits nor costs to the mynas. The association between the drongos and mynas therefore appears to be commensalistic.     

Territory quality drives intraspecific patterns of extrapair paternity

Despite the potential reproductive benefits of extrapair matings,extrapair paternity rates in many avian species often vary greatlyamong populations. Although ecological factors have been shownto influence intraspecific patterns of extrapair paternity insome species, for cooperatively breeding species living in familygroups, social/demographic factors may also play a role. Thisstudy examined how ecological factors related to territory quality(vegetation cover, insect abundance) and social/demographicfactors (group size, number of breeding pairs, genetic relatedness)influenced intraspecific patterns of extrapair paternity incooperatively breeding superb starlings, Lamprotornis superbus.Superb starlings inhabit spatiotemporally variable African savannaswhere high temporal variability drives reproductive decisions(adoption of breeding roles, offspring sex allocation) and whereterritories suitable for breeding are limited. Although extrapairpaternity rates were only 14% of offspring and 25% of nests,they varied greatly among groups, ranging from 4% to 32% ofoffspring and from 7% to 60% of nests. These among-group differencesin extrapair paternity were not related to social/demographicfactors but instead to territory quality; extrapair paternitywas higher on lower quality territories (lower vegetation coverand grasshopper abundance) than on higher quality territories(higher vegetation cover and grasshopper abundance). These resultssuggest that even in a heterogeneous landscape where suitablebreeding territories are limited, subtle differences in habitatquality can have profound effects on reproductive decisionsand patterns of extrapair paternity. Understanding the interactionbetween spatial (habitat heterogeneity) and temporal (temporalvariability) environmental variation will be important for determininghow environmental and social factors drive avian reproductiveand mating decisions.     

Bateman's principle is reversed in a cooperatively breeding bird

Bateman''s principle is not only used to explain sex differences in mating behaviour, but also to determine which sex has the greater opportunity for sexual selection. It predicts that the relationship between the number of mates and the number of offspring produced should be stronger for males than for females. Yet, it is unclear whether Bateman''s principle holds in cooperatively breeding systems where the strength of selection on traits used in intrasexual competition is high in both sexes. We tested Bateman''s principle in the cooperatively breeding superb starling (Lamprotornis superbus), finding that only females showed a significant, positive Bateman gradient. We also found that the opportunity for selection was on average higher in females, but that its strength and direction oscillated through time. These data are consistent with the hypothesis that sexual selection underlies the female trait elaboration observed in superb starlings and other cooperative breeders. Even though the Bateman gradient was steeper for females than for males, the year-to-year oscillation in the strength and direction of the opportunity for selection likely explains why cooperative breeders do not exhibit sexual role reversal. Thus, Bateman''s principle may not hold in cooperative breeders where both sexes appear to be under mutually strong sexual selection.     

Stress hormones and sociality: integrating social and environmental stressors

In cooperatively breeding species, reproductive decisions and breeding roles may be influenced by environmental (food resources) or social factors (reproductive suppression of subordinates by dominants). Studies of glucocorticoid stress hormones in cooperatively breeding species suggest that breeding roles and hormone levels are related to the relative costs of dominance and subordination, which are driven primarily by social interactions. Few studies, however, have considered how environmental factors affect glucocorticoid levels and breeding roles in cooperative breeders, even though environmental stressors modulate seasonal glucocorticoid release and often influence breeding roles. I examined baseline and stress-induced levels of the glucocorticoid corticosterone (CORT) across 4 years in the plural breeding superb starling, Lamprotornis superbus, to determine whether (i) environmental factors (namely rainfall) directly influence breeding roles or (ii) environmental factors influence social interactions, which in turn drive breeding roles. Chronic baseline and maximal stress-induced CORT changed significantly across years as a function of pre-breeding rainfall, but dominant and subordinate individuals responded differently. Pre-breeding rainfall was also correlated directly with breeding roles. The results are most consistent with the hypothesis that environmental conditions influenced the relative costs of dominance and subordination, which in turn affected the degree and intensity of social interactions and ultimately reproductive decisions and breeding roles.     

Invasive ring-necked parakeets Psittacula krameri in Belgium: habitat selection and impact on native birds

There is a large and growing number of alien species in ecosystems all over the world. In view of this, management efforts should concentrate on invasives that have detrimental effects on native biota. However, action against invasives is often hindered by a lack of relevant ecological information such as the expected distribution and impact of the invader. Our aims were to identify the habitat characteristics that influence ring-necked parakeet abundance in Belgium and to assess the effects of competition for nesting cavities with native hole-nesters. We determined the abundance of parakeets and native hole-nesters in 44 study sites using point counts. We examined the relationship between parakeet numbers and a set of habitat and landscape variables and to assess the effect of competition, we studied the relationships between the number of parakeets and the number of native hole-nesters. We found that parakeet abundance differs between vegetation types based on dominant tree species, and that abundance is higher in forests or parks surrounded by built-up areas. Parakeet numbers were also strongly associated with cavity density, suggesting that this may be a limiting factor. Of the two native hole-nesters considered to be the most vulnerable to competition (nuthatch and starling), we found a negative association with parakeet numbers for the nuthatch only. No relations were found with starlings or any of the other hole-nesting species examined. These findings should be considered in the debate concerning the possible need of action against this species.     

Large herbivores that strive mightily but eat and drink as friends

Summary Grazing in patches of Cynodon dactylon and of Sporobolus spicatus by four large herbivores, and the interaction between these sedentary herbivores was studied in Lake Manyara National Park, northern Tanzania. The herbivores were the African buffalo, Syncerus caffer; the African elephan, Loxodonta africana; the Burchell's zebra, Equus burchelli; and the wildebeest, Connochaetus taurinus. Four different hypotheses of the interactions between the herbivores were tested, viz., increased predator detection/protection through association of species, facilitation of the food intake through the influence of other species, use by other species of the food manipulation strategy of buffalo, and interspecific competition for food. On the level of a single day, zebra and wildebeest were symbiotic, which could have been caused by an increased chance of predator detection. A similar association between buffalo and wildebeest or zebra was also detected on C. dactylon grasslands. There was no indication of facilitation between any of the herbivores. Buffalo had a despotic relationship with elephant, that is the elephant's consumption was lowered when buffalo had visited a patch prior to their arrival. When elephant and buffalo arrived at the same time there appeared to be scramble competition between them.Habitat overlap was calculated for four pairs of species. In conjunction with the analyses of the patch visits, it was concluded that a small overlap was associated with interspecific competition and a large habitat overlap was associated with symbiosis.     

Food diversity and choice of white-headed langur in fragmented limestone hill habitat in Guangxi,China

We studied the diet and food choice of three groups of white-headed langurs (Trachypithecus leucocephalus) from July 2002 to June 2003 at Fusui Rare Animal Reserve, Guangxi, China. Data were collected from via focus group sampling, with continuous recording. A total of 109 plant species (including 19 unidentified species) were used as food by the white-headed langur. They spent 91.6% of total feeding time feeding on leaves, 4.2% on fruits and 3.3% on stems. Flowers and other food items accounted for only 0.3% and 0.6% of total feeding time, respectively. There were significant seasonal variations in the langurs’ diets with changes in the total number of species and the diversity of food species consumed. The langurs used significantly more plant species as food in the rainy season than in the dry season (36 species vs. 26.5 species). Dietary diversity was also significantly higher in the rainy season than in the dry season (2.6 vs. 2.4). No significant seasonal variation was found in the percentage of various plant parts in their monthly diets. Though the white-headed langur fed on many plant species, 11 species made up 84.9% of their diet. There were no significant correlations between the percentage of feeding time for main food plant species and their corresponding abundance in the habitat, content of water, crude protein, crude fiber and the ratio of crude protein to crude fiber. These results suggest that food choice of the white-headed langur is not influenced by their abundance and nutritional content.     

Interspecific signalling between mutualists: food-thieving drongos use a cooperative sentinel call to manipulate foraging partners

Interspecific communication is common in nature, particularly between mutualists. However, whether signals evolved for communication with other species, or are in fact conspecific signals eavesdropped upon by partners, is often unclear. Fork-tailed drongos (Dicrurus adsimilis) associate with mixed-species groups and often produce true alarms at predators, whereupon associating species flee to cover, but also false alarms to steal associating species'' food (kleptoparasitism). Despite such deception, associating species respond to drongo non-alarm calls by increasing their foraging and decreasing vigilance. Yet, whether these calls represent interspecific sentinel signals remains unknown. We show that drongos produced a specific sentinel call when foraging with a common associate, the sociable weaver (Philetairus socius), but not when alone. Weavers increased their foraging and decreased vigilance when naturally associating with drongos, and in response to sentinel call playback. Further, drongos sentinel-called more often when weavers were moving, and weavers approached sentinel calls, suggesting a recruitment function. Finally, drongos sentinel-called when weavers fled following false alarms, thereby reducing disruption to weaver foraging time. Results therefore provide evidence of an ‘all clear’ signal that mitigates the cost of inaccurate communication. Our results suggest that drongos enhance exploitation of a foraging mutualist through coevolution of interspecific sentinel signals.     

Long‐Distance Calling by the Willow Tit,Poecile montanus,Facilitates Formation of Mixed‐Species Foraging Flocks

The occurrence of mixed‐species foraging flocks is a worldwide phenomenon in terrestrial bird communities. Previous studies suggest that individuals participating in flocks might derive benefits in terms of improved feeding efficiency and/or reduced risk of predation. However, very little is known about how individuals establish mixed‐species flocks. Here, I provide the first experimental evidence that long‐distance calling by the willow tit, Poecile montanus, facilitates the establishment of mixed‐species flocks at a foraging patch. Observations at experimental foraging patches showed that willow tits that find a food source produce long‐distance calls, particularly when they are isolated from conspecific flockmates. The probability of long‐distance calling was negatively correlated with the number of heterospecific foraging individuals near the food source. A playback experiment confirmed that calls attract both conspecific and heterospecific members of foraging flocks. This study demonstrates that willow tits use long‐distance calls to attract conspecific flockmates to foraging patches, and these calls can also facilitate the formation of mixed‐species flocks on patches.     

Diets of adult and nestling starlings (Sturnus vulgaris) in Hawke’s Bay,New Zealand

Abstract

Gizzards were examined from 334 adult and 62 nestling starlings collected in mixed farmland during 1971–72. The birds ate insects, spiders, earthworms, snails, millipedes, centipedes, seeds, and fruits. Starlings ate fewer subterranean animals (7%) than those usually living partially hidden (45%), on the ground (31%), or on vegetation (18%). About half the invertebrates eaten were 2–5 mm long and about a quarter 6–10 mm long. Nestlings tend to be fed significantly larger items than were eaten by adults. The commonest items in adult starlings were Coleoptera adults, Lepidoptera larvae, Hemiptera, and fruits; in nestlings, Coleoptera adults and Diptera adults and larvae were important. Earthworms were found in all the nestlings. More than 50% of adult gizzards contained earthworm chaetae in wetter months, but fewer in drier months. The diet of starlings, despite considerable overlap with the foods eaten by mynas, rooks, and magpies, included somewhat different components and proportions of the food supply. Fruits were found in adults only; most were probably eaten after harvest. Although predation on two major pasture pests—Costelytra zealandica and Wiseana cervinata larvae—was insubstantial, nearly 40% of the total invertebrates eaten by adult starlings were insect pest species.     

Behavioural thermoregulation and climatic range restriction in the globally threatened Ethiopian Bush‐crow Zavattariornis stresemanni

Climate may influence the distribution and abundance of a species through a number of demographic and ecological processes, but the proximate drivers of such responses are only recently being identified. The Ethiopian Bush‐crow Zavattariornis stresemanni is a starling‐like corvid that is restricted to a small region of southern Ethiopia. It is classified as Endangered in the IUCN Red List of globally threatened species. Previous work suggested that this range restriction is almost perfectly defined by a climate envelope that is cooler than surrounding areas, but the proximate mechanism remains unexplained. The heavily altered habitats which the species inhabits are widespread across Africa, and recent work has shown that the Bush‐crow is behaviourally adaptable and has a catholic diet. We assess whether its enigmatic distribution can be explained by behavioural responses to the higher temperatures that surround its current range. Using environmental niche models and field observations of thermally mediated behaviour, we compare the range restriction and behavioural thermoregulation of the Ethiopian Bush‐crow with those of two sympatric control species that are similar in size and ecology, but have much larger ranges that include hotter environments. White‐crowned Starling Lamprotornis albicapillus and Superb Starling L. superbus occupy similar habitats to the Ethiopian Bush‐crow and all three frequently forage together. We found that the Bush‐crow's range is limited primarily by temperature, with a secondary effect of dry season rainfall, whereas the ranges of the two starling species are better predicted by wet season rainfall alone. Bush‐crows exhibited panting behaviour and moved into the shade of trees at significantly lower ambient temperatures than did the starlings, and their food intake declined more steeply with increasing temperature. These results indicate that the limited geographical range of the Bush‐crow reflects an inability to cope with higher temperatures. This suggests that a species' response to climate change might not be easily predicted by its ecological generalism, and may represent an inherited debt from its evolutionary history.     

ADAPTIVE ADVANTAGES OF MIXED-SPECIES FEEDING FLOCKS AMONG SEED-EATING FINCHES IN COSTA RICA

The feeding ecology of three Costa Rican finches occurring in mixed flocks, Tiaris olivacea, Sporophila aurita corvina and Sporophila torquella , was investigated by measuring both behavioural and ecological variables. Observations on foraging height, rate of hopping and pecking rate, as well as the identity, proximity and number of nearest neighbours were recorded. In addition the duration of all feeding and perching episodes were timed. Comparisons were also made between the abundances of food items (grass seeds) consumed and those potentially available for consumption.
The analysis of variance of the feeding behaviour revealed that the presence or absence of neighbouring birds, whether of the same or different species, influenced the duration of feeding bouts more significantly than did either differences in habitat or species-characteristic behaviour. In addition the dietary comparisons revealed overlap in both species and size of seed consumed. Such similarities suggest that these species are not partitioning fields in the classical sense.
We propose that the increase in the duration of the feeding bout associated with the presence of mixed species aggregations leads to increased feeding efficiency and is the result of intra- and inter-specific social learning. Certainly flocking is often advantageous, since searching in a group facilitates finding clumped resources; mixed species flocking, by increasing exposure to a diversity of foraging places and patterns, can further augment feeding efficiency.     

Raptors breeding on weaver nests in trees and on man-made structures

Several raptor species nest on top of large weaver nests. These weaver nests are usually sited in trees, but 11.7% of Red-billed Buffalo-Weaver Bubalornis niger and 25.7% of Sociable Weaver Philetairus socius nests occur on man-made structures. In an extensive literature search, a total of 16 raptor species were recorded as nesting on top of weaver nests. At least 10 raptor species used weaver nests built in trees. Seven raptor species used weaver nests on man-made sites and four raptor species only used weaver nests built on man-made sites. No owls have been recorded as using weaver nests on man-made sites. There are historical records of raptors nesting on top of weaver nests in trees, while nesting on top of weaver nests sited on man-made structures appears to be a more recent adaptation. Costs and benefits of nesting on man-made sites are briefly listed. Nesting on man-made sites may increase (by both weavers and raptors) and raptor researchers are encouraged to document cases of raptors nesting on weaver nests where these are placed in trees or artificial sites, so that there is a record of changes in frequency of nest site usage by raptors.     

Community ecology of helminth parasitism in an insular passerine avifauna.

Three hundred and thirty specimens of 7 species of passerine birds from South Bass Island, Ottawa County, Ohio, were examined for helminth parasites. The total number of helminth specimens collected was 4,333. Forty-one helminth taxa were identified. Ten species of helminths were identified as having foci of infection on the island. An index of association for these 10 species is presented. The low association revealed between helminth species utilizing common species of intermediate hosts indicated that a single intermediate host specimen is likely to harbor infective stages of only one species of parasitic helminth. An index of similarity is presented to express the species importance relationships of the helminth faunas of the 7 species of birds: red-winged blackbirds (Agelaius phoeniceus), blue jays (Cyanocitta cristata), brown-headed cowbirds (Molothrus ater), common grackles (Quiscalus quiscula), house sparrows (Passer domesticus), starlings (Sturnus vulgaris), and robins (Turdus migratorius). The results reveal that competition between these avian species for invertebrate food resources helps to maximize the transmission of those helminth populations which utilize these same invertebrates as intermediate hosts. The aggregation of birds into mixed feeding flocks insures a heavy loading of the feeding grounds with agents infective for the invertebrate populations.     

The fitness consequences of kin-biased dispersal in a cooperatively breeding bird

Cooperative alliances among kin may not only lead to indirect fitness benefits for group-living species, but can also provide direct benefits through access to mates or higher social rank. However, the immigrant sex in most species loses any potential benefits of living with kin unless immigrants disperse together or recruit relatives into the group in subsequent years. To look for evidence of small subgroups of related immigrants within social groups (kin substructure), we used microsatellites to assess relatedness between immigrant females of the cooperatively breeding superb starling, Lamprotornis superbus. We determined how timing of immigration led to kin subgroup formation and if being part of one influenced female fitness. Although mean relatedness in groups was higher for males than females, 26% of immigrant females were part of a kin subgroup with a sister. These immigrant sibships formed through kin recruitment across years more often than through coalitions immigrating together in the same year. Furthermore, females were more likely to breed when part of a kin subgroup than when alone, suggesting that female siblings form alliances that may positively influence their fitness. Ultimately, kin substructure should be considered when determining the role of relatedness in the evolution of animal societies.     

Changing grass height alters foraging site selection by wintering farmland birds

Grassland is a valuable habitat for many farmland bird species, some of which have declined in Europe. Successful management of this habitat is important to benefit these species. Recent studies have suggested little correlation between food abundance and bird use in grassland; instead providing access to food may be crucial. We tested the importance of manipulating sward height (via mowing) on the distribution of wintering birds in agriculturally managed grass fields using two within-field experimental designs. In experiment 1, we investigated the effect of two treatments (mown once and control). In experiment 2, we investigated the effects of four treatments (mown once early, mown once late, mown early and late, and control).Mown plots supported higher abundances of foraging kestrels, thrushes and starlings. In contrast meadow pipits and grey herons preferred unmown plots with longer grass. These differences are probably explained by differing anti-predation strategies, increased food abundance for species that prefer above-ground invertebrates (e.g. more food for meadow pipits on longer swards) and increased food accessibility on short grass swards (e.g. for starling, thrush spp.). There were limited effects of multiple mowing events on plot use suggesting one early winter mowing treatment may suffice to create favourable conditions for species that prefer shorter swards.Providing grassland heterogeneity should benefit a range of wintering farmland birds. Although we use mowing to change grass sward height, our findings may also apply at other times of year when livestock grazing may be a more practical solution.