Edge effect and structure of mixed-species bird flocks in an Afrotropical lowland forest

Avian mixed-species flocks are a dominant feature of tropical moist forests, yet their cost–benefit balance and habitat dependence in Africa are not fully documented. We recorded the composition of mixed-species bird flocks in a pristine Afrotropical lowland forest site in Salonga NP, DRC. Our data showed that at least four types of flock existed, one of which specialized on edge habitats. We used multivariate analyses to further characterize edge effect on the most documented mixed-species flock type and found a significant effect on flock composition. While neither species guild nor preferential foraging stratum played an important role in flock participation, the nucleus role was played by a different species at the edge and in the interior, and both species abundance and associations were habitat-dependant, suggesting “domino effects” on the structure of mixed-species flocks.     

Linking vegetation structure and bird organization: response of mixed-species bird flocks to forest succession in subtropical China

As forests undergo natural succession following artificial afforestation, their bird assemblages also change. However, interspecific avian social organization associated with forest succession has not been fully understood, particularly for mixed-species bird flocks. To disentangle how mixed-species flocks change as a function of local forest structure, we analyzed flock characteristics (particularly species richness, flocking frequency and propensity) and vegetation physiognomies along a presumed successional series (early, middle, and advanced) simultaneously in subtropical forests in southern China. As hypothesized, monthly point counts demonstrated that complexity of flocks increases with the progression of natural forest succession at a local scale. Advanced forests differed significantly from pioneering plantations with respect to vegetation structure, flock characteristics and constituents (especially for understory specialists). Importantly, forest succession affected flock patterns particularly in relation to the flocking propensity of regular species, and the frequency of nuclear species (Huet’s fulvetta Alcippe hueti), which in turn determined flocking occurrence at different successional stands. Canonical correspondence analysis indicated that understory flocking species (mainly Timaliidae babblers) were significantly associated with intact native canopy cover, complex DBH diversity, as well as high densities of dead trees and large trees, representing a maturity level of successional stands. Our study reveals that the effect of natural forest succession on mixed-species bird flocks is species-specific and guild-dependent. From a conservation perspective, despite a high proliferation of pine plantation in southern China, priority should be placed on protecting the advanced forest with a rich collection of understory flocking specialists.     

The influence of flocking on the foraging behaviour of the chough (Pyrrhocorax pyrrhocorax) and the Alpine chough (P. graculus) coexisting in the Alps

We studied the flocking and foraging behaviour of the chough Pyrrhocorax pyrrhocorax and the Alpine chough P. gruculus coexisting in the south-western Italian Alps in order to evaluate the costs and benefits of foraging in single- and mixed-species flocks.
In the single-species context, flock size significantly affected the foraging behaviour of the Alpine chough; in larger flocks, the birds stayed for a shorter time in a patch and fed more quickly than in smaller flocks. Flock size did not significantly affect the foraging behaviour of the chough, probably because of the small number of individuals per flock.
The propensity for mixed-species flocking was rather low. The observed frequencies of single-species flocks of choughs and Alpine choughs were significantly higher than those expected on the basis of random flocking, whereas the observed frequencies of mixed-species flocks of the two species were lower than those expected. The stay times became significantly shorter for the chough in the presence of the Alpine chough. Moreover, feeding rates of the Alpine chough were significantly lower in the presence of the dominant chough.
The present study does not confirm the hypothetical foraging advantages of flocking. In single-species flocks, the benefits for the Alpine chough (higher feeding rates in larger flocks) were roughly compensated by the costs (shorter stay times in larger flocks), whereas the chough apparently neither gained benefits nor endured costs.
In mixed-species flocks, the Alpine chough sustained costs due to a reduction of feeding rates and the chough suffered costs due to a reduction of stay times. Hence, on average, single-species flocking gives no evident foraging advantages to either the chough or the Alpine chough, whereas mixed-species flocking provides some disadvantages for both species.     

Modeling the flocking propensity of passerine birds in two Neotropical habitats

We examined the importance of mixed-species flock abundance, individual bird home range size, foraging height, and foraging patch characteristics in predicting the propensity for five Neotropical passerine bird species (Slaty Antwren, Myrmotherula schisticolor; Golden-crowned Warbler, Basileuterus culicivorus; Slate-throated Redstart, Myioborus miniatus; Wilson’s Warbler, Wilsonia pusilla; and Black-and-white Warbler, Mniotilta varia) to forage within flocks, rather than solitarily. We used study plots in primary mid-elevation forest and in shade coffee fields in western Panama. We expected that all species would spend as much time as possible flocking, but that the social and environmental factors listed above would limit compatibility between flock movements and individual bird movements, explaining variability in flocking propensity both within and among species. Flocking propensity was well predicted by home range size and flock abundance together, for four of the five species. While flock abundance was uniform across plots, home range sizes varied among species and plots, so that home range size appeared to be the principle factor limiting flocking propensity. Estimates of flock abundance were still required, however, for calculating flocking propensity values. Foraging height and patch characteristics slightly improved predictive ability for the remaining species, M. miniatus. In general, individual birds tended to join flocks whenever one was available inside their home range, regardless of a flock’s specific location within the home range. Flocking propensities of individual species were lower in shade coffee fields than in forests, and probably vary across landscapes with variations in habitat. This variability affects the stability and species composition of flocks, and may affect survival rates of individual species. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Effects of supplemental food on intra-and inter-specific behaviour of the Varied Tit Parus varius

Varied Tits Parus varius lived in flocks containing a pair, or a pair with one or two unpaired birds. To test whether the sociality of Varied Tits or their participation in mixed-species flocks is sensitive to additional food (sunflower seeds), we compared the following parameters between fed and unfed periods at the individual level: population size, degree of site fidelity, mono-specific flock size and the stability of its membership, home-range size and distribution, attendance rate with mixed-species flocks, and flock size. Neither the size of the mono-specific flock nor its membership was affected by food supply. Whether food was added or not, individuals exhibited a strong site fidelity resulting in stable population size. Supplemental feeding had no effect on home-range size or distribution. In the presence of extra food, Varied Tits were observed in mono-specific flocks but rarely in mixed-species flocks. However, when we stopped feeding, they shifted to mixed-species flocking. After adding food, mixed-species flocks were significantly smaller than in the control samples. Varied Tits were more likely to join mixed-species flocks as temperatures dropped and wind speed increased. These results suggest that intra-specific sociality of Varied Tits is relatively insensitive to food supply, but they easily shift to mixed-species flocking in relation to food and weather conditions. We conclude that Varied Tits participate in mixed-species flocks to obtain short-term benefits e.g. increased foraging efficiency but they also obtain long-term benefits from stability of pair bonds and strong site fidelity, which did not respond to supplemental food.     

Mixed-species flocking of forest birds on Little Barrier Island

Abstract

Three aspects of mixed-species flocking of forest birds on Little Barrier Island were investigated. Whiteheads, fantails, parakeets, and grey warblers occurred more often in flocks than in “non-flocking” situations. Whiteheads were the main lead species, although parakeets formed groups within flocks and occasionally appeared to lead. Whitehead clumps defined the flock centre; only fantails were found commonly in the centre with whiteheads. Birds other than whiteheads generally orientated below or to the side of their nearest neighbours. We suggest that mixed-species flocking is a significant factor influencing the structure of forest bird communities in New Zealand during winter.     

Composition and foraging behaviour of mixed-species flocks led by the Grey-cheeked Fulvetta in Fushan Experimental Forest, Taiwan

Thirty-two species were recorded in mixed-species bird flocks led by the Grey-cheeked Fulvetta Alcippe morrisonia in Fushan Experimental Forest, Taiwan. Flocks averaged (± se) 5.8 ± 0.2 species and 51.4 ± 2.7 birds. Most participants were resident species (86.3%), some were elevational migrants (12.6%) and a few were latitudinal migrants (1.1%). Flock size was determined primarily by the abundance of Grey-cheeked Fulvettas, the most abundant species (68.1%). Flocks moved at an average rate of 10.8 ± 0.7 m/min, with larger flocks moving faster than smaller flocks. In moving flocks, canopy species were usually near the front, while understorey species usually followed. Fulvettas gave higher-intensity alarm calls and dived down more frequently in response to avian threats, especially raptors, than to non-avian threats. The overall foraging niche-breadth of the fulvetta was greater than that of any attendant species. Each species in a flock had a unique foraging niche. Most attendant species exhibited low foraging niche-overlap with the Grey-cheeked Fulvetta. Both the predator avoidance and the foraging efficiency hypotheses for mixed-species flocking were supported. The Grey-cheeked Fulvetta plays a critical role in the function of mixed-species flocks. A large flock formed around the Grey-cheeked Fulvetta provides attendant species with numerous opportunities for obtaining food and protection from predators.     

Age bias in the bag of pink-footed geese <Emphasis Type="Italic">Anser brachyrhynchus</Emphasis>: influence of flocking behaviour on vulnerability

In pink-footed goose (Anser brachyrhynchus) wintering in Denmark, The Netherlands and Belgium, the proportion of juveniles in the hunting bag is consistently higher than that observed in the autumn population. Such juvenile bias in the bag is usually ascribed to young geese lacking experience with hunting or disruption of juveniles from families. An alternative explanation may be that flocking behaviour of families make juveniles more vulnerable. Observations of morning flights of pink-footed geese to the feeding grounds from two of the major autumn-staging areas showed that geese were distributed in many small flocks (median flock size = 9). This was not significantly different from the flock size distribution shot at by hunters (median = 8), suggesting that hunters targeted goose flock size in proportion to the general probability of encounter. The rate at which hunters downed geese was independent of flock size. The ratio between juveniles and adults in flocks decreased with flock size and flocks of <60 individuals primarily comprised family groups. The likelihood of being shot at was 2.4 times higher for juveniles and 3.4 times higher for older birds in small flocks (<10 individuals) compared to larger flocks. The observations suggest that both juveniles as well as successful adult breeding birds were more vulnerable than non-breeding/failed breeding birds as a result of flocking behaviour.     

Associations among Antarctic seabirds in mixed-species feeding flocks

We studied ten mixed-species feeding flocks of seabirds and seals off South Georgia (55̀S, 35̀W) to assess the factors that influenced the species composition of the flocks. The ten flocks were distributed between two oceanographic regions; four flocks were observed off the northwestern end of South Georgia, and six off the island's southeastern end. The flocks differed dramatically in size and species composition. The northwestern flocks were ten to 20 times larger and contained more seals and penguins. We tested whether these differences were a consequence of location, because the two regions were populated by different assemblages of species, and determined that this was not the case. Differences in flock size and species composition reflected differences in the depth distribution of the birds' and seals' prey. The northwestern flocks were associated with deep swarms of Antarctic krill Euphausia superba with a significant quantity of krill also present at the surface. We also tested for differential flock participation by the species found in the surrounding waters and classified species by their proclivity to participate in the flocks. Our results indicate that Black-browed Albatross Diomedea melanophris have a strong tendency to join, and diving-petrels Pelecanoides spp. to avoid, mixed-species feeding flocks. We postulate that Black-browed Albatrosses serve as visual cues to the presence of food for one another and for other species, and that diving-petrels avoid mixed-species flocks in part because of the risk of predation.     

Does mixed-species flocking influence how birds respond to a gradient of land-use intensity?

Conservation biology is increasingly concerned with preserving interactions among species such as mutualisms in landscapes facing anthropogenic change. We investigated how one kind of mutualism, mixed-species bird flocks, influences the way in which birds respond to different habitat types of varying land-use intensity. We use data from a well-replicated, large-scale study in Sri Lanka and the Western Ghats of India, in which flocks were observed inside forest reserves, in ‘buffer zones'' of degraded forest or timber plantations, and in areas of intensive agriculture. We find flocks affected the responses of birds in three ways: (i) species with high propensity to flock were more sensitive to land use; (ii) different flock types, dominated by different flock leaders, varied in their sensitivity to land use and because following species have distinct preferences for leaders, this can have a cascading effect on followers'' habitat selection; and (iii) those forest-interior species that remain outside of forests were found more inside flocks than would be expected by chance, as they may use flocks more in suboptimal habitat. We conclude that designing policies to protect flocks and their leading species may be an effective way to conserve multiple bird species in mixed forest and agricultural landscapes.     

Aggression in shorebirds in relation to flock density and composition

The aggressive interactions of Turnstones and Purple Sandpipers feeding in wintering mixed-species wader flocks could be classified into those involving food and those involving space. All observed interspecific encounters were of short duration and were initiated and won by the larger species; the majority did not involve food and were resolved by low-intensity displays. Intraspecific interactions (a greater proportion of which involved food) were also resolved quickly and were usually won by the initiator. Space-related encounters between conspecifics were more likely to be resolved than food-related encounters just by threat displays. Aggression rates increased with flock densities. However, the increase in aggression with density was dependent on the species composition of the flock: both Turnstones and Purple Sandpipers were more likely to be involved in fights (both over food and over space) with conspecifics than with other species, indicating that the aggression costs of flocking were less in mixed-species flocks.     

Mixed species flocking of tits (Parus spp.): a field experiment

Summary We tested two general models of flocking behaviour, namely the antipredation model and foraging efficiency model on mixed-species tit flocks (Parus spp.). After food addition the size of mixed-species flocks was significantly less than in the control samples. In the presence of extra food significantly more birds were observed either in monospecific flocks or solitary, than during the control observations. In the presence of a living predator the birds foraged in larger mixed-specifies flocks than during the control observations. In addition, the social behaviour of Great Spotted Woodpecker, Middle Spotted Woodpecker and Nuthatch shifted to mixed-specific flocking. The size of monospecific flocks was independent of both treatments. The density of birds increased significantly after food addition, while in the predator presence the birds tended to leave the forest. These results support the view that both the antipredation model and foraging efficiency model seem to be valid for mixed-species flocking. However, in the case of monospecific flocks, the territory maintenance could be the most important factor.     

Bird mixed‐species flock formation is driven by low temperatures between and within seasons in a Subtropical Andean‐foothill forest

According to both the predation avoidance and foraging efficiency hypotheses, birds within mixed flocks increase their foraging efficiency and/or can spend more time feeding and less time looking out for predators. These hypotheses predict that birds in mixed flocks obtain benefits. Thus, mixed flock formation could serve as a strategy to cope with difficult conditions imposed on birds such as climatic conditions that ultimately result in a change in predation pressure or food resources. We evaluate the hypotheses that forming part of a flock confers benefits to its members and the associated prediction that birds will take advantage of these benefits and flock more often under cold and dry weather conditions between and within seasons to cope with such conditions. We surveyed the presence of mixed flocks, flocking propensity, number of species and individuals in mixed flocks in the Subtropical Yungas foothill of Argentina, to examine seasonality, flocking behavior of birds and their responses to two climatic variables: temperature and humidity. Bird species presented a higher flocking propensity and mixed flocks occurred more frequently during the dry and cold seasons than during the more benign seasons, and lower values of temperature within seasons triggered the flocking behavior. Although effects between seasons were expected, birds also showed a short‐term response to small changes in temperature within seasons. These results strengthen the ideas proposed by the foraging hypothesis. Although benefits derived from flocking have yet to be determined, whatever they are should be understood in the context of seasonal variation in life‐history traits.     

山地次生林冬季鸟类集群的初步研究

通过山地次生林两冬季鸟类集群的研究表明,集群的鸟类主要是由大山雀、沼泽山雀和普通Ｓｈｉ组成;每群个体数为７．４１－１０．６３只;据群的行为（单种群、多种集群和单独个体）分析,大多数种类是单独个体活动,冬季鸟类集群大小和组成有空间的变化。气候因子影响群的形成和行为。     

Composition of mixed‐species flocks and shifts in foraging location of flocking species on Hainan Island,China

A total of 134 bird species were recorded at Jianfengling, Hainan Island, in China from May 2000 to September 2004, of which 44 participated in one or more of 134 mixed‐species flocks. These flocks averaged 3.8 ± 0.2 species and 20.3 ± 1.2 individuals. Flocking propensity in a given species ranged from 1.5 to 100%. For flocking species, frequency of flocking and number of individuals in flocks was positively correlated with frequency and number in point counts. Among all species pairs with flocking frequency above 5%, cluster and correlation analysis indicated there were two principal groups of flocking birds – canopy species and understorey species: associations were positive within a group, but negative between groups. Canopy birds had a higher flocking propensity than understorey birds. They also made significantly less use of inner branches and trunks and greater use of middle branches, and foraged at a significantly greater height when in mixed‐species flocks than when solitary. For understorey bird species, there were no significant differences in foraging locations between solitary and mixed‐species flocks. Higher flocking frequency occurred in the wet season for canopy birds, but in the dry season for understorey birds. Overall patterns were consistent with the explanation that flocking enables an expansion of foraging niche by reducing the risk of predation.     

Multi-species territoriality and dynamic of neotropical forest understorey bird flocks

1. Eleven contiguous mixed-species bird flocks, with colour-banded individuals, were monitored continuously during 3 years in a 132-ha study area of primary rainforest in French Guiana.
2. Flock members were divided into six categories according to their flocking propensity and occurrence: 10 core or permanent species and 56 regular, occasional or incidental species. Each core species was represented by a single breeding pair with their fledglings and extra 'floaters' (unmated subadults and adults).
3. Flock home ranges overlapped slightly, but were communally defended by all core species in areas of overlap. Their size varied from 3·2 to 14·3 ha and was inversely correlated with vegetation density, but not flock size or species composition.
4. Flock number, size and composition, as well as boundaries were highly stable between seasons and years. Each flock had a single permanent gathering site and bathing site in late afternoon, the latter sometimes shared by 2–3 flocks.
5. Core species produced 0·18–0·73 fledglings per pair per year, which stayed in their natal flock for 200 to over 421 days. Then, these individuals usually moved between two and six different flocks, sometimes for up to 3 years, before finding a mate and a flock where they could settle and breed. Once breeding, they probably remained for life in the same flock. The mean annual survival rate was at least 0·75.
6. This highly evolved and stable organization, associated with a low breeding success and high survival rate was a critical factor maintaining low species density, delayed reproduction and a proportion of floating individuals buffering population fluctuations.
7. These social groups with their multi-species territoriality and co-evolved roles of flock members were similar to those described elsewhere in South America. They seem to be a general phenomenon in neotropical lowland rainforests.     

Pregnancy outcome of women in the vicinity of nuclear power plants in Taiwan

The purpose of the study was to investigate whether proximity to nuclear power plants may increase the risk of abnormal pregnant outcomes among the resident women. In this ecological study, data were used from the Health Services Birth Reports Database established by the Bureau of Health Promotion, National Department of Health, Taiwan, in 2001–2004. Chi-square-tests were carried out to investigate the “Plant-vicinity” and “Non plant-vicinity” group in terms of pregnancy outcome. Additionally, logistic regression was performed to investigate whether residence in the vicinity of a nuclear power plant was related to any abnormal pregnancy results. Based on data from 5,679 included subjects, no difference was observed between pregnancy outcomes of the “Plant-vicinity” and “Non plant-vicinity” groups. After accounting for possible confounders, the adjusted odds ratios were 1.20 (95% CI = 0.56–2.56) for stillbirth, 1.21 (95% CI = 0.95–1.53) for premature birth, 1.04 (95% CI = 0.79–1.37) for low birth weight, and 1.58 (95% CI = 0.85–2.93) for congenital deficiencies, respectively, when comparing the “Plant-vicinity” with the “Non plant-vicinity” group. The results of the study indicate that residence in the vicinity of a nuclear power plant is not a significant factor which will cause abnormal health situations during pregnancy.     

The formation of “mega‐flocks” depends on vegetation structure in montane coniferous forests of Taiwan

A mixed‐species bird flock is a social assemblage where two or more bird species are moving together while foraging and might benefit from increased foraging efficiency and antipredator vigilance. A “mega‐flock,” which includes flocking species from different vegetation strata, often exhibits high species diversity. Mechanisms for the formation of mega‐flocks have not yet been explored. In this study, we evaluated the influence of vegetation structure and bird species diversity in driving the occurrence of mega‐flocks. We investigated the composition of mixed‐species flocks, local bird communities, and vegetation structure in five vegetation types of two high‐elevation sites in central Taiwan. Mega‐flocks occurred more frequently in pine woodland than later successional stages of coniferous forests. However, species richness/diversity of local bird communities increased along successional stages. Therefore, vegetation variables exhibit more influence on the occurrence of mega‐flocks than local bird communities. Besides foliage height diversity, understory coverage also showed positive effects on flock size of mixed‐species flocks. Our results indicated that pine woodlands with more evenly distributed vegetation layers could facilitate the interactions of canopy and understory flocks and increase the formation of mega‐flocks and thus the complexity of mixed‐species flocks.     

Dolichopteryx minuscula, a new species of spookfish (Argentinoidei: Opisthoproctidae) from the Indo-West Pacific

A new species Dolichopteryx minuscula is described on the basis of three specimens [49.4–59.6 mm in standard length (SL)] collected from the Indo-West Pacific. The new species is characterized by pouchlike eyes with a small lens (lens diameter 2.2% SL), an adipose fin, the anal fin base originating posterior to the dorsal fin base, and 16–17 (= 5–6 + 1 + 10–11) gill rakers. Total fecundity was relatively low, only 658 ova being obtained from one specimen, despite the ovary being mature. Ovarian eggs were clearly subdivided into “undeveloped” (0.1–0.7 mm diameter classes, n = 561) and “developed” (1.0–1.3 mm classes, n = 97) groups, based on their frequency distribution. Such relatively low fecundity and frequency distributions of ovarian eggs suggest that Dolichopteryx species spawn iteratively during spawning season.     

A common variant in the adiponectin gene and polycystic ovary syndrome risk

In this study, we explored whether polymorphisms in insulin receptor (INSR), adiponectin (ADIPOQ), parathyroid hormone (PTH), and vitamin D receptor (VDR) genes are associated with polycystic ovary syndrome (PCOS). A total of 362 subjects, including 181 women with PCOS and 181 controls were enrolled in this case-control study. Two SNPs (rs2059806 and rs1799817) in the INSR gene, two SNPs (rs2241766 and rs1501299) in the ADIPOQ gene, one SNP (rs6256) in the PTH gene, and one SNP (rs757343) in the VDR gene were analyzed using PCR-RFLP method. We observed no significant difference in genotype and allele frequencies between the women with PCOS and controls for the rs2059806, rs1799817, rs1501299, rs6256, and rs757343 polymorphisms either before or after adjustment for confounding factors including age and BMI. However, the ADIPOQ rs2241766 “TT” genotype compared with “TG and GG” genotypes was associated with a 1.93-fold increased risk for PCOS (P = 0.006, OR = 1.93, 95% CI = 1.20–3.11), and the differences remained significant after adjustment for age and BMI (P = 0.039, OR = 1.72, 95% CI = 1.03–2.86). Furthermore, the ADIPOQ rs2241766 “T” allele was significantly overrepresented in women with PCOS than controls (P = 0.006; OR = 1.80, 95% CI = 1.18–2.70), and the difference remained significant after Bonferroni correction. Our findings suggest that the ADIPOQ rs2241766 “TT” genotype is a marker of increased PCOS susceptibility. This study also indicates for the first time that there are no significant association between INSR rs2059806, PTH rs6256, and VDR rs757343 gene polymorphisms and PCOS risk. However, these data remain to be confirmed in larger studies and in other populations.