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Made available in DSpace on 2014-12-17T14:10:22Z (GMT). No. of bitstreams: 1 GlauberHBOS_DISSERT.pdf: 1092481 bytes, checksum: e18c958f4fa280552deed9ab75e1ca3b (MD5) Previous issue date: 2010-06-30 / The diet study of birds has contributed historically as a model for use to understanding ecological patterns and strategies used by several other groups of vertebrates, which are observed in season patterns and temporal availability of resources, and other. This study has as objective generate information concerning the diet of insectivorous birds during rainy season and dry season, as well as analyze Index food importance, niche overlap, niche breadth, electivity, and seasonal availability of prey. The study was conducted in a fragment of about 270 ha (center coordinates and 5 ? 53'S 35 ? 23'W). The sampling of birds occurred between March 2008 and December 2009 in three pre-established trails. Catches of birds were performed using 10 mist nets placed in line, where each trails was sampled once a month. Samples of pellets were obtained by means of tartar emetic. Sampling of availability of prey occurred between February 2009 to December 2009. We used two methods of sampling (pitfall traps and Shake cloths). We captured 269 individuals of 21 species of insectivorous birds. We collected 4116 invertebrates of which 3259 in the rainy season and 857 in the dry season. We obtained 174 samples stomach, where 10 species were exclusively insectivorous diet, nine fed on insect/plant material, an insect/plant material/vertebrate and one for insect/vertebrate. During the rainy season was observing difference between the consumption of items with higher food importance. The Coleoptera was item with higher food importance (73%), followed by Formicidae (7%) and Araneae (6%). During the dry season, no difference was found difference between the consumption of items with higher food importance. The Coleoptera was item with higher food importance (34%), followed by seeds (29%) and Formicidae (18%). The highest levels of niche overlap occurred during the rainy season, while the dry season was characterized by high levels of niche 11 segregation. This indicates that the local insectivorous birds community was structured differently between periods. No was found correlation between the values of niche breadth to the mean weight of the body size. We observed seasonal patterns in prey availability, with the peak availability of invertebrates observed seasonal patterns in rainy season. The insectivorous birds selected the same species richness during both periods, showing a specialized diet. Thamnophilus pelzelni was the only species that had their diet influenced by seasonality. Regarding the overall diet of insectivorous birds, observed a high consumption of prey, whose food availability caused the birds could invest and increase their food resources / O estudo da dieta de aves tem contribu?do, historicamente, como modelo de uso para o entendimento de padr?es e estrat?gias ecol?gicas utilizados por v?rios outros grupos de vertebrados, nos quais s?o observados padr?es sazonais e temporais na disponibilidade de recursos e outros. O presente trabalho objetivou gerar informa??es referentes ? dieta de aves inset?voras durante os per?odos chuvoso e seco, bem como analisar import?ncia alimentar, valores de sobreposi??o alimentar, amplitude de nicho, eletividade e disponibilidade de presas durantes as esta??es chuvosa e seca. O estudo foi realizado em um fragmento com cerca de 270 ha (coordenadas centrais 5?53 S e 35?23 W). O per?odo de amostragem das aves ocorreu entre mar?o de 2008 a dezembro de 2009, em tr?s trilhas pr?-estabelecidas. As capturas das aves foram realizadas por meio de 10 redes ornitol?gicas dispostas em linha, onde cada linha de rede foi colocada em uma das trilhas, sendo uma amostragem de aves realizadas em uma ?nica trilha por m?s. As amostras de regurgitos foram obtidas por meio do t?rtaro em?tico. As amostragens de disponibilidade de presas ocorreram entre fevereiro de 2009 a dezembro de 2009, sendo utilizados amostragens no solo (pitfall traps) e na folhagem (shake cloths). Capturamos 269 indiv?duos de 21 esp?cies de aves inset?voras. Foram coletados 4.116 invertebrados, dos quais 3.259 no per?odo chuvoso e 857 no per?odo seco. Obtivemos 174 amostras estomacais, onde 10 esp?cies tiveram dieta exclusivamente inset?vora, 9 alimentaram-se de inseto/material vegetal, 1 de inseto/material vegetal/vertebrado e 1 por inseto/vertebrado. Durante o per?odo chuvoso foi observada diferen?a entre o consumo dos itens com maior Import?ncia 9 Alimentar, sendo Coleoptera o item com maior Import?ncia Alimentar (73%), seguido por Formicidae (7%) e Araneae (6%). Durante o per?odo seco, n?o foi encontrada diferen?a entre o consumo dos itens com maior import?ncia Alimentar, Sendo Coleoptera o item com maior Import?ncia Alimentar (34%), seguido por sementes (29%) e Formicidae (18%). Os maiores ?ndices de sobreposi??o de nicho ocorreram durante o per?odo chuvoso, enquanto o per?odo seco foi caracterizado pelos maiores valores de segrega??o de nicho. Isto indica que a comunidade de aves inset?voras local esteve estruturada de forma diferenciada entre os per?odos. N?o foi encontrada correla??o entre os valores de Amplitude de nicho com os pesos m?dios das aves, mostrando que o tamanho do nicho ocupado pelas aves n?o tem rela??o com o seu tamanho corporal. Foram observados padr?es sazonais na disponibilidade de presas, sendo o pico da disponibilidade de invertebrados observado durante o per?odo chuvoso. As aves inset?voras selecionaram a mesma riqueza de esp?cies durante os dois per?odos, mostrando uma dieta especializada. Thamnophilus pelzelni foi a ?nica esp?cie que teve a sua dieta influenciada pela sazonalidade. No que diz respeito a dieta global das aves inset?voras, observamos um elevado consumo de presas, cuja grande disponibilidade de alimento fez com que as aves pudessem investir e aumentar os seus recursos alimentares

The question of group-living in animals has attracted a lot of research attention. Today, we have a nuanced understanding of ultimate explanations and proximate mechanisms for group-living, as well as the genetic basis for seemingly puzzling aspects of sociality such as altruism. However, our current view of sociality has been obtained almost entirely in an intraspecific context, i.e. based on groups consisting of individuals of a single species. On the other hand, heterospecific sociality, i.e. groups formed by individuals of multiple species, has been largely ignored in group-living theory and empirical research. This is despite the fact that heterospecific sociality is widely prevalent in nature and is known to occur in many different contexts, in a variety of taxa and in both marine and terrestrial habitats. My dissertation focuses on one of the best-known examples of heterospecific sociality, namely heterospecific foraging associations of terrestrial insectivorous birds (commonly called mixed-species flocks; flocks hereon). Flocks are found all over the world, in a variety of habitats, and include a unique suite of species in each area. Although flocks are the best-studied among different types of heterospecific associations known, most of the research has been descriptive. In other words, while we have descriptions of the richness, size and composition of flocks from many areas, we know little about why and how flocks form. Therefore, one broad objective of this dissertation was to better understand the causes of flock formation. Birds are thought to join flocks either to obtain direct foraging benefits or better protection from predators. I used different approaches to understand which of these two reasons is likely to be more important. Additionally, to situate heterospecific sociality within sociality in general, I also examined whether the mechanisms underlying flocks are similar to single-species groups or unique to a mixed-species condition. In other words, are flocks a different route to obtain the same benefits as single-species groups, or a route to obtain benefits that conspecifics cannot provide? In the previous section, I described why understanding flocks is important from the point of view of sociality. Flocks are also important for the consequences they might have on community organization. While each flock is a collection of interacting individuals of different species, populations of species are linked in a network of interactions across multiple flocks in an area. The theoretical framework guiding our understanding of community organization is currently dominated by the idea of interspecific competition. Mutualisms or positive interactions, if any, are only expected to occur between members of different trophic levels. Only recently has the need to incorporate positive interactions between members of the same trophic level into ecological theory been emphasized. Flocks and other heterospecific associations present an important example of within-trophic level positive interactions. Therefore, the second broad objective of this dissertation was to examine the consequences of flock formation on community organization. The objectives described above were addressed using multiple approaches in this dissertation. “Why” questions in ecology and evolution are generally tackled using manipulative experiments. Given that experiments are neither feasible nor ethical in the case of multispecies flocks in the wild, I addressed the “why” question by focusing on “who” instead. To understand “why” flocks form, I examined what kinds of species participate in flocks, who associates with whom, and who provides and who receives the benefits. This dissertation uses a combination of methods and data, including questions addressed at a global scale using descriptive information on flocks available from across the world. For other questions that were based on entirely new approaches developed in this dissertation, data were collected in a field site in the Western Ghats. In general, given the two broad objectives of this dissertation, the approaches I used were drawn from both behavioural and community ecology. The dissertation is organized in the following way: Chapter 1: General Introduction The first chapter provides the background to the two broad objectives of this thesis, namely understanding the causes and consequences of heterospecific foraging associations in terrestrial bird communities. Chapter 2 (published in The American Naturalist) Based on a global dataset (55 presence-absence matrices from 24 locations in multiple continents) on the composition of flocks, I asked if flocks largely consist of ecologically similar or dissimilar species. Using null models and randomization tests followed by meta-analysis, I found that the association strength of species in flocks was strongly related to similarity in body size and foraging behaviour, and higher for congeneric compared with non-congeneric species pairs. In other words, flocks seem to consist largely of similar species. Extending group-living and social information use theory to a heterospecific context, I discuss potential behavioral mechanisms leading to positive interactions among similar species in flocks as well as ways in which competition costs are reduced. These findings highlight the need to consider positive interactions along with competition when seeking to explain community organisation. Chapter 3 (published in Animal Behaviour) Two kinds of participants are recognized in flocks: those that join other species (‘followers’) and are therefore likely to be the recipients of the benefits of flock participation and those that are joined (‘leaders’). Through comparative analyses, using a large sample of flocks from around the world, I found (1) ‘followers’ tend to be smaller, more insectivorous, and feed in higher strata than matched species that participate in flocks to a lesser extent and (2) ‘leaders’ tend to be cooperative breeders (which are known to have well-developed anti-predatory systems) more often than matched species that are not known to lead flocks. Furthermore, meta-analyses of published results from across the world showed that bird species in terrestrial mixed-species flocks increased foraging rates and reduced vigilance compared to when they were solitary or in conspecific groups. Moreover, the increase in foraging rates was seen only in the case of flock followers and not flock leaders. These findings suggest a role for predation in the evolution of mixed-species flocking. Species that are vulnerable to predation follow species whose vigilance they can exploit. By doing so, they are able to reduce their own vigilance and forage at higher rates. Chapter 4 (to be submitted to Behavioural Ecology and Sociobiology) In this study, conducted in a tropical evergreen forest in the Western Ghats of India, I used intraflock association patterns to generate a community-wide assessment of benefits of flock participation for different species. I assumed that individuals needed to be physically proximate to particular heterospecific individuals within flocks to obtain any direct foraging benefit (flushed prey, kleptoparasitism, copying foraging locations). Alternatively, for all anti-predation benefits, physical proximity to particular heterospecifics is not required, i.e. just being in the flock vicinity will suffice. Therefore, I used the choice of locations within flocks to infer whether individual species are obtaining direct foraging or antipredation benefits. A small subset of the bird community (5/29 species), composed of all members of the sallying guild, showed non-random physical proximity to heterospecifics within flocks. All preferred associates were from non-sallying guilds, suggesting that the sallying species were likely obtaining direct foraging benefits, either in the form of flushed or snatched prey. The majority of species (24/29) chose locations randomly with respect to heterospecifics within flocks, and thus were likely obtaining anti-predation benefits. In summary, my study indicates that direct foraging benefits are important for only a small proportion of species in flocks and therefore that predation is likely to be the main driver of flocking. Chapter 5 (to be submitted to Oecologia) Two types of species – intraspecifically gregarious and sallying species – are thought to play important roles in flocks because studies have shown they attract other flock participants. However, it is not clear why these types of species are attractive, i.e. are they cues for flocks or do they directly provide benefits to other species? It is also not known whether these types are essential for flock formation. In this study, in a tropical evergreen forest in the Western Ghats of India, I used a novel approach to address these questions. Flocks contain anywhere between two and tens of species. My approach focused on the simplest, i.e. two-species, flocks. In two-species flocks, attraction between species must be based on direct benefit to at least one of the species. Therefore, only species combinations that result in benefit to at least one species will occur as two species flocks. Further, by examining leading and following behaviour in these combinations, I could also determine the direction of benefit-flow. Finally, given that all flocks pass through a two-species step, examining which two-species combinations are joined by other species helped understand species roles in flock formation. I found that intraspecifically gregarious species, but not sallying species, were disproportionately represented in two-species flocks, always provided the benefits when present, and that flocks containing them were joined significantly more often. Therefore, intraspecifically gregarious species, but not sallying species, play a role in providing flock benefits and flock formation. Moreover, given that most (7/8) intraspecifically gregarious species in my study site played these roles, the intraspecifically gregarious species guild can be considered an example of a within-trophic level “keystone”. Chapter 6 (“in press” in Oikos) There is a growing recognition of the need to integrate non-trophic interactions into ecological networks for a better understanding of whole-community organization. To achieve this, the first step is to build networks of individual non-trophic interactions. In this study, I analyzed the network of interdependencies among bird species that participated in flocks in an evergreen forest site in the Western Ghats, India. I found that flock networks contain a small core of highly important species that other species are strongly dependent on, a pattern seen in many other biological networks. Further, I found that structural importance of species in the network was strongly correlated to functional importance of species at the individual flock level. Finally, comparisons with flock networks from other Asian forests showed that the same taxonomic groups were important in general, suggesting that species importance was an intrinsic trait and not dependent on local ecological conditions. Hence, given a list of species in an area, it may be possible to predict which ones are likely to be important. Chapter 7: Conclusions In this section, I provide a summary of the main findings of this dissertation, discuss the main conclusions with regard to each of the two broad objectives and finally suggest future lines of investigation to further understand the causes and consequences of flock formation and heterospecific sociality in general. In summary, the work presented in this dissertation provides a picture of how and why flocks form and their likely consequences for community organization. The main driver of flock formation seems to be protection from predators, although a few species do obtain direct foraging benefits. Additionally, given that I found flocks to largely be groupings of similar species, the mechanisms through which benefits are obtained are likely to be similar to those in single-species groups. In other words, flocks are possibly a way for species that are unable to group with conspecifics, to obtain group-living benefits. My finding, that flocks largely consisted of similar species, also questions the stereotyping of interactions between similar species in communities as competitive. Other results from this study demonstrate the importance of intraspecifically gregarious species in flocks. Such species are the main providers of benefits to other flock participants and seem essential for flock formation. Finally, network analysis showed that a small core of species, mainly including species known to provide benefits at the individual flock level, are disproportionately important in the emergent community-level network of interdependencies. Such species are possibly one of the first examples of within-trophic level “keystones” discovered. Apart from the findings, this study also provides a set of new approaches and analytical frameworks that can be used to examine other multispecies foraging groups and heterospecific foraging associations in other contexts.

Les herbivores consomment les tissus végétaux et affectent la survie et la croissance des végétaux. Les ennemis naturels des herbivores peuvent réguler leurs populations et indirectement diminuer les dommages sur la végétation. Les relations entre plantes, herbivores et prédateurs sont liées aux caractéristiques intrinsèques des espèces végétales mais aussi influencées par la communauté végétale environnante. L'hypothèse de résistance par association stipule qu'une communauté végétale diversifiée entraîne une diminution des dégâts sur une plante cible par dilution, répulsion des herbivores et/ou favorisation des ennemis naturels. Inversement, les herbivores peuvent se concentrer sur la plante-cible dans une communauté végétale diversifiée (susceptibilité par association). L'objectif de cette thèse est d'évaluer 1) l'influence de la composition de la communauté végétale sur les herbivores et leurs dommages sur une plante-cible, 2) l'impact de la régulation des herbivores par les prédateurs et 3) les interactions entre herbivores, communauté végétale et prédateurs, et les conséquences pour les intensités d'herbivorie sur la plante-cible. Nous nous sommes focalisés sur les dommages causés par les insectes herbivores sur des plantules d'essences forestières feuillues (Betula pendula, Quercus robur et Q. ilex). La variété importante de communautés végétales, au niveau inter (essences dominantes) et intraparcellaire (présence, structure et composition), nous a permis de tester les effets du voisinage végétal sur les insectes herbivores et leurs dommages sur les plantules cibles. Les oiseaux insectivores sont les principaux prédateurs d'insectes et nous avons cherché à estimer les effets de leur exclusion sur les insectes et les conséquences pour les plantules, et ceci le long des gradients de composition du voisinage végétal. Enfin, le degré de spécialisation (spécialiste/généraliste) et le mode de vie (exo/endophyte) ont été caractérisés car ils conditionnent les réponses des insectes herbivores à la végétation voisine ainsi que la prédation avienne. Le voisinage végétal des plantules-cibles modifie la charge en herbivores et les dommages engendrés. Ces réponses sont liées à la spécialisation de l'insecte herbivore : la colonisation des plantules par les espèces spécialistes (mineuses de feuilles) est négativement affectée par la structure de la végétation locale (présence, recouvrement arbustif). Les dommages d'insectes généralistes augmentent avec la richesse spécifique des peuplements et dans les parcelles dominées par des espèces conspécifiques. Les abondances des insectes externes et leurs dommages sont régulés par les oiseaux insectivores mais cet effet indirect dépend de l'espèce de plantule-cible et de la composition de la communauté végétale. Les effets de cascade trophique sont les plus intenses dans les parcelles de pin maritime où les plantules constituent une ressource nouvelle pour les niveaux trophiques supérieurs. Plus localement, le voisinage végétal modifie la sensibilité des arthropodes à la prédation: les oiseaux insectivores bénéficient aux plantules lorsque les proies sont plus accessibles (végétation environnante supprimée).Pour conclure, une des principales richesses de ces résultats vient de la mise en évidence d'interactions très fortes existant entre les effets des oiseaux insectivores et la végétation environnante. Les effets indirects des prédateurs et directs de la végétation environnante sont très liés au degré de spécialisation des insectes phytophages et à leur mode de vie. / Herbivory is thought to depend on intrinsic plant resistance traits and negatively affect survival and growth of plants. Predators can depress herbivore populations and thereby indirectly limit the consumption of primary producers. The surrounding vegetation is also expected to modify the interactions between a focal plant, its herbivores and their natural enemies. The surrounding vegetation may disrupt the colonisation and the consumption of a focal plant by insect herbivores, providing associational resistance. Increasing diversity of neighbouring vegetation can decrease resource availability or enhance herbivores control by predators. By contrast, associational susceptibility may occur with higher levels of herbivory in a focal plant in more diverse plant communities. We tested 1) the effect of the neighbouring vegetation on the levels of herbivory on seedlings, 2) the indirect impact of predators on herbivores and subsequent herbivory, and 3) the interactions between surrounding vegetation, focal plants, herbivores and predators. We examined the damage caused by insect herbivores on three native broadleaved species. We then compared the importance of taxonomic similarity between seedlings and canopy tree species and the structure and composition of surrounding vegetation at both large- and small-scales. Birds are vertebrate predators likely to limit damage to plants and facilitate plant growth by consuming herbivorous insects. We estimated their effects by excluding them from focal plants, and compared their interactive effects with vegetation diversity or removal. The effects of surrounding vegetation and avian predation were also disentangled on concealed- vs. external-feeding guilds (predation sensitivity) and specialist- vs. generalist-insects.Surrounding vegetation of focal seedlings influences the abundance of insect herbivores and damage. Specialist insects (leaf miners) are affected by the understorey vegetation close to oak seedlings, decreasing with its presence or structural diversity. The damage caused by generalist insects depend on the large-scale composition of plant communities, and increase with tree species richness and cover or on conspecific forest habitats. Exclusion of insectivorous birds affects insect herbivory in a species-specific manner, and also greatly varies with forest habitats and presence of vegetation around focal plants. The indirect effect of bird predation on leaf damage is observed on seedlings beneath noncongeneric canopy trees. The removal of local surrounding vegetation affects the top-down effect of insectivorous birds on insect herbivores, according to their sensitivity to predation and the accessibility of preys. To conclude, our studies provide experimental evidences of interactive effects between bird predation and neighbouring vegetation on insect herbivores and levels of herbivory. These indirect effects are strongly related to the specialisation of herbivores and to their sensitivity to bird predation.

The thesis describes diversity of birds along a complete altitudinal gradient and in forest fragments in lowlands of Papua New Guinea. It focuses separately on the diversity of different feeding guilds, and discusses their links to habitat and food resources. More specifically, it focuses on forest insectivorous birds, their predation pressure on arthropods, feeding specializations and preferences, and some of the ways how insectivores search for food.

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