Early post-fledgling survival in a fragmented population of a tropical cooperative breeding passerine

During the last years tropical forest has been a target of intense study especially due to its recent big scale destruction. Although a lot still needs to be explored, we start realizing how negative can the impact of our actions be for the ecosystem. Subsequently, the living community have been developing strategies to overcome this problem avoiding bottlenecks or even extinctions. Cooperative breeding (CB) has been recently pointed out as one of those strategies. CB is a breeding system where more than two individuals raise one brood. In most of the cases, extra individuals are offspring that delay their dispersal and independent breeding what allows them to help their parents raising their siblings in the subsequent breeding season. Such behavior is believed to be due, per example, to the lack of mates or breeding territories (ecological constraints hypothesis), a consequence of habitat fragmentation and/or disturbance. From this point, CB is easily promoted by a higher reproductive success of group vs pairs or single individuals. Accordingly, during this thesis I explore the early post-fledging survival of a cooperative breeding passerine, namely the impact of individual/habitat quality in its survival probability during the dependence period of the chicks. Our study species is the Cabanis’s greenbul (Phyllastrephus cabanisi), a medium-sized, brownish passerine, classified within the Pycnonotidae family. It is found over part of Central Africa in countries such as Angola, Democratic Republic of the Congo, Mozambique and Kenya, inhabiting primary and secondary forests, as well as woodland of various types up to 2700m of altitude. Previous studies have concluded that PC is a facultative cooperative breeder. This study was conducted in Taita Hills (TH) at the Eastern Arc Mountains (EAM), a chain of mountains running from Southeast Kenya to the South of Tanzania. TH comprises an area of 430 ha and has been suffering intense deforestation reflecting 98% forest reduction over the last 200 years. Nowadays its forest is divided in fragments and our study was based in 5of those fragments. We access the post-fledging survival through radio-telemetry. The juvenile survey was done through the breeding females in which transmitters were placed with a leg-loop technique. Ptilochronology is consider to be the study of feather growth bars and has been used to study the nutritional state of a bird. This technique considers that the feather growth rate is positively proportional to the individual capability of ingesting food and to the food availability. This technique is therefore used to infer for individual/habitat quality. Survival was lowest during the first 5 days post-fledging representing 53.3%. During the next 15 days, risk of predation decreased with only 14.3% more deceased individuals. This represents a total of only 33% survived individuals in the end of the 50 days. Our results showed yet a significant positive relationship between flock size and post-fledging survival as well as between ptilochronology values and post-fledgling survival. In practice, these imply that on this population, as bigger the flock, as greater the post fledging survival and that good habitat quality or good BF quality, will lead to a higher juvenile survival rate. We believe that CB is therefore an adaptive behaviour to the lack of mates/breeding territory originated from the mass forest destruction and disturbance. Such results confirms the critical importance of habitat quality in the post-fledging survival and, for the first time, demonstrates how flock size influences the living probability of the juveniles and therefore how it impacts the (local) population dynamics of this species. In my opinion, future research should be focus in disentangle individual and habitat quality from each other and verify which relationship exist between them. Such study will allow us to understand which factor has a stronger influence in the post-fledging survival and therefore redirect our studies in that direction. In order to confirm the negative impact of human disturbance and forest fragmentation, it would be of major relevance to compare the reproductive strategies and reproductive success of populations living in intact forests and disturbed patches.

The pre-reproductive period in a tropical bird: parental care, dispersal, survival, and avian life histories

The period between offspring birth and recruitment into the breeding population is considered one of the least understood components of animal life histories. Yet, examining this period is essential for studies of parental care, dispersal, demography, and life histories. Studies of the pre-reproductive period are particularly few in tropical regions, where the organization of life histories are predicted to differ compared to northern hemisphere species. For my dissertation I used radio-telemetry, mark-resighting, and field observations to study the pre-reproductive period in a Neotropical bird, the western slaty-antshrike (Thamnophilus atrinucha), in Panama. First, I found that parental care after offspring left the nest (the post-fledging period) was greater than care during the nestling period. Prolonged care resulted in a clear trade-off for parents as they did not nest again until fledglings from the first brood were independent. Parents fed offspring for a prolonged duration during the post-fledging period and higher post-fledging survival was observed compared to many northern hemisphere species. Second, I observed that offspring that remained with parents for longer periods on the natal territory had higher survival both while on the natal territory and after dispersal compared to those dispersing earlier. Parental aggression towards offspring increased with offspring age and offspring dispersed earlier when parents renested. Contrary to other family living species, only a small proportion of antshrike offspring remained on the natal territory until the following year and all dispersed to float. Floating is when juveniles wander within other breeding pairs’ territories. These results suggest that the benefits of delayed dispersal declined with offspring age and with renesting by parents. Third, I observed that survival during the dependent period and first year was greater in slaty antshrikes compared to that of northern hemisphere species. Pre-reproductive survival relative to adult survival was equal or greater than that observed in northern hemisphere species. The date offspring left the nest, mass, and age at dispersal influenced offspring survival, whereas offspring sex and year did not. Relatively high survival during the pre-reproductive period coupled with comparatively low annual productivity clarifies how many tropical species achieve replacement. High juvenile survival appears to obtain from extended post-fledging parental care, delayed dispersal, low costs of dispersal, and a less seasonal environment. Lastly, I experimentally manipulated begging at the nest to examine changes in parental behavior. Under elevated begging, parents increased provisioning rates and reduced the time between arrival to the nest and feeding of nestlings, potentially to reduce begging sounds. Furthermore, parents switched to preferentially feed the closest offspring during the begging treatment. This suggests parents either allowed sibling competition to influence feeding decisions, or feeding the closer nestling increased the efficiency of provisioning. In summary, I found that slaty antshrikes have delayed age at reproduction, higher post-fledging and first year survival, extended post-fledging parental care, equal or greater pre-reproductive survival relative to adult survival, and delayed dispersal compared to many northern hemisphere passerines. These results suggest that this tropical species has a strategy of high investment into few offspring. Furthermore, reproductive effort is equal or greater at least in slaty antshrikes compared to northern hemisphere species, suggesting that the latitudinal gradient in clutch size is not explained by a gradient in reproductive effort.

Seasonal differences in migration patterns of a soaring bird in relation to environmental conditions: a multi-scale approach

Many studies suggest that migratory birds are expected to travel more quickly during spring, when they are en route to the breeding grounds, in order to ensure a high-quality territory. Using data recorded by means of Global Positioning System satellite tags, we analysed at three temporal scales (hourly, daily and overall journey) seasonal differences in migratory performance of the booted eagle (Aquila pennata), a soaring raptor migrating between Europe and tropical Africa, taking into account environmental conditions such as wind, thermal uplift and day length. Unexpectedly, booted eagles showed higher travel rates (hourly speed, daily distance, overall migration speed and overall straightness) during autumn, even controlling for abiotic factors, probably thanks to higher hourly speeds, more straight routes and less non-travelling days during autumn. Tailwinds were the main environmental factor affecting daily distance. During spring, booted eagles migrated more quickly when flying over the Sahara desert. Our results raise new questions about which ecological and behavioural reasons promote such unexpected faster speeds in autumn and not during spring and how events occurring in very different regions can affect migratory performance, interacting with landscape characteristics, weather conditions and flight behaviour.

Adaptation to urban environments promotes high reproductive success in the tawny frogmouth (Podargus strigoides), an endemic nocturnal bird species

The tawny frogmouth (Podargus strigoides) is an endemic, nocturnal bird species widespread throughout Australia with high numbers in urban environments but very limited information exists on its ability to cope with urban systems. We investigated the reproductive behaviour of this species in response to a continuum of urbanisation. Firstly, we asked does the degree of urbanisation influence the availability of suitable nesting locations. Secondly, does broad and/or local scale factors surrounding a nest influence reproductive success? And finally, does the degree of urbanisation influence time of breeding? We located 158 nest sites and monitored 189 breeding attempts across four breeding seasons (2010-2013). One hundred and thirty three of these attempts successfully fledged 177 chicks. We categorized 75 attempts as first known attempts for that season resulting in 77 fledged chicks. In some pairs, repeated attempts occurred after failure. We observed double brooding by three pairs, a strategy largely unknown in this species. Spatial modelling revealed that birds selected more vegetated areas of the gradient within which to locate their nests, avoiding more highly urbanized areas. We identified no association between land-use type and breeding success at both broad and local scales. Birds selected native rough-barked Eucalyptus tree species more frequently than other trees for nesting. The key drivers for the commencement of breeding were climatic variables, in particular rainfall. The ability of the species to synchronize breeding attempts to coincide with optimal environmental conditions resulted in extremely high reproductive success suggesting high individual fitness and an adaptation to local environmental conditions.

Little raven (Corvus mellori) as an egg predator of other bird species

 Little ravens were major predators of ground-nesting red-capped Plover and burrow-nesting Little penguin eggs. Ravens also congregated at the breeding sites of these two species to exploit eggs. As raven numbers increase with human habitation of land, their impact on other bird species needs to be monitored and managed.

An obligate beach bird selects sub-, inter- and supra-tidal habitat elements

Few habitat models are available for widespread, obligate, high-energy sandy shore vertebrates, such as the Eastern Hooded Plover Thinornis cucullatus cucullatus. We examined habitat attributes which determined the difference between sites where plovers breed and randomly-selected absence sites (determined from long-term systematic monitoring). A variety of habitat variables were derived from aerial photography and bathymetric and terrestrial Light Detection And Ranging (LiDAR) data. Logistic regression against eight candidate variables, in a model selection framework, revealed considerable support for four variables with respect to explaining the presence of breeding territories. In particular, the amount of unvegetated dune and foredune which was unvegetated, and the amount of intertidal and sub-tidal reef were positively associated with the presence of breeding territories. Thus, plovers apparently select certain habitat in which to breed, involving sub-tidal, intertidal and supra-tidal habitat elements. The model also helps explain the virtual absence of breeding plovers from long sections of superficially suitable habitat, such as the fourth longest continuous beach in the world.

Impact of nest sanitation on the immune system of parents and nestlings in a passerine bird

Bacterial communities are thought to have fundamental effects on the growth and development of nestling birds. The antigen exposure hypothesis suggests that, for both nestlings and adult birds, exposure to a diverse range of bacteria would select for stronger immune defences. However, there are relatively few studies that have tested the immune/bacterial relationships outside of domestic poultry. We therefore sought to examine indices of immunity (microbial killing ability in naïve birds, which is a measure of innate immunity and the antibody response to sheep red blood cells, which measures adaptive immunity) in both adult and nestling zebra finches (Taeniopygia guttata). We did this throughout breeding and between reproductive attempts in nests that were experimentally manipulated to change the intensity of bacterial exposure. Our results suggest that nest sanitation and bacterial load affected measures of the adaptive immune system, but not the innate immune parameters tested. Adult finches breeding in clean nests had a lower primary antibody response to sheep red blood cells (SRBC), particularly males, and a greater difference between primary and secondary responses. Adult microbial killing of E.coli decreased as parents moved from incubation to nestling rearing for both nest treatments; however, killing of C.albicans remained consistent throughout. In nestlings, both innate microbial killing and the adaptive antibody response did not differ between nest environments. Together, these results suggest that the exposure to microorganisms in the environment affect the adaptive immune system in nesting birds, with exposure upregulating the antibody response in adult birds.

Defining marine important bird areas: testing the foraging radius approach

Recent international initiatives have promoted a number of different approaches to identify marine Important Bird and biodiversity Areas (IBAs), which are important areas for foraging, migrating or over-wintering seabirds. The 'Foraging Radius Approach' is one of these and uses known foraging range and habitat preferences to predict the size and location of foraging areas around breeding colonies. Here we assess the performance of the Foraging Radius Approach using GPS tracking data from six seabird species with a variety of foraging modes. For each species we compared the population home-range areas of our six study species with the home-range areas defined using the Foraging Radius Approach. We also assessed whether basic information on depth preferences from tracking data could improve these home-range area estimates. Foraging Radius Approach home-range areas based on maximum foraging radii encompassed the entire population home-range of five out of six of our study species but overestimated the size of the population home-range area in every case. The mean maximum foraging radius overestimated the population home-range areas by a factor of 4-14 for five of the six species whilst the mean foraging radius overestimated the population home-range area for half of the species and underestimated for the rest. In the absence of other data, the Foraging Radius Approach appears to provide a reasonable basis for preliminary marine IBA identification. We suggest that using the mean value of all previously reported maximum foraging radii, informed by basic depth preferences provides the most appropriate prediction, balancing the needs of seabirds with efficient use of marine space.

Placement of re-nests following predation: are birds managing risk?

Nest predation is the most important source of reproductive failure for many bird species, thus placing nests in 'safe' locations that minimise predation risk is paramount to maximising fitness. After a nest predation event, some species have been shown to manage the risk of nest predation for subsequent re-nesting attempts by moving to a new location, placing re-nests in areas with increased cover, or changing the height above ground at which the re-nest is placed. The extent to which this is an adaptive behaviour for birds in general is not yet clear, as existing studies are almost exclusively restricted to northern hemisphere species and species that do not breed cooperatively. Here, we examined the re-nesting behaviour of Bell Miners (Manorina melanophrys), a species of honeyeater endemic to Australia that is both multi-brooded and also frequently re-nests soon after nesting failure; females can build up to five nests in a breeding season. We tested if these females managed within-season nest predation risk by changing nest site characteristics (height from ground and distance between nests) between successive nesting attempts. We found that female miners did indeed manage predation risk by reducing the height from the ground at which they placed re-nests following predation events, but contrary to our second prediction we found no difference in distances moved to re-nest after females experienced nest predation or successfully rearing young.

Testing hypotheses about the function of repeated nest abandonment as a life history strategy in a passerine bird

Nest structures are essential for successful reproduction in most bird species. Nest construction costs time and energy, and most bird species typically build one nest per breeding attempt. Some species, however, build more than one nest, and the reason for this behaviour is often unclear. In the Grey Fantail Rhipidura albiscapa, nest abandonment before egg-laying is very common. Fantails will build up to seven nests within a breeding season, and pairs abandon up to 71% of their nests before egg-laying. We describe multiple nest-building behaviour in the Grey Fantail and test four hypotheses explaining nest abandonment in this species: cryptic depredation, destruction of nests during storm events, and two anti-predatory responses (construction of decoy nests to confuse predators, and increasing concealment to 'hide' nests more effectively). We found support for only one hypothesis - that abandonment is related to nest concealment. Abandoned nests were significantly less concealed than nests that received eggs. Most abandoned nests were not completely built and none received eggs, thus ruling out cryptic predation. Nests were not more likely to be abandoned following storm events. The decoy nest hypothesis was refuted as abandoned nests were constructed at any point during the breeding season and some nests were dismantled and the material used to build the subsequent nest. Thus, Grey Fantails are flexible about nest-site locations during the nest-building phase and readily abandon nest locations if they are found to have deficient security.

Conserving long unburnt vegetation is important for bird species, guilds and diversity

Landscape-level wildfires have a major role in structuring faunal assemblages, particularly in fire-prone landscapes. These effects are mediated by changes to vegetation structure and composition that directly influence the availability of shelter, feeding and breeding resources. We investigated the response of a semi-arid shrubland bird community in Western Australia to the prevailing fire regime by examining the abundance, diversity and guild structure in relation to time since fire. We also examined vegetation structural attributes in relation to time since fire. We surveyed 32 sites ranging in age from 12 to 84 years since last fire. A total of 845 birds from 40 species were recorded. Vegetation structure varied with fire history with old and very old sites characterised by less bare ground, more leaf litter cover and greater canopy cover. Bird community composition varied with time since fire, driven by increased bird species richness and abundance of insectivores, granivores/frugivores, golden whistlers, grey shrike-thrush and red-capped robins with time since fire. Frequent, intense landscape-scale fires transform the landscape into homogeneous young shrublands, which may render vegetation unsuitable for several species and guilds.

Bonding beyond the pair in a monogamous bird: Impact on social structure in adult rooks (Corvus frugilegus)

The formation of social bonds outside the mated pair is not frequently reported in monogamous birds, although it may be expected in some species like rooks, living in groups all year round. Here we explore the social structure of captive adult rooks over three breeding seasons. We recorded proximities and affiliations (i.e., allofeeding, allopreening, contact-sit) to classify relationships according to their strength. Three categories of relationships emerged: primary (i.e., pairs), secondary and weak relationships. Affiliations and sexual behaviours were not restricted to pairs, and secondary relationships were clearly recognizable. Mixed-sex secondary relationships were qualitatively equivalent to pairs (i.e., same behaviours in the same proportions), although they were quantitatively less intense. Same sex pairs occurred, and were qualitatively equivalent to mixedsex pairs. Overall we found that rooks social structure is more than just an aggregation of pairs, which highlights the importance of considering extra-pair relationships in socially monogamous birds.

Exploring nest destruction and bird mortality in mown Mediterranean dry grasslands: an increasing threat to grassland bird conservation

In recent years, haying has extended to Iberian Mediterranean dry grasslands potentially impacting on grassland bird ecology. We evaluated the impact of haying on a grassland bird community of South Portugal. Our main goals were: (1) to investigate the exposure of different species to haying, (2) to investigate potential removal of nests and dead birds from hayed fields by haying machinery using the ratio (REC) between the expected number of records and the number of records collected and (3) to link clutch destruction and bird mortality with haying management practices. Hayed fields were surveyed for signs of breeding and birds censused prior to mowing. Linear models were computed, linking the REC with haying machinery and sward properties. GLMs and model averaging were used to obtain models linking clutch destruction, bird mortality and haying management variables. Only 4 % of records evidenced successful nesting attempts (N = 177). REC evaluation suggested high nest or dead bird removal by the machinery, particularly in fields with lower vegetation biomass prior to cutting. Sickle bar mowers and one-rotor rotary rakes returned higher REC but lower probability of found nests removed from the original nesting sites comparatively to discs mowers and wheel rakes. Higher probabilities of mortality events were found in fields mown earlier (but not in all years). On the other hand, lower mortality was found in fields raked with two-rotor rotary rakes. Delayed haying, silage production in temporary crops and the use haying machinery enabling simultaneously mowing and gathering hay in lines are discussed as management alternatives.

Between and within-year effects of haying on grassland bird

In recent years, haying has extended to Iberian Mediterranean dry grasslands potentially threatening grassland birds. We evaluate the between and within-year effects of haying on grassland birds in Alentejo region, Portugal. Our main goals were: (1) to investigate variations on bird abundance and species richness in the fields hayed, with respect to past haying events occurred in a field and its surroundings and (2) to investigate the shifts in bird abundance, species richness and spatial dynamics resulting from haying a field and its surrounding area in a given year. We conducted grassland bird censuses during the breeding season through point counts from 2012 to 2015. The relationship between bird abundance/richness and past haying events was investigated using Generalized Linear Models whereas within-year effects of haying were analysed using Generalized Additive Models. Bird abundance in a field was positively related with the surface hayed in the vicinity of that field in the previous year. However, contrasting yearly effects were found for non passerines. Also, some species prefer fields with less haying events or surface hayed, whereas others occur mostly in fields frequently managed for haying. Haying a field leads, in the short term, to its abandonment by birds, and thus to a decrease in bird abundance and, for some species, to spatial concentration in surrounding fields offering suitable habitat. We conclude that within-year effects of haying have higher impact on grassland birds than between-year effects. Maintaining haying at low levels by rotating haying yearly through the different fields in each farm and using partial haying may be an adequate way to ensure an effective management of grassland bird populations.