Divergence with gene flow and fine-scale phylogeographical structure in the wedge-billed woodcreeper, Glyphorynchus spirurus, a Neotropical rainforest bird.

Determining the relative roles of vicariance and selection in restricting gene flow between populations is of central importance to the evolutionary process of population divergence and speciation. Here we use molecular and morphological data to contrast the effect of isolation (by mountains and geographical distance) with that of ecological factors (altitudinal gradients) in promoting differentiation in the wedge-billed woodcreeper, Glyphorynchus spirurus, a tropical forest bird, in Ecuador. Tarsus length and beak size increased relative to body size with altitude on both sides of the Andes, and were correlated with the amount of moss on tree trunks, suggesting the role of selection in driving adaptive divergence. In contrast, molecular data revealed a considerable degree of admixture along these altitudinal gradients, suggesting that adaptive divergence in morphological traits has occurred in the presence of gene flow. As suggested by mitochondrial DNA sequence data, the Andes act as a barrier to gene flow between ancient subspecific lineages. Genome-wide amplified fragment length polymorphism markers reflected more recent patterns of gene flow and revealed fine-scale patterns of population differentiation that were not detectable with mitochondrial DNA, including the differentiation of isolated coastal populations west of the Andes. Our results support the predominant role of geographical isolation in driving genetic differentiation in G. spirurus, yet suggest the role of selection in driving parallel morphological divergence along ecological gradients.

Assessment of Coastal Bird Populations and Habitats on the Finnish Coast of the Baltic Sea: Implications for Monitoring and Management

Coastal birds are an integral part of coastal ecosystems, which nowadays are subject to severe environmental pressures. Effective measures for the management and conservation of seabirds and their habitats call for insight into their population processes and the factors affecting their distribution and abundance. Central to national and international management and conservation measures is the availability of accurate data and information on bird populations, as well as on environmental trends and on measures taken to solve environmental problems. In this thesis I address different aspects of the occurrence, abundance, population trends and breeding success of waterbirds breeding on the Finnish coast of the Baltic Sea, and discuss the implications of the results for seabird monitoring, management and conservation. In addition, I assess the position and prospects of coastal bird monitoring data, in the processing and dissemination of biodiversity data and information in accordance with the Convention on Biological Diversity (CBD) and other national and international commitments. I show that important factors for seabird habitat selection are island area and elevation, water depth, shore openness, and the composition of island cover habitats. Habitat preferences are species-specific, with certain similarities within species groups. The occurrence of the colonial Arctic Tern (Sterna paradisaea) is partly affected by different habitat characteristics than its abundance. Using long-term bird monitoring data, I show that eutrophication and winter severity have reduced the populations of several Finnish seabird species. A major demographic factor through which environmental changes influence bird populations is breeding success. Breeding success can function as a more rapid indicator of sublethal environmental impacts than population trends, particularly for long-lived and slowbreeding species, and should therefore be included in coastal bird monitoring schemes. Among my target species, local breeding success can be shown to affect the populations of the Mallard (Anas platyrhynchos), the Eider (Somateria mollissima) and the Goosander (Mergus merganser) after a time lag corresponding to their species-specific recruitment age. For some of the target species, the number of individuals in late summer can be used as an easier and more cost-effective indicator of breeding success than brood counts. My results highlight that the interpretation and application of habitat and population studies require solid background knowledge of the ecology of the target species. In addition, the special characteristics of coastal birds, their habitats, and coastal bird monitoring data have to be considered in the assessment of their distribution and population trends. According to the results, the relationships between the occurrence, abundance and population trends of coastal birds and environmental factors can be quantitatively assessed using multivariate modelling and model selection. Spatial data sets widely available in Finland can be utilised in the calculation of several variables that are relevant to the habitat selection of Finnish coastal species. Concerning some habitat characteristics field work is still required, due to a lack of remotely sensed data or the low resolution of readily available data in relation to the fine scale of the habitat patches in the archipelago. While long-term data sets exist for water quality and weather, the lack of data concerning for instance the food resources of birds hampers more detailed studies of environmental effects on bird populations. Intensive studies of coastal bird species in different archipelago areas should be encouraged. The provision and free delivery of high-quality coastal data concerning bird populations and their habitats would greatly increase the capability of ecological modelling, as well as the management and conservation of coastal environments and communities. International initiatives that promote open spatial data infrastructures and sharing are therefore highly regarded. To function effectively, international information networks, such as the biodiversity Clearing House Mechanism (CHM) under the CBD, need to be rooted at regional and local levels. Attention should also be paid to the processing of data for higher levels of the information hierarchy, so that data are synthesized and developed into high-quality knowledge applicable to management and conservation.

Foraging ecology and parental behaviour in the common tern (Sterna hirundo)

One component of successful parenting is related to efficiency in foraging behaviour. The relationships among chick feeding, the size and type of food package, and length of parental foraging trips has not been well studied in seabirds. In addition, relatively few data have been collected on the activities of seabirds when foraging away from the nest site. The objectives of this study were: (1) to contrast productivity, feeding rate, and attendance patterns of individuals carrying a novel transmitter with a control group of birds; (2) to use radio-telemetry to assess the variability in foraging locations within and between individual male Common Terns; (3) to determine the seasonal variation in chick diet; (4) to determine for each transmittered bird, the relationships among the foraging patterns, parental behaviour, and seasonal reproductive success. The study took place over two years (1990-91) on a concrete breakwater 1 km offshore on Lake Erie near Port Colbome, Ontario. Ten pairs of terns in 1990 and 12 pairs in 1991 were radio-tracked by boat or car during the chick rearing stage. Concurrent behavioural observations documented the time each sex spent foraging or at the nest. The frequency and prey species composition of feeds to chicks were also recorded. The transmitters had negligible effects on the feeding frequency and brood attendance patterns of transmitter carrying birds. Peak nesting transmittered birds in 1990 and 1991 exhibited some inter-individual variability in foraging locations, however intraindividual variability was low. Birds foraged primarily to the west and northwest of the colony. Late nesters exhibited greater inter-individual variability, however intra-individual variability remained low for most birds. Neither group demonstrated sufficient variability to support the regular use of this colony as an "information centre". Individual transmittered birds had unique and predictable foraging patterns, and corresponding differences in feeding frequencies and brood attendance patterns, yet productivity was essentially equal between nests due to the impact and importance of stochastic events. Individuals that were recaptured in 1991 exhibited very similar foraging patterns to 1990, suggesting littie variability between years. Conservation of foraging patterns between years may have potential implications for mate choice decisions in future breeding seasons. Prey species delivered to chicks differed between morning and evening for peak and late nesters in 1990, but not 1991. Peak nesters in 1990 fed significantiy more Rainbow Smelt fOsmerus mordM) than Emerald Shiner (Notropis atherinoidesV this trend was reversed for late nesters who also fed large numbers of unidentified larval fish. No significant differences were found in 1991. Seasonal changes in prey species delivered to chicks is believed to be attributable to the temperature tolerances of the smelt and shiners, and the presence of large schools of larval fish during the late nesting season.

The Influence of Body Condition on the Stopover Ecology of Least Sandpipers in the Lower Mississippi Alluvial Valley during Fall Migration

Many shorebirds are long-distance migrants and depend on the energy gained at stopover sites to complete migration. Competing hypotheses have described strategies used by migrating birds; the energy-selection hypothesis predicts that shorebirds attempt to maximize energy gained at stopover sites, whereas the time-selection hypothesis predicts that shorebirds attempt to minimize time spent at stopover sites. The energy- and time-selection hypotheses both predict that birds in better condition will depart sites sooner. However, numerous studies of stopover duration have found little support for this prediction, leading to the suggestion that migrating birds operate under energy and time constraints for only a small portion of the migratory season. During fall migration 2002, we tested the prediction that birds in better condition depart stopover sites sooner by examining the relationship between stopover duration and body condition for migrating Least Sandpipers (Calidris minutilla) at three stopover sites in the Lower Mississippi Alluvial Valley. We also tested the assumption made by the Lower Mississippi Alluvial Valley Migratory Bird Science Team that shorebirds stay in the Mississippi Valley for 10 d. The assumption of 10 d was used to estimate the amount of habitat required by shorebirds in the Mississippi Valley during fall migration; a period longer than 10 d would increase the estimate of the amount habitat required. We used multiple-day constancy models of apparent survival and program MARK to estimate stopover duration for 293 individually color-marked and resighted Least Sandpipers. We found that a four-day constancy interval and a site x quadratic time trend interaction term best modeled apparent survival. We found only weak support for body condition as a factor explaining length of stopover duration, which is consistent with findings from similar work. Stopover duration estimates were 4.1 d (95% CI = 2.8–6.1) for adult Least Sandpipers at Bald Knob National Wildlife Refuge, Arkansas, 6.5 d (95% CI = 4.9–8.7) for adult and 6.1 d (95% CI =4.2–9.1) for juvenile Least Sandpipers at Yazoo National Wildlife Refuge, Mississippi, and 6.9 d (95% CI = 5.5–8.7) for juvenile Least Sandpipers at Morgan Brake National Wildlife Refuge, Mississippi. Based on our estimates of stopover duration and the assumption made by the Lower Mississippi Alluvial Valley Migratory Bird Science Team, there is sufficient habitat in the lower Mississippi Valley to support shorebirds during fall migration.

Population Change in a Marine Bird Colony is Driven By Changes in Recruitment

The population dynamics of long-lived birds are thought to be very sensitive to changes in adult survival. However, where natal philopatry is low, recruitment from the larger metapopulation may have the strongest effect on population growth rate even in long-lived species. Here, we illustrate such a situation where changes in a seabird colony size appeared to be the consequence of changes in recruitment. We studied the population dynamics of a declining colony of Ancient Murrelets (Synthliboramphus antiquus) at East Limestone Island, British Columbia. During 1990-2010, Ancient Murrelet chicks were trapped at East Limestone Island while departing to sea, using a standard trapping method carried on throughout the departure period. Adult murrelets were trapped while departing from the colony during 1990-2003. Numbers of chicks trapped declined during 1990-1995, probably because of raccoon predation, increased slightly from 1995-2000 and subsequently declined again. Reproductive success was 30% lower during 2000-2003 than in earlier years, mainly because of an increase in desertions. The proportion of nonbreeders among adult birds trapped at night also declined over the study period. Mortality of adult birds, thought to be mainly prebreeders, from predators more than doubled over the same period. Apparent adult survival of breeders remained constant during 1991-2002 once the first year after banding was excluded, but the apparent survival rates in the first year after banding fell and the survival of birds banded as chicks to age three halved over the same period. A matrix model of population dynamics suggested that even during the early part of the study immigration from other breeding areas must have been substantial, supporting earlier observations that natal philopatry in this species is low. The general colony decline after 2000 probably was related to diminished recruitment, as evidenced by the lower proportion of nonbreeders in the trapped sample. Hence the trend is determined by the recruitment decisions of externally reared birds, rather than demographic factors operating on the local breeding population, an unusual situation for a colonial marine bird. Because of the contraction in the colony it may now be subject to a level of predation pressure from which recovery will be impossible without some form of intervention.

Estimated Mortality of Selected Migratory Bird Species from Mowing and Other Mechanical Operations in Canadian Agriculture

Mechanical operations such as mowing, tilling, seeding, and harvesting are well-known sources of direct avian mortality in agricultural fields. However, there are currently no mortality rate estimates available for any species group or larger jurisdiction. Even reviews of sources of mortality in birds have failed to address mechanical disturbance in farm fields. To overcome this information gap we provide estimates of total mortality rates by mechanical operations for five selected species across Canada. In our step-by-step modeling approach we (i) quantified the amount of various types of agricultural land in each Bird Conservation Region (BCR) in Canada, (ii) estimated population densities by region and agricultural habitat type for each selected species, (iii) estimated the average timing of mechanical agricultural activities, egg laying, and fledging, (iv) and used these values and additional demographical parameters to derive estimates of total mortality by species within each BCR. Based on our calculations the total annual estimated incidental take of young ranged from ~138,000 for Horned Lark (Eremophila alpestris) to as much as ~941,000 for Savannah Sparrow (Passerculus sandwichensis). Net losses to the fall flight of birds, i.e., those birds that would have fledged successfully in the absence of mechanical disturbance, were, for example ~321,000 for Bobolink (Dolichonyx oryzivorus) and ~483,000 for Savannah Sparrow. Although our estimates are subject to an unknown degree of uncertainty, this assessment is a very important first step because it provides a broad estimate of incidental take for a set of species that may be particularly vulnerable to mechanical operations and a starting point for future refinements of model parameters if and when they become available.

Off-road sampling reveals a different grassland bird community than roadside sampling: implications for survey design and estimates to guide conservation

Grassland bird species continue to decline steeply across North America. Road-based surveys such as the North American Breeding Bird Survey (BBS) are often used to estimate trends and population sizes and to build species distribution models for grassland birds, although roadside survey counts may introduce bias in estimates because of differences in habitats along roadsides and in off-road surveys. We tested for differences in land cover composition and in the avian community on 21 roadside-based survey routes and in an equal number of adjacent off-road walking routes in the grasslands of southern Alberta, Canada. Off-road routes (n = 225 point counts) had more native grassland and short shrubs and less fallow land and road area than the roadside routes (n = 225 point counts). Consequently, 17 of the 39 bird species differed between the two route types in frequency of occurrence and relative abundance, measured using an indicator species analysis. Six species, including five obligate grassland species, were more prevalent at off-road sites; they included four species listed under the Canadian federal Species At Risk Act or listed by the Committee on the Status of Endangered Wildlife in Canada: Sprague’s Pipit (Anthus spragueii), Baird’s Sparrow (Ammodramus bairdii), the Chestnut-collared Longspur (Calcarius ornatus), and McCown’s Longspur (Rhynchophanes mccownii). The six species were as much as four times more abundant on off-road sites. Species more prevalent along roadside routes included common species and those typical of farmland and other human-modified habitats, e.g., the European Starling (Sturnus vulgaris), the Black-billed Magpie (Pica hudsonia), and the House Sparrow (Passer domesticus). Differences in avian community composition between roadside and off-road surveys suggest that the use of BBS data when generating population estimates or distribution models may overestimate certain common species and underestimate others of conservation concern. Our results highlight the need to develop appropriate corrections for bias in estimates derived from roadside sampling, and the need to design surveys that sample bird communities across a more representative cross-section of the landscape, both near and far from roads. 

How well do regional or national Breeding Bird Survey data predict songbird population trends at an intact boreal site?

A study to monitor boreal songbird trends was initiated in 1998 in a relatively undisturbed and remote part of the boreal forest in the Northwest Territories, Canada. Eight years of point count data were collected over the 14 years of the study, 1998-2011. Trends were estimated for 50 bird species using generalized linear mixed-effects models, with random effects to account for temporal (repeat sampling within years) and spatial (stations within stands) autocorrelation and variability associated with multiple observers. We tested whether regional and national Breeding Bird Survey (BBS) trends could, on average, predict trends in our study area. Significant increases in our study area outnumbered decreases by 12 species to 6, an opposite pattern compared to Alberta (6 versus 15, respectively) and Canada (9 versus 20). Twenty-two species with relatively precise trend estimates (precision to detect > 30% decline in 10 years; observed SE ≤ 3.7%/year) showed nonsignificant trends, similar to Alberta (24) and Canada (20). Precision-weighted trends for a sample of 19 species with both reliable trends at our site and small portions of their range covered by BBS in Canada were, on average, more negative for Alberta (1.34% per year lower) and for Canada (1.15% per year lower) relative to Fort Liard, though 95% credible intervals still contained zero. We suggest that part of the differences could be attributable to local resource pulses (insect outbreak). However, we also suggest that the tendency for BBS route coverage to disproportionately sample more southerly, developed areas in the boreal forest could result in BBS trends that are not representative of range-wide trends for species whose range is centred farther north.

Predicting origins of passerines migrating through Canadian migration monitoring stations using stable-hydrogen isotope analyses of feathers: a new tool for bird conservation

The Canadian Migration Monitoring Network (CMMN) consists of standardized observation and migration count stations located largely along Canada’s southern border. A major purpose of CMMN is to detect population trends of migratory passerines that breed primarily in the boreal forest and are otherwise poorly monitored by the North American Breeding Bird Survey (BBS). A primary limitation of this approach to monitoring is that it is currently not clear which geographic regions of the boreal forest are represented by the trends generated for each bird species at each station or group of stations. Such information on “catchment areas” for CMMN will greatly enhance their value in contributing to understanding causes of population trends, as well as facilitating joint trend analysis for stations with similar catchments. It is now well established that naturally occurring concentrations of deuterium in feathers grown in North America can provide information on their approximate geographic origins, especially latitude. We used stable hydrogen isotope analyses of feathers (δ²Hf) from 15 species intercepted at 22 CMMN stations to assign approximate origins to populations moving through stations or groups of stations. We further constrained the potential catchment areas using prior information on potential longitudinal origins based upon bird migration trajectories predicted from band recovery data and known breeding distributions. We detected several cases of differences in catchment area of species passing through sites, and between seasons within species. We discuss the importance of our findings, and future directions for using this approach to assist conservation of migratory birds at continental scales.

Shifts in resident bird communities associated with cloud forest patch size in Central Veracruz, Mexico

Avian communities in cloud forests have high levels of endemism and are at major risk given the accelerated rate of habitat fragmentation. Nevertheless, the response of these communities to changes in fragment size remains poorly understood. We evaluated species richness, bird community density, community composition, and dominance as indicators of the response to fragment size in a fragmented cloud forest landscape in central Veracruz, Mexico. Medium-sized fragments had statistically higher than expected species richness and more even communities, which may be a reflection of the intermediate disturbance hypothesis, in which medium-sized fragments are exploited by both forest and disturbance-associated species. Bird density also reached higher values in medium-sized fragments, which may indicate a carrying capacity in this habitat. However, large cloud forest fragments had a distinct taxonomic and functional composition, attributable to an increased number of understory insectivore species and canopy frugivores. By comparison, omnivorous species associated with human-altered habitats were more abundant in smaller fragments. Hence, although medium-sized cloud forest fragments had higher species richness and high bird density, large forest tracts maintained a distinct avian community composition, particularly of insectivorous and frugivorous species. Furthermore, the underlying response to fragmentation can only be properly addressed when contrasting several community attributes, such as richness, density, composition, and species dominance. Therefore, cloud forest conservation should aim to preserve the remaining large forest fragments to maintain comprehensive avian communities and avoid local extinctions.

Bird feeders and their effects on bird-window collisions at residential houses

Feeding wild birds creates an important link between homeowners and conservation. The effects of bird feeders and year-round feeding on birds have not been well studied, however, particularly in relationship to bird-window collisions. We determined effects of bird feeder presence and placement on bird-window collisions at residential homes. Paired month-long trials in which a feeder was either present or absent for one month and then removed or added for the second month were completed at 55 windows at 43 houses. In each trial, homeowners were asked to search their study window daily for evidence of a bird-window collision. During the study there were 51 collisions when there was no bird feeder and 94 when the feeder was present. The season when each trial was set up was the best individual predictor of bird-window collisions. The largest number of collisions was observed during fall migration and the lowest during the winter months. There were no collisions at 26 of the study windows. High variance was observed in the number of collisions at different houses, indicating that effects of bird feeders are context dependent. Changing the occurrence, timing, and placement of feeders can alter collision rates but is only one of many factors that influence whether a residential house is likely to have a bird window-collision or not.

Comparing the results of recall surveys and standardized searches in understanding bird-window collisions at houses

Every year a large number of birds die when they collide with windows. The actual number is difficult to ascertain. Previous attempts to estimate bird-window collision rates in Canada relied heavily on a prior citizen-science study that used memory-based surveys. Such an approach to data collection has many potential biases. We built upon this study and its recommendations for future research by creating a citizen-science program that actively searched for collision evidence at houses and apartments for an extended period with the objective to see how standardized approaches to data collection compared with memory recall. Absolute collision estimates as well as relative differences were compared between residence types in the two studies, and we found considerable differences in absolute values for collisions but similar rankings of collision rates between residence types. Collision recall rates in our study (56.5%) were very similar those in the prior 2012 study, where 50.5% of participants remembered a bird colliding with a window at some time in the past. Fatality estimates, however, were 1.4 times higher in the 2012 study than in our study based on standardized searches. Rural houses with a bird feeder consistently had the highest number of collisions. This suggests that memory recall surveys may be a useful tool for understanding the relative importance of different risk factors causing bird-window collisions.