Population Assessment of an Endangered Shorebird: the Piping Plover (Charadrius melodus melodus) in Eastern Canada

Small, at-risk populations are those for which accurate demographic information is most crucial to conservation and recovery, but also where data collection is constrained by logistical challenges and small sample sizes. Migratory animals in particular may experience a wide range of threats to survival and reproduction throughout each annual cycle, and identification of life stages most critical to persistence may be especially difficult for these populations. The endangered eastern Canadian breeding population of Piping Plover (Charadrius melodus melodus) was estimated at only 444 adults in 2005, and extensive effort has been invested in conservation activities, reproductive monitoring, and marking of individual birds, providing a comprehensive data set on population dynamics since 1998. We used these data to build a matrix projection model for two Piping Plover population segments that nest in eastern Canada in order to estimate both deterministic and stochastic rates of population growth (λd and λs, respectively). Annual population censuses suggested moderate growth in abundance between 1998–2003, but vital rate estimates indicated that this temporary growth may be replaced by declines in the long term, both in southern Nova Scotia (λd = 1.0043, λs = 0.9263) and in the Gulf of St. Lawrence (λd = 0.9651, λs = 0.8214). Nonetheless, confidence intervals on λ estimates were relatively wide, highlighting remaining uncertainty in future population trajectories. Differences in projected growth between regions appear to be driven by low estimated juvenile post-fledging survival in the Gulf, but threats to juveniles of both population segments following departure from nesting beaches remain unidentified. Similarly, λ in both population segments was particularly sensitive to changes in adult survival as expected for most migratory birds, but very little is understood about the threats to Piping Plover survival during migration and overwintering. Consequently, we suggest that future recovery efforts for these and other vulnerable migrants should quantify and manage the largely unknown sources of both adult and juvenile mortality during non-breeding seasons while maintaining current levels of nesting habitat protection.

Understanding Demographic and Behavioral Mechanisms that Guide Responses of Neotropical Migratory Birds to Urbanization: a Simulation Approach

Although studies often report that densities of many forest birds are negatively related to urbanization, the mechanisms guiding this pattern are poorly understood. Our objective was to use a population simulation to examine the relative influence of six demographic and behavioral processes on patterns of avian abundance in urbanizing landscapes. We constructed an individual-based population simulation model representing the annual cycle of a Neotropical migratory songbird. Each simulation was performed under two landscape scenarios. The first scenario had similar proportions of high- and low-quality habitat across the urban to rural gradient. Under the first scenario, avian density was negatively related to urbanization only when rural habitats were perceived to be of higher quality than they actually were. The second landscape scenario had declining proportions of high-quality habitat as urbanization increased. Under the second scenario, each mechanism generated a negative relationship between density and urbanization. The strongest effect on density resulted when birds preferentially selected habitats in landscapes from which they fledged or were constrained from dispersing. The next strongest patterns occurred when birds directly evaluated habitat quality and accurately selected the highest-quality available territories. When birds selected habitats based on the presence of conspecifics, the density–urbanization relationship was only one-third the strength of other habitat selection mechanisms and only occurred under certain levels of population survival. Although differences in adult or nest survival in the face of random habitat selection still elicited reduced densities in urban landscapes, the relationships between urbanization and density were weaker than those produced by the conspecific attraction mechanism. Results from our study identify key predictions and areas for future research, including assessing habitat quality in urban and rural areas in order to determine if habitats in urban areas are underutilized.

A Demographic Model to Evaluate Population Declines in the Endangered Streaked Horned Lark

The Streaked Horned Lark (Eremophila alpestris strigata) is listed as endangered by the State of Washington, USA and by Canada under the Species at Risk Act and is also classified as a federal candidate for listing under the Endangered Species Act in the USA. A substantial portion of Streaked Horned Lark habitat has been lost or degraded, and range contraction has occurred in Oregon, Washington, and British Columbia. We estimate the vital rates (fecundity, adult and juvenile survival) and population growth rate (λ) for Streaked Horned Larks breeding in Washington, USA and conduct a Life-Stage Simulation Analysis (LSA) to evaluate which vital rate has the greatest influence on λ. We simulated changes in the three vital rates to examine how much they would need to be adjusted either independently or in concert to achieve a stable Streaked Horned Lark population (λ = 1). We also evaluated which fecundity component (the number of fledglings per egg laid or renesting interval) had the greatest impact on λ. The estimate of population growth suggests that Streaked Horned Larks in Washington are declining rapidly (λ = 0.62 ± 0.10) and that local breeding sites are not sustainable without immigration. The LSA results indicate that adult survival had the greatest influence on λ, followed by juvenile survival and fecundity. However, increases in vital rates led to λ = 1 only when adult survival was raised from 0.47 to 0.85, juvenile survival from 0.17 to 0.58, and fecundity from 0.91 to 3.09. Increases in breeding success and decreases in the renesting interval influenced λ similarly; however, λ did not reach 1 even when breeding success was raised to 100% or renesting intervals were reduced to 1 day. Only when all three vital rates were increased simultaneously did λ approach 1 without requiring highly unrealistic increases in each vital rate. We conclude that conservation activities need to target all or multiple vital rates to be successful. The baseline data presented here and subsequent efforts to manage Streaked Horned Larks will provide valuable information for management of other declining Horned Lark subspecies and other grassland songbirds across North America.

Lesser Scaup Population Dynamics: What Can Be Learned from Available Data?

Populations of Lesser Scaup (Aythya affinis) have declined markedly in North America since the early 1980s. When considering alternatives for achieving population recovery, it would be useful to understand how the rate of population growth is functionally related to the underlying vital rates and which vital rates affect population growth rate the most if changed (which need not be those that influenced historical population declines). To establish a more quantitative basis for learning about life history and population dynamics of Lesser Scaup, we summarized published and unpublished estimates of vital rates recorded between 1934 and 2005, and developed matrix life-cycle models with these data for females breeding in the boreal forest, prairie-parklands, and both regions combined. We then used perturbation analysis to evaluate the effect of changes in a variety of vital-rate statistics on finite population growth rate and abundance. Similar to Greater Scaup (Aythya marila), our modeled population growth rate for Lesser Scaup was most sensitive to unit and proportional change in adult female survival during the breeding and non-breeding seasons, but much less so to changes in fecundity parameters. Interestingly, population growth rate was also highly sensitive to unit and proportional changes in the mean of nesting success, duckling survival, and juvenile survival. Given the small samples of data for key aspects of the Lesser Scaup life cycle, we recommend additional research on vital rates that demonstrate a strong effect on population growth and size (e.g., adult survival probabilities). Our life-cycle models should be tested and regularly updated in the future to simultaneously guide science and management of Lesser Scaup populations in an adaptive context.

Evaluating the Small Population Paradigm for Rare Large-Bodied Woodpeckers, with Implications for the Ivory-billed Woodpecker

Six large-bodied, ≥ 120 g, woodpecker species are listed as near-threatened to critically endangered by the International Union for Conservation of Nature (IUCN). The small population paradigm assumes that these populations are likely to become extinct without an increase in numbers, but the combined influences of initial population size and demographic rates, i.e., annual adult survival and fecundity, may drive population persistence for these species. We applied a stochastic, stage-based single-population model to available demographic rates for Dryocopus and Campephilus woodpeckers. In particular, we determined the change in predicted extinction rate, i.e., proportion of simulated populations that went extinct within 100 yr, to concomitant changes in six input parameters. To our knowledge, this is the first study to evaluate the combined importance of initial population size and demographic rates for the persistence of large-bodied woodpeckers. Under a worse-case scenario, the median time to extinction was 7 yr (range: 1–32). Across the combinations of other input values, increasing initial population size by one female induced, on average, 0.4%–3.2% (range: 0%–28%) reduction in extinction rate. Increasing initial population size from 5–30 resulted in extinction rates < 0.05 under limited conditions: (1) all input values were intermediate, or (2) Allee effect present and annual adult survival ≥ 0.8. Based on our model, these species can persist as rare, as few as five females, and thus difficult-to-detect, populations provided they maintain ≥ 1.1 recruited females annually per adult female and an annual adult survival rate ≥ 0.8. Athough a demographic-based population viability analysis (PVA) is useful to predict how extinction rate changes across scenarios for life-history attributes, the next step for modeling these populations should incorporate more easily acquired data on changes in patch occupancy to make predictions about patch colonization and extinction rates.

Identification of Putative Wintering Areas and Ecological Determinants of Population Dynamics of Common House-Martin (Delichon urbicum) and Common Swift (Apus apus) Breeding in Northern Italy

To identify the causes of population decline in migratory birds, researchers must determine the relative influence of environmental changes on population dynamics while the birds are on breeding grounds, wintering grounds, and en route between the two. This is problematic when the wintering areas of specific populations are unknown. Here, we first identified the putative wintering areas of Common House-Martin (Delichon urbicum) and Common Swift (Apus apus) populations breeding in northern Italy as those areas, within the wintering ranges of these species, where the winter Normalized Difference Vegetation Index (NDVI), which may affect winter survival, best predicted annual variation in population indices observed in the breeding grounds in 1992–2009. In these analyses, we controlled for the potentially confounding effects of rainfall in the breeding grounds during the previous year, which may affect reproductive success; the North Atlantic Oscillation Index (NAO), which may account for climatic conditions faced by birds during migration; and the linear and squared term of year, which account for nonlinear population trends. The areas thus identified ranged from Guinea to Nigeria for the Common House-Martin, and were located in southern Ghana for the Common Swift. We then regressed annual population indices on mean NDVI values in the putative wintering areas and on the other variables, and used Bayesian model averaging (BMA) and hierarchical partitioning (HP) of variance to assess their relative contribution to population dynamics. We re-ran all the analyses using NDVI values at different spatial scales, and consistently found that our population of Common House-Martin was primarily affected by spring rainfall (43%–47.7% explained variance) and NDVI (24%–26.9%), while the Common Swift population was primarily affected by the NDVI (22.7%–34.8%). Although these results must be further validated, currently they are the only hypotheses about the wintering grounds of the Italian populations of these species, as no Common House-Martin and Common Swift ringed in Italy have been recovered in their wintering ranges.

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.

Stability of a Seabird Population in the Presence of an Introduced Predator

We hypothesized that although large populations may appear able to withstand predation and disturbance, added stochasticity in population growth rate (λ) increases the risk of dramatic population declines. Approximately half of the Aleutian Islands' population of Least Auklets (Aethia pusilla) breed at one large colony at Kiska Island in the presence of introduced Norway rats (Rattus norvegicus) whose population erupts periodically. We evaluated two management plans, do nothing or eradicate rats, for this colony, and performed stochastic elasticity analysis to focus future research and management. Our results indicated that Least Auklets breeding at Kiska Island had the lowest absolute value of growth rate and more variable λ's (neither statistically significant) during 2001-2010, when compared with rat-free colonies at Buldir and Kasatochi islands. We found variability in the annual proportional change in population size among islands with Kiska Island having the fastest rate of decline, 78% over 20 years. Under the assumption that the eradication of rats would result in vital rates similar to those observed at rat-free Buldir and Kasatochi islands, we found the projected population decline decreased from 78% to 24% over 20 years. Overall, eradicating rats at Kiska Island is not likely to increase Least Auklet vital rates, but will decrease the amount of variation in λ, resulting in a significantly slower rate of population decline. We recommend the eradication of rats from Kiska Island to decrease the probability of dramatic population declines and ensure the future persistence of this important colony.

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.