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The effects of harvest on waterfowl populations

Change in the size of populations over space and time is, arguably, the motivation for much of pure and applied ecological research. The fundamental model for the dynamics of any population is straightforward: the net change in the abundance is the simple difference between the number of individuals entering the population and the number leaving the population, either or both of which may change in response to factors intrinsic and extrinsic to the population. While harvest of individuals from a population constitutes a clear extrinsic source of removal of individuals, the response of populations to harvest is frequently complex, reflecting an interaction of harvest with one or more population processes. Here we consider the role of these interactions, and factors influencing them, on the effective harvest management of waterfowl populations. We review historical ideas concerning harvest and discuss the relationship(s) between waterfowl life histories and the development and application of population models to inform harvest management. The influence of population structure (age, spatial) on derivation of optimal harvest strategies (with and without explicit consideration of various sources of uncertainty) is considered. In addition to population structure, we discuss how the optimal harvest strategy may be influenced by: 1) patterns of density-dependence in one or more vital rates, and 2) heterogeneity in vital rates among individuals within an age-sex-size class. Although derivation of the optimal harvest strategy for simple population models (with or without structure) is generally straightforward, there are several potential difficulties in application. In particular, uncertainty concerning the population structure at the time of harvest, and the ability to regulate the structure of the harvest itself, are significant complications. We therefore review the evidence of effects of harvest on waterfowl populations. Some of this evidence has focussed on correspondence of data with more phenomenological models and other evidence relates to specific mechanisms, including densitydependence and heterogeneity. An important part of this evidence is found in the evolution of model weights under various adaptive harvest management programmes of the U.S. Fish and Wildlife Service for North American waterfowl. Overall, there is substantial uncertainty about system dynamics, about the impacts of potential management and conservation decisions on those dynamics, and how to optimise management decisions in the presence of such uncertainties. Such relationships are unlikely to be stationary over space or time, and selective harvest of some individuals can potentially alter life history allocation of resources over time – both of which will potentially influence optimal harvest strategies. These sources of variation and uncertainty argue for the use of adaptive approaches to waterfowl harvest management.

Wildfowl

Spring migration of waterfowl in the Northern Hemisphere: a management and conservation perspective

Spring migration is a key part of the annual cycle for waterfowl populations in the northern hemisphere, due to its temporal proximity to the breeding season and because resources may be limited at one or more staging sites. Research based on field observations during spring lags behind other periods of the year, despite the potential for fitness consequences through diminished survival or cross-seasonal effects of conditions experienced during migration. Consequently, conservation strategies for waterfowl on spring migration are often only refined versions of practices used during autumn and winter. Here we discuss the current state of knowledge of habitat requirements for waterfowl at their spring migratory sites and the intrinsic and extrinsic factors that lead to variability in those requirements. The provision of plant foods has become the main conservation strategy during spring because of the birds’ energy requirements at this time, not only to fuel migration but to facilitate early clutch formation on arrival at the breeding grounds. Although energy sources are important to migrants, there is little evidence on the extent to which the availability of carbohydrate-based food is limiting for many migratory waterfowl populations. Such limitation is relatively unlikely among populations that exploit agricultural grain during migration (e.g. arctic-nesting geese), suggesting that conservation strategies for these populations may be misplaced. In general, however, we found few cases in which an ecological understanding of spring-migrating waterfowl was sufficient to indicate true resource limitation during migration, and still fewer cases where conservation efforts ameliorated these limitations. We propose a framework that aims to address knowledge gaps and apply empirical research results to conservation strategies based on documented limitations and associated fitness impacts on migrating waterfowl. Such a strategy would improve allocation of scarce conservation resources during spring migration and greatly improve ecological understanding of migratory waterfowl and their habitats in the northern hemisphere.

Wildfowl

Waterfowl populations of conservation concern: learning from diverse challenges, models, and conservation strategies

There are 30 threatened or endangered species of waterfowl worldwide, and several sub-populations are also threatened. Some of these species occur in North America, and others there are also of conservation concern due to declining population trends and their importance to hunters. Here we review conservation initiatives being undertaken for several of these latter species, along with conservation measures in place in Europe, to seek common themes and approaches that could be useful in developing broad conservation guidelines. While focal species may vary in their life histories, population threats and geopolitical context, most conservation efforts have used a systematic approach to understand factors limiting populations and o identify possible management or policy actions. This approach generally includes a priori identification of plausible hypotheses about population declines or status, incorporation of hypotheses into conceptual or quantitative planning models, and the use of some form of structured decision making and adaptive management to develop and implement conservation actions in the face of many uncertainties. A climate of collaboration among jurisdictions sharing these birds is important to the success of a conservation or management programme. The structured conservation approach exemplified herein provides an opportunity to involve stakeholders at all planning stages, allows for all views to be examined and incorporated into model structures, and yields a format for improved communication, cooperation and learning, which may ultimately be one of the greatest benefits of this strategy.

Wildfowl

Managing harvest and habitat as integrated components

In 2007, several important initiatives in the North American waterfowl management community called for an integrated approach to habitat and harvest management. The essence of the call for integration is that harvest and habitat management affect the same resources, yet exist as separate endeavours with very different regulatory contexts. A common modelling framework could help these management streams to better understand their mutual effects. Particularly, how does successful habitat management increase harvest potential? Also, how do regional habitat programmes and large-scale harvest strategies affect continental population sizes (a metric used to express habitat goals)? In the ensuing five years, several projects took on different aspects of these challenges. While all of these projects are still on-going, and are not yet sufficiently developed to produce guidance for management decisions, they have been influential in expanding the dialogue and producing some important emerging lessons. The first lesson has been that one of the more difficult aspects of integration is not the integration across decision contexts, but the integration across spatial and temporal scales. Habitat management occurs at local and regional scales. Harvest management decisions are made at a continental scale. How do these actions, taken at different scales, combine to influence waterfowl population dynamics at all scales? The second lesson has been that consideration of the interface of habitat and harvest management can generate important insights into the objectives underlying the decision context. Often the objectives are very complex and trade-off against one another. The third lesson follows from the second – if an understanding of the fundamental objectives is paramount, there is no escaping the need for a better understanding of human dimensions, specifically the desires of hunters and nonhunters and the role they play in conservation. In the end, the compelling question is how to better understand, guide and justify decisions about conservation investments in waterfowl management. Future efforts to integrate harvest and habitat management will include completion of the species-specific case-studies, initiation of policy discussions around how to integrate the decision contexts and governing institutions, and possible consideration of a new level of integration – integration of harvest and habitats management decisions across waterfowl stocks.

Wildfowl

Waterfowl habitat use and selection during the remigial moult period in the northern hemisphere

This paper reviews factors affecting site selection amongst waterfowl (Anatidae) during the flightless remigial moult, emphasising the roles of predation and food supply (especially protein and energy). The current literature suggests survival during flightless moult is at least as high as at other times of the annual cycle, but documented cases of predation of flightless waterfowl under particular conditions lead us to infer that habitat selection is generally highly effective in mitigating or avoiding predation. High energetic costs of feather replacement and specific amino-acid requirements for their construction imply adoption of special energetic and nutritional strategies at a time when flightlessness limits movements. Some waterfowl meet their energy needs from endogenous stores accumulated prior to remigial moult, others rely on exogenous supply, but this varies with species, age, reproductive status and site. Limited evidence suggests feather proteins are derived from endogenous and exogenous sources which may affect site selection. Remigial moult does not occur independently of other annual cycle events and is affected by reproductive investment and success. Hence, moult strategies are affected by age, sex and reproductive history, and may be influenced by the need to attain a certain internal state for the next stage in the annual cycle (e.g. autumn migration). We know little about habitat selection during moult and urge more research of this poorly known part of the annual cycle, with particular emphasis on identifying key concentrations and habitats for specific flyway populations and the effects of disturbance upon these. This knowledge will better inform conservation actions and management actions concerning waterfowl during moult and the habitats that they exploit.

Wildfowl

Drivers of waterfowl population dynamics: from teal to swans

Waterfowl are among the best studied and most extensively monitored species in the world. Given their global importance for sport and subsistence hunting, viewing and ecosystem functioning, great effort has been devoted since the middle part of the 20th century to understanding both the environmental and demographic mechanisms that influence waterfowl population and community dynamics. Here we use comparative approaches to summarise and contrast our understanding of waterfowl population dynamics across species as short-lived as the teal Anas discors and A.crecca to those such as the swans Cygnus sp. which have long life-spans. Specifically, we focus on population responses to vital rate perturbations across life history strategies, discuss bottom-up and top-down responses of waterfowl populations to global change, and summarise our current understanding of density dependence across waterfowl species. We close by identifying research needs and highlight ways to overcome the challenges of sustainably managing waterfowl populations in the 21st century.

Wildfowl

Delineation of Tundra Swan Cygnus c. columbianus populations in North America: geographic boundaries and interchange

North American Tundra Swans Cygnus c. columbianus are composed of two wellrecognised populations: an Eastern Population (EP) that breeds across northern Canada and north of the Brooks Range in Alaska, which migrates to the eastern seaboard of the United States, and a Western Population (WP) that breeds in coastal regions of Alaska south of the Brooks Range and migrates to western North America. We present results of a recent major ringing effort from across the breeding range in Alaska to provide a better definition of the geographic extent of the migratory divide in Alaska. We also reassess the staging and winter distributions of these populations based on locations of birds tracked using satellite transmitters, and recent recoveries and sightings of neck-collared birds. Summer sympatry of EP and WP Tundra Swans is very limited, and largely confined to a small area in northwest Alaska. Autumn migration pathways of EP and WP Tundra swans abut in southwest Saskatchewan, a region where migrating WP birds turn west, and EP birds deviate abruptly eastward. Overall, from 1989 to 2013 inclusive, 2.6% of recoveries or resightings reported to the USGS Bird Banding Laboratory were of birds that moved from the domain of the population in which they were initially captured to within the range of the other population; a proportion roughly comparable to the results of Limpert et al . (1991) for years before 1990. Of the 70 cross-boundary movements reported since 1989, 39% were of birds marked on breeding areas and 61% were of birds marked on wintering areas. Dispersing swans (i.e. those that made crossboundary movements) did not differ with respect to age or sex from those that did not move between populations. The Brooks Range in northern Alaska effectively separates the two populations within Alaska, but climate-induced changes in tundra breeding habitats and losses of wetlands on staging areas may alter the distribution for both of these populations.

Wildfowl

Evaluating predictors of local dabbling duck abundance during migration: Managing the spectrum of conditions faced by migrants

The development of robust modelling techniques to derive inferences from large-scale migratory bird monitoring data at appropriate scales has direct relevance to their management. The Integrated Waterbird Management and Monitoring programme (IWMM) represents one of the few attempts to monitor migrating waterbirds across entire flyways using targeted local surveys. This dataset included 13,208,785 waterfowl (eight Anas species) counted during 28,000 surveys at nearly 1,000 locations across the eastern United States between autumn 2010 and spring 2013 and was used to evaluate potential predictors of waterfowl abundance at the wetland scale. Mixed-effects, log-linear models of local abundance were built for the Atlantic and Mississippi flyways during spring and autumn migration to identify factors relating to habitat structure, forage availability, and migration timing that influence target dabbling duck species abundance. Results indicated that migrating dabbling ducks responded differently to environmental factors. While the factors identified demonstrated a high degree of importance, they were inconsistent across species, flyways and seasons. Furthermore, the direction and magnitude of the importance of each covariate group considered here varied across species. Given our results, actionable policy recommendations are likely to be most effective if they consider species-level variation within targeted taxonomic units and across management areas. The methods implemented here can easily be applied to other contexts, and serve as a novel investigation into local-level population patterns using data from broad-scale monitoring programmes.

Wildfowl

Observations of wing-feather molt and summer feeding ecology of Steller's Eiders at Nelson Lagoon, Alaska

The population size, moult chronology, food habits, and feeding behaviour of Steller's Eiders Polysticta stelleri were studied at Nelson Lagoon, Alaska from May to October 1977. Sub-adults were flightless from late July to late August and the adult males were flightless from late August to mid September. Adult females were rarely flightless at Nelson Lagoon but commonly flightless at Izembek Bay. Steller's Eiders ate primarily bivalve mollusca and amphipod crustacea, with Mytilus edulis and Anisogammarus pugettensis the most important foods. Eiders took amphipoda prior to the wing-feather moult, bivalves during the wing-feather moult, and both types of invertebrates after the wing-feather moult. Steller's Eiders fed primarily at the low tide by diving (flock-feeding) or head-dipping, both during the day and at night. There was no difference in feeding behaviour between ages or sexes. Eiders fed almost exclusively by diving after the wing-feather moult and apparently fed more as the season progressed.

Alaska

Estimating habitat carrying capacity for migrating and wintering waterfowl: Considerations, pitfalls and improvements

Population-based habitat conservation planning for migrating and wintering waterfowl in North America is carried out by habitat Joint Venture (JV) initiatives and is based on the premise that food can limit demography (i.e. food limitation hypothesis). Consequently, planners use bioenergetic models to estimate food (energy) availability and population-level energy demands at appropriate spatial and temporal scales, and translate these values into regional habitat objectives. While simple in principle, there are both empirical and theoretical challenges associated with calculating energy supply and demand including: 1) estimating food availability, 2) estimating the energy content of specific foods, 3) extrapolating site-specific estimates of food availability to landscapes for focal species, 4) applicability of estimates from a single species to other species, 5) estimating resting metabolic rate, 6) estimating cost of daily behaviours, and 7) estimating costs of thermoregulation or tissue synthesis. Most models being used are daily ration models (DRMs) whose set of simplifying assumptions are well established and whose use is widely accepted and feasible given the empirical data available to populate such models. However, DRMs do not link habitat objectives to metrics of ultimate ecological importance such as individual body condition or survival, and largely only consider food-producing habitats. Agent-based models (ABMs) provide a possible alternative for creating more biologically realistic models under some conditions; however, ABMs require different types of empirical inputs, many of which have yet to be estimated for key North American waterfowl. Decisions about how JVs can best proceed with habitat conservation would benefit from the use of sensitivity analyses that could identify the empirical and theoretical uncertainties that have the greatest influence on efforts to estimate habitat carrying capacity. Development of ABMs at restricted, yet biologically relevant spatial scales, followed by comparisons of their outputs to those generated from more simplistic, deterministic models can provide a means of assessing degrees of dissimilarity in how alternative models describe desired landscape conditions for migrating and wintering waterfowl.

Wildfowl

Waterfowl in Cuba: Current status and distribution

Cuba and its satellite islands represent the largest landmass in the Caribbean archipelago and a major repository of the region’s biodiversity. Approximately 13.4% of the Cuban territory is covered by wetlands, encompassing approximately 1.48 million ha which includes mangroves, flooded savannas, peatlands, freshwater swamp forests and various types of managed wetlands. Here, we synthesise information on the distribution and abundance of waterfowl on the main island of Cuba, excluding the numerous surrounding cays and the Isla de la Juventud (Isle of Youth), and report on band recoveries from wintering waterfowl harvested in Cuba by species and location. Twenty-nine species of waterfowl occur in Cuba, 24 of which are North American migrants. Of the five resident Anatid species, three are of conservation concern: the West Indian Whistling-duck Dendrocygna arborea (globally vulnerable), White-cheeked Pintail Anas bahamensis (regional concern) and Masked Duck Nomonyx dominicus (regional concern). The most abundant species of waterfowl wintering in Cuba include Blue-winged Teal A. discors , Northern Pintail A. acuta , and Northern Shoveler A. clypeata . Waterfowl banded in Canada and the United States and recovered in Cuba included predominantly Blue-winged Teal, American Wigeon and Northern Pintail. Banding sites of recovered birds suggest that most of the waterfowl moving through and wintering in Cuba are from the Atlantic and Mississippi flyways. Threats to wetlands and waterfowl in Cuba include: 1) egg poaching of resident species, 2) illegal hunting of migratory and protected resident species, 3) mangrove deforestation, 4) reservoirs for irrigation, 5) periods of pronounced droughts, and 6) hurricanes. Wetland and waterfowl conservation efforts continue across Cuba’s extensive system of protected areas. Expanding collaborations with international conservation organisations, researchers and governments in North America will enhance protection of waterfowl and wetlands in Cuba.

Wildfowl

Population and status of Emperor Geese along the north side of the Alaska Peninsula

We gathered information on the timing of spring and autumn migration of Emperor Geese Anser canagicus from Nelson Lagoon, the age ratios of geese during autumn migration, and the numbers of geese in estuaries along the north side of the Alaska Peninsula and Bristol Bay during spring and autumn migration. Birds staged in most of the lagoons and bays along the north side of the Alaska Peninsula, but Nelson Lagoon and Seal Islands Lagoon were the most heavily used. Geese concentrated in Nelson Lagoon for about six to eight weeks in both spring and autumn. We suspect that the population of Emperor Geese in Alaska may have declined by as much as 34% between the 1960's and 1981. Although the average number of young per successful breeding pair has remained high during this period, the overall proportion of young in the population has been extremely low, at least during the past five years. More studies are necessary to determine factors influencing the status of this species.

Alaska

Autumn staging of cackling Canada geese on the Alaska Peninsula

Cackling Canada Geese Branta canadensis minima undergo a long (2,800 km) migration between their autumn staging areas on the Alaska Peninsula and wintering areas in Oregon and California. Adults and young lost an average of 543 g during this migratory flight. Weight gains between fledging and autumn migration are essential for the successful completion of the flight. Autumn staging is important for the acquisition or maintenance of energy stores, particularly for young of the year, as evidenced by the intense feeding of this age class at Ugashik Bay. Factors associated with the arrival and departure of Cackling Geese at Ugashik Bay are presently unknown.

Alaska

Current status and recent dynamics of the Black Brant Branta bernicla breeding population

We summarize current knowledge about the distribution of Pacific Black Brant and recent dynamics of colonies, particularly on the Yukon-Kuskokwim (Y-K) Delta, Alaska. About 20,000 nests are required to produce the number of young in the autumn flight using estimates of clutch size, hatching success and gosling survival based on colonies on the Y-K Delta. More than 80% of the nests in the population can be accounted for currently on the Y-K Delta. Most moulting individuals that did not breed, or were unsuccessful, are unaccounted for in late summer. Numbers of Black Brant nesting in major colonies on the Y-K Delta declined >60% in the early 1980s, most likely as a result of local subsistence harvest combined with predation by arctic foxes. Effective management of this population requires a better understanding of the distribution of breeding and moulting birds, the importance of breeding habitat to colony dynamics and the role of both sport and subsistence harvest in population dynamics.

Wildfowl

How many Laysan Teal Anas laysanensis are on Midway Atoll? Methods for monitoring abundance after reintroduction

Wildlife managers often request a simple approach to monitor the status of species of concern. In response to that need, we used eight years of monitoring data to estimate population size and test the validity of an index for monitoring accurately the abundance of reintroduced, endangered Laysan Teal Anas laysanensis. The population was established at Midway Atoll in the Hawaiian archipelago after 42 wild birds were translocated from Laysan Island during 2004–2005. We fitted 587 birds with unique markers during 2004–2015, recorded 21,309 sightings until March 2016, and conducted standardised survey counts during 2007–2015. A modified Lincoln-Petersen mark-resight estimator and ANCOVA models were used to test the relationship between survey counts, seasonal detectability, and population abundance. Differences were found between the breeding and non-breeding seasons in detection and how maximum counts recorded related to population estimates. The results showed strong, positive correlations between the seasonal maximum counts and population estimates. The ANCOVA models supported the use of standardised bi-monthly counts of unmarked birds as a valid index to monitor trends among years within a season at Midway Atoll. The translocated population increased to 661 adult and juvenile birds (95% CI = 608–714) by 2010, then declined by 38% between 2010 and 2012 after the Toˉhoku Japan earthquake-generated tsunami inundated 41% of the atoll and triggered an Avian Botulism type C Clostridium botulinum outbreak. Following another severe botulism outbreak during 2015, the population experienced a 37% decline. Data indicated that the Midway Atoll population, like the founding Laysan Island population, is susceptible to catastrophic population declines. Consistent standardised monitoring using simple counts, in place of mark-recapture and resightings surveys, can be used to evaluate population status over the long-term. We estimate there were 314–435 Laysan Teal (95% CI for population estimate; point estimate = 375 individuals) at Midway Atoll in 2015; c. 50% of the global population. In comparison, the most recent estimate for numbers on Laysan Island was of 339 individuals in 2012 (95% CI = 265–413). We suggest that this approach can be used to validate a survey index for any marked, reintroduced resident wildlife population.

Wildfowl

Capture myopathy in a captive Black-bellied Whistling Duck

A female captive-raised Black-bellied Whistling Duck exhibited signs of capture myopathy within 24 hours after a strenuous capture. Serum creatine kinase activity 48 hrs after capture was 23,680 IU/L and aspartate aminotransferase activity was 898 IU/L. Postmortem lesions included large, pale areas of muscle tissue in the hind limbs and heart. Histologic lesions of skeletal muscle included severe myofiber degeneration and necrosis with mineralization, interstitial hypercellularity, and occasional areas of dense fibrous connective tissue scarring.

Wildfowl

A leg-hold noose capture method for Brent Geese Branta bernicla at staging or wintering sites

Effective and efficient capture methods are needed for marking and monitoring individuals in studies of demography, migration and habitat use. We describe a novel use of leg-hold nooses aligned on lines and mats to capture non-breeding Brent Geese Branta bernicla in water at a staging and wintering site in Japan. A total of 24 Brent Geese were caught in autumn 2017 and 2018. The traps, which were easy to set up and transport, were effective at catching small numbers of Brent Goose at intertidal roosting and gritting sites. Leg-hold noose lines and mats may be a suitable alternative to other standard catching techniques, such as cannon-netting and flat net traps, in locations where it is not practicable to use these other methods.

Wildfowl

Leech parasitism of waterfowl in North America

Leech parasitism of waterfowl is widespread in North America. Twenty species of ducks, geese, and swans have been infested by leeches, particularly Theromyzon rude and Placobdella ornata. Sites of attachment include the eyes, nasal passages, and body. Information is lacking on the biology and ecology of duck leeches. Their significance as morbidity and mortality factors and their continental distribution should be determined in relation to waterfowl habitats and populations.

Wildfowl