Nathan R. Senner

An ontogenetic perspective on individual differences

Phenotypic differences among individuals can arise during any stage of life. Although several distinct processes underlying individual differences have been defined and studied (e.g. parental effects, senescence), we lack an explicit, unified perspective for understanding how these processes contribute separately and synergistically to observed variation in functional traits. We propose a conceptual framework based on a developmental view of life-history variation, linking each ontogenetic stage with the types of individual differences originating during that period. In our view, the salient differences among these types are encapsulated by three key criteria: timing of onset, when fitness consequences are realized, and potential for reversibility. To fill a critical gap in this framework, we formulate a new term to refer to individual differences generated during adulthood—reversible state effects. We define these as ‘reversible changes in a functional trait resulting from life-history trade-offs during adulthood that affect fitness’, highlighting how the adult phenotype can be repeatedly altered in response to environmental variation. Defining individual differences in terms of trade-offs allows explicit predictions regarding when and where fitness consequences should be expected. Moreover, viewing individual differences in a developmental context highlights how different processes can work in concert to shape phenotype and fitness, and lays a foundation for research linking individual differences to ecological and evolutionary theory.

An exception to the rule: carry-over effects do not accumulate in a long-distance migratory bird.

Recent years have seen a growing consensus that events during one part of an animals annual cycle can detrimentally affect its future fitness. Notably, migratory species have been shown to commonly display such carry-over effects, facing severe time constraints and physiological stresses that can influence events across seasons. However, to date, no study has examined a full annual cycle to determine when these carry-over effects arise and how long they persist within and across years. Understanding when carry-over effects are created and how they persist is critical to identifying those periods and geographic locations that constrain the annual cycle of a population and determining how selection is acting upon individuals throughout the entire year. Using three consecutive years of migration tracks and four consecutive years of breeding success data, we tested whether carry-over effects in the form of timing deviations during one migratory segment of the annual cycle represent fitness costs that persist or accumulate across the annual cycle for a long-distance migratory bird, the Hudsonian godwit, Limosa haemastica. We found that individual godwits could migrate progressively later than population mean over the course of an entire migration period, especially southbound migration, but that these deviations did not accumulate across the entire year and were not consistently detected among individuals across years. Furthermore, neither the accumulation of lateness during previous portions of the annual cycle nor arrival date at the breeding grounds resulted in individuals suffering reductions in their breeding success or survival. Given their extreme life history, such a lack of carry-over effects suggests that strong selection exists on godwits at each stage of the annual cycle and that carry-over effects may not be able to persist in such a system, but also emphasizes that high-quality stopover and wintering sites are critical to the maintenance of long-distance migratory populations.

When Siberia came to the Netherlands : The response of continental black-tailed godwits to a rare spring weather event

1. Extreme weather events have the potential to alter both short- and long-term population dynamics as well as community- and ecosystem-level function. Such events are rare and stochastic, making it difficult to fully document how organisms respond to them and predict the repercussions of similar events in the future. 2. To improve our understanding of the mechanisms by which short-term events can incur long-term consequences, we documented the behavioural responses and fitness consequences for a long-distance migratory bird, the continental black-tailed godwit Limosa limosa limosa, resulting from a spring snowstorm and three-week period of record low temperatures. 3. The event caused measurable responses at three spatial scales - continental, regional and local - including migratory delays (+19 days), reverse migrations (>90 km), elevated metabolic costs (+8·8% maintenance metabolic rate) and increased foraging rates (+37%). 4. There were few long-term fitness consequences, however, and subsequent breeding seasons instead witnessed high levels of reproductive success and little evidence of carry-over effects. 5. This suggests that populations with continued access to food, behavioural flexibility and time to dissipate the costs of the event can likely withstand the consequences of an extreme weather event. For populations constrained in one of these respects, though, extreme events may entail extreme ecological consequences.

Unexpected diversity in socially synchronized rhythms of shorebirds

The behavioural rhythms of organisms are thought to be under strong selection, influenced by the rhythmicity of the environment. Such behavioural rhythms are well studied in isolated individuals under laboratory conditions, but free-living individuals have to temporally synchronize their activities with those of others, including potential mates, competitors, prey and predators. Individuals can temporally segregate their daily activities (for example, prey avoiding predators, subordinates avoiding dominants) or synchronize their activities (for example, group foraging, communal defence, pairs reproducing or caring for offspring). The behavioural rhythms that emerge from such social synchronization and the underlying evolutionary and ecological drivers that shape them remain poorly understood. Here we investigate these rhythms in the context of biparental care, a particularly sensitive phase of social synchronization where pair members potentially compromise their individual rhythms. Using data from 729 nests of 91 populations of 32 biparentally incubating shorebird species, where parents synchronize to achieve continuous coverage of developing eggs, we report remarkable within- and between-species diversity in incubation rhythms. Between species, the median length of one parent’s incubation bout varied from 1–19 h, whereas period length—the time in which a parent’s probability to incubate cycles once between its highest and lowest value—varied from 6–43 h. The length of incubation bouts was unrelated to variables reflecting energetic demands, but species relying on crypsis (the ability to avoid detection by other animals) had longer incubation bouts than those that are readily visible or who actively protect their nest against predators. Rhythms entrainable to the 24-h light–dark cycle were less prevalent at high latitudes and absent in 18 species. Our results indicate that even under similar environmental conditions and despite 24-h environmental cues, social synchronization can generate far more diverse behavioural rhythms than expected from studies of individuals in captivity. The risk of predation, not the risk of starvation, may be a key factor underlying the diversity in these rhythms.

Post-Breeding Migration of Dutch-Breeding Black-Tailed Godwits: Timing, Routes, Use of Stopovers, and Nonbreeding Destinations

Conservation of long-distance migratory shorebirds is complex because these species use habitats spread across continents and hemispheres, making identification of critical habitats and potential bottlenecks in the annual cycle especially difficult. The population of Black-tailed Godwits that breeds in Western Europe, Limosa limosa limosa, has declined precipitously over the past few decades. Despite significant efforts to identify the root causes of this decline, much remains unclear. To better understand the migratory timing, use of stopover and nonbreeding sites, and the potential impact of breeding success on these parameters, we attached 15 Argos satellite transmitters and 10 geolocation tracking devices to adult godwits nearing completion of incubation at breeding sites in southwest Friesland, The Netherlands during the spring of 2009. We successfully tracked 16 adult godwits for their entire southward migration and two others for part of it. Three migration patterns and four regions of use were apparent. Most godwits left their breeding sites and proceeded south directly to stopover sites in the Mediterranean — e.g. Spain, Portugal, and Morocco — before flying on to non-breeding sites in West Africa. Other individuals spent the entire nonbreeding season in the Mediterranean. A third pattern included a few individuals that flew nonstop from their Dutch breeding sites to nonbreeding sites in West Africa. Tracking data from this study will be immediately useful for conservation efforts focused on preserving the dispersed network of sites used by godwits during their southward migration.

EFFECTS OF COLOR BANDS ON SEMIPALMATED SANDPIPERS BANDED AT HATCH

Abstract Effects of color bands on adult birds have been investigated in many studies, but much less is known about the effects of bands on birds banded at hatch. We captured Semipalmated Sandpiper (Calidris pusilla) chicks at hatch on the Alaskan North Slope and attached 0–3 bands to them. The chicks were resighted and reweighed during the subsequent two weeks. The number of chicks banded varied from 18 to 21 among treatments; 6–9 were resighted, and 6–7 were reweighed, per treatment. The proportion resighted varied from 0.33 to 0.45. The estimated resighting probability, given that we encountered a brood, was 82%. We tested for effects of the bands on survival and mass gain by analyzing whether the proportion of chicks resighted, or their mass, varied with the number of bands. We found no evidence that bands affected the chicks and were able to rule out (with 95% confidence) a decline in survivorship of more than 13% and a loss of mass of more than 10%. Although bands had little if any effect on chicks in our study, we believe their effects should be evaluated whenever survivorship or mass gain are estimated using color-marked chicks.

Estimating the size of the Dutch breeding population of continental black-tailed godwits from 2007-2015 using resighting data from spring staging sites

Over the past 50 years, the population of Continental Black-tailed Godwits Limosa limosa limosa breeding of the East Atlantic Flyway has been in steep decline. This decline has previously been documented in trend analyses and six Netherlands-wide count-based population estimates, the last of which was completed in 1999. We provide an updated population size estimate and describe inter-annual fluctuations in the population between 2007 and 2015. To generate these estimates, we integrated a mark-recapture survival analysis with estimates of the densities of colour-marked individuals at migratory staging sites with known proportions of Continental and Icelandic L. l. islandica Black-tailed Godwits within a Bayesian framework. The use of these analytical techniques means that, in contrast with earlier efforts, our estimates are accompanied with confidence intervals, allowing us to estimate the population size with known precision. Using additional information on the breeding destination of 43 godwits equipped with satellite transmitters at Iberian staging areas, we found that 87% (75–95% 95% CI) of the nominate subspecies in the East Atlantic Flyway breed in The Netherlands. We estimated that the number of breeding pairs in The Netherlands has declined from 47,000 (38,000–56,000) pairs in 2007 to 33,000 (26,000–41,000) in 2015. Despite a temporary increase in 2010 and 2011, the population declined by an average of 3.7% per year over the entire period from 2007–2015, and by 6.3% from 2011–2015. We conclude that investing in an intensive demographic programme at a regional scale, when combined with targeted resightings of marked individuals elsewhere, can yield population estimates at the flyway scale.

One Species but Two Patterns: Populations of the Hudsonian Godwit (Limosa haemastica) Differ in Spring Migration Timing

ABSTRACT. Climate change can cause mismatches between the breeding phenology and peak abundance of food resources of migratory species. Moreover, asynchronously changing climate regimes across their ranges may constrain the ability of migratory species to adapt to all the regimes they encounter. To understand the potential effect of asynchronous changes, I examined the influences of both large- and local-scale weather and climate on the timing of arrival of two disjunct breeding populations of Hudsonian Godwits (Limosa haemastica). I used arrival data from two study sites—Beluga River, Alaska, and Churchill, Manitoba—combined with 37 years of weather and climate data from both winter and stopover sites and the breeding grounds. The Alaskan population now arrives ∼9 days earlier than it did in the early 1970s, and the Churchill population arrives >10 days later. A model-selection process using linear regression models suggested that these divergent trends result from different suites of environmental factors affecting the timing of migration for the two populations. The cues used by the Alaskan population have remained reliable indicators of the timing of the onset of spring on their breeding grounds, but this is not the case for the Churchill population. Conflicting warming regimes in midcontinental North America cause the Churchill population to arrive later to their breeding grounds and limit their ability to properly time their breeding efforts. These results suggest that ecological and phenological limitations, not just evolutionary constraints, are critical to determining how populations respond to climate change.

A global threats overview for Numeniini populations: Synthesising expert knowledge for a group of declining migratory birds

The Numeniini is a tribe of 13 wader species (Scolopacidae, Charadriiformes) of which seven are Near Threatened or globally threatened, including two Critically Endangered. To help inform conservation management and policy responses, we present the results of an expert assessment of the threats that members of this taxonomic group face across migratory flyways. Most threats are increasing in intensity, particularly in non-breeding areas, where habitat loss resulting from residential and commercial development, aquaculture, mining, transport, disturbance, problematic invasive species, pollution and climate change were regarded as having the greatest detrimental impact. Fewer threats (mining, disturbance, problematic native species and climate change) were identified as widely affecting breeding areas. Numeniini populations face the greatest number of non-breeding threats in the East Asian-Australasian Flyway, especially those associated with coastal reclamation; related threats were also identified across the Central and Atlantic Americas, and East Atlantic flyways. Threats on the breeding grounds were greatest in Central and Atlantic Americas, East Atlantic and West Asian flyways. Three priority actions were associated with monitoring and research: to monitor breeding population trends (which for species breeding in remote areas may best be achieved through surveys at key non-breeding sites), to deploy tracking technologies to identify migratory connectivity, and to monitor land-cover change across breeding and non-breeding areas. Two priority actions were focused on conservation and policy responses: to identify and effectively protect key non-breeding sites across all flyways (particularly in the East Asian- Australasian Flyway), and to implement successful conservation interventions at a sufficient scale across human-dominated landscapes for species’ recovery to be achieved. If implemented urgently, these measures in combination have the potential to alter the current population declines of many Numeniini species and provide a template for the conservation of other groups of threatened species.

Does wintering north or south of the Sahara correlate with timing and breeding performance in black-tailed godwits?

Abstract Migrating long distances requires time and energy, and may interact with an individuals performance during breeding. These seasonal interactions in migratory animals are best described in populations with disjunct nonbreeding distributions. The black‐tailed godwit (Limosa limosa limosa), which breeds in agricultural grasslands in Western Europe, has such a disjunct nonbreeding distribution: The majority spend the nonbreeding season in West Africa, while a growing number winters north of the Sahara on the Iberian Peninsula. To test whether crossing the Sahara has an effect on breeding season phenology and reproductive parameters, we examined differences in the timing of arrival, breeding habitat quality, lay date, egg volume, and daily nest survival among godwits (154 females and 157 males), individually marked in a breeding area in the Netherlands for which wintering destination was known on the basis of resightings. We also examined whether individual repeatability in arrival date differed between birds wintering north or south of the Sahara. Contrary to expectation, godwits wintering south of the Sahara arrived two days earlier and initiated their clutch six days earlier than godwits wintering north of the Sahara. Arrival date was equally repeatable for both groups, and egg volume larger in birds wintering north of the Sahara. Despite these differences, we found no association between wintering location and the quality of breeding habitat or nest survival. This suggests that the crossing of an important ecological barrier and doubling of the migration distance, twice a year, do not have clear negative reproductive consequences for some long‐distance migrants.