1. Carry-over effects occur when processes in one season influence the success of an individual in the following season. This phenomenon has the potential to explain a large amount of variation in individual fitness, but so far has only been described in a limited number of species. This is largely due to difficulties associated with tracking individuals between periods of the annual cycle, but also because of a lack of research specifically designed to examine hypotheses related to carry-over effects. 2. We review the known mechanisms that drive carry-over effects, most notably macronutrient supply, and highlight the types of life histories and ecological situations where we would expect them to most often occur. We also identify a number of other potential mechanisms that require investigation, including micronutrients such as antioxidants. 3. We propose a series of experiments designed to estimate the relative contributions of extrinsic and intrinsic quality effects in the pre-breeding season, which in turn will allow an accurate estimation of the magnitude of carry-over effects. To date this has proven immensely difficult, and we hope that the experimental frameworks described here will stimulate new avenues of research vital to advancing our understanding of how carry-over effects can shape animal life histories. 4. We also explore the potential of state-dependent modelling as a tool for investigating carry-over effects, most notably for its ability to calculate optimal rates of acquisition of a multitude of resources over the course of the annual cycle, and also because it allows us to vary the strength of density-dependent relationships which can alter the magnitude of carry-over effects in either a synergistic or agonistic fashion. 5. In conclusion carry-over effects are likely to be far more widespread than currently indicated, and they are likely to be driven by a multitude of factors including both macro- and micronutrients. For this reason they could feasibly be responsible for a large amount of the observed variation in performance among individuals, and consequently warrant a wealth of new research designed specifically to decompose components of variation in fitness attributes related to processes across and within seasons.

https://besjournals.onlinelibrary.wiley.com/doi/epdf/10.1111/j.1365-2656.2010.01740.x

Carry‐over effects as drivers of fitness differences in animals

Populations of migratory birds are usually considered to be limited by conditions in breeding or wintering areas, but some might be limited by conditions encountered on migration. This could occur at stopover sites where competition for restricted food supplies can reduce subsequent survival or breeding success, or during the flights themselves, when adverse weather can occasionally kill large numbers of individuals. Competition for food could act in a density-dependent manner and help to regulate populations, whereas weather effects are more likely to act in a density-independent manner. The evidence for these views is explored in this paper. When preparing for migration, birds must normally obtain more food per day than usual, in order to accumulate the body reserves that fuel their flights. Birds often concentrate in large numbers at particular stopover sites, where food can become scarce, thus affecting migratory performance. Rates of weight gain, departure weights, and stopover durations often correlate with food supplies at stopover sites, sometimes influencing the subsequent survival and reproductive success of individuals, which can in turn affect subsequent breeding numbers. Many studies have provided evidence for interference and depletion competition at stopover sites, relatively few for migration conditions influencing the subsequent breeding or survival of individuals, and even fewer for effects on subsequent breeding numbers. Migrants in flight occasionally suffer substantial mortality in storms, especially over water, sometimes involving many thousands of birds at a time. Other mass mortalities have resulted from atypical ‘winter-like’ weather, occurring soon after the arrival of summer migrants in their breeding areas or just before their departure in autumn. Again, many thousands of birds at a time have been killed in such incidents, causing reductions of 30–90% in local breeding densities. In some bird species, migration-related events can at times have substantial effects on the year-to-year changes in breeding population levels. Nonetheless, the difficulties involved in investigating migrating birds at different points on their migration routes have so far limited the number of studies on the influence of events during migration periods on population levels.

Can conditions experienced during migration limit the population levels of birds

Unlike resident bird species, the population sizes of migratory species can be influenced by conditions in more than one part of the world. Changes in the numbers of migrant birds, either long-term or year-to-year, may be caused by changes in conditions in the breeding or wintering areas or both. The strongest driver of numerical change is provided in whichever area the per capita effects of adverse factors on survival or fecundity are greatest. Examples are given of some species whose numbers have changed in association with conditions in breeding areas, and of others whose numbers have changed in association with conditions in wintering areas. In a few such species, the effects of potential limiting factors have been confirmed locally by experiment. In theory, population sizes might also be limited by severe competition at restricted stopover sites, where bird densities are often high and food supplies heavily depleted, but (with one striking exception) the evidence is as yet no more than suggestive. In some species, habitats occupied in wintering and migration areas, and their associated food supplies, can influence the body condition, migration dates and subsequent breeding success of migrants. Body reserves accumulated in spring by large waterfowl serve for migration and for subsequent breeding, and females with the largest reserves are most likely to produce young. Hence, the conditions experienced by individuals in winter in one region can affect their subsequent breeding success in another region. Such effects are apparent at the level of the individual and at the level of the population. Similarly, the numbers of young produced in one region could, through density-dependent processes, affect subsequent overall mortality in another region. Events in breeding, migration and wintering areas are thus interlinked in their effects on bird numbers. Although in the last 30–40 years the numbers of some tropical wintering birds have declined in western Europe and others in eastern North America, the causes seem to differ. In Europe, declines have mainly involved species that winter in the arid savannas of tropical Africa, which have suffered from the effects of drought and increasing desertification. In several species, annual fluctuations in numbers and adult survival rates were correlated with annual fluctuations in rainfall, and by implication in winter food supplies. In North America, by contrast, numerical declines have affected many species that breed and winter in forest, especially those eastern species favouring the forest interior. Declines have been attributed ultimately to human-induced changes in the breeding range, particularly forest fragmentation, which have led to increases in the densities of nest predators and parasitic cowbirds. These in turn are thought to have caused declines in the breeding success of some neotropical migrants, which is now too low to offset the usual adult mortality, but as yet convincing evidence is available for only a minority of species. The breeding rates and population changes of some migratory species have been influenced by natural changes in the availability of defoliating caterpillars. In other species, tropical deforestation is likely to have played the major role in population decline, and if recent rates of tropical deforestation continue, it is likely to affect an increasing range of migratory species in the future. Not all such species are likely to be affected adversely by deforestation, however, and some may benefit from the resulting habitat changes.

https://onlinelibrary.wiley.com/doi/pdf/10.1111/j.1474-919X.2004.00293.x

Population limitation in migrants

Long-distance migrations are among the wonders of the natural world, but this multi-taxon review shows that the characteristics of species that undertake such movements appear to make them particularly vulnerable to detrimental impacts of climate change. Migrants are key components of biological systems in high latitude regions, where the speed and magnitude of climate change impacts are greatest. They also rely on highly productive seasonal habitats, including wetlands and ocean upwellings that, with climate change, may become less food-rich and predictable in space and time. While migrants are adapted to adjust their behaviour with annual changes in the weather, the decoupling of climatic variables between geographically separate breeding and non-breeding grounds is beginning to result in mistimed migration. Furthermore, human land-use and activity patterns will constrain the ability of many species to modify their migratory routes and may increase the stress induced by climate change. Adapting conservation strategies for migrants in the light of climate change will require substantial shifts in site designation policies, flexibility of management strategies and the integration of forward planning for both people and wildlife. While adaptation to changes may be feasible for some terrestrial systems, wildlife in the marine ecosystem may be more dependent on the degree of climate change mitigation that is achievable.

/pdf/travelling-through-a-warming-world-climate-change-and-2p836ukauk.pdf

Travelling through a warming world: climate change and migratory species.

Summary An estimated 19% of the world’s 9,856 extant bird species are migratory, including some 1,600 species of land- and waterbirds. In 2008, 11% of migratory land- and waterbirds were classed by BirdLife International as threatened or near-threatened on the IUCN Red List. Red List indices show that these migrants have become more threatened since 1988, with 33 species deteriorating and just six improving in status. There is also increasing evidence of regional declines. Population trend data show that more Nearctic–Neotropical migrants have declined than increased in North America since the 1980s, and more Palearctic–Afrotropical migrants breeding in Europe declined than increased during 1970–2000. Reviews of the status of migratory raptors show unfavourable conservation status for 51% of species in the African–Eurasian region (in 2005), and 33% of species in Central, South and East Asia (in 2007). Land-use change owing to agriculture is the most frequently cited threat affecting nearly 80% of all threatened and nearthreatened species. However, while agricultural intensification on the breeding grounds is often proposed as the major driver of declines in Palearctic–Afrotropical migrants, some species appear to be limited by the quantity and quality of available habitat in non-breeding areas, notably the drylands of tropical Africa. Forest fragmentation in breeding areas has contributed to the declines of Nearctic–Neotropical migrants with deforestation in non-breeding areas another possible factor. Infrastructure development including wind turbines, cables, towers and masts can also be a threat. Over-harvesting and persecution remain serious threats, particularly at key migration locations. Climate change is affecting birds already, is expected to exacerbate all these pressures, and may also increase competition between migratory and non-migratory species. The conservation of migratory birds thus requires a multitude of approaches. Many migratory birds require effective management of their critical sites, and Important Bird Areas (IBAs) provide an important foundation for such action; however to function effectively in conserving migratory species, IBAs need to be protected and the coherence of the network requires regular review. Since many migratory species (c. 55%) are widely dispersed across their breeding or nonbreeding ranges, it is essential to address the human-induced changes at the wider landscape scale, a very considerable challenge. Efforts to conserve migratory birds in one part of the range are less effective if unaddressed threats are reducing these species’ populations and habitats elsewhere. International collaboration and coordinated action along migration flyways as a whole are thus key elements in any strategy for the conservation of migratory birds.

http://www.globalraptors.org/grin/researchers/uploads/302/kirby_et_al._2008.pdf

Key conservation issues for migratory land- and waterbird species on the world's major flyways

Most populations of migrant shorebirds around the world are in serious decline, suggesting that vital condition-dependent rates such as fecundity and annual survival are being affected globally. A striking example is the red knot (Calidris canutus rufa) population wintering in Tierra del Fuego, which undertakes marathon 30,000 km hemispheric migrations annually. In spring, migrant birds forage voraciously on horseshoe crab eggs in Delaware Bay in the eastern USA before departing to breed in Arctic polar deserts. From 1997 to 2002 an increasing proportion of knots failed to reach threshold departure masses of 180-200 g, possibly because of later arrival in the Bay and food shortage from concurrent over-harvesting of crabs. Reduced nutrient storage, especially in late-arriving birds, possibly combined with reduced sizes of intestine and liver during refuelling, had severe fitness consequences for adult survival and recruitment of young in 2000-2002. From 1997 to 2002 known survivors in Delaware Bay were heavier at initial capture than birds never seen again, annual survival of adults decreased by 37% between May 2000 and May 2001, and the number of second-year birds in wintering flocks declined by 47%. Population size in Tierra del Fuego declined alarmingly from 51,000 to 27,000 in 2000-2002, seriously threatening the viability of this subspecies. Demographic modelling predicts imminent endangerment and an increased risk of extinction of the subspecies without urgent risk-averse management.

/pdf/rapid-population-decline-in-red-knots-fitness-consequences-3yewqlwre8.pdf

Rapid population decline in red knots: fitness consequences of decreased refuelling rates and late arrival in Delaware Bay

RESUMO 
O presente estudo apresenta dados a respeito da variabilidade morfometrica de ovos, filhotes e adultos de Sterna sandvicensis eurygnatha no Brasil, nos sitios reprodutivos no Estado do Espirito Santo, bem como em areas de alimentacao como o Parque Nacional da Lagoa do Peixe, no Rio Grande do Sul e na Ilha Coroa Vermelha, na Bahia. Foram verificados tres padroes de cores em relacao a quantidade
de manchas existentes nos ovos. Para a plumagem dos filhotes foram verificados seis padroes de cores e para a variacao na coloracao do culmem foram registrados sete padroes de cores. A constante de crescimento K foi de 0,951 e o tempo para a especie atingir 50% da massa assintotica foi de 1,8 semanas. Foram definidos tres padroes de cores do culmem para adultos. A coloracao do tarso apresentou dois padroes. Os dados morfometricos dos adultos mostraram-se significativamente diferentes entre as aves das tres areas, apoiando a hipotese de que elas pertencam a populacoes diferentes. Palavras-chaves: morfologia, coloracao, crescimento, trinta-reis-de-bico-amarelo, Brasil. ABSTRACT Morphologic variations and growth patterns in Sterna sandvicenis eurygnatha in Brazil 
The present study presents data regarding the morphometric variability of eggs, chicks and adults of Sterna sandvicensis eurygnatha in Brazil, in the reproductive sites in Espirito Santo State, as well as in feeding areas as the National Park of the Lagoa do Peixe, in Rio Grande do Sul State and in the Coroa Vermelha Island, in Bahia State. Three patterns of colors were verified in relation to the amount of existent stains in the eggs. For the plumage of the chicks six patterns of colors were verified and for the variation in the coloration of the bill seven patterns of colors were registered. The growth constant K was of 0,951 and the time for the species to reach 50% of the asymptotic mass it was of 1,8 weeks. They were defined three patterns of colors of the bill for adults. The coloration of the tarsus presented two patterns. The adults’ morphometric data were shown significantly different among the birds of the three areas, being able to therefore, to accept the hypothesis that they belong to different populations. Key words: morphology variability, coloration, growth, Cayenne Tern, Brazil.

Inês de Lima Serrano do Nascimento

Papers

Rapid population decline in red knots: fitness consequences of decreased refuelling rates and late arrival in Delaware Bay

VARIAÇÕES MORFOLÓGICAS E PADRÕES DE CRESCIMENTO DE FILHOTES DE Sterna sandvicensis eurygnatha NO BRASIL