Keith A. Hobson

Heterogeneity in stable isotope profiles predicts coexistence of populations of barn swallows Hirundo rustica differing in morphology and reproductive performance

Population studies assume that individuals belonging to a study population are homogeneous for natal and breeding origin, although this assumption is rarely tested. We tested for heterogeneity in stable–isotope profiles (δ15N, δ13C, δD) of feathers grown in the African winter quarters from a Danish breeding population of adult barn swallows, Hirundo rustica. Deuterium isotope values did not provide useful information on population segregation of wintering swallows in Africa. However, both δ15N and δ13C values showed a clearly bimodal distribution with 6% belonging to one category and the remaining birds belonging to another category, resulting in this population comprising three categories of birds. Adults belonging to the two categories of δ13C isotope profiles differed weakly in morphology for several different characters. The frequency and the size of second broods differed between categories of δ13C isotope profiles. Phenotypes of nestlings from the first brood in terms of tarsus length, body mass and T–cell response differed significantly between the two δ15N isotope categories, suggesting that conditions during winter carried over to the breeding season at least as late as the first brood. Polymorphism can be maintained only if fitness is similar for birds from categories of isotope profiles. We suggest that fluctuating selection or migration–selection balance may maintain the observed polymorphism.

Do Healthy Monarchs Migrate Farther? Tracking Natal Origins of Parasitized vs. Uninfected Monarch Butterflies Overwintering in Mexico

Long-distance migration can lower parasite prevalence if strenuous journeys remove infected animals from wild populations. We examined wild monarch butterflies (Danaus plexippus) to investigate the potential costs of the protozoan Ophryocystis elektroscirrha on migratory success. We collected monarchs from two wintering sites in central Mexico to compare infection status with hydrogen isotope (δ 2H) measurements as an indicator of latitude of origin at the start of fall migration. On average, uninfected monarchs had lower δ 2H values than parasitized butterflies, indicating that uninfected butterflies originated from more northerly latitudes and travelled farther distances to reach Mexico. Within the infected class, monarchs with higher quantitative spore loads originated from more southerly latitudes, indicating that heavily infected monarchs originating from farther north are less likely to reach Mexico. We ruled out the alternative explanation that lower latitudes give rise to more infected monarchs prior to the onset of migration using citizen science data to examine regional differences in parasite prevalence during the summer breeding season. We also found a positive association between monarch wing area and estimated distance flown. Collectively, these results emphasize that seasonal migrations can help lower infection levels in wild animal populations. Our findings, combined with recent declines in the numbers of migratory monarchs wintering in Mexico and observations of sedentary (winter breeding) monarch populations in the southern U.S., suggest that shifts from migratory to sedentary behavior will likely lead to greater infection prevalence for North American monarchs.

Expanding the Isotopic Toolbox: Applications of Hydrogen and Oxygen Stable Isotope Ratios to Food Web Studies

The measurement of stable carbon (δ13C) and nitrogen (δ15N) isotopes in tissues of organisms has formed the foundation of isotopic food web reconstructions, as these values directly reflect assimilated diet. In contrast, stable hydrogen (δ2H) and oxygen (δ18O) isotope measurements have typically been reserved for studies of migratory origin and paleoclimate reconstruction based on systematic relationships between organismal tissue and local environmental water. Recently, innovative applications using δ2H and, to a lesser extent, δ18O values have demonstrated potential for these elements to provide novel insights in modern food web studies. We explore the advantages and challenges associated with three applications of δ2H and δ18O values in food web studies. First, large δ2H differences between aquatic and terrestrial ecosystem end members can permit the quantification of energy inputs and nutrient fluxes between these two sources, with potential applications for determining allochthonous vs. autochthonous nutrient sources in freshwater systems and relative aquatic habitat utilization by terrestrial organisms. Next, some studies have identified a relationship between δ2H values and trophic position, which suggests that this marker may serve as a trophic indicator, in addition to the more commonly used δ15N values. Finally, coupled measurements of δ2H and δ18O values are increasing as a result of reduced analytical challenges to measure both simultaneously and may provide additional ecological information over single element measurements. In some organisms, the isotopic ratios of these two elements are tightly coupled, whereas the isotopic disequilibrium in other organisms may offer insight into the diet and physiology of individuals. Although a coherent framework for interpreting δ2H and δ18O data in the context of food web studies is emerging, many fundamental uncertainties remain. We highlight directions for targeted research that will increase our understanding of how these markers move through food webs and reflect ecological processes.

Long-distance autumn migration across the Sahara by painted lady butterflies: exploiting resource pulses in the tropical savannah

The painted lady, Vanessa cardui, is a migratory butterfly that performs an annual multi-generational migration between Europe and North Africa. Its seasonal appearance south of the Sahara in autumn is well known and has led to the suggestion that it results from extremely long migratory flights by European butterflies to seasonally exploit the Sahel and the tropical savannah. However, this possibility has remained unproven. Here, we analyse the isotopic composition of butterflies from seven European and seven African countries to provide new support for this hypothesis. Each individual was assigned a geographical natal origin, based on its wing stable hydrogen isotope (δ2Hw) value and a predicted δ2Hw basemap for Europe and northern Africa. Natal assignments of autumn migrants collected south of the Sahara confirmed long-distance movements (of 4000 km or more) starting in Europe. Samples from Maghreb revealed a mixed origin of migrants, with most individuals with a European origin, but others having originated in the Sahel. Therefore, autumn movements are not only directed to northwestern Africa, but also include southward and northward flights across the Sahara. Through this remarkable behaviour, the productive but highly seasonal region south of the Sahara is incorporated into the migratory circuit of V. cardui.

Fuel loads acquired at a stopover site influence the pace of intercontinental migration in a boreal songbird

Long-distance migratory organisms are under strong selection to migrate quickly. Stopovers demand more time than flying and are used by individuals to refuel during migration, but the effect of fuel loads (fat) acquired at stopover sites on the subsequent pace of migration has not been quantified. We studied stopover behaviour of Grey-cheeked Thrush (Catharus minimus) at a site in northern Colombia and then tracked their migration using an intercontinental radio-telemetry array. Tracking confirmed long-distance flights of more than 3000 km, highlighting the key importance of a single stopover site to the migration strategy of this species. Our results suggest that these songbirds behave as time-minimizers as predicted by optimal migration theory, and that fuel loads acquired at this South American stopover site, together with departure date, carry-over to influence the pace of migration, contributing to differences in travel time of up to 30 days in birds subsequently detected in the U. S. and Canada. Such variation in the pace of migration arising from a single stopover site, likely has important fitness consequences and suggests that identifying important fuelling sites will be essential to effectively conserve migratory species.

Density-dependent resource selection by a terrestrial herbivore in response to sea-to-land nutrient transfer by seals

Sea-to-land nutrient transfers can connect marine food webs to those on land, creating a dependence on marine webs by opportunistic species. We show how nitrogen, imported by gray seals, Halichoerus grypus, and traced through stable isotope (δ15 N) measurements in marram grass, Ammophila breviligulata, significantly alters foraging behavior of a free-roaming megaherbivore (feral horses, Equus ferus caballus) on Sable Island, Canada. Values of δ15 N correlated with protein content of marram and strongly related to pupping-seal densities, and positively influenced selective foraging by horses. The latter was density dependent, consistent with optimal foraging theory. We present the first demonstration of how sea-to-land nutrient transfers can affect the behavioral process of resource selection (resource use relative to availability) of terrestrial consumers. We hypothesize that persistence of horses on Sable Island is being facilitated by N subsidies. Our results have relevance to advancing theory on trophic dynamics in island biogeography and metaecosystem ecology.

Chronological and Isotopic data support a revision for the timing of cave bear extinction in Mediterranean Europe

Abstract The Cave Bear, Ursus spelaeus (sensu lato), was one of many megafaunal species that became extinct during the Late Pleistocene in Europe. With new data we revisit the debate about the extinction and paleoecology of this species by presenting new chronometric, isotopic and taphonomic evidence from two Palaeolithic cave bear sites in northeastern Italy: Paina Cave and Trene Cave. Two direct radiocarbon dates on well-preserved collagen have yielded ages around 24,200–23,500 cal yr BP, which make them the latest known representatives of the species in Europe. The carbon (δ13C) and nitrogen (δ15N) isotopic values of bone collagen exhibit values similar to those of older cave bears from Swabian Jura and France, suggesting that the feedings preferences of cave bears remained unchanged until the disappearance of this species in Europe. Several bear remains preserved traces of human modification such as cut marks, which enables a reconstruction of the main steps of fur recovery and the butchering process. The broad range of plant types available and the favorable location of Berici Hills may have played an important role in the range expansion of cave bears and their interaction with the Paleolithic hunters settled the same area.

Patterns of parasitism in monarch butterflies during the breeding season in eastern North America

1. Migratory behaviour can result in reduced prevalence of pathogens in host populations. Two hypotheses have been proposed to explain this relationship: (i) ‘migratory escape’, where migrants benefit from escaping pathogen accumulation in contaminated environments; and (ii) ‘migratory culling’, where the selective removal of infected individuals occurs during migration.

State-dependent capital and income breeding: a novel approach to evaluating individual strategies with stable isotopes

BackgroundSpecies-specific strategies for financing the costs of reproduction are well understood, forming a continuum ranging from high to low reliance on stored nutrients. Animals relying mostly on stored reserves are termed ‘capital breeders’, whereas ‘income breeders’ rely mostly on concurrent intake when financing the costs of reproduction. The role and adaptive value of individual variation in these strategies remain elusive. Life-history theory posits that capital breeding should be favoured when offspring reproductive value peaks, typically occurring early in the season, and that current income should increasingly be used with progressing season. Because resource limitation may hamper flexible resource allocation, a corollary prediction is that only good-condition individuals may show the expected seasonal shift in resource use. To test this prediction, we examined stable isotopes (δ13C and δ15N) in blood and lipid-free egg yolk of breeding eider females (Somateria mollissima) from the Baltic Sea to assess the role of individual variation in the use of proteins from local diet vs. stored reserves.ResultsWe show for the first time that individuals from a single population differ in their utilization of stored reserves and concurrent intake to finance the costs of reproduction. Consistent with our prediction, heavy females predominantly used stored reserves for producing egg yolks early in the season, increasingly relying on local feeding with later onset of breeding, whereas light females showed no seasonal change in allocation strategy.ConclusionsStable isotope profiling at the individual level is a powerful tool for monitoring relative changes in investment strategies through time, showing promise as an early warning indicator of ecological change in food webs.

Environmental conditions during winter predict age- and sex-specific differences in reproductive success of a trans-Saharan migratory bird

Previous studies have linked winter habitat with subsequent breeding phenology and physical condition of migratory birds, but few have found delayed effects of winter habitat on subsequent reproductive success. The aim of this study was to test if African winter habitat is related to subsequent reproductive success of house martins (Delichon urbicum) breeding at a colony in Spain. We measured stable isotope (δ2H, δ13C, δ15N) values from feathers moulted in West Africa and used confirmatory path analysis to test if isotopic values of winter-grown feathers were related to reproductive success through the mediation of breeding phenology and body condition. We conducted separate analyses for males, females and age classes (yearlings vs ≥ 2 years old). Experienced males wintering in habitats of higher rainfall (as inferred from lower feather δ2H values) were in better body condition and produced more offspring during the subsequent breeding season. In contrast, we did not find any effect of winter habitat on reproductive success of young males or females. These findings provide evidence consistent with a complex causal link between winter habitat quality and subsequent breeding success of long-distance migratory songbirds.