Dynamics of stored lipids in fall migratory monarch butterflies (Danaus plexippus): Nectaring in northern Mexico allows recovery from droughts at higher latitudes.

The eastern population of the North American monarch butterfly (Danaus plexippus) overwinters from November through March in the high-altitude (3000 m+) forests of central Mexico during which time they rely largely on stored lipids. These are acquired during larval development and the conversion of sugars from floral nectar by adults. We sampled fall migrant monarchs from southern Canada through the migratory route to two overwintering sites in 2019 (n = 10 locations), 2020 (n = 8 locations) and 2021 (n = 7 locations). Moderate to extreme droughts along the migratory route were expected to result in low lipid levels in overwintering monarchs but our analysis of lipid levels of monarchs collected at overwintering sites indicated that in all years most had high levels of lipids prior to winter. Clearly, a significant proportion of lipids were consistently acquired in Mexico during the last portion of the migration. Drought conditions in Oklahoma, Texas and northern Mexico in 2019 resulted in the lowest levels of lipid mass and wing loading observed in that year but with higher levels at locations southward in Mexico to the overwintering sites. Compared with 2019, lipid levels increased during the 2020 and 2021 fall migrations but were again higher during the Mexican portion of the migration than for Oklahoma and Texas samples, emphasizing a recovery of lipids as monarchs advanced toward the overwintering locations. In all 3 years, body water was highest during the Canada-USA phase of migration but then declined during the nectar foraging phase in Mexico before recovering again at the overwintering sites. The increase in mass and lipids from those in Texas to the overwintering sites in Mexico indicates that nectar availability in Mexico can compensate for poor conditions experienced further north. Our work emphasizes the need to maintain the floral and therefore nectar resources that fuel both the migration and storage of lipids throughout the entire migratory route.

A feather hydrogen (δ2H) isoscape for Brazil.

Spatial patterns of stable isotopes in animal tissues or "isoscapes" can be used to investigate animal origins in a range of ecological and forensic investigations. Here, we developed a feather hydrogen isotope (δ2Hf) isoscape for Brazil based on 192 samples of feathers from the family Thraupidae from scientific collections. Raw values of δ2Hf ranged from -107.3 to +5.0‰, with higher values at the Caatinga biome (northeast Brazil) and lower values at the Amazon and Pantanal. A Random Forest (RF) method was used to model the spatial surface, using a range of environmental data as auxiliary variables. The RF model indicated a negative relationship between δ2Hf and Mean Annual Precipitation, Precipitation in the Warmest Quarter, and Annual Temperature Range and positive relationships for amount-weighted February-April precipitation δ2H (δ2Hp(Feb-April)) and Mean Annual Solar Radiation. Modelled δ2Hf values ranged from -85.7 to -13.6‰. Ours is the first δ2Hf isoscape for Brazil that can greatly assist our understanding of both ecological and biogeochemical processes controlling spatial variation in δ2H for this region. This isoscape can be used with caution, due to its poor predictive power (as found in other tropical regions) and can benefit from new sample input, new GNIP data, ecological and physiological studies, and keratin standard material better encompassing the range in feather samples from Brazil. So, we encourage new attempts to build more precise feather H isoscapes, as well as isoscapes based on other elements.

Rapid adjustments of migration and life history in hemisphere-switching cliff swallows.

Many aspects of bird migration are necessarily innate.1 However, the extent of deterministic genetic control, environmental influence, and individual decision making in the control of migration remains unclear.2-8 Globally, few cases of rapid and dramatic life-history changes resulting in novel migration strategies are known. An example is latitudinal trans-hemispheric breeding colonization, whereby a subpopulation suddenly begins breeding on its non-breeding range.9-13 These life-history reversals demand concomitant changes in the timing of migration, feather molt, and breeding if the population is to remain viable.13 Cliff swallows, Petrochelidon pyrrhonota, are long-distance migrants that breed in North America and spend the non-breeding season mostly in South America.14 However, in 2015, a small population switched hemispheres by breeding successfully in Argentina,9 over 8,000 km from the nearest potential source, after presumably failed attempts.15,16 This provided a unique chance to characterize the early mechanisms of change in migratory behavior and phenology and to assess the possibility of double breeding. We tracked cliff swallows with geolocators following their second and fourth breeding seasons in Argentina, documenting inverted seasonality, three new migratory patterns and non-breeding areas (North America, Mesoamerica, and South America), and a shift of molt phenology by approximately 6 months, all possibly arising within a single generation. These birds did not practice migratory double breeding, although some spent the boreal summer in the traditional breeding range. Our data show that fundamental phenological changes occurred very rapidly during colonization and that phenotypic plasticity can underlie profound changes in the life histories of migratory birds.

Migratory connectivity then and now: a northward shift in breeding origins of a long-distance migratory bird wintering in the tropics.

Temporal variation in the connectivity of populations of migratory animals has not been widely documented, despite having important repercussions for population ecology and conservation. Because the long-distance movements of migratory animals link ecologically distinct and geographically distant areas of the world, changes in the abundance and migratory patterns of species may reflect differential drivers of demographic trends acting over various spatial scales. Using stable hydrogen isotope analyses (δ2H) of feathers from historical museum specimens and contemporary samples obtained in the field, we provide evidence for an approximately 600 km northward shift over 45 years in the breeding origin of a species of songbird of major conservation concern (blackpoll warbler, Setophaga striata) wintering in the foothills of the eastern Andes of Colombia. Our finding mirrors predictions of range shifts for boreal-breeding species under warming climate scenarios and habitat loss in the temperate zone, and underscores likely drivers of widespread declines in populations of migratory birds. Our work also highlights the value of natural history collections to document the effects of global change on biodiversity.

Earlier and slower or later and faster: Spring migration pace linked to departure time in a Neotropical migrant songbird.

Migratory birds travel vast distances and the timing of migratory flights can affect survival and the ability to reproduce. For Neotropical migrant songbirds, early spring departure from wintering sites, early arrival to the breeding grounds and higher reproductive success have been related to the use of suitable habitats and environmental conditions during the non-breeding season. However, how migratory strategies are shaped by winter habitat choice is largely unknown due to the general inability to track birds from specific wintering habitats to stopovers or breeding destinations. We assessed how winter habitat (native forest vs. shade-grown coffee plantations) relates to spring departure date and migration pace in Swainson's Thrush Catharus ustulatus. We also determined the effect of departure date and total migration duration on the arrival date of birds detected near or within their breeding range. We used a novel application of Motus radiotelemetry arrays to track individuals from their wintering grounds in the Andes of South America along their migratory journey to North America. We found variation in migratory strategies between habitats, with birds wintering in native forest departing later than birds in coffee. We present isotopic evidence for native forest being of higher quality than shade-coffee for Swainson's Thrush and hypothesize that moister conditions in forest, as shown by stable isotope (δ13 C) analysis of thrush whole blood, provides favourable pre-migratory conditions allowing birds to delay departure from wintering grounds. Habitat, between-site and -year variation in departure date, suggests that birds made facultative adjustments to winter habitat quality and environmental conditions. Independent of habitat, birds that departed later migrated faster and this pattern was maintained along the migration route (n = 44). Migrating earlier and slower or later and faster was unlikely to result in significant differences in arrival time to breeding destinations. Our findings reveal underappreciated complexity in migratory decisions by long-distance migrants that contrast with the current paradigm of earlier departures and arrival from optimal habitats. The next step is to understand the relative fitness benefits of early versus late schedules or whether each strategy is an equally good response to experienced conditions.

Las aves migratorias viajan largas distancias y la fecha de sus vuelos migratorios puede afectar su supervivencia y habilidad para reproducirse. Para las aves migratorias Neotropicales canoras, salir más temprano de los sitios de invernada, llegar más temprano a los sitios de reproducción y tener mayor éxito reproductivo, está relacionado con el uso de hábitats y condiciones medioambientales adecuadas durante la época no reproductiva. Sin embargo, la forma en que las estrategias migratorias son moldeadas por la selección del hábitat invernal es en gran parte desconocida debido a la inhabilidad general de rastrear las aves desde hábitats de invernada específicos hasta los sitios de descanso o de reproducción. Determinamos la relación entre el hábitat de invierno (bosque nativo vs. plantaciones de café con sombra), la fecha de salida durante la migración de primavera y el número de días usados durante la migración por la mirla buchipecosa Catharus ustulatus. También determinamos el efecto de la fecha de salida y la duración total de la migración en la fecha de llegada de las aves detectadas cerca o dentro de su rango de reproducción. Usamos la novedosa red de radio telemetría Motus para rastrear individuos desde sus sitios de invernada en las montañas de los Andes en Sudamérica a lo largo de su viaje migratorio hasta Norte América. Encontramos variación en las estrategias migratorias entre hábitats, las aves invernando en bosque nativo salieron más tarde que las aves invernando en cafetales. Presentamos evidencia isotópica, que sugiere que el bosque nativo es de mejor calidad que las plantaciones de café con sombra. Hipotetizamos que condiciones más húmedas en el bosque, indicadas por análisis de isotopos estables (δ13 C) de la sangre de las mirlas, proveen condiciones favorables antes de la migración permitiendo a las aves retrasar su salida de los territorios de invernada. La variación en la fecha de salida entre hábitats, años y sitios, sugiere que las aves hicieron ajustes facultativos a la calidad del hábitat de invierno y a las condiciones medio ambientales. Independientemente del hábitat, las aves que salieron más tarde migraron más rápido, y este patrón fue mantenido a lo largo de la ruta migratoria (n = 44). Fue poco probable que migrar más temprano y más lento o más tarde y más rápido resultara en diferencias significativas en la fecha de llegada a los sitios de reproducción. Nuestros resultados revelan una subestimada complejidad en las decisiones migratorias de las aves y contrastan con el paradigma actual de salida y llegada más temprana en aves que usan hábitats óptimos. El siguiente paso es entender los beneficios relativos de migrar temprano versus tarde en el desempeño o si cada estrategia es una respuesta igualmente buena a las condiciones experimentadas.

Tracking data and retrospective analyses of diet reveal the consequences of loss of marine subsidies for an obligate scavenger, the Andean condor.

Over the last century, marine mammals have been dramatically reduced in the world's oceans. We examined evidence that this change caused dietary and foraging pattern shifts of the Andean condor (Vultur gryphus) in Patagonia. We hypothesized that, after the decrease in marine mammals and the increase in human use of coastlines, condor diet changed to a more terrestrial diet, which in turn influenced their foraging patterns. We evaluated the diet by means of stable isotope analysis (δ13C, δ15N and δ34S) of current (last decade) and historical (1841-1933) feathers. We further evaluated the movement patterns of 23 condors using satellite tracking of individuals. Condors reduced their use of marine-derived prey in recent compared with historical times from 33 ± 13% to less than 8 ± 3% respectively; however, they still breed close to the coast. The average distance between the coast and nests was 62.5 km, but some nests were located close to the sea (less than 5 km). Therefore, some birds must travel up to 86 km from nesting sites, crossing over the mountain range to find food. The worldwide reduction in marine mammal carcasses, especially whales, may have major consequences on the foraging ecology of scavengers, as well as on the flux of marine inputs within terrestrial ecosystems.

Assessing seasonal changes in animal diets with stable-isotope analysis of amino acids: a migratory boreal songbird switches diet over its annual cycle.

Tools to study seasonal changes in animal diets are needed to address a wide range of ecological questions. This is especially true of migratory animals that experience distinct environments where diets may be substantially different. However, tracking diets of individuals that move vast distances has proven difficult. Compound-specific isotope analysis has emerged as a valuable tool to study diets but has been little used to study dietary changes of migratory animals. Using this technique, we quantify seasonal variation in the annual diet of a migratory songbird (gray-cheeked thrush, Catharus minimus) and test the hypothesis that migrants change their diet in response to the energetic requirements of different periods of the annual cycle. By measuring δ13C and δ15N values of amino acids from feathers grown on the breeding grounds, blood formed during migration and claw grown on the wintering grounds, we found that migration is associated with greater consumption of fruit, compared to the breeding or wintering periods. This was confirmed by the lower trophic position of blood compared to feather and claw, by a decrease in the δ15N value of the source amino acid phenylalanine in blood as a function of days of stopover, and by the positive correlation between δ15N and δ13C values of phenylalanine in blood, and not in feather or claw. This study illustrates how isotopic analysis of amino acids can contribute to understand food webs, seasonal dietary changes and metabolic routing of nutrients in migratory animals.

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.

Long-Distance Range Expansion and Rapid Adjustment of Migration in a Newly Established Population of Barn Swallows Breeding in Argentina.

When bird populations spread, long-distance pioneering populations are often backfilled by a more slowly advancing front [1-3]. The Barn Swallow Hirundo rustica, a globally distributed passerine [4, 5], expanded its breeding range an exceptional 7,000 km when it began breeding 35 years ago in its regular wintering range in Argentina [6], subsequently expanding over 500 km from its starting point [7-11]. Trans-hemispheric breeding attempts have occurred previously in related swallows [12-14], but only this colonization has lasted. Comparative studies of birds show a remarkable diversity in patterns of change in migratory habits [15-21], and these Argentine-breeding swallows might retain ancestral patterns, breeding in Argentina but returning to North America for the austral winter. Feather isotopes from these birds are consistent with the alternative possibility that they migrate no farther than northern South America [22]. Because isotopic patterns cannot definitively distinguish these alternatives, we pursued a solar geolocator study [23, 24] to do so. Data from nine tagged birds show conclusively that Barn Swallows breeding in Argentina have rapidly changed their movements to migrate no farther north in austral winter than northern South America. The phenology of the annual cycles of molt, migration, and breeding for these Argentine-breeding swallows have all shifted by about 6 months, and we suggest that stimulatory day lengths and the proliferation of nesting substrates facilitated their colonization.

Regional climate on the breeding grounds predicts variation in the natal origin of monarch butterflies overwintering in Mexico over 38 years.

Addressing population declines of migratory insects requires linking populations across different portions of the annual cycle and understanding the effects of variation in weather and climate on productivity, recruitment, and patterns of long-distance movement. We used stable H and C isotopes and geospatial modeling to estimate the natal origin of monarch butterflies (Danaus plexippus) in eastern North America using over 1000 monarchs collected over almost four decades at Mexican overwintering colonies. Multinomial regression was used to ascertain which climate-related factors best-predicted temporal variation in natal origin across six breeding regions. The region producing the largest proportion of overwintering monarchs was the US Midwest (mean annual proportion = 0.38; 95% CI: 0.36-0.41) followed by the north-central (0.17; 0.14-0.18), northeast (0.15; 0.11-0.16), northwest (0.12; 0.12-0.16), southwest (0.11; 0.08-0.12), and southeast (0.08; 0.07-0.11) regions. There was no evidence of directional shifts in the relative contributions of different natal regions over time, which suggests these regions are comprising the same relative proportion of the overwintering population in recent years as in the mid-1970s. Instead, interannual variation in the proportion of monarchs from each region covaried with climate, as measured by the Southern Oscillation Index and regional-specific daily maximum temperature and precipitation, which together likely dictate larval development rates and food plant condition. Our results provide the first robust long-term analysis of predictors of the natal origins of monarchs overwintering in Mexico. Conservation efforts on the breeding grounds focused on the Midwest region will likely have the greatest benefit to eastern North American migratory monarchs, but the population will likely remain sensitive to regional and stochastic weather patterns.

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 (δ2H) measurements as an indicator of latitude of origin at the start of fall migration. On average, uninfected monarchs had lower δ2H 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.

Large-scale spatial variation in feather corticosterone in invasive house sparrows (Passer domesticus) in Mexico is related to climate.

Ecologists frequently use physiological tools to understand how organisms cope with their surroundings but rarely at macroecological scales. This study describes spatial variation in corticosterone (CORT) levels in feathers of invasive house sparrows (Passer domesticus) across their range in Mexico and evaluates CORT-climate relationships with a focus on temperature and precipitation. Samples were collected from 49 sites across Mexico. Feather CORT (CORTf) was measured using methanol-based extraction and radioimmunoassay. Relationships between CORTf and spatial and climate variables were examined using simple linear regressions. Ordination was used on climate data, CORTf was plotted against the resulting axes, and univariate regression trees were used to identify important predictors of CORTf. Universal kriging interpolation was used to illustrate spatial variation in CORTf across Mexico. Correlations with ordination axes showed that high CORTf was associated with low precipitation during the rainy season and low dry season temperatures. Specifically, CORTf was negatively related to May precipitation and January and July minimum temperatures, and positively related to April deuterium excess and June minimum temperatures. CORTf was higher in second-year birds compared to after-hatch years and after-second years. House sparrows had higher CORTf levels in the hot, dry, north-central region of Mexico, and CORTf was negatively related to temperature and precipitation. House sparrows molt primarily from August-September but climate conditions throughout the year were important predictors of CORTf, suggesting that conditions outside of molt can carry over to influence energetics during feather growth. These data suggest that dry conditions are challenging for house sparrows in Mexico, supporting previous work showing that precipitation is an important predictor of broad-scale CORT variation. This work highlights the utility of CORTf for evaluating the influence of physiology on current avian range limits; furthermore, these data may allow us to predict future changes in species distributions.

Switching hemispheres: a new migration strategy for the disjunct Argentinean breeding population of Barn Swallow (Hirundo rustica).

BACKGROUND: Barn Swallows (Hirundo rustica) breed almost exclusively in the Northern Hemisphere. However, since the early 1980's, a small disjunct breeding population has become established in eastern Argentina, presumably by birds previously derived from those breeding in North America. Currently, it is unknown where these individuals go following breeding and how they have adjusted to a reversal in phenology. Their austral wintering period corresponds to the breeding period of the northern ancestral population and so they can potentially return to these more traditional breeding sites or they may occupy other South American wintering regions left vacant by conspecifics returning to the Northern Hemisphere. PRINCIPAL FINDINGS: We used a three-isotope (δ(13)C, δ(15)N, δ(2)H) approach to investigate potential wintering areas in Central and South America of individuals breeding in Argentina. Feather isotope values differed from those expected and measured at local breeding sites in Argentina indicating molt after the austral breeding period and away from the breeding grounds. Potential molting origins were identified applying likelihood-based assignment methods to a δ(2)H isoscape for South America and dichotomous prior information on the distribution of C3 and C4 vegetation types based on modeled vegetation-δ(13)C values. Barn Swallows now breeding in Argentina have changed their migratory behavior but presumably use the same cues as those used by the ancestral population, molting their feathers during the austral winter, likely in north-eastern South America.

Lekking birds in a tropical forest forego sex for migration.

Facultative, partially migratory animals provide a contemporary window into the evolution of migration, offering rare opportunities to examine the life-history trade-offs associated with migration. For the first time, to our knowledge, we describe the nature of these trade-offs, using a lek-breeding tropical bird, the white-ruffed manakin (Corapipo altera). Previous evidence indicated that weather drives post-breeding migration to lower elevations bringing condition-related benefits. Using elevation-sensitive stable isotope measurements and more than 1200 h of behavioural observations, we show that male manakins which migrate incur costs of diminished social status and matings with females the following breeding season. Because migratory tendency depends on inter-annual variation in weather, physical costs of displays and breeding prospects the following year, migratory decisions are subject to both natural and sexual selection, with the outcome of such decisions linked to changing climatic regimes.

Migratory double breeding in Neotropical migrant birds.

Neotropical migratory songbirds typically breed in temperate regions and then travel long distances to spend the majority of the annual cycle in tropical wintering areas. Using stable-isotope methodology, we provide quantitative evidence of dual breeding ranges for 5 species of Neotropical migrants. Each is well known to have a Neotropical winter range and a breeding range in the United States and Canada. However, after their first bout of breeding in the north, many individuals migrate hundreds to thousands of kilometers south in midsummer to breed a second time during the same summer in coastal west Mexico or Baja California Sur. They then migrate further south to their final wintering areas in the Neotropics. Our discovery of dual breeding ranges in Neotropical migrants reveals a hitherto unrealized flexibility in life-history strategies for these species and underscores that demographic models and conservation plans must consider dual breeding for these migrants.

Trophic partitioning in tropical rain forest birds: insights from stable isotope analysis.

Bird communities reach their highest taxonomic and trophic diversity in tropical rain forest, but the use of different foraging strategies to meet food requirements in such competitive environments is poorly understood. Conventional dietary analyses are poorly suited to investigate dietary patterns in complex systems. We used stable carbon ((13)C/(12)C) and nitrogen ((15)N/(14)N) isotope analysis of whole blood to examine avian trophic patterns and sources of diet in the tropical rain forest of Los Tuxtlas, Veracruz, Mexico. We used stable nitrogen isotope analysis to delineate trophic levels, and stable carbon isotope analysis to distinguish the relative contribution of C-3 and CAM/C-4 ultimate sources of proteins to diets. There was large inter- and intraspecific variation in whole blood delta(13)C and delta(15)N values in 23 species of birds. Stable nitrogen isotope analysis separated birds into several trophic levels, including species that obtained their dietary protein mostly from plants, insects or a combination of both food sources. Stable carbon isotope analysis showed that most birds fed on C3-based foods but Stub-tailed Spadebills (Platyrinchus cancrominus) included C-3- and C-4/CAM-specialist individuals. Our analyses provided insights into the nutritional contribution of plant and animal sources of protein and distinguish their photosynthetic origin over relatively long average time periods.

Stable isotopes as indicators of altitudinal distributions and movements in an Ecuadorean hummingbird community.

Altitudinal migration and dispersal is an important component of the life history of several temperate and tropical birds but remains poorly understood due to the limited success of mark and recapture techniques. Stable isotopes of hydrogen (deltaD) in rainfall, and to a lesser extent, carbon (delta13C) in plants are known to change with altitude and hence may provide the basis of a technique for tracking the altitudinal movements in birds and other wildlife. We investigated the potential for this technique by measuring delta13C, deltaD, and delta15N values in tail feathers of eight species of hummingbirds ( Phaethornis malaris, P. syrmatophorus, P. guy, Adelomyia melanogenys, Coeligena torquata, C. lutetiae, Metallura baroni, M. williami) along an altitudinal gradient (300-3,290 m asl) in the Andes Mountains of Ecuador. Feather delta13C and deltaD values were correlated and each changed significantly with elevation above 400 m. In general, we found good agreement between feather deltaD values and those predicted from a generalized relationship of precipitation and surface water deltaD with altitude. Similarly, feather delta13C values showed an enrichment of approximately 1.5 per thousand per 1,000 m over the linear portion of the elevational response. Stable-nitrogen isotope values were variable, and so did not provide useful information on elevation in birds, apart from trophic effects. Overall there appears to be good potential for using the (deltaD, delta13C) stable isotope approach to track altitudinal movements and to elucidate previously unrecognized patterns of life history variation in both temperate and tropical species that migrate across elevational isotopic gradients.