Low migratory connectivity and similar migratory strategies in a shorebird with contrasting wintering population trends in Europe and West Africa.

Migratory shorebird populations are declining worldwide, showing an apparent inability to respond to the interplaying challenges emerging along their flyways. Within the East Atlantic Flyway, non-breeding populations show moderate to strong declines in Sub-Saharan Africa, contrasting with stable or increasing trends in Europe. Local factors are insufficient to explain the opposite tendencies and, therefore, investigating migratory strategies and connectivity of these populations may help identifying the drivers of their demography. We followed the migratory journeys of 20 grey plovers (Pluvialis squatarola) from their wintering grounds in Guinea-Bissau (West Africa), Portugal and France (Europe) using tracking devices. Grey plovers wintering in Africa and Europe were found to share breeding grounds at European Russia and Western Siberia, revealing low migratory connectivity in the Eastern Atlantic population. All individuals followed a "skipping" migratory strategy, flying mostly mid-distance bouts, and using an unexpected large network of stopover sites to re-fuel usually for short periods. We identified 66 different stopover sites along the West African, European and Russian/Siberian coasts. All birds stopped at the Wadden Sea in both migratory periods, highlighting the importance of this region and the risk for a potential bottleneck. Low migratory connectivity and similar migratory strategies shared by grey plovers wintering in Europe and West Africa do not support their contrasting population trends.

GPS tracking analyses reveal finely-tuned shorebird space use and movement patterns throughout the non-breeding season in high-latitude austral intertidal areas.

BACKGROUND: Long-distance migratory birds spend most of their annual cycle in non-breeding areas. During this period birds must meet their daily nutritional needs and acquire additional energy intake to deal with future events of the annual cycle. Therefore, patterns of space use and movement may emerge as an efficient strategy to maintain a trade-off between acquisition and conservation of energy during the non-breeding season. However, there is still a paucity of research addressing this issue, especially in trans-hemispheric migratory birds. METHODS: Using GPS-tracking data and a recently developed continuous-time stochastic process modeling framework, we analyzed fine-scale movements in a non-breeding population of Hudsonian godwits (Limosa haemastica), a gregarious long-distance migratory shorebird. Specifically, we evaluated if these extreme migrants exhibit restricted, shared, and periodic patterns of space use on one of their main non-breeding grounds in southern South America. Finally, via a generalized additive model, we tested if the observed patterns were consistent within a circadian cycle. RESULTS: Overall, godwits showed finely-tuned range-residence and periodic movements (each 24-72 h), being similar between day and night. Remarkably, range-resident individuals segregated spatially into three groups. In contrast, a smaller fraction of godwits displayed unpredictable and irregular movements, adding functional connectivity within the population. CONCLUSIONS: In coastal non-breeding areas where resource availability is highly predictable due to tidal cycles, range-resident strategies during both the day and night are the common pattern in a long-distance shorebird population. Alternative patterns exhibited by a fraction of non-resident godwits provide functional connectivity and suggest that the exploratory tendency may be essential for information acquisition and associated with individual traits. The methodological approach we have used contributes to elucidate how the composition of movement phases operates during the non-breeding season in migratory species and can be replicated in non-migratory species as well. Finally, our results highlight the importance of considering movement as a continuum within the annual cycle.

Mediterranean songbirds show pronounced seasonal variation in thermoregulatory traits.

Addressing the patterns of variation in thermal traits is crucial to better predict the potential effects of climate change on organisms. Here, we assessed seasonal (winter vs summer) adjustments in key thermoregulatory traits in eight Mediterranean-resident songbirds. Overall, songbirds increased whole-animal (by 8%) and mass-adjusted (by 9%) basal metabolic rate and decreased (by 56%) thermal conductance below the thermoneutral zone during winter. The magnitude of these changes was within the lower values found in songbirds from northern temperate areas. Moreover, songbirds increased (by 11%) evaporative water loss within the thermoneutral zone during summer, while its rate of increase above the inflection point of evaporative water loss (i.e., the slope of evaporative water loss versus temperature) decreased by 35% during summer - a value well above that reported for other temperate and tropical songbirds. Finally, body mass increased by 5% during winter, a pattern similar to that found in many northern temperate species. Our findings support the idea that physiological adjustments might enhance the resilience of Mediterranean songbirds to environmental changes, with short-term benefits by saving energy and water under thermally stressful conditions. Nevertheless, not all species showed the same patterns, suggesting different strategies in their thermoregulatory adaptations to seasonal environments.

Human activities favour prolific life histories in both traded and introduced vertebrates.

Species' life histories determine population demographics and thus the probability that introduced populations establish and spread. Life histories also influence which species are most likely to be introduced, but how such 'introduction biases' arise remains unclear. Here, we investigate how life histories affect the probability of trade and introduction in phylogenetic comparative analyses across three vertebrate classes: mammals, reptiles and amphibians. We find that traded species have relatively high reproductive rates and long reproductive lifespans. Within traded species, introduced species have a more extreme version of this same life history profile. Species in the pet trade also have long reproductive lifespans but lack 'fast' traits, likely reflecting demand for rare species which tend to have slow life histories. We identify multiple species not yet traded or introduced but with life histories indicative of high risk of future trade, introduction and potentially invasion. Our findings suggest that species with high invasion potential are favoured in the wildlife trade and therefore that trade regulation is crucial for preventing future invasions.

Heat tolerance limits of Mediterranean songbirds and their current and future vulnerabilities to temperature extremes.

Songbirds are one of the groups most vulnerable to extreme heat events. Although several recent studies have assessed their physiological responses to heat, most of them have focused solely on arid-zone species. We investigated thermoregulatory responses to heat in eight small-sized songbirds occurring in the Mediterranean Basin, where heatwaves are becoming more frequent and intense. Specifically, we determined their heat tolerance limits (HTLs) and evaporative cooling efficiency, and evaluated their current and future vulnerabilities to heat in southwestern Iberia, a Mediterranean climate warming hotspot. To do this, we exposed birds to an increasing profile of air temperatures (Ta) and measured resting metabolic rate (RMR), evaporative water loss (EWL), evaporative cooling efficiency (the ratio between evaporative heat loss and metabolic heat production) and body temperature (Tb). HTL ranged between 40 and 46°C across species, and all species showed rapid increases in RMR, EWL and Tb in response to increasing Ta. However, only the crested lark (Galerida cristata) achieved an evaporative cooling efficiency greater than 1. The studied songbirds currently experience summer Ta maxima that surpass the upper critical temperatures of their thermoneutral zone and even their HTL. Our estimates indicate that five of the eight species will experience moderate risk of lethal dehydration by the end of the century. We argue that the limited heat tolerance and evaporative cooling efficiency of small-sized Mediterranean songbirds make them particularly vulnerable to heatwaves, which will be exacerbated under future climate change scenarios.

Climate change and lithium mining influence flamingo abundance in the Lithium Triangle.

The development of technologies to slow climate change has been identified as a global imperative. Nonetheless, such 'green' technologies can potentially have negative impacts on biodiversity. We explored how climate change and the mining of lithium for green technologies influence surface water availability, primary productivity and the abundance of three threatened and economically important flamingo species in the 'Lithium Triangle' of the Chilean Andes. We combined climate and primary productivity data with remotely sensed measures of surface water levels and a 30-year dataset on flamingo abundance using structural equation modelling. We found that, regionally, flamingo abundance fluctuated dramatically from year-to-year in response to variation in surface water levels and primary productivity but did not exhibit any temporal trends. Locally, in the Salar de Atacama-where lithium mining is focused-we found that mining was negatively correlated with the abundance of two of the three flamingo species. These results suggest continued increases in lithium mining and declines in surface water could soon have dramatic effects on flamingo abundance across their range. Efforts to slow the expansion of mining and the impacts of climate change are, therefore, urgently needed to benefit local biodiversity and the local human economy that depends on it.

Urohidrosis as an overlooked cooling mechanism in long-legged birds.

Behavioural thermoregulation could buffer the impacts of climate warming on vertebrates. Specifically, the wetting of body surfaces and the resulting evaporation of body fluids serves as a cooling mechanism in a number of vertebrates coping with heat. Storks (Ciconiidae) frequently excrete onto their legs to prevent overheating, a phenomenon known as urohidrosis. Despite the increasingly recognised role of bare and highly vascularised body parts in heat exchange, the ecological and evolutionary determinants of urohidrosis have been largely ignored. We combine urohidrosis data from a scientifically curated media repository with microclimate and ecological data to investigate the determinants of urohidrosis in all extant stork species. Our phylogenetic generalised linear mixed models show that high temperature, humidity and solar radiation, and low wind speed, promote the use of urohidrosis across species. Moreover, species that typically forage in open landscapes exhibit a more pronounced use of urohidrosis than those mainly foraging in waterbodies. Substantial interspecific variation in temperature thresholds for urohidrosis prevalence points to different species vulnerabilities to high temperatures. This integrated approach that uses online data sources and methods to model microclimates should provide insight into animal thermoregulation and improve our capacity to make accurate predictions of climate change's impact on biodiversity.

Bird migration: Flying high to avoid overheating?

Acts of flight have always baffled humans. Recent tracking studies of birds only add to the awe and mystery. A new study shows that migrating great snipes fly high - over two kilometres high - during the day and lower at night. Why?

Migrant birds and mammals live faster than residents.

Billions of vertebrates migrate to and from their breeding grounds annually, exhibiting astonishing feats of endurance. Many such movements are energetically costly yet there is little consensus on whether or how such costs might influence schedules of survival and reproduction in migratory animals. Here we provide a global analysis of associations between migratory behaviour and vertebrate life histories. After controlling for latitudinal and evolutionary patterns, we find that migratory birds and mammals have faster paces of life than their non-migratory relatives. Among swimming and walking species, migrants tend to have larger body size, while among flying species, migrants are smaller. We discuss whether pace of life is a determinant, consequence, or adaptive outcome, of migration. Our findings have important implications for the understanding of the migratory phenomenon and will help predict the responses of bird and mammal species to environmental change.

Avian major histocompatibility complex copy number variation is associated with helminth richness.

Genes of the major histocompatibility complex (MHC) play a key role in the adaptive immunity of vertebrates, as they encode receptors responsible for antigen recognition. Evolutionary history of the MHC proceeded through numerous gene duplications, which increase the spectrum of pathogens recognized by individuals. Although pathogen-mediated selection is believed to be a primary driver of MHC expansion over evolutionary times, empirical evidence for this association is virtually lacking. Here, we used an extensive dataset on MHC class II copy number variation in non-passerine birds to test for an evolutionary correlation with helminth parasite richness. As expected, our phylogenetically-informed modelling revealed a positive association between MHC copy number and total helminth richness, even after controlling for a broad spectrum of ecological and life-history traits. Thus, total helminth richness appears to be the most important correlate of MHC copy number, supporting a leading role of pathogen-mediated selection in the evolution of MHC in birds. Our results provide some of the first, although correlative, evidence linking parasitism to interspecific variation in MHC copy number among birds.

Oxidative status and metabolic profile in a long-lived bird preparing for extreme endurance migration.

The high metabolic activity associated with endurance flights and intense fuelling of migrant birds may produce large quantities of reactive oxygen species, which cause oxidative damage. Yet it remains unknown how long-lived birds prepare for oxidative challenges prior to extreme flights. We combined blood measurements of oxidative status and enzyme and fat metabolism in Hudsonian godwits (Limosa haemastica, a long-lived shorebird) before they embarked on non-stop flights longer than 10,000 km during their northbound migrations. We found that godwits increased total antioxidant capacity (TAC) and reduced oxidative damage (TBARS) as the pre-migratory season progressed, despite higher basal metabolic rates before departure. Elevations in plasma ß-hydroxybutyrate and uric acid suggest that lipid and protein breakdown supports energetic requirements prior to migration. Significant associations between blood mitochondrial cytochrome-c oxidase and plasma TAC (negative) and TBARS (positive) during winter indicate that greater enzyme activity can result in greater oxidative damage and antioxidant responses. However enzyme activity remained unchanged between winter and premigratory stages, so birds may be unable to adjust metabolic enzyme activity in anticipation of future demands. These results indicate that godwits enhance their oxidative status during migratory preparation, which might represent an adaptation to diminish the physiological costs of long-distance migration.

Day and night use of habitats by northern pintails during winter in a primary rice-growing region of Iberia.

Loss of natural wetlands is a global phenomenon that has severe consequences for waterbird populations and their associated ecosystem services. Although agroecosystems can reduce the impact of natural habitat loss, drivers of use of such artificial habitats by waterbirds remain poorly understood. Using the cosmopolitan northern pintail Anas acuta as a model species, we monitored home-range and fine-scale resource selection across the agricultural landscape. Individuals were tracked using GPS-GSM transmitters, and a suite of environmental and landscape features were measured throughout the winter seasons. Spatial patterns of habitat use were analysed using generalized linear mixed effect models by integrating field-observations with GPS telemetry. All birds used rice fields as foraging grounds at night and commuted to an adjacent reservoir to roost during daylight. Home-ranges and maximum foraging distances of nocturnally foraging birds increased with decreasing availability of flooded fields, and were positively correlated with moonlight levels. Birds selected flooded rice paddies (water depth range: 9-21 cm) with standing stubble and substrate with pebbles smaller than 0.5 cm in diameter. Density of rice seeds, rice paddy size, and other environmental and landscape features did not emerge as significant predictors. Our findings indicate that nocturnal foraging of northern pintails within rice fields is driven primarily by straw manipulation, water level and substrate pebble size. Thus, the presence of standing stubble in flooded paddies with soft bottoms should be prioritized to improve foraging areas for dabbling ducks. These management procedures in themselves would not increase economic costs or affect rice production and could be applied for dabbling-duck conservation throughout the world.

Micro- and macroparasite species richness in birds: The role of host life history and ecology.

Identifying the factors shaping variation in parasite diversity among host species is crucial to understand wildlife diseases. Although micro- and macroparasites may exert different selective pressures on their hosts, studies investigating the determinants of parasite species richness in animals have rarely considered this divide. Here, we investigated the role of host life history and ecology in explaining the species richness of helminths (macroparasites) and haemosporidians (microparasites) in birds world-wide. We collated data from multiple global datasets on diverse bird traits (longevity, body mass, coloniality, migration distance/tendency, geographic range size and dietary and habitat breadths) and the species richness of their helminth and haemosporidian parasites. We tested predictors of helminth and haemosporidian parasite richness using phylogenetic generalized linear mixed models in a Bayesian framework. We found that, after controlling for research effort and host phylogeny, the richness of helminths, but not of haemosporidians, increased with host longevity, range size, migration distance and dietary breadth. Overall, these correlates were also important across different helminth groups (acanthocephalans, cestodes, nematodes and trematodes), and two additional ones (body mass, coloniality) emerged as important for cestodes and acanthocephalans. We propose that long life spans may promote the diversity of helminth parasite assemblages over evolutionary time, thus resulting in richer helminth faunas. Similarly, longer-distance migrations, larger ranges and broader dietary breadths are likely to lead to greater encounter rates and the accumulation of trophically transmitted helminths. In contrast, vector-borne haemosporidians may be influenced more by factors related to vector ecology than by the host traits included in the analyses. The lack of strong associations between haemosporidian species richness and host characteristics emphasizes the need to find appropriate traits to model the distribution and diversity of parasites with different environmental preferences in order to anticipate disease emergence risks associated with global change.

Assessing the effects of human activities on the foraging opportunities of migratory shorebirds in Austral high-latitude bays.

Human presence at intertidal areas could impact coastal biodiversity, including migratory waterbird species and the ecosystem services they provide. Assessing this impact is therefore essential to develop management measures compatible with migratory processes and associated biodiversity. Here, we assess the effects of human presence on the foraging opportunities of Hudsonian godwits (Limosa haemastica, a trans-hemispheric migratory shorebird) during their non-breeding season on Chiloé Island, southern Chile. We compared bird density and time spent foraging in two similar bays with contrasting disturbance levels: human presence (mostly seaweed harvesters accompanied by dogs) was on average 0.9±0.4 people per 10 ha in the disturbed bay, whereas it was negligible (95% days absent) in the non-disturbed bay. Although overall abundances were similar between bays, godwit density was higher in the non-disturbed bay throughout the low tide period. Both days after the start of the non-breeding season and tidal height significantly affected godwit density, with different effects in either bay. Time spent foraging was significantly higher in the non-disturbed bay (86.5±1.1%) than in the disturbed one (81.3±1.4%). As expected, godwit density significantly decreased with the number of people and accompanying dogs in the disturbed bay. Our results indicate that even a low density of people and dogs can significantly reduce the foraging opportunities of shorebirds. These constraints, coupled with additional flushing costs, may negatively affect godwits' pre-migratory fattening. Hence, as a first step we suggest limiting human presence within bays on Chiloé to 1 person per 10 ha and banning the presence of accompanying dogs in sensitive conservation areas.

Wetland salinity induces sex-dependent carry-over effects on the individual performance of a long-distance migrant.

Salinization is having a major impact on wetlands and its biota worldwide. Specifically, many migratory animals that rely on wetlands are increasingly exposed to elevated salinity on their nonbreeding grounds. Experimental evidence suggests that physiological challenges associated with increasing salinity may disrupt self-maintenance processes in these species. Nonetheless, the potential role of salinity as a driver of ecological carry-over effects remains unstudied. Here, we investigated the extent to which the use of saline wetlands during winter - inferred from feather stable isotope values - induces residual effects that carry over and influence physiological traits relevant to fitness in black-tailed godwits Limosa limosa limosa on their northward migration. Overwintering males and females were segregated by wetland salinity in West Africa, with females mostly occupying freshwater wetlands. The use of these wetlands along a gradient of salinities was associated with differences in immune responsiveness to phytohaemagglutinin and sized-corrected body mass in godwits staging in southern Europe during northward migration - 3,000 km from the nonbreeding grounds - but in males only. These findings provide a window onto the processes by which wetland salinity can induce carry-over effects and can help predict how migratory species should respond to future climate-induced increases in salinity.