Migration tactics and connectivity of a Nearctic-Neotropical migratory shorebird.

During long-distance spring migrations, birds may rest and refuel at numerous stopover sites while minimizing the time to reach the breeding grounds. If habitat is limited along the migration route, pre-breeding birds optimize flight range by having longer stopovers at higher quality sites compared to poorer quality sites. Stopover duration also depends on distance remaining to breeding grounds, ecological barriers and individual characteristics. We assessed spring migration tactics and connectivity of a Nearctic-Neotropical migratory shorebird, the semipalmated sandpiper Calidris pusilla, at two sites with known relative habitat quality on the northern Gulf of Mexico (NGOM) coast, the first land encountered after crossing the Gulf of Mexico (GOM). We used automated radio telemetry (Motus) to estimate stopover duration and probability of departure. Migration speed was estimated for individuals detected at subsequent receivers on the Motus network. To measure migratory connectivity, we used morphometrics and the Motus network to assign general breeding regions. Additionally, feather stable isotope ratios of C and N provided coarse information about overwintering regions. Stopover duration declined with higher fuel loads at capture as expected under a time-minimizing strategy. After accounting for fuel load, stopover duration was approximately 40% longer at the higher quality site. We found no detectable effect of age, sex or breeding location on stopover behaviour. The probability of departure was strongly affected by humidity and also by tailwind and weather conditions. Birds stopping at the higher quality site had earlier apparent arrival to the breeding grounds. The Louisiana coast is an apparent stopover hub for this species, since the individuals were departing to range-wide breeding regions and isotope values suggested birds were also using widespread wintering regions. Our study shows how high-quality, coastal wetlands along the NGOM coast serve a critical role in the annual cycle of a migratory shorebird. Stopover behaviour indicated that high-quality habitat may be limited for this species during spring migration. As threats to the GOM coast increase, protection of these already limited wetlands is vitally important.

Light-level geolocator analyses: A user's guide.

Light-level geolocator tags use ambient light recordings to estimate the whereabouts of an individual over the time it carried the device. Over the past decade, these tags have emerged as an important tool and have been used extensively for tracking animal migrations, most commonly small birds. Analysing geolocator data can be daunting to new and experienced scientists alike. Over the past decades, several methods with fundamental differences in the analytical approach have been developed to cope with the various caveats and the often complicated data. Here, we explain the concepts behind the analyses of geolocator data and provide a practical guide for the common steps encompassing most analyses - annotation of twilights, calibration, estimating and refining locations, and extraction of movement patterns - describing good practices and common pitfalls for each step. We discuss criteria for deciding whether or not geolocators can answer proposed research questions, provide guidance in choosing an appropriate analysis method and introduce key features of the newest open-source analysis tools. We provide advice for how to interpret and report results, highlighting parameters that should be reported in publications and included in data archiving. Finally, we introduce a comprehensive supplementary online manual that applies the concepts to several datasets, demonstrates the use of open-source analysis tools with step-by-step instructions and code and details our recommendations for interpreting, reporting and archiving.

A genoscape-network model for conservation prioritization in a migratory bird.

Migratory animals are declining worldwide and coordinated conservation efforts are needed to reverse current trends. We devised a novel genoscape-network model that combines genetic analyses with species distribution modeling and demographic data to overcome challenges with conceptualizing alternative risk factors in migratory species across their full annual cycle. We applied our method to the long distance, Neotropical migratory bird, Wilson's Warbler (Cardellina pusilla). Despite a lack of data from some wintering locations, we demonstrated how the results can be used to help prioritize conservation of breeding and wintering areas. For example, we showed that when genetic, demographic, and network modeling results were considered together it became clear that conservation recommendations will differ depending on whether the goal is to preserve unique genetic lineages or the largest number of birds per unit area. More specifically, if preservation of genetic lineages is the goal, then limited resources should be focused on preserving habitat in the California Sierra, Basin Rockies, or Coastal California, where the 3 most vulnerable genetic lineages breed, or in western Mexico, where 2 of the 3 most vulnerable lineages overwinter. Alternatively, if preservation of the largest number of individuals per unit area is the goal, then limited conservation dollars should be placed in the Pacific Northwest or Central America, where densities are estimated to be the highest. Overall, our results demonstrated the utility of adopting a genetically based network model for integrating multiple types of data across vast geographic scales and better inform conservation decision-making for migratory animals.

Un Modelo de Redes de Panorama Poblacional para la Priorización de la Conservación de un Ave Migratoria Resumen Los animales migratorios están pasando por una declinación mundial y se requieren esfuerzos coordinados de conservación para revertir las tendencias actuales. Diseñamos un modelo novedoso de redes de panorama poblacional que combina el análisis genético con el modelado de la distribución de especies y los datos demográficos para sobreponerse a los obstáculos con la conceptualización de los factores alternativos de riesgo en las especies migratorias durante su ciclo anual completo. Aplicamos nuestro método al chipe de corona negra (Cardellina pusilla), un ave migratoria neotropical que recorre largas distancias. A pesar de la falta de datos de algunas localidades de invernación, mostramos cómo pueden usarse los resultados para ayudar a priorizar la conservación de las áreas de reproducción y de invernación. Por ejemplo, mostramos que cuando se consideraron en conjunto los resultados del modelado genético, demográfico y de redes queda claro que las recomendaciones de conservación diferirán dependiendo de si el objetivo es preservar linajes genéticos únicos o el mayor número de aves por unidad de área. Más específicamente, si el objetivo es la conservación de los linajes genéticos, entonces los recursos limitados deberían enfocarse en preservar el hábitat en la Sierra de California, la Cuenca de las Rocallosas, la costa de California (lugares en donde se reproducen los tres linajes genéticos más vulnerables) o en el oeste de México (en donde dos de los tres linajes más vulnerables pasan el invierno). Alternativamente, si el objetivo es la conservación del mayor número de individuos por unidad de área, entonces el financiamiento limitado debería aplicarse en el noroeste del Pacífico o en América Central, en donde se estima que las densidades poblacionales son las más altas. En general, nuestros resultados demostraron la utilidad de adoptar un modelo de redes basadas en la genética para la integración de datos a lo largo de escalas geográficas amplias y para informar de mejor manera la toma de decisiones de conservación para los animales migratorios.

A range-wide domino effect and resetting of the annual cycle in a migratory songbird.

Latitudinal differences in timing of breeding are well documented but how such differences carry over to influence timing of events in the annual cycle of migratory birds is not well understood. We examined geographical variation in timing of events throughout the year using light-level geolocator tracking data from 133 migratory tree swallows ( Tachycineta bicolor) originating from 12 North American breeding populations. A swallow's breeding latitude influenced timing of breeding, which then carried over to affect breeding ground departure. This resulted in subsequent effects on the arrival and departure schedules at autumn stopover locations and timing of arrival at non-breeding locations. This 'domino effect' between timing events was no longer apparent by the time individuals departed for spring migration. Our range-wide analysis demonstrates the lasting impact breeding latitude can have on migration schedules but also highlights how such timing relationships can reset when individuals reside at non-breeding sites for extended periods of time.

Sublethal Toxicity of Crude Oil Exposure in The Blue Crab, Callinectes sapidus, at Two Life History Stages.

The 2010 Deepwater Horizon oil spill occurred during peak spawning season for many Gulf of Mexico fish and invertebrates. Early life stages of important fishery species were at risk to encounter crude oil. In this study, we investigated the effect of crude oil exposure on two life stages of the blue crab (Callinectes sapidus). We tested the effects of oil exposure on the survival and growth of larval (pelagic) and juvenile (estuarine) blue crabs as well as the effects of oil on the molt frequency of juveniles. Larval crabs exposed to crude oil showed no discernable growth or mortality differences when compared to non-exposed controls. Juvenile crabs exposed to oil also showed no differences in size but exhibited increased intermolt duration (time between molts). Our study suggests that different life-stages may respond differently when exposed to crude oil and that oil exposure negatively affects growth rate of juvenile blue crabs.

Effects of breeding versus winter habitat loss and fragmentation on the population dynamics of a migratory songbird.

Many migratory species are in decline and understanding these declines is challenging because individuals occupy widely divergent and geographically distant habitats during a single year and therefore populations across the range are interconnected in complex ways. Network modeling has been used to show, theoretically, that shifts in migratory connectivity patterns can occur in response to habitat or climate changes and that habitat loss in one region can affect sub-populations in regions that are not directly connected. Here, we use a network model, parameterized by integrating long-term monitoring data with direct tracking of -100 individuals, to explain population trends in the rapidly declining Wood Thrush (Hylocichla mustelina) and to predict future trends. Our model suggests that species-level declines in Wood Thrush are driven primarily by tropical deforestation in Central America but that protection of breeding habitat in some regions is necessary to prevent shifts in migratory connectivity and to sustain populations in all breeding regions. The model illustrates how shifts in migratory connectivity may lead to unexpected population declines in key regions. We highlight current knowledge gaps that make modeling full life-cycle population demographics in migratory species challenging but also demonstrate that modeling can inform conservation while these gaps are being filled.

The response of migratory populations to phenological change: a Migratory Flow Network modelling approach.

Declines in migratory species have been linked to anthropogenic climate change through phenological mismatch, which arises due to asynchronies between the timing of life-history events (such as migration) and the phenology of available resources. Long-distance migratory species may be particularly vulnerable to phenological change in their breeding ranges, since the timing of migration departure is based on environmental cues at distant non-breeding sites. Migrants may, however, be able to adjust migration speed en route to the breeding grounds, and thus, ability of migrants to update their timing of migration may depend critically on stopover frequency during migration; however, understanding how migratory strategy influences population dynamics is hindered by a lack of predictive models explicitly linking habitat quality to demography and movement patterns throughout the migratory cycle. Here, we present a novel modelling framework, the Migratory Flow Network (MFN), in which the seasonally varying attractiveness of breeding, winter and stopover regions drives the direction and timing of migration based on a simple general flux law. We use the MFN to investigate how populations respond to shifts in breeding site phenology based on their frequency of stopover and ability to detect and adapt to these changes. With perfect knowledge of advancing phenology, 'jump' migrants (low-frequency stopover) require more adaptation for populations to recover than 'hop' and 'skip' (high or medium frequency stopover) migrants. If adaptation depends on proximity, hop and skip migrants' populations can recover but jump migrants cannot adjust and decline severely. These results highlight the importance of understanding migratory strategies and maintaining high-quality stopover habitat to buffer migratory populations from climate-induced mismatch. We discuss how MFNs could be applied to diverse migratory taxa and highlight the potential of MFNs as a tool for exploring how migrants respond to other environmental changes such as habitat loss.

Metapopulation models for seasonally migratory animals.

Metapopulation models are widely used to study species that occupy patchily distributed habitat, but are rarely applied to migratory species, because of the difficulty of identifying demographically independent subpopulations. Here, we extend metapopulation theory to describe the directed seasonal movement of migratory populations between two sets of habitat patches, breeding and non-breeding, with potentially different colonization and extinction rates between patch types. By extending the classic metapopulation model, we show that migratory metapopulations will persist if the product of the two colonization rates exceeds the product of extinction rates. Further, we develop a spatially realistic migratory metapopulation model and derive a landscape metric-the migratory metapopulation capacity-that determines persistence. This new extension to metapopulation theory introduces an important tool for the management and conservation of migratory species and may also be applicable to model the dynamics of two host-parasite systems.

Host plant specificity of the monarch butterfly Danaus plexippus: A systematic review and meta-analysis.

The preference-performance hypothesis explains host specificity in phytophagous insects, positing that host plants chosen by adults confer the greatest larval fitness. However, adults sometimes oviposit on plants supporting low larval success because the components of host specificity (adult preference, plant palatability, and larval survival) are non-binary and not necessarily correlated. Palatability (willingness to eat) is governed by chemical cues and physical barriers such as trichomes, while survival (ability to complete development) depends upon nutrition and toxicity. Absence of a correlation between the components of host specificity results in low-performance hosts supporting limited larval development. Most studies of specificity focus on oviposition behavior leaving the importance and basis of palatability and survival under-explored. We conducted a comprehensive review of 127 plant species that have been claimed or tested to be hosts for the monarch butterfly Danaus plexippus to classify them as non-hosts, low performance, or high performance. We performed a meta-analysis to test if performance status could be explained by properties of neurotoxic cardenolides or trichome density. We also conducted a no-choice larval feeding experiment to identify causes of low performance. We identified 34 high performance, 42 low performance, 33 non-hosts, and 18 species with unsubstantiated claims. Mean cardenolide concentration was greater in high- than low-performance hosts and a significant predictor of host status, suggesting possible evolutionary trade-offs in monarch specialization. Other cardenolide properties and trichome density were not significant predictors of host status. In the experiment, we found, of the 62% of larvae that attempted to eat low-performance hosts, only 3.5% survived to adult compared to 85% of those on the high-performance host, demonstrating that multiple factors affect larval host plant specificity. Our study is the first to classify all known host plants for monarchs and has conservation implications for this threatened species.

The evolution of migration in a seasonal environment.

Despite the fact that migration occurs in a wide variety of taxa worldwide, little is known about the conditions under which migration is expected to evolve from an ancestral resident population. We develop a model that focuses on ecological factors affecting the evolution of migration in a seasonal environment within a genetically explicit framework. We model the evolution of migration for two common types of migration: 'shared breeding' where migrants share a breeding ground with residents and migrate to a separate non-breeding area, versus 'shared non-breeding', where migrants share a non-breeding ground with residents and migrate to a separate breeding area. Ecologically, migration is more easily established in the shared-breeding case versus the shared-non-breeding case. Genetically, the additive effect of a migratory allele affects its establishment more in the shared-non-breeding case versus the shared-breeding case, whereas the dominance effect of the allele affects its establishment more in the shared-breeding case versus the shared-non-breeding case. Generally, migratory alleles can invade even when residents are competitively superior to migrants during the shared season. Partial migration occurs when the population is polymorphic for migratory and non-migratory alleles, and is dependent upon which season is shared and the additive and dominance behaviour of the migratory allele.

Effects of crude oil on survival and development in embryonated eggs in Callinectes sapidus Rathbun, 1896 (Decapoda, Portunidae).

Blue crabs, Callinectes sapidus Rathbun, 1896, are ubiquitous along the Atlantic and Gulf coasts of the USA. These organisms play an integral role in the ecosystems of the Gulf of Mexico (GOM), where not only are they a keystone species, but are also socioeconomically important. The survival of embryonated eggs is necessary to ensure adequate recruitment into the next generation. Because the 2010 Deepwater Horizon oil spill (DWH) occurred during the peak of the blue crab spawning season, the incident likely impacted blue crab embryos. In order to assess the effect of oil on embryonic growth and development, we collected embryonated eggs from seven different female blue crabs from the GOM throughout the spawning season and exposed them to an oil concentration of 500 ppb (the approximate concentration of oil at the surface water near the site of the Deepwater Horizon oil rig). Exposure to oil at this concentration caused a significantly larger proportion of prezoeae vs. zoeae to hatch from embryonated eggs in experiments lasting longer than 4 days. Exposure to oil did not significantly affect overall survival or development rate. The prezoeal stage is a little-studied stage of blue crab development. Though it may or may not be a normal stage of development, this stage has been found to occur in suboptimal conditions and has lower survival than zoeal stages. The larger proportion of prezoeae following prolonged exposure to oil thus indicates that crude oil at concentrations likely to be experienced by crabs after the DWH spill negatively impacted the development of blue crab embryos. In addition to providing insight into the effects of the DWH, this study sheds light on embryonic development in blue crabs, a critical, but poorly investigated phase of this important species' life cycle.

Inherent limits of light-level geolocation may lead to over-interpretation.

In their 2015 Current Biology paper, Streby et al.[1] reported that Golden-winged Warblers (Vermivora chrysoptera), which had just migrated to their breeding location in eastern Tennessee, performed a facultative and up to ">1,500 km roundtrip" to the Gulf of Mexico to avoid a severe tornadic storm. From light-level geolocator data, wherein geographical locations are estimated via the timing of sunrise and sunset, Streby et al.[1] concluded that the warblers had evacuated their breeding area approximately 24 hours before the storm and returned about five days later. The authors presented this finding as evidence that migratory birds avoid severe storms by temporarily moving long-distances. However, the tracking method employed by Streby et al.[1] is prone to considerable error and uncertainty. Here, we argue that this interpretation of the data oversteps the limits of the used tracking technique. By calculating the expected geographical error range for the tracked birds, we demonstrate that the hypothesized movements fell well within the geolocators' inherent error range for this species and that such deviations in latitude occur frequently even if individuals remain stationary.

The shape of density dependence in fragmented landscapes explains an inverse buffer effect in a migratory songbird.

It is well known that forest fragmentation reduces fecundity in several avian species, including wood thrush, Hylocichla mustelina, a migratory songbird that has been declining for several decades. However, I found that landscape-scale density in wood thrush was lower and population declines steeper in higher quality, less-fragmented landscapes (an inverse buffer effect) than in poor quality landscapes. These patterns suggest that wood thrush was not limited by availability of breeding habitat but that declines were primarily driven by non-breeding season events. A two-season model predicts that if this hypothesis is correct, breeding population trends will be negatively related to the strength of density dependence (b') in the breeding season. To test this, a site-dependence model was used to construct fecundity-density curves and showed that landscape fragmentation affected the shape of density dependence. In good quality landscapes, the onset of strong density dependence was much more abrupt than in poorer quality landscapes and the realized strength of density dependence, b', was lower in good quality landscapes. Population trends were negatively associated with b', providing support for the non-breeding limitation hypothesis. The combination of the negative associations of trends with b' and b' with landscape quality explain the inverse buffer effect.

Effects of crude oil and oil/dispersant mixture on growth and expression of vitellogenin and heat shock protein 90 in blue crab, Callinectes sapidus, juveniles.

The 2010 Deepwater Horizon (DWH) oil spill in the northern Gulf of Mexico (NGOM) resulted in over 780million liters of crude oil spilling into Gulf waters. In an effort to disperse the oil, nearly 7.6million liters of dispersant was applied. Many commercially and recreationally important species reside in or near the area of the spill. The blue crab, Callinectes sapidus, is common in the NGOM and is both economically and ecologically important in this region. In this study, after exposing juvenile blue crabs to oil or a mixture of oil and dispersant we tested for relative expression of heat shock protein 90 (hsp90) and vitellogenin (vtg) by measuring their corresponding mRNA expression. We also monitored crabs over two molts to test for effects on growth. Expression of hsp90 was significantly downregulated, and we did not detect any effects of exposure to oil or oil/dispersant mixture on growth or vtg expression.

Evaluation of Blue Crab, Callinectes sapidus, Megalopal Settlement and Condition during the Deepwater Horizon Oil Spill.

The Blue Crab, Callinectes sapidus, is a commercially, culturally, and ecologically significant species in the Gulf of Mexico (GOM), whose offshore stages were likely impacted by the Deepwater Horizon oil spill (DWH). To test for DWH effects and to better understand the planktonic ecology of this species, we monitored Callinectes spp. megalopal settlement and condition at sites within and outside of the spill extent during and one year after the DWH. We tested for DWH effects by comparing 2010 settlement against baseline data available for two sites, and by testing for differences in settlement and condition inside and outside of the spill extent. We also developed time series models to better understand natural drivers of daily settlement variation (seasonal and lunar trends, hydrodynamics, wind) during 2010 and 2011. Overall, we found that neither megalopal settlement nor body weight were significantly reduced at oiled sites, but that high unexplained variation and low statistical power made detection of even large effects unlikely. Time series models revealed remarkably consistent and relatively strong seasonal and lunar trends within sites (explaining on average 28% and 9% of variation, respectively), while wind and hydrodynamic effects were weak (1-5% variation explained) and variable among sites. This study provides insights into DWH impacts as well as the natural drivers of Callinectes spp. megalopal settlement across the northern GOM.

Influence of sediment characteristics on the composition of soft-sediment intertidal communities in the northern Gulf of Mexico.

Benthic infaunal communities are important components of coastal ecosystems. Understanding the relationships between the structure of these communities and characteristics of the habitat in which they live is becoming progressively more important as coastal systems face increasing stress from anthropogenic impacts and changes in climate. To examine how sediment characteristics and infaunal community composition were related along the northern Gulf of Mexico coast, we sampled intertidal infaunal communities at seven sites covering common habitat types at a regional scale. Across 69 samples, the communities clustered into four distinct groups on the basis of faunal composition. Nearly 70% of the variation in the composition of the communities was explained by salinity, median grain size, and total organic content. Our results suggest that at a regional level coarse habitat characteristics are able to explain a large amount of the variation among sites in infaunal community structure. By examining the relationships between infaunal communities and their sedimentary habitats, we take a necessary first step that will allow the exploration of how changes in habitat and community composition influence higher trophic levels and ecosystem scale processes.

Assessing costs of carrying geolocators using feather corticosterone in two species of aerial insectivore.

Despite benefits of using light-sensitive geolocators to track animal movements and describe patterns of migratory connectivity, concerns have been raised about negative effects of these devices, particularly in small species of aerial insectivore. Geolocators may act as handicaps that increase energetic expenditure, which could explain reported effects of geolocators on survival. We tested this 'Energetic Expenditure Hypothesis' in 12 populations of tree swallows (Tachycineta bicolor) and barn swallows (Hirundo rustica) from North America and Europe, using measurements of corticosterone from feathers (CORTf) grown after deployment of geolocators as a measure of physiology relevant to energetics. Contrary to predictions, neither among- (both species) nor within-individual (tree swallows only) levels of CORTf differed with respect to instrumentation. Thus, to the extent that CORTf reflects energetic expenditure, geolocators apparently were not a strong handicap for birds that returned post-deployment. While this physiological evidence suggests that information about migration obtained from returning geolocator-equipped swallows is unbiased with regard to levels of stress, we cannot discount the possibility that corticosterone played a role in reported effects of geolocators on survival in birds, and suggest that future studies relate corticosterone to antecedent factors, such as reproductive history, and to downstream fitness costs.

Modeling activity rhythms in fiddler crabs.

Burrowing crabs of the genus Uca inhabit tidal mudflats and beaches. They feed actively during low tide and remain in their burrows when the tide is high. The timing of this activity has been shown to persist in the absence of external light and tidal cues, indicating the presence of an internal timing mechanism. Researchers report the persistence of several variations in locomotor activity under laboratory conditions that cannot be explained by a single circatidal clock. Previous studies supported two alternative hypotheses: the presence of either two circalunidian clocks, or a circadian and circatidal clock to regulate these activity rhythms. In this paper, we formulate mathematical models to describe and test these hypotheses. The models suggested by the literature contain some important differences beyond the frequency of proposed clocks, and these are reflected in the mathematical formulations and simulation results. One hypothesis suggests independent phase oscillators, while the other hypothesis suggests that they are coupled in anti-phase. Neither model is able to recover all of the variations in locomotor acitivity observed under laboratory conditions. However, we propose a new model that incorporates aspects of both existing hypotheses and is able to reproduce all laboratory observations.

Predicting conditions for migration: effects of density dependence and habitat quality.

Migration is widespread among animals, but the factors that influence the decision to migrate are poorly understood. Within a single species, populations may be completely migratory, completely sedentary or partially migratory. We use a population model to derive conditions for migration and demonstrate how migratory survival, habitat quality and density dependence on both the breeding and non-breeding grounds influence conditions for migration and the proportion of migrants within a population. Density dependence during the season in which migratory and sedentary individuals use separate sites is necessary for partial migration. High levels of density dependence at the non-shared sites widen the range of survival values within which we predict partial migration, whereas increasing the strength of density dependence at the shared sites narrows the range of survival values within which we predict partial migration. Our results have important implications for predicting how contemporary populations with variable migration strategies may respond to changes in the quality or quantity of habitat.

Predicting the consequences of carry-over effects for migratory populations.

Migratory animals present a unique challenge for predicting population size because they are influenced by events in multiple stages of the annual cycle that are separated by large geographic distances. Here, we develop a model that incorporates non-fatal carry-over effects to predict changes in population size and show how this can be integrated with predictive models of habitat loss and deterioration. Examples from Barn swallows, Greater snow geese and American redstarts show how carry-over effects can be estimated and integrated into the model. Incorporation of carry-over effects should increase the predictive power of models. However, the challenge for developing accurate predictions rests both on the ability to estimate parameters from multiple stages of the annual cycle and to understand how events between these periods interact to influence individual success.