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Stephen Blake

Publications and source records attributed to Stephen Blake.

11 recordsLinked to original sources

Environmental variation structures reproduction and recruitment in long-lived mega-herbivores: Galapagos giant tortoises

Migratory, long-lived animals are an important focus for life-history theory because they manifest extreme trade-offs in life-history traits: delayed maturity, low fecundity, variable recruitment rates, long generation times, and vital rates that respond to variation across environments. Galapagos tortoises are an iconic example: they are long-lived, migrate seasonally, face multiple anthropogenic threats, and have cryptic early life-history stages for which vital rates are unknown. From 2012 to 2021, we studied the reproductive ecology of two species of Galapagos tortoises ( Chelonoidis porteri and C. donfaustoi ) along elevation gradients that coincided with substantial changes in climate and vegetation productivity. Specifically, we (1) measured the body and reproductive condition of 166 adult females, (2) tracked the movements of 33 adult females using global positioning system telemetry, and monitored their body condition seasonally, (3) recorded nest temperatures, clutch characteristics, and egg survival from 107 nests, and (4) used radiotelemetry to monitor growth, survival, and movements of 104 hatchlings. We also monitored temperature and rainfall from field sites, and remotely sensed primary productivity along the elevation gradient. Our study showed that environmental variability, mediated by elevation, influenced vital rates of giant tortoises, specifically egg production by adult females and juvenile recruitment. Adult females were either elevational migrants or year-round lowland residents. Migrants had higher body condition than residents, and body condition was positively correlated with the probability of being gravid. Nests occurred in the hottest, driest parts of the tortoise's range, between 6 and 165 m elevation. Clutch size increased with elevation, whereas egg survival decreased. Hatchling survival and growth were highest at intermediate elevations. Hatchlings dispersed rapidly to 100–750 m from their nests before becoming sedentary (ranging over <0.2 ha). Predicted future climates may impact the relationships between elevation and vital rates of Galapagos tortoises and other species living across elevation gradients. Resilience will be maximized by ensuring the connectivity of foraging and reproductive areas within the current and possible future elevational ranges of these species.

Galapagos Islands, Santa Cruz Island

A greener future for the Galapagos: Forecasting ecosystem productivity by finding climate analogs in time

Forecasting ecosystem response to climate change is critical for guiding policymaking but challenging due to: complicated relationships between microclimates and regional climates; species’ responses that are driven by extremes rather than averages; the multifaceted nature of species’ interactions; and the lack of historical analogs to future climates. Given these challenges, even model systems such as the Galapagos Islands, a world-famous biodiversity hotspot and World Heritage Site, lack clear forecasts for future environmental change. Here, we developed a novel nonparametric method for simulating the ecosystem futures based on observed vegetation productivity (NDVI) during analogous weather observed historically. Using satellite images taken from the past to piece together a simulated future, we projected that productivity of terrestrial vegetation of the Galapagos will increase over the next century by approximately one standard deviation archipelago-wide, with largest increases during the wet season (January to June) and in the arid zones. Such greening would impact a variety of ecological and evolutionary processes, species of conservation concern, and agricultural practices. Our straightforward approach can be applied to many other regions, particularly those with rapid ecosystem responses to stochastic inter-annual climatic fluctuations that provide appropriate climate analogs for forecasting.

Galápagos Islands

Movement ecology

(Yackulic) At first glance, the decision to study movement in Galapagos tortoises seems curious. Given the slow speed of tortoises and tendency to forage and rest as they move, it seems implausible that tortoises would string their slow bursts of activity together to accomplish large-scale movements. Nonetheless, as early as 1815 (Porter 1815), visitors to Galapagos have noted the propensity for tortoises to walk long distances along well used trails, leading to seasonal changes in the distribution of tortoises on the various islands of the Archipelago. In recent years, advances in the technology used to track animals have led to a better understanding of movement and the diversity of movement strategies among many animal taxa. This chapter focuses on the application of these technologies to study of movement in Galapagos tortoises. Recent work has shown not only the diversity of movement strategies employed by Galapagos tortoises, but also illustrated how movement can both cause and be a consequence of interaction between reproductive and foraging ecologies of tortoises and the presence of strong environmental gradients. Understanding how critical unimpeded movement is to the persistence of Galapagos tortoises also informs efforts to maintain connectivity to, and suitable habitat within, tortoise range outside of protected areas.

Book chapter

Migration triggers in a large herbivore: Galapagos giant tortoises navigating resources gradients on volcanoes

To understand how migratory behavior evolved and to predict the future of migratory species in the face of global environmental change it is important to quantify intra- and inter-individual variation in migratory behavior. Intra-individual variation includes behavioral response to changing environmental conditions and hence behavioral plasticity in the context of novel conditions. Inter-individual variation determines the degree of variation on which selection can act and the rate of evolutionary response to changes in average and extreme environmental conditions. Here we focus on variation in the partial migratory behavior of Galapagos giant tortoises ( Chelonoidis spp.), which exhibit high fidelity to migratory routes over many years. We evaluate the extent and mechanisms by which tortoises adjust migration timing in response to varying annual environmental conditions, integrating movement data within a bioenergetic model of tortoise migration to quantify the fitness consequences of migration timing. We find strong inter-individual variation in the timing of migration, which was not affected by environmental conditions prevailing at the time of migration but rather by marginal expectations estimated from multi-annual averaged conditions, leading to an average annual loss in efficiency of ~15% relative to optimal timing based on year-specific conditions. These results point towards a limited ability of tortoises to adjust the timing of their migrations based on prevailing (and, by extension, future) conditions, suggesting that the adaptability of tortoise migratory behavior to changing conditions is predicated more on past “normal” conditions than responsive to current, changing conditions. Our work offers insights into the level of environmental-tuning in migratory behavior and a general framework for future research across taxa.

Galapagos Islands

Moving in the Anthropocene: Global reductions in terrestrial mammalian movements

Animal movement is fundamental for ecosystem functioning and species survival, yet the effects of the anthropogenic footprint on animal movements have not been estimated across species. Using a unique GPS-tracking database of 803 individuals across 57 species, we found that movements of mammals in areas with a comparatively high human footprint were on average one-half to one-third the extent of their movements in areas with a low human footprint. We attribute this reduction to behavioral changes of individual animals and to the exclusion of species with long-range movements from areas with higher human impact. Global loss of vagility alters a key ecological trait of animals that affects not only population persistence but also ecosystem processes such as predator-prey interactions, nutrient cycling, and disease transmission.

Science

Benefits of the destinations, not costs of the journeys, shape partial migration patterns

1. The reasons that lead some animals to seasonally migrate, and others to remain in the same area year-round, are poorly understood. Associations between traits, such as body size, and migration provide clues. For example, larger species and individuals are more likely to migrate. 2. One explanation for this size bias in migration is that larger animals are capable of moving faster (movement hypothesis). However, body size is linked to many other biological processes. For instance, the energetic balances of larger animals are generally more sensitive to variation in food density because of body size effects on foraging and metabolism and this sensitivity could drive migratory decisions (forage hypothesis). 3. Identifying the primary selective forces that drive migration ultimately requires quantifying fitness impacts over the full annual migratory cycle. Here, we develop a full annual migratory cycle model from metabolic and foraging theory to compare the importance of the forage and movement hypotheses. We parameterize the model for Galapagos tortoises, which were recently discovered to be size-dependent altitudinal migrants. 4. The model predicts phenomena not included in model development including maximum body sizes, the body size at which individuals begin to migrate, and the seasonal timing of migration and these predictions generally agree with available data. Scenarios strongly support the forage hypothesis over the movement hypothesis. Furthermore, male Galapagos tortoises on Santa Cruz Island would be unable to grow to their enormous sizes without access to both highlands and lowlands. 5. Whereas recent research has focused on links between traits and the migratory phases of the migratory cycle, we find that effects of body size on the non-migratory phases are far more important determinants of the propensity to migrate. Larger animals are more sensitive to changing forage conditions than smaller animals with implications for maintenance of migration and body size in the face of environmental change.

Journal of Animal Ecology

Ecosystem implications of conserving endemic versus eradicating introduced large herbivores in the Galapagos Archipelago

Restoration of damaged ecosystems through invasive species removal and native species conservation is an increasingly common practice in biodiversity conservation. Estimating the degree of ecosystem response attributable specifically to eradication of exotic herbivores versus restoration of native herbivores is often difficult and is complicated by concurrent temporal changes in other factors, especially climate. We investigated the interactive impacts of native mega-herbivores (giant tortoises) and the eradication of large alien herbivores (goats) on vegetation productivity across the Galapagos Archipelago. We examined archipelago-wide patterns of Normalized Difference Vegetation Index (NDVI) as a proxy for vegetation productivity between 2001 and 2015 and evaluated how goat and historical and current tortoise occurrence influenced productivity. We used a breakpoint analysis to detect change in trends in productivity from five targeted areas following goat eradication. We found a positive association between tortoise occurrence and vegetation productivity and a negative association with goat occurrence. We also documented an increase in plant productivity following goat removal with recovery higher in moister regions than in arid region, potentially indicating an alternate stable state has been created in the latter. Climate variation also contributed to the detected improvement in productivity following goat eradication, sometimes obscuring the effect of eradication but more usually magnifying it by up to 300%. Our work offers perspectives regarding the effectiveness and outcomes of eradicating introduced herbivores and re-introducing native herbivores, and the merits of staging them simultaneously in order to restore critical ecosystem processes such as vegetation productivity.

Galapagos Islands

Allometric and temporal scaling of movement characteristics in Galapagos tortoises

Understanding how individual movement scales with body size is of fundamental importance in predicting ecological relationships for diverse species. One-dimensional movement metrics scale consistently with body size yet vary over different temporal scales. Knowing how temporal scale influences the relationship between animal body size and movement would better inform hypotheses about the efficiency of foraging behaviour, the ontogeny of energy budgets, and numerous life-history trade-offs. We investigated how the temporal scaling of allometric patterns in movement varies over the course of a year, specifically during periods of motivated (directional and fast movement) and unmotivated (stationary and tortuous movement) behaviour. We focused on a recently diverged group of species that displays wide variation in movement behaviour – giant Galapagos tortoises ( Chelonoidis spp.) – to test how movement metrics estimated on a monthly basis scaled with body size. We used state-space modelling to estimate seven different movement metrics of Galapagos tortoises. We used log-log regression of the power law to evaluate allometric scaling for these movement metrics and contrasted relationships by species and sex. Allometric scaling of movement was more apparent during motivated periods of movement. During this period, allometry was revealed at multiple temporal intervals (hourly, daily and monthly), with values observed at daily and monthly intervals corresponding most closely to the expected one-fourth scaling coefficient, albeit with wide credible intervals. We further detected differences in the magnitude of scaling among taxa uncoupled from observed differences in the temporal structuring of their movement rates. Our results indicate that the definition of temporal scales is fundamental to the detection of allometry of movement and should be given more attention in movement studies. Our approach not only provides new conceptual insights into temporal attributes in one-dimensional scaling of movement, but also generates valuable insights into the movement ecology of iconic yet poorly understood Galapagos giant tortoises.

Journal of Animal Ecology

Flexible characterization of animal movement pattern using net squared displacement and a latent state model

Background Characterizing the movement patterns of animals is an important step in understanding their ecology. Various methods have been developed for classifying animal movement at both coarse (e.g., migratory vs. sedentary behavior) and fine (e.g., resting vs. foraging) scales. A popular approach for classifying movements at coarse resolutions involves fitting time series of net-squared displacement (NSD) to models representing different conceptualizations of coarse movement strategies (i.e., migration, nomadism, sedentarism, etc.). However, the performance of this method in classifying actual (as opposed to simulated) animal movements has been mixed. Here, we develop a more flexible method that uses the same NSD input, but relies on an underlying discrete latent state model. Using simulated data, we first assess how well patterns in the number of transitions between modes of movement and the duration of time spent in a mode classify movement strategies. We then apply our approach to elucidate variability in the movement strategies of eight giant tortoises ( Chelonoidis sp.) using a multi-year (2009&ndash;2014) GPS dataset from three different Galapagos Islands. Results With respect to patterns of time spent and the number of transitions between modes, our approach out-performed previous efforts to distinguish among migration, dispersal, and sedentary behavior. We documented marked inter-individual variation in giant tortoise movement strategies, with behaviors indicating migration, dispersal, nomadism and sedentarism, as well as hybrid behaviors such as &ldquo;exploratory residence&rdquo;. Conclusions Distilling complex animal movement into discrete modes remains a fundamental challenge in movement ecology, a problem made more complex by the ever-longer duration, ever-finer resolution, and gap-ridden trajectories recorded by GPS devices. By clustering into modes, we derived information on the time spent within one mode and the number of transitions between modes which enabled finer differentiation of movement strategies over previous methods. Ultimately, the techniques developed here address limitations of previous approaches and provide greater insights with respect to characterization of movement strategies across scales by more fully utilizing long-term GPS telemetry datasets.

Movement Ecology

The dominance of introduced plant species in the diets of migratory Galapagos tortoises increases with elevation on a human-occupied island

The distribution of resources and food selection are fundamental to the ecology, life history, physiology, population dynamics, and conservation of animals. Introduced plants are changing foraging dynamics of herbivores in many ecosystems often with unknown consequences. Galapagos tortoises, like many herbivores, undertake migrations along elevation gradients driven by variability in vegetation productivity which take them into upland areas dominated by introduced plants. We sought to characterize diet composition of two species of Galapagos tortoises, focussing on how the role of introduced forage species changes over space and the implications for tortoise conservation. We quantified the distribution of tortoises with elevation using GPS telemetry. Along the elevation gradient, we quantified the abundance of introduced and native plant species, estimated diet composition by recording foods consumed by tortoises, and assessed tortoise physical condition from body weights and blood parameter values. Tortoises ranged between 0 and 429 m in elevation over which they consumed at least 64 plant species from 26 families, 44 percent of which were introduced species. Cover of introduced species and the proportion of introduced species in tortoise diets increased with elevation. Introduced species were positively selected for by tortoises at all elevations. Tortoise physical condition was either consistent or increased with elevation at the least biologically productive season on Galapagos. Santa Cruz tortoises are generalist herbivores that have adapted their feeding behavior to consume many introduced plant species that has likely made a positive contribution to tortoise nutrition. Some transformed habitats that contain an abundance of introduced forage species are compatible with tortoise conservation.

Galapagos Islands

The 2006 eruption of Augustine Volcano - Combined analyses of thermal satellite data and reduced displacement: Chapter 23 in The 2006 eruption of Augustine Volcano, Alaska

Augustine Volcano erupted explosively after 20 years of quiescence on January 11, 2006, followed by approximately 2 months of dome building and lava extrusion. This is the best monitored eruption in Alaska to date; the diverse complementary datasets gathered enable an interdisciplinary interpretation of volcanic activity. An analysis of reduced displacement (continuous measure of seismic tremor amplitude) and thermal energy output (from satellite imagery) observed between January 1 and April 30, 2006, shows relationships linked to the type of eruptive activity. Three different types of volcanic behavior can be identified as they show specific patterns in the combined data sets: (1) explosive activity, (2) lava extrusion (dome growth), and (3) cooling of erupted products. Explosive activity was characterized by high reduced displacement values but relatively low radiative thermal flux. Lava extrusion occurred in three distinct sequences characterized by increased values of reduced displacement and increased thermal emissions. Two periods of elevated thermal energy output and reduced displacement coincided with times of deflation, suggesting an increase in extrusion rate. Periods of cooling were marked by decreasing thermal emissions and reduced displacement. This work highlights the value of combined observations, which reveal more about the status of an active volcano than individual methods alone.

Alaska