Phoebe L. Zarnetske

Climate change and the past, present, and future of biotic interactions.

Biotic interactions drive key ecological and evolutionary processes and mediate ecosystem responses to climate change. The direction, frequency, and intensity of biotic interactions can in turn be altered by climate change. Understanding the complex interplay between climate and biotic interactions is thus essential for fully anticipating how ecosystems will respond to the fast rates of current warming, which are unprecedented since the end of the last glacial period. We highlight episodes of climate change that have disrupted ecosystems and trophic interactions over time scales ranging from years to millennia by changing species’ relative abundances and geographic ranges, causing extinctions, and creating transient and novel communities dominated by generalist species and interactions. These patterns emerge repeatedly across disparate temporal and spatial scales, suggesting the possibility of similar underlying processes. Based on these findings, we identify knowledge gaps and fruitful areas for research that will further our understanding of the effects of climate change on ecosystems.

Moving forward: dispersal and species interactions determine biotic responses to climate change

We need accurate predictions about how climate change will alter species distributions and abundances around the world. Most predictions assume simplistic dispersal scenarios and ignore biotic interactions. We argue for incorporating the complexities of dispersal and species interactions. Range expansions depend not just on mean dispersal, but also on the shape of the dispersal kernel and the populations growth rate. We show how models using species‐specific dispersal can produce more accurate predictions than models applying all‐or‐nothing dispersal scenarios. Models that additionally include species interactions can generate distinct outcomes. For example, species interactions can slow climate tracking and produce more extinctions than models assuming no interactions. We conclude that (1) just knowing mean dispersal is insufficient to predict biotic responses to climate change, and (2) considering interspecific dispersal variation and species interactions jointly will be necessary to anticipate future changes to biological diversity. We advocate for collecting key information on interspecific dispersal differences and strong biotic interactions so that we can build the more robust predictive models that will be necessary to inform conservation efforts as climates continue to change.

Biophysical feedback mediates effects of invasive grasses on coastal dune shape

Vegetation at the aquatic-terrestrial interface can alter landscape features through its growth and interactions with sediment and fluids. Even similar species may impart different effects due to variation in their interactions and feedbacks with the environment. Consequently, replacement of one engineering species by another can cause significant change in the physical environment. Here we investigate the species-specific ecological mechanisms influencing the geomorphology of U.S. Pacific Northwest coastal dunes. Over the last century, this system changed from open, shifting sand dunes with sparse vegetation (including native beach grass, Elymus mollis), to densely vegetated continuous foredune ridges resulting from the introduction and subsequent invasions of two nonnative grass species (Ammophila arenaria and Ammophila breviligulata), each of which is associated with different dune shapes and sediment supply rates along the coast. Here we propose a biophysical feedback responsible for differences in dune shape, and we investigate two, non-mutually exclusive ecological mechanisms for these differences: (1) species differ in their ability to capture sand and (2) species differ in their growth habit in response to sand deposition. To investigate sand capture, we used a moveable bed wind tunnel experiment and found that increasing tiller density increased sand capture efficiency and that, under different experimental densities, the native grass had higher sand capture efficiency compared to the Ammophila congeners. However, the greater densities of nonnative grasses under field conditions suggest that they have greater potential to capture more sand overall. We used a mesocosm experiment to look at plant growth responses to sand deposition and found that, in response to increasing sand supply rates, A. arenaria produced higher-density vertical tillers (characteristic of higher sand capture efficiency), while A. breviligulata and E. mollis responded with lower-density lateral tiller growth (characteristic of lower sand capture efficiency). Combined, these experiments provide evidence for a species-specific effect on coastal dune shape. Understanding how dominant ecosystem engineers, especially nonnative ones, differ in their interactions with abiotic factors is necessary to better parameterize coastal vulnerability models and inform management practices related to both coastal protection ecosystem services and ecosystem restoration.

Invasive grasses, climate change, and exposure to storm-wave overtopping in coastal dune ecosystems.

The worlds coastal habitats are critical to human well-being, but are also highly sensitive to human habitat alterations and climate change. In particular, global climate is increasing sea levels and potentially altering storm intensities, which may result in increased risk of flooding in coastal areas. In the Pacific Northwest (USA), coastal dunes that protect the coast from flooding are largely the product of a grass introduced from Europe over a century ago (Ammophila arenaria). An introduced congener (A. breviligulata) is displacing A. arenaria and reducing dune height. Here we quantify the relative exposure to storm-wave induced dune overtopping posed by the A. breviligulata invasion in the face of projected multi-decadal changes in sea level and storm intensity. In our models, altered storm intensity was the largest driver of overtopping extent, however the invasion by A. breviligulata tripled the number of areas vulnerable to overtopping and posed a fourfold larger exposure than sea-level rise over multi-decadal time scales. Our work demonstrates the importance of a transdisciplinary approach that draws on insights from ecology, geomorphology, and civil engineering to assess the vulnerability of ecosystem services in light of global change.

HABITAT CLASSIFICATION MODELING WITH INCOMPLETE DATA: PUSHING THE HABITAT ENVELOPE

Habitat classification models (HCMs) are invaluable tools for species conservation, land-use planning, reserve design, and metapopulation assessments, particularly at broad spatial scales. However, species occurrence data are often lacking and typically limited to presence points at broad scales. This lack of absence data precludes the use of many statistical techniques for HCMs. One option is to generate pseudo-absence points so that the many available statistical modeling tools can bb used. Traditional techniques generate pseudo-absence points at random across broadly defined species ranges, often failing to include biological knowledge concerning the species-habitat relationship. We incorporated biological knowledge of the species-habitat relationship into pseudo-absence points by creating habitat envelopes that constrain the region from which points were randomly selected. We define a habitat envelope as an ecological representation of a species, or species features (e.g., nest) observed distribution (i.e., realized niche) based on a single attribute, or the spatial intersection of multiple attributes. We created HCMs for Northern Goshawk (Accipiter gentilis atricapillus) nest habitat during the breeding season across Utah forests with extant nest presence points and ecologically based pseudo-absence points using logistic regression. Predictor variables were derived from 30-m USDA Landfire and 250-m Forest Inventory and Analysis (FIA) map products. These habitat-envelope-based models were then compared to null envelope models which use traditional practices for generating pseudo-absences. Models were assessed for fit and predictive capability using metrics such as kappa, threshold-independent receiver operating characteristic (ROC) plots, adjusted deviance (D(adj)2), and cross-validation, and were also assessed for ecological relevance. For all cases, habitat envelope-based models outperformed null envelope models and were more ecologically relevant, suggesting that incorporating biological knowledge into pseudo-absence point generation is a powerful tool for species habitat assessments. Furthermore, given some a priori knowledge of the species-habitat relationship, ecologically based pseudo-absence points can be applied to any species, ecosystem, data resolution, and spatial extent.

Non-target effects of invasive species management: beachgrass, birds, and bulldozers in coastal dunes.

Alteration of ecosystem processes by invasive species can lead to the decline of native species. Management actions targeted at removing these invaders and restoring native populations may have knock-on effects on non-target native species and ecosystems. For example, coastal dunes in the Pacific Northwest of North America are nearly monocultures of the introduced beach grasses, Ammophila arenaria and Ammophila breviligulata. These invasive grasses have converted open, low-lying sand dunes with a sparse covering of native plants to tall, densely-vegetated ridges dominated by the two invaders. As a result, the critical open-sand habitat of the federally threatened Western Snowy plover (Charadrius alexandrinus nivosus) has declined along with populations of several native dune plant species. Here we investigate how nearly 20 years of management targeted at the removal of Ammophila for plover recovery are impacting native plant species and dune morphology along 500 km of coastline in Oregon and Washington, U...

Indirect effects and facilitation among native and non‐native species promote invasion success along an environmental stress gradient

Summary 1. The spatial distribution of species is mediated by a combination of biotic interactions and environmental conditions. Understanding the relative importance of these factors and how they interact is particularly important for predicting the spread of non-native species and their impact on resident communities. 2. We used a 3-species Lotka–Volterra model parameterized with field and experimental data to understand the potential for continued spread by an introduced, non-native, dune-building beach grass (Ammophila breviligulata) and whether this invasion will result in coexistence or displacement in the resident beach grass communities (native Elymus mollis and introduced, non-native Ammophila arenaria) of the US Pacific Northwest. 3. We also used the model to investigate to what extent different rates of ocean-driven sand supply mediate the outcomes of beach grass species interactions. 4. Our model suggests that A. breviligulata could invade and dominate dunes across the range of sand supply rates observed in the region. We found that sand supply influenced intra- vs. interspecific interactions, the strength of positive and indirect effects among beach grasses and the long-term abundances of the beach grass species themselves. 5. Of the two non-natives, A. breviligulata is the inferior dune-building species. Thus, our results suggest that further invasions by A. breviligulata could reduce the coastal protection services afforded by tall dunes currently dominated by A. arenaria. 6. Synthesis: In systems with strong environmental forcing and stressful conditions such as coastal dunes, environmentally mediated positive and indirect species interactions can govern invasion success and long-term native–non-native coexistence. In doing so, these interactions ultimately shape community structure and ecosystem function. Understanding the joint effects of environmental forcing and species interactions on community assembly is particularly important in cases where species introductions can alter ecosystem services, such as coastal protection, which are vulnerable to the effects of climate change.

Coastal foredune evolution: the relative influence of vegetation and sand supply in the US Pacific Northwest.

Biophysical feedbacks between vegetation and sediment are important for forming and modifying landscape features and their ecosystem services. These feedbacks are especially important where landscape features differ in their provision of ecosystem services. For example, the shape of coastal foredunes, a product of both physical and biological forces, determines their ability to protect communities from rising seas and changing patterns of storminess. Here we assessed how sand supply and changes in vegetation over interannual (3 year) and decadal (21 year) scales influenced foredune shape along 100 km of coastline in the US Pacific Northwest. Across 21 years, vegetation switched from one congeneric non-native beachgrass to another (Ammophila arenaria to A. breviligulata) while sand supply rates were positive. At interannual timescales, sand supply rates explained the majority of change in foredune height (64–69%) and width (56–80%). However, at decadal scales, change in vegetation explained the majority of the change in foredune width (62–68%), whereas sand supply rates explained most of the change in foredune height (88–90%). In areas with lower shoreline change rates (±2 m yr−1), the change in vegetation explained the majority of decadal changes in foredune width (56–57%) and height (59–76%). Foredune shape directly impacts coastal protection, thus our findings are pertinent to coastal management given pressures of development and climate change.

Invasive congeners differ in successional impacts across space and time

Invasive species can alter the succession of ecological communities because they are often adapted to the disturbed conditions that initiate succession. The extent to which this occurs may depend on how widely they are distributed across environmental gradients and how long they persist over the course of succession. We focus on plant communities of the USA Pacific Northwest coastal dunes, where disturbance is characterized by changes in sediment supply, and the plant community is dominated by two introduced grasses – the long-established Ammophila arenaria and the currently invading A. breviligulata. Previous studies showed that A. breviligulata has replaced A. arenaria and reduced community diversity. We hypothesize that this is largely due to A. breviligulata occupying a wider distribution across spatial environmental gradients and persisting in later-successional habitat than A. arenaria. We used multi-decadal chronosequences and a resurvey study spanning 2 decades to characterize distributions of both species across space and time, and investigated how these distributions were associated with changes in the plant community. The invading A. breviligulata persisted longer and occupied a wider spatial distribution across the dune, and this corresponded with a reduction in plant species richness and native cover. Furthermore, backdunes previously dominated by A. arenaria switched to being dominated by A. breviligulata, forest, or developed land over a 23-yr period. Ammophila breviligulata likely invades by displacing A. arenaria, and reduces plant diversity by maintaining its dominance into later successional backdunes. Our results suggest distinct roles in succession, with A. arenaria playing a more classically facilitative role and A. breviligulata a more inhibitory role. Differential abilities of closely-related invasive species to persist through time and occupy heterogeneous environments allows for distinct impacts on communities during succession.

Incorporating Context Dependency of Species Interactions in Species Distribution Models

SYNOPSIS Species distribution models typically use correlative approaches that characterize the species-environment relationship using occurrence or abundance data for a single species. However, species distributions are determined by both abiotic conditions and biotic interactions with other species in the community. Therefore, climate change is expected to impact species through direct effects on their physiology and indirect effects propagated through their resources, predators, competitors, or mutualists. Furthermore, the sign and strength of species interactions can change according to abiotic conditions, resulting in context-dependent species interactions that may change across space or with climate change. Here, we incorporated the context dependency of species interactions into a dynamic species distribution model. We developed a multi-species model that uses a time-series of observational survey data to evaluate how abiotic conditions and species interactions affect the dynamics of three rocky intertidal species. The model further distinguishes between the direct effects of abiotic conditions on abundance and the indirect effects propagated through interactions with other species. We apply the model to keystone predation by the sea star Pisaster ochraceus on the mussel Mytilus californianus and the barnacle Balanus glandula in the rocky intertidal zone of the Pacific coast, USA. Our method indicated that biotic interactions between P. ochraceus and B. glandula affected B. glandula dynamics across >1000 km of coastline. Consistent with patterns from keystone predation, the growth rate of B. glandula varied according to the abundance of P. ochraceus in the previous year. The data and the model did not indicate that the strength of keystone predation by P. ochraceus varied with a mean annual upwelling index. Balanus glandula cover increased following years with high phytoplankton abundance measured as mean annual chlorophyll-a. M. californianus exhibited the same pattern to a lesser degree, although this pattern was not significant. This work bridges the disciplines of biogeography and community ecology to develop tools to better understand the direct and indirect effects of abiotic conditions on ecological communities.