Integrated ecosystems: linking food webs through reciprocal resource reliance.

Ecosystems are defined, studied, and managed according to boundaries constructed to conceptualize patterns of interest at a certain scale and scope. The distinction between ecosystems becomes obscured when resources from multiple origins cross porous boundaries and are assimilated into food webs through repeated trophic transfers. Ecosystem compartments can define bounded localities in a heterogeneous landscape that simultaneously retain and exchange energy in the form of organic matter. Here we developed and tested a framework to quantify reciprocal reliance on cross-boundary resource exchange and calculate the contribution of primary production from adjacent ecosystem compartments cycling through food webs to support consumers at different trophic levels. Under this framework, an integrated ecosystem can be measured and designated when the boundary between spatially distinct compartments is permeable and the bidirectional exchange of resources contributes significantly to sustaining both food webs. Using a desert river and riparian zone as a case study, we demonstrate that resources exchanged across the aquatic-riparian boundary cycle through multiple trophic levels. Furthermore, predators on both sides of the boundary were supported by externally produced resources to a similar extent, indicating this is a tightly integrated river-riparian ecosystem and that changes to either compartment will substantially impact the other. Using published data on lake ecosystems, we demonstrated that benthic and pelagic ecosystem compartments are likely not fully integrated, but differences between lakes could be used to test ecological hypotheses. Finally, we discuss how the integrated ecosystem framework could be applied in urban-preserve and field-forest ecosystems to address a broad range of ecological concepts. Because few systems function in complete isolation, this novel approach has application to research and management strategies globally as ecosystems continue to face novel pressures that precipitate cascading ecological repercussions well beyond a bounded system of focus.

Do latitudinal gradients exist in New Zealand stream invertebrate metacommunities?

That biodiversity declines with latitude is well known, but whether a metacommunity process is behind this gradient has received limited attention. We tested the hypothesis that dispersal limitation is progressively replaced by mass effects with increasing latitude, along with a series of related hypotheses. We explored these hypotheses by examining metacommunity structure in stream invertebrate metacommunities spanning the length of New Zealand's two largest islands (∼1,300 km), further disentangling the role of dispersal by deconstructing assemblages into strong and weak dispersers. Given the highly dynamic nature of New Zealand streams, our alternative hypothesis was that these systems are so unpredictable (at different stages of post-flood succession) that metacommunity structure is highly context dependent from region to region. We rejected our primary hypotheses, pinning this lack of fit on the strong unpredictability of New Zealand's dynamic stream ecosystems and fauna that has evolved to cope with these conditions. While local community structure turned over along this latitudinal gradient, metacommunity structure was highly context dependent and dispersal traits did not elucidate patterns. Moreover, the emergent metacommunity types exhibited no trends, nor did the important environmental variables. These results provide a cautionary tale for examining singular metacommunities. The considerable level of unexplained contingency suggests that any inferences drawn from one-off snapshot sampling may be misleading and further points to the need for more studies on temporal dynamics of metacommunity processes.

Flow regime alteration degrades ecological networks in riparian ecosystems.

Riverine ecosystems are governed by patterns of temporal variation in river flows. This dynamism will change due to climate change and the near-ubiquitous human control of river flows globally, which may have severe effects on species distributions and interactions. We employed a combination of population modelling and network theory to explore the consequences of possible flow regime futures on riparian plant communities, including scenarios of increased drought, flooding and flow homogenization (removal of flow variability). We found that even slight modifications to the historic natural flow regime had significant consequences for the structure of riparian plant networks. Networks of emergent interactions between plant guilds were most connected at the natural flow regime and became simplified with increasing flow alteration. The most influential component of flow alteration was flood reduction, with drought and flow homogenization both having greater simplifying community-wide consequences than increased flooding. These findings suggest that maintaining floods under future climates will be needed to overcome the negative long-term consequences of flow modification on riverine ecosystems.

High mortality and enhanced recovery: modelling the countervailing effects of disturbance on population dynamics.

Disturbances cause high mortality in populations while simultaneously enhancing population growth by improving habitats. These countervailing effects make it difficult to predict population dynamics following disturbance events. To address this challenge, we derived a novel form of the logistic growth equation that permits time-varying carrying capacity and growth rate. We combined this equation with concepts drawn from disturbance ecology to create a general model for population dynamics in disturbance-prone systems. A river flooding example using three insect species (a fast life-cycle mayfly, a slow life-cycle dragonfly and an ostracod) found optimal tradeoffs between disturbance frequency vs. magnitude and a close fit to empirical data in 62% of cases. A savanna fire analysis identified fire frequencies of 3-4 years that maximised population size of a perennial grass. The model shows promise for predicting population dynamics after multiple disturbance events and for management of river flows and fire regimes.

Importance of neutral processes varies in time and space: Evidence from dryland stream ecosystems.

Many ecosystems experience strong temporal variability in environmental conditions; yet, a clear picture of how niche and neutral processes operate to determine community assembly in temporally variable systems remains elusive. In this study, we constructed neutral metacommunity models to assess the relative importance of neutral processes in a spatially and temporally variable ecosystem. We analyzed macroinvertebrate community data spanning multiple seasons and years from 20 sites in a Sonoran Desert river network in Arizona. The model goodness-of-fit was used to infer the importance of neutral processes. Averaging over eight stream flow conditions across three years, we found that neutral processes were more important in perennial streams than in non-perennial streams (intermittent and ephemeral streams). Averaging across perennial and non-perennial streams, we found that neutral processes were more important during very high flow and in low flow periods; whereas, at very low flows, the relative importance of neutral processes varied greatly. These findings were robust to the choice of model parameter values. Our study suggested that the net effect of disturbance on the relative importance of niche and neutral processes in community assembly varies non-monotonically with the severity of disturbance. In contrast to the prevailing view that disturbance promotes niche processes, we found that neutral processes could become more important when the severity of disturbance is beyond a certain threshold such that all organisms are adversely affected regardless of their biological traits and strategies.

Linking river flow regimes to riparian plant guilds: a community-wide modeling approach.

Modeling riparian plant dynamics along rivers is complicated by the fact that plants have different edaphic and hydrologic requirements at different life stages. With intensifying human demands for water and continued human alteration of rivers, there is a growing need for predicting responses of vegetation to flow alteration, including responses related to climate change and river flow management. We developed a coupled structured population model that combines stage-specific responses of plant guilds with specific attributes of river hydrologic regime. The model uses information on the vital rates of guilds as they relate to different hydrologic conditions (flood, drought, and baseflow), but deliberately omits biotic interactions from the structure (interaction neutral). Our intent was to (1) consolidate key vital rates concerning plant population dynamics and to incorporate these data into a quantitative framework, (2) determine whether complex plant stand dynamics, including biotic interactions, can be predicted from basic vital rates and river hydrology, and (3) project how altered flow regimes might affect riparian communities. We illustrated the approach using five flow-response guilds that encompass much of the river floodplain community: hydroriparian tree, xeroriparian shrub, hydroriparian shrub, mesoriparian meadow, and desert shrub. We also developed novel network-based tools for predicting community-wide effects of climate-driven shifts and deliberately altered flow regimes. The model recovered known patterns of hydroriparian tree vs. xeroriparian shrub dominance, including the relative proportion of these two guilds as a function of river flow modification. By simulating flow alteration scenarios ranging from increased drought to shifts in flood timing, the model predicted that mature hydroriparian forest should be most abundant near the observed natural flow regime. Multiguild sensitivity analysis identified substantial network connectivity (many connected nodes) and biotic linkage (strong pairwise connections between nodes) under natural flow regime conditions. Both connectivity and linkage were substantially reduced under drought and other flow-alteration scenarios, suggesting that community structure is destabilized under such conditions. This structured population modeling approach provides a useful tool for understanding the community-wide effects of altered flow regimes due to climate change and management actions that influence river flow regime.

Seasonality and predictability shape temporal species diversity.

Temporal environmental fluctuations, such as seasonality, exert strong controls on biodiversity. While the effects of seasonality are well known, the predictability of fluctuations across years may influence seasonality in ways that are less well understood. The ability of a habitat to support unique, non-nested assemblages of species at different times of the year should depend on both seasonality (occurrence of events at specific periods of the year) and predictability (the reliability of event recurrence) of characteristic ecological conditions. Drawing on tools from wavelet analysis and information theory, we developed a framework for quantifying both seasonality and predictability of habitats, and applied this using global long-term rainfall data. Our analysis predicted that temporal beta diversity should be maximized in highly predictable and highly seasonal climates, and that low degrees of seasonality, predictability, or both would lower diversity in characteristic ways. Using stream invertebrate communities as a case study, we demonstrated that temporal species diversity, as exhibited by community turnover, was determined by a balance between temporal environmental variability (seasonality) and the reliability of this variability (predictability). Communities in highly seasonal mediterranean environments exhibited strong oscillations in community structure, with turnover from one unique community type to another across seasons, whereas communities in aseasonal New Zealand environments fluctuated randomly. Understanding the influence of seasonal and other temporal scales of environmental oscillations on diversity is not complete without a clear understanding of their predictability, and our framework provides tools for examining these trends at a variety of temporal scales, seasonal and beyond. Given the uncertainty of future climates, seasonality and predictability are critical considerations for both basic science and management of ecosystems (e.g., dam operations, bioassessment) spanning gradients of climatic variability.

Herbivory enhances the diversity of primary producers in pond ecosystems.

Diversity of primary producer is often surprisingly high, despite few limiting factors such as nutrients and light to facilitate species coexistence. In theory, the presence of herbivores could increase the diversity of primary producers, resolving this "paradox of the plankton." Little experimental evidence supports this natural enemies hypothesis, but previous tests suffer from several deficiencies. Previous experiments often did not allow for multigeneration effects; utilized low diversity assemblages of herbivores; and limited opportunities for new primary producer and herbivore species to colonize and undergo species sorting that favors some species over others. Using pond plankton, we designed a mesocosm experiment that overcame these problems by allowing more time for interactions over multiple generations, openness to allow new colonists, and manipulated higher diversity of primary producers and grazers than have previous studies. With this design, the presence of zooplankton grazers doubled phytoplankton richness. The additional phytoplankton species in grazed mesocosms were larger, and therefore likely more grazer resistant. Furthermore, phytoplankton richness in grazed mesocosms was similar to that observed in natural ponds whereas it was much lower in mesocosms without grazers. However, stoichiometric imbalance caused by variation in nitrogen : phosphorus ratios and light supply did not alter phytoplankton richness. Therefore, grazers enhanced primary producer richness more strongly than ratios of nutrient supply (even though both grazing and ratios of resource supply altered composition of primary producer assemblages). Taken together, these experimental and field data show that grazing from a diverse assemblage of herbivores greatly elevated richness of phytoplankton producers in pond ecosystems.

The puzzle of partial migration: Adaptive dynamics and evolutionary game theory perspectives.

We consider the phenomenon of partial migration which is exhibited by populations in which some individuals migrate between habitats during their lifetime, but others do not. First, using an adaptive dynamics approach, we show that partial migration can be explained on the basis of negative density dependence in the per capita fertilities alone, provided that this density dependence is attenuated for increasing abundances of the subtypes that make up the population. We present an exact formula for the optimal proportion of migrants which is expressed in terms of the vital rates of migrant and non-migrant subtypes only. We show that this allocation strategy is both an evolutionary stable strategy (ESS) as well as a convergence stable strategy (CSS). To establish the former, we generalize the classical notion of an ESS because it is based on invasion exponents obtained from linearization arguments, which fail to capture the stabilizing effects of the nonlinear density dependence. These results clarify precisely when the notion of a "weak ESS", as proposed in Lundberg (2013) for a related model, is a genuine ESS. Secondly, we use an evolutionary game theory approach, and confirm, once again, that partial migration can be attributed to negative density dependence alone. In this context, the result holds even when density dependence is not attenuated. In this case, the optimal allocation strategy towards migrants is the same as the ESS stemming from the analysis based on the adaptive dynamics. The key feature of the population models considered here is that they are monotone dynamical systems, which enables a rather comprehensive mathematical analysis.

Linking multidimensional functional diversity to quantitative methods: a graphical hypothesis--evaluation framework.

Functional trait analysis is an appealing approach to study differences among biological communities because traits determine species' responses to the environment and their impacts on ecosystem functioning. Despite a rapidly expanding quantitative literature, it remains challenging to conceptualize concurrent changes in multiple trait dimensions ("trait space") and select quantitative functional diversity methods to test hypotheses prior to analysis. To address this need, we present a widely applicable framework for visualizing ecological phenomena in trait space to guide the selection, application, and interpretation of quantitative functional diversity methods. We describe five hypotheses that represent general patterns of responses to disturbance in functional community ecology and then apply a formal decision process to determine appropriate quantitative methods to test ecological hypotheses. As a part of this process, we devise a new statistical approach to test for functional turnover among communities. Our combination of hypotheses and metrics can be applied broadly to address ecological questions across a range of systems and study designs. We illustrate the framework with a case study of disturbance in freshwater communities. This hypothesis-driven approach will increase the rigor and transparency of applied functional trait studies.

Ecological strategies predict associations between aquatic and genetic connectivity for dryland amphibians.

The study of how population genetic structure is shaped by attributes of the environment is a central scientific pursuit in ecology and conservation. But limited resources may prohibit landscape genetics studies for many threatened species, particularly given the pace of current environmental change. Understanding the extent to which species' ecological strategies--their life histories, biology, and behavior-predict patterns and drivers of population connectivity is a critical step in evaluating the potential of multi-taxa inference in landscape genetics. We present results of a landscape genetic study of three dryland amphibians: the canyon treefrog (Hyla arenicolor), red-spotted toad (Anaxyrus punctatus), and Mexican spadefoot (Spea multiplicata). These species characterize a range of ecological strategies, driven primarily by different water dependencies, enabling amphibian survival in arid and semiarid environments. We examined a suite of hypothesized relationships between genetic connectivity and landscape connectivity across species. We found a positive relationship between population differentiation and water dependency, e.g., longer larval development periods and site fidelity for reliable water sources. We also found that aquatic connectivity is important for all species, particularly when considered with topography (slope). The effect of spatial scale varied by species, with canyon treefrogs and Mexican spadefoots characterized by relatively consistent results at different scales in contrast to the stark differences in results for red-spotted toads at different scales. Using ecological information to predict relationships between genetic and landscape connectivity is a promising approach for multi-taxa inference and may help inform conservation efforts where single-species genetic studies are not possible.

Two new species of Grylloblatta Walker, 1914 (Grylloblattodea: Grylloblattidae) from western North America, and a neotype designation for G. rothi Gurney 1953.

Grylloblatta rothi Gurney, 1953 is redescribed and a neotype is designated from Cultus Mountain in the Oregon Cascades, U.S.A. Two new species of Grylloblatta are described, bringing the total number of Grylloblatta species to 15. Grylloblatta chintimini new species is described from Marys Peak in the Coast Range of Western Oregon, where it occurs on snowpack near the 1250 m summit. Grylloblatta newberryensis new species is described from Newberry Volcano in Central Oregon, where it is associated with snowfields overlying geologically-young lava flows. Morphological characters, primarily derived from male genitalia, are presented to diagnose these species and differentiate them from other Grylloblatta spp. in Oregon, Washington, and California. Molecular sequences from the cytochrome oxidase subunit II gene suggest that significant divergence has occurred among these species and provide a tool to aid identification of juvenile and female specimens.

Dispersal ability and habitat requirements determine landscape-level genetic patterns in desert aquatic insects.

Species occupying the same geographic range can exhibit remarkably different population structures across the landscape, ranging from highly diversified to panmictic. Given limitations on collecting population-level data for large numbers of species, ecologists seek to identify proximate organismal traits-such as dispersal ability, habitat preference and life history-that are strong predictors of realized population structure. We examined how dispersal ability and habitat structure affect the regional balance of gene flow and genetic drift within three aquatic insects that represent the range of dispersal abilities and habitat requirements observed in desert stream insect communities. For each species, we tested for linear relationships between genetic distances and geographic distances using Euclidean and landscape-based metrics of resistance. We found that the moderate-disperser Mesocapnia arizonensis (Plecoptera: Capniidae) has a strong isolation-by-distance pattern, suggesting migration-drift equilibrium. By contrast, population structure in the flightless Abedus herberti (Hemiptera: Belostomatidae) is influenced by genetic drift, while gene flow is the dominant force in the strong-flying Boreonectes aequinoctialis (Coleoptera: Dytiscidae). The best-fitting landscape model for M. arizonensis was based on Euclidean distance. Analyses also identified a strong spatial scale-dependence, where landscape genetic methods only performed well for species that were intermediate in dispersal ability. Our results highlight the fact that when either gene flow or genetic drift dominates in shaping population structure, no detectable relationship between genetic and geographic distances is expected at certain spatial scales. This study provides insight into how gene flow and drift interact at the regional scale for these insects as well as the organisms that share similar habitats and dispersal abilities.

Quantifying invertebrate resistance to floods: a global-scale meta-analysis.

Floods are a key component of the ecology and management of riverine ecosystems around the globe, but it is not clear whether floods have predictable effects on organisms that can allow us to generalize across regions and continents. To address this, we conducted a global-scale meta-analysis to investigate effects of natural and managed floods on invertebrate resistance, the ability of invertebrates to survive flood events. We considered 994 studies for inclusion in the analysis, and after evaluation based on a priori criteria, narrowed our analysis to 41 studies spanning six of the seven continents. We used the natural-log-ratio of invertebrate abundance before and within 10 days after flood events because this measure of effect size can be directly converted to estimates of percent survival. We conducted categorical and continuous analyses that examined the contribution of environmental and study design variables to effect size heterogeneity, and examined differences in effect size among taxonomic groups. We found that invertebrate abundance was lowered by at least one-half after flood events. While natural vs. managed floods were similar in their effect, effect size differed among habitat and substrate types, with pools, sand, and boulders experiencing the strongest effect. Although sample sizes were not sufficient to examine all taxonomic groups, floods had a significant, negative effect on densities of Coleoptera, Eumalacostraca, Annelida, Ephemeroptera, Diptera, Plecoptera, and Trichoptera. Results from this study provide guidance for river flow regime prescriptions that will be applicable across continents and climate types, as well as baseline expectations for future empirical studies of freshwater disturbance.

Demographic stability metrics for conservation prioritization of isolated populations.

Systems of geographically isolated habitat patches house species that occur naturally as small, disjunct populations. Many of these species are of conservation concern, particularly under the interacting influences of isolation and rapid global change. One potential conservation strategy is to prioritize the populations most likely to persist through change and act as sources for future recolonization of less stable localities. We propose an approach to classify long-term population stability (and, presumably, future persistence potential) with composite demographic metrics derived from standard population-genetic data. Stability metrics can be related to simple habitat measures for a straightforward method of classifying localities to inform conservation management. We tested these ideas in a system of isolated desert headwater streams with mitochondrial sequence data from 16 populations of a flightless aquatic insect. Populations exhibited a wide range of stability scores, which were significantly predicted by dry-season aquatic habitat size. This preliminary test suggests strong potential for our proposed method of classifying isolated populations according to persistence potential. The approach is complementary to existing methods for prioritizing local habitats according to diversity patterns and should be tested further in other systems and with additional loci to inform composite demographic stability scores.

Evolution of aquatic insect behaviours across a gradient of disturbance predictability.

Natural disturbance regimes--cycles of fire, flood, drought or other events--range from highly predictable (disturbances occur regularly in time or in concert with a proximate cue) to highly unpredictable. While theory predicts how populations should evolve under different degrees of disturbance predictability, there is little empirical evidence of how this occurs in nature. Here, we demonstrate local adaptation in populations of an aquatic insect occupying sites along a natural gradient of disturbance predictability, where predictability was defined as the ability of a proximate cue (rainfall) to signal a disturbance (flash flood). In controlled behavioural experiments, populations from predictable environments responded to rainfall events by quickly exiting the water and moving sufficiently far from the stream to escape flash floods. By contrast, populations from less predictable environments had longer response times and lower response rates, reflecting the uncertainty inherent to these environments. Analysis with signal detection theory showed that for 13 out of 15 populations, observed response times were an optimal compromise between the competing risks of abandoning versus remaining in the stream, mediated by the rainfall-flood correlation of the local environment. Our study provides the first demonstration that populations can evolve in response to differences in disturbance predictability, and provides evidence that populations can adapt to among-stream differences in flow regime.

Grazers, producer stoichiometry, and the light : nutrient hypothesis revisited.

The stoichiometric light:nutrient hypothesis (LNH) links the relative supplies of key resources with the nutrient content of tissues of producers. This resource-driven variation in producer stoichiometry, in turn, can mediate the efficiency of grazing. Typically, discussions of the LNH attribute this resource-stoichiometry link to bottom-up effects of light and phosphorus, which are mediated through producer physiology. Emphasis on bottom-up effects implies that grazers must consume food of quality solely determined by resource supply to ecosystems (i.e., they eat what they are served). Here, we expand upon this largely bottom-up interpretation with evidence from pond surveys, a mesocosm experiment, and a model. Data from shallow ponds showed the "LNH pattern" (positive correlation of an index of light : phosphorus supply with algal carbon : phosphorus content). However, algal carbon : phosphorus content also declined as zooplankton biomass increased in the ponds. The experiment and model confirmed that this latter correlation was partially caused by the various bottom-up and top-down roles of grazers: the LNH pattern emerged only in treatments with crustacean grazers, not those without them. Furthermore, model and experiment clarified that another bottom-up factor, natural covariation of nitrogen : phosphorus ratios with light : phosphorus supply (as seen in ponds), does not likely contribute to the LNH pattern. Finally, the experiment produced correlations between shifts in species composition of algae, partially driven by grazing effects of crustaceans, and algal stoichiometry. These shifts in species composition might shape stoichiometric response of producer assemblages to resource supply and grazing, but their consequences remain largely unexplored. Thus, this study accentuated the importance of grazing for the LNH; de-emphasized a potentially confounding, bottom-up factor (covarying nitrogen : phosphorus supply); and highlighted an avenue for future research for the LNH (grazer-mediated shifts in producer composition).

Inedible producers in food webs: controls on stoichiometric food quality and composition of grazers.

Ecological stoichiometry and food web theories focus on distinct mechanisms that shape communities. These mechanisms, however, likely interact in ways that neither theory alone addresses. To illustrate, we show how a model that tracks flow of energy and nutrients through two producers and two grazers reveals two indirect, interrelated roles for "neutrally inedible" producers. First, inedible producers can exert controls over the nutrient content of edible producers and indirectly influence whether grazers are nutrient or energy limited. Second, through these controls, inedible producers can shape community assembly by excluding grazers that are weak competitors for nutrients contained in edible producers. A mesocosm experiment revealed patterns consistent with both predictions: high abundances of inedible algae were accompanied by low phosphorus contents of edible algae and low abundances of the grazer Daphnia. Both lines of inference suggest that interactions between stoichiometry and plant heterogeneity may shape plankton communities.

Adaptation to natural flow regimes.

Floods and droughts are important features of most running water ecosystems, but the alteration of natural flow regimes by recent human activities, such as dam building, raises questions related to both evolution and conservation. Among organisms inhabiting running waters, what adaptations exist for surviving floods and droughts? How will the alteration of the frequency, timing and duration of flow extremes affect flood- and drought-adapted organisms? How rapidly can populations evolve in response to altered flow regimes? Here, we identify three modes of adaptation (life history, behavioral and morphological) that plants and animals use to survive floods and/or droughts. The mode of adaptation that an organism has determines its vulnerability to different kinds of flow regime alteration. The rate of evolution in response to flow regime alteration remains an open question. Because humans have now altered the flow regimes of most rivers and many streams, understanding the link between fitness and flow regime is crucial for the effective management and restoration of running water ecosystems.