A flexible consistent framework for modelling multiple interacting environmental responses to management in space and time.

Management of resources is often a large-scale task addressed using many small-scale interventions. The range of scales at which organisms respond to those interventions, along with the many outcomes which management aims to achieve can make determining the success of management complex. Environmental flow is an example of management where there is a recognized need for managers to demonstrate the impact of their actions by integrating different types of environmental responses. Here, we aim to support decision making in environmental management via the development of a new modelling framework (eFlowEval). It has the capacity to capture best-available knowledge, to scale it in space and time, explore interactions among species, compare scenarios, and account for uncertainty. Thus, it provides a basis for including multiple target groups in a common system. The framework is readily updatable as new information becomes available and can identify where data are insufficient to be scientifically robust. We demonstrate the eFlowEval framework using three very different environmental responses: 1) metabolism, which is a measure of the energy produced and then used in an ecosystem, 2) favorability for a bird species of interest (royal spoonbill Platalea regia), and 3) competing wetland plants (Centipeda cunninghamii and lippia Phyla canescens). These demonstrations illustrate the capability of the eFlowEval framework but the specific outputs shown here should not be used to assess environmental responses to management. Using these demonstrations, we illustrate the capacity of the eFlowEval framework to provide assessments across a range of scales (local to landscape) and from short time frames (weeks to months) to multi-year assessments. Further, we illustrate the ability to: i) scale responses from local to basin scales, ii) vary driver-response model types, iii) represent uncertainty, iv) compare scenarios, v) accommodate variable parameter values at different locations, and vi) incorporate spatial and temporal dependencies and dependencies among species. We also illustrate the framework's ability to capture inter- and intraspecific interactions and their impact in space and time. The eFlowEval framework extends the capacity of the component response models to provide novel modeling capabilities for management at scale. It allows for interactions among species or processes to be incorporated, as well as in space and time. A large degree of flexibility is offered by the framework, in terms of driver-response model types, input data, and aggregation methods. Thus, the eFlowEval framework provides a mechanism to enhance the transparency of environmental watering decision making, capture institutional knowledge, enhance adaptive management and undertake evaluation of the impact of environmental watering at a range of spatial and temporal scales.

Stream and ocean hydrodynamics mediate partial migration strategies in an amphidromous Hawaiian goby.

Partial migration strategies, in which some individuals migrate but others do not, are widely observed in populations of migratory animals. Such patterns could arise via variation in migratory behaviors made by individual animals, via genetic variation in migratory predisposition, or simply by variation in migration opportunities mediated by environmental conditions. Here we use spatiotemporal variation in partial migration across populations of an amphidromous Hawaiian goby to test whether stream or ocean conditions favor completing its life cycle entirely within freshwater streams rather than undergoing an oceanic larval migration. Across 35 watersheds, microchemical analysis of otoliths revealed that most adult Awaous stamineus were freshwater residents (62% of n = 316 in 2009, 83% of n = 274 in 2011), but we found considerable variation among watersheds. We then tested the hypothesis that the prevalence of freshwater residency increases with the stability of stream flows and decreases with the availability of dispersal pathways arising from ocean hydrodynamics. We found that streams with low variation of daily discharge were home to a higher incidence of freshwater residents in each survey year. The magnitude of the shift in freshwater residency between survey years was positively associated with predicted interannual variability in the success of larval settlement in streams on each island based on passive drift in ocean currents. We built on these findings by developing a theoretical model of goby life history to further evaluate whether mediation of migration outcomes by stream and ocean hydrodynamics could be sufficient to explain the range of partial migration frequency observed across populations. The model illustrates that the proportion of larvae entering the ocean and differential survival of freshwater-resident versus ocean-going larvae are plausible mechanisms for range-wide shifts in migration strategies. Thus, we propose that hydrologic variation in both ocean and stream environments contributes to spatiotemporal variation in the prevalence of migration phenotypes in A. stamineus. Our empirical and theoretical results suggest that the capacity for partial migration could enhance the persistence of metapopulations of diadromous fish when confronted with variable ocean and stream conditions.

Scale-Dependent Community Theory for Streams and Other Linear Habitats.

The maintenance of species diversity occurs at the regional scale but depends on interacting processes at the full range of lower scales. Although there is a long history of study of regional diversity as an emergent property, analyses of fully multiscale dynamics are rare. Here, we use scale transition theory for a quantitative analysis of multiscale diversity maintenance with continuous scales of dispersal and environmental variation in space and time. We develop our analysis with a model of a linear habitat, applicable to streams or coastlines, to provide a theoretical foundation for the long-standing interest in environmental variation and dispersal, including downstream drift. We find that the strength of regional coexistence is strongest when local densities and local environmental conditions are strongly correlated. Increasing dispersal and shortening environmental correlations weaken the strength of coexistence regionally and shift the dominant coexistence mechanism from fitness-density covariance to the spatial storage effect, while increasing local diversity. Analysis of the physical and biological determinants of these mechanisms improves understanding of traditional concepts of environmental filters, mass effects, and species sorting. Our results highlight the limitations of the binary distinction between local communities and a species pool and emphasize species coexistence as a problem of multiple scales in space and time.

Variation in moisture duration as a driver of coexistence by the storage effect in desert annual plants.

Temporal environmental variation is a leading hypothesis for the coexistence of desert annual plants. Environmental variation is hypothesized to cause species-specific patterns of variation in germination, which then generates the storage effect coexistence mechanism. However, it has never been shown how sufficient species differences in germination patterns for multispecies coexistence can arise from a shared fluctuating environment. Here we show that nonlinear germination responses to a single fluctuating physical environmental factor can lead to sufficient differences between species in germination pattern for the storage effect to yield coexistence of multiple species. We derive these nonlinear germination responses from experimental data on the effects of varying soil moisture duration. Although these nonlinearities lead to strong species asymmetries in germination patterns, the relative nonlinearity coexistence mechanism is minor compared with the storage effect. However, these asymmetries mean that the storage effect can be negative for some species, which then only persist in the face of interspecific competition through average fitness advantages. This work shows how a low dimensional physical environment can nevertheless stabilize multispecies coexistence when the species have different nonlinear responses to common conditions, as supported by our experimental data.

Population structure influences sexual conflict in wild populations of water striders.

In sexual conflict, aggressive males frequently diminish the long-term reproductive success of females in efforts to gain a short-term advantage over rival males. This short-term advantage can selectively favour high-exploitation males. However, just as the over-exploitation of resources can lead to local extinction, the over-exploitation of females in the form of harassment by aggressive males can yield similar consequences resulting in reduced female fecundity, increased female mortality and overall decline in mating activity. This outcome may often be prevented by selection acting at multiple levels of biological organization. Directional selection favouring aggressive exploitation within groups can be balanced by directional selection amongst groups opposing exploitation. Such between-group selection has recently been demonstrated in laboratory studies of water striders, where the conditional dispersal of individuals increased variation amongst groups and influenced the balance of selection toward reduced male aggression. This multilevel selection (MLS) framework also provides predictive value when investigating natural populations differing in their relative strength of selection within versus among groups. For water striders, the consequences of local exploitation cause fitness differences between groups, favouring less aggressive males. Inconsistently flowing ephemeral streams consist of isolated pools that prevent aggressive male water striders from escaping the consequences of local exploitation. We, therefore, predicted that inconsistently flowing ephemeral streams would favour the evolution of less aggressive males than would perennial streams, which allow aggressive males to move more freely and to escape the group-level costs of their aggression. Comparing two neighbouring streams during the mating season, we found that males dispersed naturally between pools at much higher rates in the perennial stream than in the ephemeral stream. As predicted, we found that males from the perennial stream were significantly more aggressive than those from the ephemeral stream. We also found that dispersers were significantly more aggressive than non-dispersers within each stream. These field results illustrate the relevance of the MLS framework in our understanding of the evolution of sexual conflict.