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Hawai‘i forest review: Synthesizing the ecology, evolution, and conservation of a model system

As the most remote archipelago in the world, the Hawaiian Islands are home to a highly endemic and disharmonic biota that has fascinated biologists for centuries. Forests are the dominant terrestrial biome in Hawai‘i, spanning complex, heterogeneous climates across substrates that vary tremendously in age, soil structure, and nutrient availability. Species richness is low in Hawaiian forests compared to other tropical forests, as a consequence of dispersal limitation from continents and adaptive radiations in only some lineages, and forests are dominated by the widespread Metrosideros species complex. Low species richness provides a relatively tractable model system for studies of community assembly, local adaptation, and species interactions. Moreover, Hawaiian forests provide insights into predicted patterns of evolution on islands, revealing that while some evidence supports “island syndromes,” there are exceptions to them all. For example, Hawaiian plants are not as a whole less defended against herbivores, less dispersible, more conservative in resource use, or more slow-growing than their continental relatives. Clearly, more work is needed to understand the drivers, sources, and constraints on phenotypic variation among Hawaiian species, including both widespread and rare species, and to understand the role of this variation for ecological and evolutionary processes, which will further contribute to conservation of this unique biota. Today, Hawaiian forests are among the most threatened globally. Resource management failures – the proliferation of non-native species in particular – have led to devastating declines in native taxa and resulted in dominance by novel species assemblages. Conservation and restoration of Hawaiian forests now rely on managing threats including climate change, ongoing species introductions, novel pathogens, lost mutualists, and altered ecosystem dynamics through the use of diverse tools and strategies grounded in basic ecological, evolutionary, and biocultural principles. The future of Hawaiian forests thus depends on the synthesis of ecological and evolutionary research, which will continue to inform future conservation and restoration practices.

Hawaii↗

Conceptualizing ecological responses to dam removal: If you remove it, what's to come?

One of the desired outcomes of dam decommissioning and removal is the recovery of aquatic and riparian ecosystems. To investigate this common objective, we synthesized information from empirical studies and ecological theory into conceptual models that depict key physical and biological links driving ecological responses to removing dams. We define models for three distinct spatial domains: upstream of the former reservoir, within the reservoir, and downstream of the removed dam. Emerging from these models are response trajectories that clarify potential pathways of ecological transitions in each domain. We illustrate that the responses are controlled by multiple causal pathways and feedback loops among physical and biological components of the ecosystem, creating recovery trajectories that are dynamic and nonlinear. In most cases, short-term effects are typically followed by longer-term responses that bring ecosystems to new and frequently predictable ecological condition, which may or may not be similar to what existed prior to impoundment.

BioScience↗

The relative importance of intrinsic and extrinsic drivers to population growth vary among local populations of Greater Sage-Grouse: An integrated population modeling approach

Consideration of ecological scale is fundamental to understanding and managing avian population growth and decline. Empirically driven models for population dynamics and demographic processes across multiple spatial scales can be powerful tools to help guide conservation actions. Integrated population models (IPMs) provide a framework for better parameter estimation by unifying multiple sources of data (e.g., count and demographic data). Hierarchical structure within such models that include random effects allow for varying degrees of data sharing across different spatiotemporal scales. We developed an IPM to investigate Greater Sage-Grouse ( Centrocercus urophasianus ) on the border of California and Nevada, known as the Bi-State Distinct Population Segment. Our analysis integrated 13 years of lek count data ( n > 2,000) and intensive telemetry (VHF and GPS; n > 350 individuals) data across 6 subpopulations. Specifically, we identified the most parsimonious models among varying random effects and density-dependent terms for each population vital rate (e.g., nest survival). Using a joint likelihood process, we integrated the lek count data with the demographic models to estimate apparent abundance and refine vital rate parameter estimates. To investigate effects of climatic conditions, we extended the model to fit a precipitation covariate for instantaneous rate of change ( r ). At a metapopulation extent (i.e. Bi-State), annual population rate of change λ ( e r ) did not favor an overall increasing or decreasing trend through the time series. However, annual changes in λ were driven by changes in precipitation (one-year lag effect). At subpopulation extents, we identified substantial variation in λ and demographic rates. One subpopulation clearly decoupled from the trend at the metapopulation extent and exhibited relatively high risk of extinction as a result of low egg fertility. These findings can inform localized, targeted management actions for specific areas, and status of the species for the larger Bi-State.

The Auk↗

Evaluating model structure adequacy: The case of the Maggia Valley groundwater system, southern Switzerland

Model adequacy is evaluated with alternative models rated using model selection criteria (AICc, BIC, and KIC) and three other statistics. Model selection criteria are tested with cross-validation experiments and insights for using alternative models to evaluate model structural adequacy are provided. The study is conducted using the computer codes UCODE_2005 and MMA (MultiModel Analysis). One recharge alternative is simulated using the TOPKAPI hydrological model. The predictions evaluated include eight heads and three flows located where ecological consequences and model precision are of concern. Cross-validation is used to obtain measures of prediction accuracy. Sixty-four models were designed deterministically and differ in representation of river, recharge, bedrock topography, and hydraulic conductivity. Results include: (1) What may seem like inconsequential choices in model construction may be important to predictions. Analysis of predictions from alternative models is advised. (2) None of the model selection criteria consistently identified models with more accurate predictions. This is a disturbing result that suggests to reconsider the utility of model selection criteria, and/or the cross-validation measures used in this work to measure model accuracy. (3) KIC displayed poor performance for the present regression problems; theoretical considerations suggest that difficulties are associated with wide variations in the sensitivity term of KIC resulting from the models being nonlinear and the problems being ill-posed due to parameter correlations and insensitivity. The other criteria performed somewhat better, and similarly to each other. (4) Quantities with high leverage are more difficult to predict. The results are expected to be generally applicable to models of environmental systems.

Water Resources Research↗

Estimating abundance: Chapter 27

This chapter provides a non-technical overview of ‘closed population capture–recapture’ models, a class of well-established models that are widely applied in ecology, such as removal sampling, covariate models, and distance sampling. These methods are regularly adopted for studies of reptiles, in order to estimate abundance from counts of marked individuals while accounting for imperfect detection. Thus, the chapter describes some classic closed population models for estimating abundance, with considerations for some recent extensions that provide a spatial context for the estimation of abundance, and therefore density. Finally, the chapter suggests some software for use in data analysis, such as the Windows-based program MARK, and provides an example of estimating abundance and density of reptiles using an artificial cover object survey of Slow Worms ( Anguis fragilis ).

Book chapter↗

The Everglades Depth Estimation Network (EDEN) surface-water interpolation model, version 3

The Everglades Depth Estimation Network (EDEN) is an integrated network of water-level gages, interpolation models that estimate daily water-level data at ungaged locations, and applications that generate derived hydrologic data across the freshwater part of the Greater Everglades landscape. Version 3 (V3) of the EDEN interpolation surface-water model is the most recent update, replacing the version 2 (V2) model released in 2011. The primary revision for the V3 model is the switch to the R programming language to create a more efficient and portable EDEN code relative to V2, without reliance on proprietary software. Using R, the interpolation script runs over 10 times faster and is more easily updated, for example, to accommodate changes in the gage network or to incorporate R software updates. Additional revisions made for the V3 model include updates to the interpolation model, the gage network, and groundwater-level estimations. The EDEN model domain in the Greater Everglades and Big Cypress National Preserve is divided into subdomains that are based on hydrologic boundaries. In the V3 model, the number of subdomains was increased from five to eight, which allows hydrologic boundaries, such as levees and canals, to be better represented in the interpolation scheme. Five pseudogages were added to constrain the water-level surface at subdomain boundaries. Changes made to the water-level gage network between the implementation of the V2 and V3 models are incorporated, and groundwater-level estimations are added, which are important information for hydrologic and ecological studies. Summary model performance statistics indicate similar accuracy in water-level surfaces generated by the V3 and V2 models, with a root mean square error of 4.78 centimeters for both interpolation models against independent water-level measurements. Providing stability and continuity for the EDEN user community, the V3 model closely replicates the V2 model, with a root mean square difference of 3.87 centimeters for interpolated surfaces from April 1, 2014, to March 31, 2018. The additional groundwater levels provide a realistic estimate of the saturated groundwater surface continuous with the surface-water surface for Water Conservation Areas 2A and 2B from 2000 to 2011. This continuous surface is a more accurate estimation of the spatial distribution of water in the hydrologic system than before, providing needed information for ecological studies in areas where depth to water table affects habitats. Development of the EDEN V3 model advances the tools available to scientists and resource managers for guiding large-scale field operations, describing hydrologic changes, and supporting biological and ecological assessments.

Florida↗

Nonlinear relationships can lead to bias in biomass calculations and drift-foraging models when using summaries of invertebrate drift data

Drift-foraging models offer a mechanistic description of how fish feed in flowing water and the application of drift-foraging bioenergetics models to answer both applied and theoretical questions in aquatic ecology is growing. These models typically include nonlinear descriptions of ecological processes and as a result may be sensitive to how model inputs are summarized because of a mathematical property of nonlinear equations known as Jensen’s inequality. In particular, we show that the way in which continuous size distributions of invertebrate prey are represented within foraging models can lead to biases within the modeling process. We begin by illustrating how different equations common to drift-foraging models are sensitive to invertebrate inputs. We then use two case studies to show how different representations of invertebrate prey can influence predictions of energy intake and lifetime growth. Greater emphasis should be placed on accurate characterizations of invertebrate drift, acknowledging that inferences from drift-foraging models may be influenced by how invertebrate prey are represented.

Environmental Biology of Fishes↗

Rayleigh step-selection functions and connections to continuous-time mechanistic movement models

Background The process known as ecological diffusion emerges from a first principles view of animal movement, but ecological diffusion and other partial differential equation models can be difficult to fit to data. Step-selection functions (SSFs), on the other hand, have emerged as powerful practical tools for ecologists studying the movement and habitat selection of animals. Methods SSFs typically involve comparing resources between a set of used and available points at each step in a sequence of observed positions. We use change of variables to show that ecological diffusion implies certain distributions for available steps that are more flexible than others commonly used. We then demonstrate advantages of these distributions with SSF models fit to data collected for a mountain lion in Colorado, USA. Results We show that connections between ecological diffusion and SSFs imply a Rayleigh step-length distribution and uniform turning angle distribution, which can accommodate data collected at irregular time intervals. The results of fitting an SSF model with these distributions compared to a set of commonly used distributions revealed how precision and inference can vary between the two approaches. Conclusions Our new continuous-time step-length distribution can be integrated into various forms of SSFs, making them applicable to data sets with irregular time intervals between successive animal locations.

Movement Ecology↗

Identifying legal, ecological and governance obstacles and opportunities for adapting to climate change

Current governance of regional scale water management systems in the United States has not placed them on a path toward sustainability, as conflict and gridlock characterize the social arena and ecosystem services continue to erode. Changing climate may continue this trajectory, but it also provides a catalyst for renewal of ecosystems and a window of opportunity for change in institutions. Resilience provides a bridging concept that predicts that change in ecological and social systems is often dramatic, abrupt, and surprising. Adapting to the uncertainty of climate driven change must be done in a manner perceived as legitimate by the participants in a democratic society. Adaptation must begin with the current hierarchical and fragmented social-ecological system as a baseline from which new approaches must be applied. Achieving a level of integration between ecological concepts and governance requires a dialogue across multiple disciplines, including ecologists with expertise in ecological resilience, hydrologists and climate experts, with social scientists and legal scholars. Criteria and models that link ecological dynamics with policies in complex, multi-jurisdictional water basins with adaptive management and governance frameworks may move these social-ecological systems toward greater sustainability.

Sustainability↗

The accuracy of ecological flow metrics derived using a physics-based distributed rainfall-runoff model in the Great Plains, USA

The development of a hydrologic foundation, essential for advancing our understanding of flow-ecology relationships, was developed using the high-resolution physics-based distributed rainfall–runoff model V flo in a semi-arid region. We compared the accuracy and bias associated with flow metrics that were generated using V flo , gauge data, and drainage area ratios at both a daily and monthly time step in the Canadian River basin, USA. First, we calibrated and applied bias correction to the V flo model to simulate streamflow at ungauged catchment locations. Next, flow metrics were calculated using simulated and observed data from stream gauge locations. We found discharge predictions using V flo were more accurate than drainage area ratios. General correspondence between predicted discharge and the gauge data was apparent; however, flow metrics calculated using the V flo output did not accurately represent flow variability. Results from the V flo model showed systematic discharge over-predictions in the upper basin and isolated over-predictions in the lower basin, likely due to hail events and sparse rainfall data across the large catchment. Goodness-of-fit statistics (Nash–Sutcliffe efficiency, root-mean square error, and the coefficient of variation) indicated the drainage area ratio and V flo were more accurate at a monthly rather than daily time step, even after quantile mapping. This finding limits the number of streamflow metrics available to develop ecological models, but more importantly, the coarser resolution may hinder our understanding of ecological processes that occur at a submonthly time step. Our approach provides a framework for selecting flow metrics that best represent hydrologic patterns across a large semi-arid catchment with the necessary accuracy to address the ecological questions of interest.

New Mexico, Oklahoma, Texas↗

Modeling avian abundance from replicated counts using binomial mixture models

Abundance estimation in ecology is usually accomplished by capture–recapture, removal, or distance sampling methods. These may be hard to implement at large spatial scales. In contrast, binomial mixture models enable abundance estimation without individual identification, based simply on temporally and spatially replicated counts. Here, we evaluate mixture models using data from the national breeding bird monitoring program in Switzerland, where some 250 1-km 2 quadrats are surveyed using the territory mapping method three times during each breeding season. We chose eight species with contrasting distribution (wide–narrow), abundance (high–low), and detectability (easy–difficult). Abundance was modeled as a random effect with a Poisson or negative binomial distribution, with mean affected by forest cover, elevation, and route length. Detectability was a logit-linear function of survey date, survey date-by-elevation, and sampling effort (time per transect unit). Resulting covariate effects and parameter estimates were consistent with expectations. Detectability per territory (for three surveys) ranged from 0.66 to 0.94 (mean 0.84) for easy species, and from 0.16 to 0.83 (mean 0.53) for difficult species, depended on survey effort for two easy and all four difficult species, and changed seasonally for three easy and three difficult species. Abundance was positively related to route length in three high-abundance and one low-abundance (one easy and three difficult) species, and increased with forest cover in five forest species, decreased for two nonforest species, and was unaffected for a generalist species. Abundance estimates under the most parsimonious mixture models were between 1.1 and 8.9 (median 1.8) times greater than estimates based on territory mapping; hence, three surveys were insufficient to detect all territories for each species. We conclude that binomial mixture models are an important new approach for estimating abundance corrected for detectability when only repeated-count data are available. Future developments envisioned include estimation of trend, occupancy, and total regional abundance.

Ecological Applications↗

Assessing tiger population dynamics using photographic capture-recapture sampling

Although wide-ranging, elusive, large carnivore species, such as the tiger, are of scientific and conservation interest, rigorous inferences about their population dynamics are scarce because of methodological problems of sampling populations at the required spatial and temporal scales. We report the application of a rigorous, noninvasive method for assessing tiger population dynamics to test model-based predictions about population viability. We obtained photographic capture histories for 74 individual tigers during a nine-year study involving 5725 trap-nights of effort. These data were modeled under a likelihood-based, ?robust design? capture?recapture analytic framework. We explicitly modeled and estimated ecological parameters such as time-specific abundance, density, survival, recruitment, temporary emigration, and transience, using models that incorporated effects of factors such as individual heterogeneity, trap-response, and time on probabilities of photo-capturing tigers. The model estimated a random temporary emigration parameter of =K' =Y' 0.10 ? 0.069 (values are estimated mean ? SE). When scaled to an annual basis, tiger survival rates were estimated at S = 0.77 ? 0.051, and the estimated probability that a newly caught animal was a transient was = 0.18 ? 0.11. During the period when the sampled area was of constant size, the estimated population size Nt varied from 17 ? 1.7 to 31 ? 2.1 tigers, with a geometric mean rate of annual population change estimated as = 1.03 ? 0.020, representing a 3% annual increase. The estimated recruitment of new animals, Bt, varied from 0 ? 3.0 to 14 ? 2.9 tigers. Population density estimates, D, ranged from 7.33 ? 0.8 tigers/100 km2 to 21.73 ? 1.7 tigers/100 km2 during the study. Thus, despite substantial annual losses and temporal variation in recruitment, the tiger density remained at relatively high levels in Nagarahole. Our results are consistent with the hypothesis that protected wild tiger populations can remain healthy despite heavy mortalities because of their inherently high reproductive potential. The ability to model the entire photographic capture history data set and incorporate reduced-parameter models led to estimates of mean annual population change that were sufficiently precise to be useful. This efficient, noninvasive sampling approach can be used to rigorously investigate the population dynamics of tigers and other elusive, rare, wide-ranging animal species in which individuals can be identified from photographs or other means.

Ecology↗

A generalized model for estimating the energy density of invertebrates

Invertebrate energy density (ED) values are traditionally measured using bomb calorimetry. However, many researchers rely on a few published literature sources to obtain ED values because of time and sampling constraints on measuring ED with bomb calorimetry. Literature values often do not account for spatial or temporal variability associated with invertebrate ED. Thus, these values can be unreliable for use in models and other ecological applications. We evaluated the generality of the relationship between invertebrate ED and proportion of dry-to-wet mass (pDM). We then developed and tested a regression model to predict ED from pDM based on a taxonomically, spatially, and temporally diverse sample of invertebrates representing 28 orders in aquatic (freshwater, estuarine, and marine) and terrestrial (temperate and arid) habitats from 4 continents and 2 oceans. Samples included invertebrates collected in all seasons over the last 19 y. Evaluation of these data revealed a significant relationship between ED and pDM ( r 2 = 0.96, p < 0.0001), where ED (as J/g wet mass) was estimated from pDM as ED = 22,960pDM &minus; 174.2. Model evaluation showed that nearly all (98.8%) of the variability between observed and predicted values for invertebrate ED could be attributed to residual error in the model. Regression of observed on predicted values revealed that the 97.5% joint confidence region included the intercept of 0 (&minus;103.0 &plusmn; 707.9) and slope of 1 (1.01 &plusmn; 0.12). Use of this model requires that only dry and wet mass measurements be obtained, resulting in significant time, sample size, and cost savings compared to traditional bomb calorimetry approaches. This model should prove useful for a wide range of ecological studies because it is unaffected by taxonomic, seasonal, or spatial variability.

Freshwater Science↗

A laboratory-calibrated model of coho salmon growth with utility for ecological analyses

We conducted a meta-analysis of laboratory- and hatchery-based growth data to estimate broadly applicable parameters of mass- and temperature-dependent growth of juvenile coho salmon ( Oncorhynchus kisutch ). Following studies of other salmonid species, we incorporated the Ratkowsky growth model into an allometric model and fit this model to growth observations from eight studies spanning ten different populations. To account for changes in growth patterns with food availability, we reparameterized the Ratkowsky model to scale several of its parameters relative to ration. The resulting model was robust across a wide range of ration allocations and experimental conditions, accounting for 99% of the variation in final body mass. We fit this model to growth data from coho salmon inhabiting tributaries and constructed ponds in the Klamath Basin by estimating habitat-specific indices of food availability. The model produced evidence that constructed ponds provided higher food availability than natural tributaries. Because of their simplicity (only mass and temperature are required as inputs) and robustness, ration-varying Ratkowsky models have utility as an ecological tool for capturing growth in freshwater fish populations.

Canadian Journal of Fisheries and Aquatic Sciences↗

Spatial occupancy models for predicting metapopulation dynamics and viability following reintroduction

The reintroduction of a species into its historic range is a critical component of conservation programmes designed to restore extirpated metapopulations. However, many reintroduction efforts fail, and the lack of rigorous monitoring programmes and statistical models have prevented a general understanding of the factors affecting metapopulation viability following reintroduction. Spatially explicit metapopulation theory provides the basis for understanding the dynamics of fragmented populations linked by dispersal, but the theory has rarely been used to guide reintroduction programmes because most spatial metapopulation models require presence&ndash;absence data from every site in the network, and they do not allow for observation error such as imperfect detection. We develop a spatial occupancy model that relaxes these restrictive assumptions and allows for inference about metapopulation extinction risk and connectivity. We demonstrate the utility of the model using six years of data on the Chiricahua leopard frog Lithobates chiricahuensis , a threatened desert-breeding amphibian that was reintroduced to a network of sites in Arizona USA in 2003. Our results indicate that the model can generate precise predictions of extinction risk and produce connectivity maps that can guide conservation efforts following reintroduction. In the case of L. chiricahuensis , many sites were functionally isolated, and 82% of sites were characterized by intermittent water availability and high local extinction probabilities (0&middot;84, 95% CI: 0&middot;64&ndash;0&middot;99). However, under the current hydrological conditions and spatial arrangement of sites, the risk of metapopulation extinction is estimated to be <3% over a 50-year time horizon. Low metapopulation extinction risk appears to result from the high dispersal capability of the species, the high density of sites in the region and the existence of predator-free permanent wetlands with low local extinction probabilities. Should management be required, extinction risk can be reduced by either increasing the hydroperiod of existing sites or by creating new sites to increase connectivity. Synthesis and applications . This work demonstrates how spatio-temporal statistical models based on ecological theory can be applied to forecast the outcomes of conservation actions such as reintroduction. Our spatial occupancy model should be particularly useful when management agencies lack the funds to collect intensive individual-level data.

Arizona, New Mexico↗

Creating multithemed ecological regions for macroscale ecology: Testing a flexible, repeatable, and accessible clustering method

Understanding broad-scale ecological patterns and processes often involves accounting for regional-scale heterogeneity. A common way to do so is to include ecological regions in sampling schemes and empirical models. However, most existing ecological regions were developed for specific purposes, using a limited set of geospatial features and irreproducible methods. Our study purpose was to: (1) describe a method that takes advantage of recent computational advances and increased availability of regional and global data sets to create customizable and reproducible ecological regions, (2) make this algorithm available for use and modification by others studying different ecosystems, variables of interest, study extents, and macroscale ecology research questions, and (3) demonstrate the power of this approach for the research question—How well do these regions capture regional-scale variation in lake water quality? To achieve our purpose we: (1) used a spatially constrained spectral clustering algorithm that balances geospatial homogeneity and region contiguity to create ecological regions using multiple terrestrial, climatic, and freshwater geospatial data for 17 northeastern U.S. states (~1,800,000 km 2 ); (2) identified which of the 52 geospatial features were most influential in creating the resulting 100 regions; and (3) tested the ability of these ecological regions to capture regional variation in water nutrients and clarity for ~6,000 lakes. We found that: (1) a combination of terrestrial, climatic, and freshwater geospatial features influenced region creation, suggesting that the oft-ignored freshwater landscape provides novel information on landscape variability not captured by traditionally used climate and terrestrial metrics; and (2) the delineated regions captured macroscale heterogeneity in ecosystem properties not included in region delineation—approximately 40% of the variation in total phosphorus and water clarity among lakes was at the regional scale. Our results demonstrate the usefulness of this method for creating customizable and reproducible regions for research and management applications.

Ecology and Evolution↗