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Charles B. Yackulic

Publications and source records attributed to Charles B. Yackulic.

At least 127 records · Page 7Linked to original sources

Taxonomic and compositional differences of ground-dwelling arthropods in riparian habitats in Glen Canyon, Arizona, USA

The disturbance history, plant species composition, productivity, and structural complexity of a site can exert bottom-up controls on arthropod diversity, abundance, and trophic structure. Regulation alters the hydrology and disturbance regimes of rivers and affects riparian habitats by changing plant quality parameters. Fifty years of regulation along the Colorado River downstream of Glen Canyon Dam has created a no-analog, postdam “lower” riparian zone close to the water's edge that includes tamarisk ( Tamarix sp.), a nonnative riparian shrub. At the same time, the predam “upper” facultative riparian zone has persisted several meters above the current flood stage. In summer 2009, we used pitfall traps within these 2 riparian zones that differ in plant composition, productivity, and disturbance frequency to test for differences in arthropod community (Hymenoptera, Arachnida, and Coleoptera) structure. Arthropod community structure differed substantially between the 2 zones. Arthropod abundance and species richness was highest in the predam upper riparian zone, even though there was a greater amount of standing plant biomass in the postdam lower riparian zone. Omnivore abundance was proportionately greater in the upper riparian zone and was associated with lower estimated productivity values. Predators and detritivores were proportionately greater in the postdam lower riparian zone. In this case, river regulation may create habitats that support species of spiders and carabid beetles, but few other species that are exclusive to this zone. The combined richness found in both zones suggests a small increase in total richness and functional diversity for the Glen Canyon reach of the Colorado River.

Arizona↗

Competitive exclusion over broad spatial extents is a slow process: Evidence and implications for species distribution modeling

There is considerable debate about the role of competition in shaping species distributions over broad spatial extents. This debate has practical implications because predicting changes in species' geographic ranges in response to ongoing environmental change would be simpler if competition could be ignored. While this debate has been the subject of many reviews, recent literature has not addressed the rates of relevant processes. This omission is surprising in that ecologists hypothesized decades ago that regional competitive exclusion is a slow process. The goal of this review is to reassess the debate under the hypothesis that competitive exclusion over broad spatial extents is a slow process. Available evidence, including simulations presented for the first time here, suggests that competitive exclusion over broad spatial extents occurs slowly over temporal extents of many decades to millennia. Ecologists arguing against an important role for competition frequently study modern patterns and/or range dynamics over periods of decades, while much of the evidence for competition shaping geographic ranges at broad spatial extents comes from paleoecological studies over time scales of centuries or longer. If competition is slow, as evidence suggests, the geographic distributions of some, perhaps many species, would continue to change over time scales of decades to millennia, even if environmental conditions did not continue to change. If the distributions of competing species are at equilibrium it is possible to predict species distributions based on observed species–environment relationships. However, disequilibrium is widespread as a result of competition and many other processes. Studies whose goal is accurate predictions over intermediate time scales (decades to centuries) should focus on factors associated with range expansion (colonization) and loss (local extinction), as opposed to current patterns. In general, understanding of modern range dynamics would be enhanced by considering the rates of relevant processes.

Ecography↗

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↗

Allometric and temporal scaling of movement characteristics in Galapagos tortoises

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

Journal of Animal Ecology↗

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

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

Movement Ecology↗

Flow management for hydropower extirpates aquatic insects, undermining river food webs

Dams impound the majority of rivers and provide important societal benefits, especially daily water releases that enable on-peak hydroelectricity generation. Such “hydropeaking” is common worldwide, but its downstream impacts remain unclear. We evaluated the response of aquatic insects, a cornerstone of river food webs, to hydropeaking using a life history–hydrodynamic model. Our model predicts that aquatic-insect abundance will depend on a basic life-history trait—adult egg-laying behavior—such that open-water layers will be unaffected by hydropeaking, whereas ecologically important and widespread river-edge layers, such as mayflies, will be extirpated. These predictions are supported by a more-than-2500-sample, citizen-science data set of aquatic insects from the Colorado River in the Grand Canyon and by a survey of insect diversity and hydropeaking intensity across dammed rivers of the Western United States. Our study reveals a hydropeaking-related life history bottleneck that precludes viable populations of many aquatic insects from inhabiting regulated rivers.

BioScience↗

Demographic response of northern spotted owls to barred owl removal

Federally listed as threatened in 1990 primarily because of habitat loss, the northern spotted owl ( Strix occidentalis caurina ) has continued to decline despite conservation efforts resulting in forested habitat being reserved throughout its range. Recently, there is growing evidence the congeneric invasive barred owl ( Strix varia ) may be responsible for the continued decline primarily by excluding spotted owls from their preferred habitat. We used a long-term demographic study for spotted owls in coastal northern California as the basis for a pilot barred owl removal experiment. Our demography study used capture–recapture, reproductive output, and territory occupancy data collected from 1990 to 2013 to evaluate trends in vital rates and populations. We used a classic before-after-control-impact (BACI) experimental design to investigate the demographic response of northern spotted owls to the lethal removal of barred owls. According to the best 2-species dynamic occupancy model, there was no evidence of differences in barred or northern spotted owl occupancy prior to the initiation of the treatment (barred owl removal). After treatment, barred owl occupancy was lower in the treated relative to the untreated areas and spotted owl occupancy was higher relative to the untreated areas. Barred owl removal decreased spotted owl territory extinction rates but did not affect territory colonization rates. As a result, spotted owl occupancy increased in the treated area and continued to decline in the untreated areas. Prior to and after barred owl removal, there was no evidence that average fecundity differed on the 2 study areas. However, the greater number of occupied spotted owl sites on the treated areas resulted in greater productivity in the treated areas based on empirical counts of fledged young. Prior to removal, survival was declining at a rate of approximately 0.2% per year for treated and untreated areas. Following treatment, estimated survival was 0.859 for the treated areas and 0.822 for the untreated areas. Derived estimates of population change on both study areas showed the same general decline before removal with an estimated slope of –0.0036 per year. Following removal, the rate of population change on the treated areas increased to an average of 1.029 but decreased to an average of 0.870 on the untreated areas. The results from this first experiment demonstrated that lethal removal of barred owls allowed the recovery of northern spotted owl populations in the treated portions of our study area. If additional federally funded barred owl removal experiments provide similar results, this could be the foundation for development of a long-term conservation strategy for northern spotted owls.

California↗

Decision analysis to support development of the Glen Canyon Dam long-term experimental and management plan

The U.S. Geological Survey, in cooperation with the Bureau of Reclamation, National Park Service, and Argonne National Laboratory, completed a decision analysis to use in the evaluation of alternatives in the Environmental Impact Statement concerning the long-term management of water releases from Glen Canyon Dam and associated management activities. Two primary decision analysis methods, multicriteria decision analysis and the expected value of information, were used to evaluate the alternative strategies against the resource goals and to evaluate the influence of uncertainty. A total of 18 performance metrics associated with 8 out of 12 resource goals (fundamental objectives) were developed by the Bureau of Reclamation and National Park Service in partnership with subject-matter teams composed of Federal, State, tribal, and private experts. A total of 19 long-term strategies associated with 7 alternatives were developed by the Bureau of Reclamation, National Park Service, Argonne National Laboratory, U.S. Geological Survey, and Cooperating Agencies. The 19 long-term strategies were evaluated against the 18 performance metrics using a series of coupled simulation models, taking into account the effects of several important sources of uncertainty. A total of 27 Federal, State, tribal, and nongovernmental agencies were invited by the Assistant Secretary of Interior to participate in a swing-weighting exercise to understand the range of perspectives about how to place relative value on the resource goals and performance metrics; 14 of the 27 chose to participate. The results of the swing-weighting exercise were combined with the evaluation of the alternatives to complete a multicriteria decision analysis. The effects of uncertainty on the ranking of long-term strategies were evaluated through calculation of the value of information. The alternatives and their long-term strategies differed across performance metrics, producing unavoidable tradeoffs; thus, there was no long-term strategy that was dominated by another across all performance metrics. When the performance of each alternative was weighted across performance metrics, three alternatives (B, D, and G) were top-ranked depending on the set of weights proposed: Alternative B was favored by those stakeholders that placed a high value on hydropower; Alternative G was favored by those stakeholders that placed a high value on the restoration of natural processes, like beachbuilding and natural vegetation; and Alternative D was favored by the remaining stakeholders. Surprisingly, these rankings were not sensitive to the critical uncertainties that were evaluated; that is, the choice of a preferred long-term strategy was sensitive to the value-based judgment about how to place relative weight on the resource goals but was not sensitive to the uncertainties in the system dynamics that were evaluated in this analysis. The one area of uncertainty that did slightly affect the ranking of alternatives was the long-term pattern of hydrological input; because of this sensitivity, some attention to the possible effects of climate change is warranted. The results of the decision analysis are meant to serve as only one of many sources of information that can be used to evaluate the alternatives proposed in the Environmental Impact Statement. These results only focus on those resource goals for which quantitative performance metrics could be formulated and evaluated; there are other important aspects of the resource goals that also need to be considered. Not all the stakeholders who were invited to participate in the decision analysis chose to do so; thus, the Bureau of Reclamation, National Park Service, and U.S. Department of Interior may want to consider other input.

Arizona↗

Prey size and availability limits maximum size of rainbow trout in a large tailwater: insights from a drift-foraging bioenergetics model

The cold and clear water conditions present below many large dams create ideal conditions for the development of economically important salmonid fisheries. Many of these tailwater fisheries have experienced declines in the abundance and condition of large trout species, yet the causes of these declines remain uncertain. Here, we develop, assess, and apply a drift-foraging bioenergetics model to identify the factors limiting rainbow trout (Oncorhynchus mykiss) growth in a large tailwater. We explored the relative importance of temperature, prey quantity, and prey size by constructing scenarios where these variables, both singly and in combination, were altered. Predicted growth matched empirical mass-at-age estimates, particularly for younger ages, demonstrating that the model accurately describes how current temperature and prey conditions interact to determine rainbow trout growth. Modeling scenarios that artificially inflated prey size and abundance demonstrate that rainbow trout growth is limited by the scarcity of large prey items and overall prey availability. For example, shifting 10% of the prey biomass to the 13 mm (large) length class, without increasing overall prey biomass, increased lifetime maximum mass of rainbow trout by 88%. Additionally, warmer temperatures resulted in lower predicted growth at current and lower levels of prey availability; however, growth was similar across all temperatures at higher levels of prey availability. Climate change will likely alter flow and temperature regimes in large rivers with corresponding changes to invertebrate prey resources used by fish. Broader application of drift-foraging bioenergetics models to build a mechanistic understanding of how changes to habitat conditions and prey resources affect growth of salmonids will benefit management of tailwater fisheries.

Arizona↗

The scaling of geographic ranges: implications for species distribution models

Context The geographic ranges of many species are responding to ongoing environmental change. Processes operating at different levels of biological organization, with corresponding spatial extents and grains and temporal rates, interact with the evolving configuration of environmental conditions to determine range dynamics. Objectives To synthesize understanding of scales and scaling, including relevant biological levels of organization, focusing on the processes that mediate species-environment relationships and the models used to make inferences about species distributions. Methods We review concepts related to the scaling of geographic ranges and implications for the most commonly used analytic methods, using simple simulations to illustrate important issues. Results Many processes lead to species distributions being dependent on environmental conditions within sites and within a neighborhood. Studies with large extents and fine grains can cut across several levels of biological organization (individual, within-population, and metapopulation processes) complicating interpretation. Many geographic ranges are not in dynamic equilibrium, but common models used for inference assume equilibrium. Interspecific interactions shape species distributions at multiple scales, and arguments for ignoring species interactions also assume equilibrium. Conclusions There is a need for timely science to inform policy and management decisions; however, we must also strive to provide predictions that best reflect our understanding of ecological systems. Species distributions evolve through time and reflect responses to environmental conditions that are mediated through individual and population processes. Species distribution models that reflect this understanding, and explicitly model dynamics, are likely to give more accurate predictions.

Landscape Ecology↗

The effects of habitat, climate, and Barred Owls on long-term demography of Northern Spotted Owls

Estimates of species' vital rates and an understanding of the factors affecting those parameters over time and space can provide crucial information for management and conservation. We used mark–recapture, reproductive output, and territory occupancy data collected during 1985–2013 to evaluate population processes of Northern Spotted Owls ( Strix occidentalis caurina ) in 11 study areas in Washington, Oregon, and northern California, USA. We estimated apparent survival, fecundity, recruitment, rate of population change, and local extinction and colonization rates, and investigated relationships between these parameters and the amount of suitable habitat, local and regional variation in meteorological conditions, and competition with Barred Owls ( Strix varia ). Data were analyzed for each area separately and in a meta-analysis of all areas combined, following a strict protocol for data collection, preparation, and analysis. We used mixed effects linear models for analyses of fecundity, Cormack-Jolly-Seber open population models for analyses of apparent annual survival (ϕ), and a reparameterization of the Jolly-Seber capture–recapture model (i.e. reverse Jolly-Seber; RJS) to estimate annual rates of population change (λ RJS ) and recruitment. We also modeled territory occupancy dynamics of Northern Spotted Owls and Barred Owls in each study area using 2-species occupancy models. Estimated mean annual rates of population change (λ) suggested that Spotted Owl populations declined from 1.2% to 8.4% per year depending on the study area. The weighted mean estimate of λ for all study areas was 0.962 (± 0.019 SE; 95% CI: 0.925–0.999), indicating an estimated range-wide decline of 3.8% per year from 1985 to 2013. Variation in recruitment rates across the range of the Spotted Owl was best explained by an interaction between total winter precipitation and mean minimum winter temperature. Thus, recruitment rates were highest when both total precipitation (29 cm) and minimum winter temperature (−9.5°C) were lowest. Barred Owl presence was associated with increased local extinction rates of Spotted Owl pairs for all 11 study areas. Habitat covariates were related to extinction rates for Spotted Owl pairs in 8 of 11 study areas, and a greater amount of suitable owl habitat was generally associated with decreased extinction rates. We observed negative effects of Barred Owl presence on colonization rates of Spotted Owl pairs in 5 of 11 study areas. The total amount of suitable Spotted Owl habitat was positively associated with colonization rates in 5 areas, and more habitat disturbance was associated with lower colonization rates in 2 areas. We observed strong declines in derived estimates of occupancy in all study areas. Mean fecundity of females was highest for adults (0.309 ± 0.027 SE), intermediate for 2-yr-olds (0.179 ± 0.040 SE), and lowest for 1-yr-olds (0.065 ± 0.022 SE). The presence of Barred Owls and habitat covariates explained little of the temporal variation in fecundity in most study areas. Climate covariates occurred in competitive fecundity models in 8 of 11 study areas, but support for these relationships was generally weak. The fecundity meta-analysis resulted in 6 competitive models, all of which included the additive effects of geographic region and annual time variation. The 2 top-ranked models also weakly supported the additive negative effects of the amount of suitable core area habitat, Barred Owl presence, and the amount of edge habitat on fecundity. We found strong support for a negative effect of Barred Owl presence on apparent survival of Spotted Owls in 10 of 11 study areas, but found few strong effects of habitat on survival at the study area scale. Climate covariates occurred in top or competitive survival models for 10 of 11 study areas, and in most cases the relationships were as predicted; however, there was little consistency among areas regarding the relative importance of specific climate covariates. In contrast, meta-analysis results suggested that Spotted Owl survival was higher across all study areas when the Pacific Decadal Oscillation (PDO) was in a warming phase and the Southern Oscillation Index (SOI) was negative, with a strongly negative SOI indicative of El Niño events. The best model that included the Barred Owl covariate (BO) was ranked 4 th and also included the PDO covariate, but the BO effect was strongly negative. Our results indicated that Northern Spotted Owl populations were declining throughout the range of the subspecies and that annual rates of decline were accelerating in many areas. We observed strong evidence that Barred Owls negatively affected Spotted Owl populations, primarily by decreasing apparent survival and increasing local territory extinction rates. However, the amount of suitable owl habitat, local weather, and regional climatic patterns also were related to survival, occupancy (via colonization rate), recruitment, and, to a lesser extent, fecundity, although there was inconsistency in regard to which covariates were important for particular demographic parameters or across study areas. In the study areas where habitat was an important source of variation for Spotted Owl demographics, vital rates were generally positively associated with a greater amount of suitable owl habitat. However, Barred Owl densities may now be high enough across the range of the Northern Spotted Owl that, despite the continued management and conservation of suitable owl habitat on federal lands, the long-term prognosis for the persistence of Northern Spotted Owls may be in question without additional management intervention. Based on our study, the removal of Barred Owls from the Green Diamond Resources (GDR) study area had rapid, positive effects on Northern Spotted Owl survival and the rate of population change, supporting the hypothesis that, along with habitat conservation and management, Barred Owl removal may be able to slow or reverse Northern Spotted Owl population declines on at least a localized scale.

California, Oregon, Washington↗

Not putting all their eggs in one basket: bet-hedging despite extraordinary annual reproductive output of desert tortoises

Bet-hedging theory makes the counter-intuitive prediction that, if juvenile survival is low and unpredictable, organisms should consistently reduce short-term reproductive output to minimize the risk of reproductive failure in the long-term. We investigated the long-term reproductive output of an Agassiz's desert tortoise ( Gopherus agassizii ) population and conformance to a bet-hedging strategy of reproduction in an unpredictable but comparatively productive environment. Most females reproduced every year, even during periods of low precipitation and poor germination of food plants, and the mean percentage of reproducing females did not differ significantly on an annual basis. Although mean annual egg production (clutch size × clutch frequency) differed significantly among years, mean clutch size and mean clutch frequency remained relatively constant. During an El Niño year, mean annual egg production and mean annual clutch frequency were the highest ever reported for this species. Annual egg production was positively influenced by maternal body size but clutch size and clutch frequency were not. Our long-term results confirm earlier conclusions based on short-term research that desert tortoises have a bet-hedging strategy of producing small clutches almost every year. The risk of long-term reproductive failure is minimized in unpredictable environments, both through time by annually producing multiple small clutches over a long reproductive lifespan, even in years of low resource availability, and through space by depositing multiple annual clutches in different locations. The extraordinary annual reproductive output of this population appears to be the result of a typically high but unpredictable biomass of annual food plants at the site relative to tortoise habitat in dryer regions. Under the comparatively productive but unpredictable conditions, tortoises conform to predictions of a bet-hedging strategy of reproduction with relatively small but consistent clutch sizes.

Biological Journal of the Linnean Society↗

Assessing juvenile native fish demographic responses to a steady flow experiment in a large regulated river

The Colorado River below Glen Canyon Dam, Arizona, is part of an adaptive management programme which optimizes dam operations to improve various resources in the downstream ecosystem within Grand Canyon. Understanding how populations of federally endangered humpback chub Gila cypha respond to these dam operations is a high priority. Here, we test hypotheses concerning temporal variation in juvenile humpback chub apparent survival rates and abundance by comparing estimates between hydropeaking and steady discharge regimes over a 3-year period (July 2009–July 2012). The most supported model ignored flow type (steady vs hydropeaking) and estimated a declining trend in daily apparent survival rate across years (99.90%, 99.79% and 99.67% for 2009, 2010 and 2011, respectively). Corresponding abundance of juvenile humpback chub increased temporally; open population model estimates ranged from 615 to 2802 individuals/km, and closed model estimates ranged from 94 to 1515 individuals/km. These changes in apparent survival and abundance may reflect broader trends, or simply represent inter-annual variation. Important findings include (i) juvenile humpback chub are currently surviving and recruiting in the mainstem Colorado River with increasing abundance; (ii) apparent survival does not benefit from steady fall discharges from Glen Canyon Dam; and (iii) direct assessment of demographic parameters for juvenile endangered fish are possible and can rapidly inform management actions in regulated rivers.

River Research and Applications↗

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

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

Galapagos Islands↗

Turbidity, light, temperature, and hydropeaking control primary productivity in the Colorado River, Grand Canyon

Dams and river regulation greatly alter the downstream environment for gross primary production (GPP) because of changes in water clarity, flow, and temperature regimes. We estimated reach-scale GPP in five locations of the regulated Colorado River in Grand Canyon using an open channel model of dissolved oxygen. Benthic GPP dominates in Grand Canyon due to fast transport times and low pelagic algal biomass. In one location, we used a 738 days time series of GPP to identify the relative contribution of different physical controls of GPP. We developed both linear and semimechanistic time series models that account for unmeasured temporal covariance due to factors such as algal biomass dynamics. GPP varied from 0 g O 2 m −2 d −1 to 3.0 g O 2 m −2 d −1 with a relatively low annual average of 0.8 g O 2 m −2 d −1 . Semimechanistic models fit the data better than linear models and demonstrated that variation in turbidity primarily controlled GPP. Lower solar insolation during winter and from cloud cover lowered GPP much further. Hydropeaking lowered GPP but only during turbid conditions. Using the best model and parameter values, the model accurately predicted seasonal estimates of GPP at 3 of 4 upriver sites and outperformed the linear model at all sites; discrepancies were likely from higher algal biomass at upstream sites. This modeling approach can predict how changes in physical controls will affect relative rates of GPP throughout the 385 km segment of the Colorado River in Grand Canyon and can be easily applied to other streams and rivers.

Arizona↗

Factors controlling the abundance of rainbow trout in the Colorado River in Grand Canyon in a reach utilized by endangered humpback chub

We estimated the abundance, survival, movement, and recruitment of non-native rainbow trout in the Colorado River in Grand Canyon to determine what controls their abundance near the Little Colorado River (LCR) confluence where endangered humpback chub rear. Over a 3-year period, we tagged more than 70,000 trout and recovered over 8,200 tagged fish. Trout density was highest (10,000-25,000 fish/km) in the reach closest to Glen Canyon Dam where the majority of trout recruitment occurs, and was 30-50-fold lower (200-800 fish/km) in reaches near the LCR confluence ~100 km downstream. The extent of rainbow trout movement was limited with less than 1% of recaptures making movements greater than 20 km. However, due to high trout densities in upstream source areas, this small dispersal rate was sufficient to explain the 3-fold increase in the relatively small population near the LCR. Reducing dispersal rates of trout from upstream sources is the most feasible solution to maintain low densities near the LCR to minimize negative effects of competition and predation on humpback chub.

Arizona↗

To predict the niche, model colonization and extinction

Ecologists frequently try to predict the future geographic distributions of species. Most studies assume that the current distribution of a species reflects its environmental requirements (i.e., the species' niche). However, the current distributions of many species are unlikely to be at equilibrium with the current distribution of environmental conditions, both because of ongoing invasions and because the distribution of suitable environmental conditions is always changing. This mismatch between the equilibrium assumptions inherent in many analyses and the disequilibrium conditions in the real world leads to inaccurate predictions of species' geographic distributions and suggests the need for theory and analytical tools that avoid equilibrium assumptions. Here, we develop a general theory of environmental associations during periods of transient dynamics. We show that time-invariant relationships between environmental conditions and rates of local colonization and extinction can produce substantial temporal variation in occupancy–environment relationships. We then estimate occupancy–environment relationships during three avian invasions. Changes in occupancy–environment relationships over time differ among species but are predicted by dynamic occupancy models. Since estimates of the occupancy–environment relationships themselves are frequently poor predictors of future occupancy patterns, research should increasingly focus on characterizing how rates of local colonization and extinction vary with environmental conditions.

Ecology↗