Spatial variogram estimation from temporally aggregated seabird count data

Seabird abundance is an important indicator for assessing impact of human activities on the marine environment. However, data collection at sea is time consuming and surveys are carried out over several consecutive days for efficiency reasons. This study investigates the validity of aggregating those data over time to estimate a spatial variogram that is representative for spatial correlation in species abundance. For this purpose we simulate four-day surveys of seabird count data that contain spatial and temporal correlation arising from temporal changes in the spatial pattern of environmental conditions. Estimates of the aggregated spatial variogram are compared to a variogram that would arise when data were collected over a single day. The study reveals that, under changing environmental conditions over surveys days, aggregating data over a four-day survey increases both the non-spatial variation in the data and the scale of spatial correlation in seabird data. Next, the effect of using an aggregated variogram on the statistical power to test the significance of an impact is investigated. The impact concerns a case of establishing an offshore wind farm resulting in seabird displacement. The study shows that both overestimation and underestimation of statistical power occurs, with power estimates differing up to a factor of two. We conclude that the spatial variation in seabird abundance can be misrepresented by using temporally aggregated data. In impact studies, such misrepresentation can lead to erroneous assessments of the ability to detect impact.     

Fishing the line near marine reserves in single and multispecies fisheries.

Throughout the world "fishing the line" is a frequent harvesting tactic in communities where no-take marine reserves are designated. This practice of concentrating fishing effort at the boundary of a marine reserve is predicated upon the principle of spillover, the net export of stock from the marine reserve to the surrounding unprotected waters. We explore the consequences and optimality of fishing the line using a spatially explicit theoretical model. We show that fishing the line: (1) is part of the optimal effort distribution near no-take marine reserves with mobile species regardless of the cooperation level among harvesters; (2) has a significant impact on the spatial patterns of catch per unit effort (CPUE) and fish density both within and outside of the reserve; and (3) can enhance total population size and catch simultaneously under a limited set of conditions for overexploited populations. Additionally, we explore the consequences of basing the spatial distribution of fishing effort for a multispecies fishery upon the optimality of the most mobile species that exhibits the greatest spillover. Our results show that the intensity of effort allocated to fishing the line should instead be based upon more intermediate rates of mobility within the targeted community. We conclude with a comparison between model predictions and empirical findings from a density gradient study of two important game fish in the vicinity of a no-take marine-life refuge on Santa Catalina Island, California (USA). These results reveal the need for empirical studies to account for harvester behavior and suggest that the implications of spatial discontinuities such as fishing the line should be incorporated into marine-reserve design.     

Accounting for enforcement costs in the spatial allocation of marine zones

Marine fish stocks are in many cases extracted above sustainable levels, but they may be protected through restricted‐use zoning systems. The effectiveness of these systems typically depends on support from coastal fishing communities. High management costs including those of enforcement may, however, deter fishers from supporting marine management. We incorporated enforcement costs into a spatial optimization model that identified how conservation targets can be met while maximizing fishers’ revenue. Our model identified the optimal allocation of the study area among different zones: no‐take, territorial user rights for fisheries (TURFs), or open access. The analysis demonstrated that enforcing no‐take and TURF zones incurs a cost, but results in higher species abundance by preventing poaching and overfishing. We analyzed how different enforcement scenarios affected fishers’ revenue. Fisher revenue was approximately 50% higher when territorial user rights were enforced than when they were not. The model preferentially allocated area to the enforced‐TURF zone over other zones, demonstrating that the financial benefits of enforcement (derived from higher species abundance) exceeded the costs. These findings were robust to increases in enforcement costs but sensitive to changes in species’ market price. We also found that revenue under the existing zoning regime in the study area was 13–30% lower than under an optimal solution. Our results highlight the importance of accounting for both the benefits and costs of enforcement in marine conservation, particularly when incurred by fishers. Justificación de los Costos de Aplicación en la Asignación Espacial de Zonas Marinas     

Understanding abundance patterns of a declining seabird: implications for monitoring.

The Kittlitz's Murrelet (Brachyramphus brevirostris) is a rare, non-colonial seabird often associated with tidewater glaciers and a recent candidate for listing under the Endangered Species Act. We estimated abundance of Kittlitz's Murrelets across space and time from at-sea surveys along the coast of Alaska (USA) and then used these data to develop spatial models to describe abundance patterns and identify environmental factors affecting abundance. Over a five-week period in the summer of 2005, we recorded 794 Kittlitz's Murrelets, 16 Marbled Murrelets (B. marmoratus), and 70 unidentified murrelets. The overall population estimate (N, mean +/- SE) during the peak period (3-9 July) was 1317 +/- 294 birds, decreasing to 68 +/- 37 by the last survey period (31 July-6 August). Density of Kittlitz's Murrelets was highest in pelagic waters of Taan Fjord (18.6 +/- 7.8 birds/km2, mean +/- SE) during 10-16 July. Spatial models identified consistent "hotspots" of Kittlitz's Murrelets, including several small areas where high densities of murrelets were found throughout the survey period. Of the explanatory variables that we evaluated, tidal current strength influenced murrelet abundance most consistently, with higher abundance associated with strong tidal currents. Simulations based on the empirically derived estimates of variation demonstrated that spatial variation strongly influenced power to detect trend, although power changed little across the threefold difference in the coefficient of variation on detection probability. We include recommendations for monitoring Kittlitz's Murrelets (or other marine species) when there is a high degree of uncertainty about factors affecting abundance, especially spatial variability.     

The Importance of Being Spatial (and Reserved): Assessing Northern Spotted Owl Habitat Relationships with Hierarchical Bayesian Models

Abstract:  Regional conservation planning increasingly draws on habitat suitability models to support decisions regarding land allocation and management. Nevertheless, statistical techniques commonly used for developing such models may give misleading results because they fail to account for 3 factors common in data sets of species distribution: spatial autocorrelation, the large number of sites where the species is absent (zero inflation), and uneven survey effort. We used spatial autoregressive models fit with Bayesian Markov Chain Monte Carlo techniques to assess the relationship between older coniferous forest and the abundance of Northern Spotted Owl nest and activity sites throughout the species' range. The spatial random-effect term incorporated in the autoregressive models successfully accounted for zero inflation and reduced the effect of survey bias on estimates of species–habitat associations. Our results support the hypothesis that the relationship between owl distribution and older forest varies with latitude. A quadratic relationship between owl abundance and older forest was evident in the southern portion of the range, and a pseudothreshold relationship was evident in the northern portion of the range. Our results suggest that proposed changes to the network of owl habitat reserves would reduce the proportion of the population protected by up to one-third, and that proposed guidelines for forest management within reserves underestimate the proportion of older forest associated with maximum owl abundance and inappropriately generalize threshold relationships among subregions. Bayesian spatial models can greatly enhance the utility of habitat analysis for conservation planning because they add the statistical flexibility necessary for analyzing regional survey data while retaining the interpretability of simpler models.     

Identifying and mapping local bycatch hotspots of loggerhead sea turtles using a GIS-based method: implications for conservation

Fisheries bycatch of marine megafauna is a worldwide conservation issue. In this study, we propose a method for generating reliable maps of sea turtle bycatch hotspots. Based on a well-defined area of fishing effort determined from the longline sets monitored in 2007 and 2010, we calculated the fishing area with the highest turtle bycatch density for both years. Our results show that it is important to consider all the components of fishing effort (area, number of hooks and soak time) in order to standardize the bycatch events, and thus the spatial density of the captures can be considered an index of abundance and aggregation of the species. Moreover, the high-resolution level of the analyses was useful for investigating the cumulative effect of the longline sets, which often overlap, and made it possible to correctly map the capture density and the intensity of the fishing effort at any given location.     

Defining ecologically relevant scales for spatial protection with long‐term data on an endangered seabird and local prey availability

Human activities are important drivers of marine ecosystem functioning. However, separating the synergistic effects of fishing and environmental variability on the prey base of nontarget predators is difficult, often because prey availability estimates on appropriate scales are lacking. Understanding how prey abundance at different spatial scales links to population change can help integrate the needs of nontarget predators into fisheries management by defining ecologically relevant areas for spatial protection. We investigated the local population response (number of breeders) of the Bank Cormorant (Phalacrocorax neglectus), a range‐restricted endangered seabird, to the availability of its prey, the heavily fished west coast rock lobster (Jasus lalandii). Using Bayesian state‐space modeled cormorant counts at 3 colonies, 22 years of fisheries‐independent data on local lobster abundance, and generalized additive modeling, we determined the spatial scale pertinent to these relationships in areas with different lobster availability. Cormorant numbers responded positively to lobster availability in the regions with intermediate and high abundance but not where regime shifts and fishing pressure had depleted lobster stocks. The relationships were strongest when lobsters 20–30 km offshore of the colony were considered, a distance greater than the Bank Cormorant's foraging range when breeding, and may have been influenced by prey availability for nonbreeding birds, prey switching, or prey ecology. Our results highlight the importance of considering the scale of ecological relationships in marine spatial planning and suggest that designing spatial protection around focal species can benefit marine predators across their full life cycle. We propose the precautionary implementation of small‐scale marine protected areas, followed by robust assessment and adaptive‐management, to confirm population‐level benefits for the cormorants, their prey, and the wider ecosystem, without negative impacts on local fisheries.     

Chilean Blue Whales as a Case Study to Illustrate Methods to Estimate Abundance and Evaluate Conservation Status of Rare Species

Abstract: Often abundance of rare species cannot be estimated with conventional design‐based methods, so we illustrate with a population of blue whales (Balaenoptera musculus) a spatial model‐based method to estimate abundance. We analyzed data from line‐transect surveys of blue whales off the coast of Chile, where the population was hunted to low levels. Field protocols allowed deviation from planned track lines to collect identification photographs and tissue samples for genetic analyses, which resulted in an ad hoc sampling design with increased effort in areas of higher densities. Thus, we used spatial modeling methods to estimate abundance. Spatial models are increasingly being used to analyze data from surveys of marine, aquatic, and terrestrial species, but estimation of uncertainty from such models is often problematic. We developed a new, broadly applicable variance estimator that showed there were likely 303 whales (95% CI 176–625) in the study area. The survey did not span the whales' entire range, so this is a minimum estimate. We estimated current minimum abundance relative to pre‐exploitation abundance (i.e., status) with a population dynamics model that incorporated our minimum abundance estimate, likely population growth rates from a meta‐analysis of rates of increase in large baleen whales, and two alternative assumptions about historic catches. From this model, we estimated that the population was at a minimum of 9.5% (95% CI 4.9–18.0%) of pre‐exploitation levels in 1998 under one catch assumption and 7.2% (CI 3.7–13.7%) of pre‐exploitation levels under the other. Thus, although Chilean blue whales are probably still at a small fraction of pre‐exploitation abundance, even these minimum abundance estimates demonstrate that their status is better than that of Antarctic blue whales, which are still <1% of pre‐exploitation population size. We anticipate our methods will be broadly applicable in aquatic and terrestrial surveys for rarely encountered species, especially when the surveys are intended to maximize encounter rates and estimate abundance.     

Currents connecting communities: nearshore community similarity and ocean circulation

Understanding the mechanisms that create spatial heterogeneity in species distributions is fundamental to ecology. For nearshore marine systems, most species have a pelagic larval stage where dispersal is strongly influenced by patterns of ocean circulation. Concomitantly, nearshore habitats and the local environment are also influenced by ocean circulation. Because of the shared dependence on the seascape, distinguishing the relative importance of the local environment from regional patterns of dispersal for community structure remains a challenge. Here, we quantify the "oceanographic distance" and "oceanographic asymmetry" between nearshore sites using ocean circulation modeling results. These novel metrics quantify spatial separation based on realistic patterns of ocean circulation, and we explore their explanatory power for intertidal and subtidal community similarity in the Southern California Bight. We find that these metrics show significant correspondence with patterns of community similarity and that their combined explanatory power exceeds that of the thermal structure of the domain. Our approach identifies the unique influence of ocean circulation on community structure and provides evidence for oceanographically mediated dispersal limitation in nearshore marine communities.     

Spatially Explicit Power Analyses for Occupancy‐Based Monitoring of Wolverine in the U.S. Rocky Mountains

Conservation scientists and resource managers often have to design monitoring programs for species that are rare or patchily distributed across large landscapes. Such programs are frequently expensive and seldom can be conducted by one entity. It is essential that a prospective power analysis be undertaken to ensure stated monitoring goals are feasible. We developed a spatially based simulation program that accounts for natural history, habitat use, and sampling scheme to investigate the power of monitoring protocols to detect trends in population abundance over time with occupancy‐based methods. We analyzed monitoring schemes with different sampling efforts for wolverine (Gulo gulo) populations in 2 areas of the U.S. Rocky Mountains. The relation between occupancy and abundance was nonlinear and depended on landscape, population size, and movement parameters. With current estimates for population size and detection probability in the northern U.S. Rockies, most sampling schemes were only able to detect large declines in abundance in the simulations (i.e., 50% decline over 10 years). For small populations reestablishing in the Southern Rockies, occupancy‐based methods had enough power to detect population trends only when populations were increasing dramatically (e.g., doubling or tripling in 10 years), regardless of sampling effort. In general, increasing the number of cells sampled or the per‐visit detection probability had a much greater effect on power than the number of visits conducted during a survey. Although our results are specific to wolverines, this approach could easily be adapted to other territorial species. Poder de Análisis Espacialmente Explícito para el Monitoreo Basado en Ocupación del Glotón (Gulo gulo) en las Montañas Rocallosas de Estados Unidos     

The effectiveness of marine reserve systems constructed using different surrogates of biodiversity

Biological sampling in marine systems is often limited, and the cost of acquiring new data is high. We sought to assess whether systematic reserves designed using abiotic domains adequately conserve a comprehensive range of species in a tropical marine inter‐reef system. We based our assessment on data from the Great Barrier Reef, Australia. We designed reserve systems aiming to conserve 30% of each species based on 4 abiotic surrogate types (abiotic domains; weighted abiotic domains; pre‐defined bioregions; and random selection of areas). We evaluated each surrogate in scenarios with and without cost (cost to fishery) and clumping (size of conservation area) constraints. To measure the efficacy of each reserve system for conservation purposes, we evaluated how well 842 species collected at 1155 sites across the Great Barrier Reef seabed were represented in each reserve system. When reserve design included both cost and clumping constraints, the mean proportion of species reaching the conservation target was 20–27% higher for reserve systems that were biologically informed than reserves designed using unweighted environmental data. All domains performed substantially better than random, except when there were no spatial or economic constraints placed on the system design. Under the scenario with no constraints, the mean proportion of species reaching the conservation target ranged from 98.5% to 99.99% across all surrogate domains, whereas the range was 90–96% across all domains when both cost and clumping were considered. This proportion did not change considerably between scenarios where one constraint was imposed and scenarios where both cost and clumping constraints were considered. We conclude that representative reserve systems can be designed using abiotic domains; however, there are substantial benefits if some biological information is incorporated.     

Predicting abundance of desert riparian birds: validation and calibration of the Effective Area Model.

Reliable prediction of the effects of landscape change on species abundance is critical to land managers who must make frequent, rapid decisions with long-term consequences. However, due to inherent temporal and spatial variability in ecological systems, previous attempts to predict species abundance in novel locations and/or time frames have been largely unsuccessful. The Effective Area Model (EAM) uses change in habitat composition and geometry coupled with response of animals to habitat edges to predict change in species abundance at a landscape scale. Our research goals were to validate EAM abundance predictions in new locations and to develop a calibration framework that enables absolute abundance predictions in novel regions or time frames. For model validation, we compared the EAM to a null model excluding edge effects in terms of accurate prediction of species abundance. The EAM outperformed the null model for 83.3% of species (N=12) for which it was possible to discern a difference when considering 50 validation sites. Likewise, the EAM outperformed the null model when considering subsets of validation sites categorized on the basis of four variables (isolation, presence of water, region, and focal habitat). Additionally, we explored a framework for producing calibrated models to decrease prediction error given inherent temporal and spatial variability in abundance. We calibrated the EAM to new locations using linear regression between observed and predicted abundance with and without additional habitat covariates. We found that model adjustments for unexplained variability in time and space, as well as variability that can be explained by incorporating additional covariates, improved EAM predictions. Calibrated EAM abundance estimates with additional site-level variables explained a significant amount of variability (P < 0.05) in observed abundance for 17 of 20 species, with R2 values >25% for 12 species, >48% for six species, and >60% for four species when considering all predictive models. The calibration framework described in this paper can be used to predict absolute abundance in sites different from those in which data were collected if the target population of sites to which one would like to statistically infer is sampled in a probabilistic way.     

Spatial anatomy of species survival: effects of predation and climate-driven environmental variability

The majority of survival analyses focus on temporal scales. Consequently, there is a limited understanding of how species survival varies over space and, ultimately, how spatial variability in the environment affects the temporal dynamics of species abundance. Using data from the Barents Sea, we study the spatiotemporal variability of the juvenile Atlantic cod (Gadus morhua) survival. We develop an index of spatial survival based on changes of juvenile cod distribution through their first winter of life (from age-0 to age-1) and study its variability in relation to biotic and abiotic factors. Over the 25 years analyzed (1980-2004), we found that, once the effect of passive drift due to dominant currents is accounted for, the area where age-0 cod survival was lowest coincided with the area of highest abundance of older cod. Within this critical region, the survival of age-0 cod was negatively affected by its own abundance, by that of older cod, and by bottom depth. Furthermore, during cold years, age-0 cod survival increased in the eastern and coldest portion of the examined area, which was typically avoided by older conspecifics. Based on these results we propose that within the examined area top-down mechanisms and predation-driven density dependence can strongly affect the spatial pattern of age-0 cod survival. Climate-related variables can also influence the spatial survival of age-0 cod by affecting their distribution and that of their predators. Results from these and similar studies, focusing on the spatial variability of survival rates, can be used to characterize species habitat quality of marine renewable resources.     

Shifting elasmobranch community assemblage at Cocos Island—an isolated marine protected area

Fishing pressure has increased the extinction risk of many elasmobranch (shark and ray) species. Although many countries have established no‐take marine reserves, a paucity of monitoring data means it is still unclear if reserves are effectively protecting these species. We examined data collected by a small group of divers over the past 21 years at one of the world's oldest marine protected areas (MPAs), Cocos Island National Park, Costa Rica. We used mixed effects models to determine trends in relative abundance, or probability of occurrence, of 12 monitored elasmobranch species while accounting for variation among observers and from abiotic factors. Eight of 12 species declined significantly over the past 2 decades. We documented decreases in relative abundance for 6 species, including the iconic scalloped hammerhead shark (Sphyrna lewini) (?45%), whitetip reef shark (Triaenodon obesus) (?77%), mobula ray (Mobula spp.) (?78%), and manta ray (Manta birostris) (?89%), and decreases in the probability of occurrence for 2 other species. Several of these species have small home ranges and should be better protected by an MPA, which underscores the notion that declines of marine megafauna will continue unabated in MPAs unless there is adequate enforcement effort to control fishing. In addition, probability of occurrence at Cocos Island of tiger (Galeocerdo cuvier), Galapagos (Carcharhinus galapagensis), blacktip (Carcharhinus limbatus), and whale (Rhincodon typus) sharks increased significantly. The effectiveness of MPAs cannot be evaluated by examining single species because population responses can vary depending on life history traits and vulnerability to fishing pressure.     

Bridging information domains to improve ecological understanding of biological invasions in a marine ecosystem

We devised a practical method for integrating information on 2 marine invasive species using 3 different approaches: standardized ecological monitoring, online-reporting databases, and surveys of anglers and crabbers. Focusing on 2 recently introduced species with different characteristics, the Asian shore crab (Hemigrapsus sanguineus) and Chinese mitten crab (Eriocheir sinensis), in the Hudson-Raritan watershed of New York and New Jersey, we used sensitivity analyses to explore the relative contribution of each information source to knowledge of species abundance and distribution. All 3 information sources contributed something unique to understanding abundance and distribution of the introduced crabs. Online and survey data on Asian shore crabs significantly affected predictions of abundance, whereas monitoring data did not. When survey data were omitted, abundance estimates were unchanged over time, but when they were included, the model predicted an increased abundance in 2012. All 3 data sets for the Asian shore crab significantly affected estimates of species coverage; surveys had the biggest influence, increasing range size by 4097.25 km². For the catadromous Chinese mitten crab, ecological monitoring data collected in freshwater shortly after the original sighting significantly shaped model estimates for abundance and documented the establishment phase of the mitten crab in an area outside the spatial scope of the surveyed resource users. However, the survey data significantly enlarged mitten crab range-size estimates by 6498.01 km². By demonstrating that data integration produced an image of the invasion process that would not have emerged had we used any 1 method individually, model results provide evidence for the advantages of an interdisciplinary approach.     

Global conservation status of sponges

Sponges are important for maintaining ecosystem function and integrity of marine and freshwater benthic communities worldwide. Despite this, there has been no assessment of their current global conservation status. We assessed their status, accounting for the distribution of research effort; patterns of temporal variation in sponge populations and assemblages; the number of sponges on threatened species lists; and the impact of environmental pressures. Sponge research effort has been variable; marine sponges in the northeastern Atlantic and Mediterranean and freshwater sponges in Europe and North America have received the most attention. Although sponge abundance has increased in some locations since 1990, these were typically on coral reefs, in response to declines in other benthic organisms, and restricted to a few species. Few data were available on temporal trends in freshwater sponge abundance. Despite over 8500 described sponge species, only 20 are on threatened species lists, and all are marine species from the northeastern Atlantic and Mediterranean. Of the 202 studies identified, the effects of temperature, suspended sediment, substratum loss, and microbial pathogens have been studied the most intensively for marine sponges, although responses appear to be variable. There were 20 studies examining environmental impacts on freshwater sponges, and most of these were on temperature and heavy metal contamination. We found that most sponges do not appear to be threatened globally. However, little information is available for most species and more data are needed on the impacts of anthropogenic‐related pressures. This is a critical information gap in understanding sponge conservation status. Estado Global de la Conservación de Esponjas     

Evaluating the coastal environment for marine birds

The marine environment is being increasingly exploited by fisheries and the oil and gas industry. Conservationists urgently need the ability to identify the processes that determine patterns of abundance of marine species. We describe a preliminary Geographic Information System (GIS) in which spatial data on environmental variables (seabird colony locations, sea depth and seabed sediments) are integrated with realistic energy constraints faced by marine birds during the breeding season. A simple foraging model predicts the spatial variation in the quality of given locations as potential feeding sites under different feeding conditions and stages of the breeding cycle. We show how the approach can be used to help managers identify key marine areas and assess the impacts of environmental change or damage.     

Use of Abundance of One Species as a Surrogate for Abundance of Others

Abstract: Indicator species concepts have a long history in conservation biology. Arguments in favor of these approaches generally stress expediency and assume efficacy. We tested the premise that the abundance patterns of one species can be used to infer those of other species. Our data consisted of 72,495 bird observations on 55 species across 1046 plots distributed across 30 sub basins. We analyzed abundance patterns at two spatial scales (plot and sub basin) and for empirical and a priori grouping. There were few significant indicator relationships at either scale or under either grouping rule, and those few we found did not explain a substantial portion of the abundance of other species. Coupled with the lack of proven efficacy for species surrogacy in the literature, our results indicate the utility of indicators and similar types of surrogate approaches must be demonstrated rather than assumed.     

Allee Effect and the Uncertainty of Population Recovery

Recovery of depleted populations is fundamentally important for conservation biology and sustainable resource harvesting. At low abundance, population growth rate, a primary determinant of population recovery, is generally assumed to be relatively fast because competition is low (i.e., negative density dependence). But population growth can be limited in small populations by an Allee effect. This is particularly relevant for collapsed populations or species that have not recovered despite large reductions in, or elimination of, threats. We investigated how an Allee effect can influence the dynamics of recovery. We used Atlantic cod (Gadus morhua) as the study organism and an empirically quantified Allee effect for the species to parameterize our simulations. We simulated recovery through an individual‐based mechanistic simulation model and then compared recovery among scenarios incorporating an Allee effect, negative density dependence, and an intermediate scenario. Although an Allee effect significantly slowed recovery, such that population increase could be negligible even after 100 years or more, it also made the time required for biomass rebuilding much less predictable. Our finding that an Allee effect greatly increased the uncertainty in recovery time frames provides an empirically based explanation for why the removal of threat does not always result in the recovery of depleted populations or species. El Efecto Allee y la Incertidumbre de la Recuperación de Poblaciones