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Quantifying source and sink habitats and pathways in spatially structured populations: A generalized modelling approach

The ability to classify habitats and movement pathways as sources or sinks is an important part of the decision making process for the conservation of spatially structured populations. Diverse approaches have been used to quantify the importance of habitats and pathways in a spatial network, however these approaches have been limited by a lack of general applicability across life histories and movement strategies. In this paper we develop a generalized per-capita contribution metric, the C-metric, for quantifying habitat and pathway quality. This metric is novel in that it can be applied broadly to both metapopulations and migratory species. It allows for any number of age and sex classes, unlimited number of seasons or time intervals within the annual cycle, and for density-dependent parameters. We demonstrate the flexibility of the metric with four case studies: a hypothetical metapopulation, elk of the Greater Yellowstone Ecosystem, northern pintail ducks in North America, and the eastern population of the monarch butterfly. General computer code to calculate the per-capita contribution metric is provided. We demonstrate that the C-metric is useful for identifying source and sink habitats in a network and suggest that the C-metric could be supplemented by some measure of network structure for a more robust description of habitat or pathway importance.

Ecological Modelling

Population age and initial density in a patchy environment affect the occurrence of abrupt transitions in a birth-and-death model of Taylor's law

Taylor's power law describes an empirical relationship between the mean and variance of population densities in field data, in which the variance varies as a power, b , of the mean. Most studies report values of b varying between 1 and 2. However, Cohen (2014a) showed recently that smooth changes in environmental conditions in a model can lead to an abrupt, infinite change in b . To understand what factors can influence the occurrence of an abrupt change in b , we used both mathematical analysis and Monte Carlo samples from a model in which populations of the same species settled on patches, and each population followed independently a stochastic linear birth-and-death process. We investigated how the power relationship responds to a smooth change of population growth rate, under different sampling strategies, initial population density, and population age. We showed analytically that, if the initial populations differ only in density, and samples are taken from all patches after the same time period following a major invasion event, Taylor's law holds with exponent b =1, regardless of the population growth rate. If samples are taken at different times from patches that have the same initial population densities, we calculate an abrupt shift of b , as predicted by Cohen (2014a). The loss of linearity between log variance and log mean is a leading indicator of the abrupt shift. If both initial population densities and population ages vary among patches, estimates of b lie between 1 and 2, as in most empirical studies. But the value of b declines to ~1 as the system approaches a critical point. Our results can inform empirical studies that might be designed to demonstrate an abrupt shift in Taylor's law.

Ecological Modelling

Effects of future urban and biofuel crop expansions on the riverine export of phosphorus to the Laurentian Great Lakes

Increased phosphorus (P) loadings threaten the health of the world’s largest freshwater resource, the Laurentian Great Lakes (GL). To understand the linkages between land use and P delivery, we coupled two spatially explicit models, the landscape-scale SPARROW P fate and transport watershed model and the Land Transformation Model (LTM) land use change model, to predict future P export from nonpoint and point sources caused by changes in land use. According to LTM predictions over the period 2010–2040, the GL region of the U.S. may experience a doubling of urbanized areas and agricultural areas may increase by 10%, due to biofuel feedstock cultivation. These land use changes are predicted to increase P loadings from the U.S. side of the GL basin by 3.5–9.5%, depending on the Lake watershed and development scenario. The exception is Lake Ontario, where loading is predicted to decrease by 1.8% for one scenario, due to population losses in the drainage area. Overall, urban expansion is estimated to increase P loadings by 3.4%. Agricultural expansion associated with predicted biofuel feedstock cultivation is predicted to increase P loadings by an additional 2.4%. Watersheds that export P most efficiently and thus are the most vulnerable to increases in P sources tend to be found along southern Lake Ontario, southeastern Lake Erie, western Lake Michigan, and southwestern Lake Superior where watershed areas are concentrated along the coastline with shorter flow paths. In contrast, watersheds with high soil permeabilities, fractions of land underlain by tile drains, and long distances to the GL are less vulnerable.

Laurentian Great Lakes

Understanding predator-prey-competitor dynamics between Lower Missouri River Macrhybopsis and Scaphirhynchus using a population—bioenergetics model ensemble

The pallid sturgeon Scaphirhynchus albus is a long-lived, endangered fish in the Missouri River. Individuals become piscivorous as adults, so recruitment from stocking or reproduction could reduce populations of prey, including Macrhybopsis chubs. We constructed an individual- and age-based, multi-species, predator-prey-competitor model (IAMP) to represent the benthic community (sturgeons, chubs, and chironomids) of the Lower Missouri River (LMR) to explore scenarios of potential predator-prey-competitor dynamics. Our simulations suggest that chubs alone are unlikely able to support a level of LMR pallid sturgeon similar to historical or current populations. These simulations also suggest that adult pallid sturgeon may need to shift to non-chub prey fish to achieve the greater sizes observed in the Upper Missouri River. When annual hydrologic regimes were included, we found a negative relationship between chub relative abundance and previous year 30-day minimum flows. Inclusion of temporal environmental variability made it clear that large chub populations may be necessary to support LMR pallid sturgeon. When full stochasticity was included in the IAMP, chub population sizes needed to increase further to ensure continued reproduction and recruitment of both chubs and pallid sturgeon. These results support the hypothesis that the pallid sturgeon population in the Lower Missouri River may be food-limited. However, the full extent of this limitation and the management changes needed to address this will require more research on the biology and population dynamics of this fish community, on pallid sturgeon interactions with prey species, and on how sympatric species may be affected during the pallid sturgeon recovery process.

Colorado, Kansas, Minnesota, Missouri, Montana, Ne

A food web modeling analysis of a Midwestern, USA eutrophic lake dominated by non-native Common Carp and Zebra Mussels

Food web modeling is recognized as fundamental to understanding the complexities of aquatic systems. Ecopath is the most common mass-balance model used to represent food webs and quantify trophic interactions among groups. We constructed annual Ecopath models for four consecutive years during the first half-decade of a zebra mussel invasion in shallow, eutrophic Clear Lake, Iowa, USA, to evaluate changes in relative biomass and total system consumption among food web groups, evaluate food web impacts of non-native common carp and zebra mussels on food web groups, and to interpret food web impacts in light of on-going lake restoration. Total living biomass increased each year of the study; the majority of the increase due to a doubling in planktonic blue green algae, but several other taxa also increased including a more than two-order of magnitude increase in zebra mussels. Common carp accounted for the largest percentage of total fish biomass throughout the study even with on-going harvest. Chironomids, common carp, and zebra mussels were the top-three ranking consumer groups. Non-native common carp and zebra mussels accounted for an average of 42% of the total system consumption. Despite the relatively high biomass densities of common carp and zebra mussel, food web impacts was minimal due to excessive benthic and primary production in this eutrophic system. Consumption occurring via benthic pathways dominated system consumption in Clear Lake throughout our study, supporting the argument that benthic food webs are significant in shallow, eutrophic lake ecosystems and must be considered if ecosystem-level understanding is to be obtained.

Iowa

Simulating the effects of management practices on cropland soil organic carbon changes in the Temperate Prairies Ecoregion of the United States from 1980 to 2012

Understanding the effects of management practices on soil organic carbon (SOC) is important for designing effective policies to mitigate greenhouse gas emissions in agriculture. In the Midwest United States, management practices in the croplands have been improved to increase crop production and reduce SOC loss since the 1980s. Many studies of SOC dynamics in croplands have been performed to understand the effects of management, but the results are still not conclusive. This study quantified SOC dynamics in the Midwest croplands from 1980 to 2012 with the General Ensemble Biogeochemical Modelling System (GEMS) and available management data. Our results showed that the total SOC in the croplands decreased from 1190 Tg C in 1980 to 1107 TgC in 1995, and then increased to 1176 TgC in 2012. Continuous cropping and intensive tillage may have driven SOC loss in the early period. The increase of crop production and adoption of conservation tillage increased the total SOC so that the decrease in the total SOC stock after 32 years was only 1%. The small change in average SOC did not reflect the large spatial variations of SOC change in the region. Major SOC losses occurred in the north and south of the region, where SOC baseline values were high and cropland production was low. The SOC gains took place in the central part of the region where SOC baseline values were moderate and cropland production was higher than the other areas. We simulated multiple land-use land-cover (LULC) change scenarios and analyzed the results. The analysis showed that among all the LULC changes, agricultural technology that increased cropland production had the greatest impact on SOC changes, followed by the tillage practices, changes in crop species, and the conversions of cropland to other land use. Information on management practice induced spatial variation in SOC can be useful for policy makers and farm managers to develop long-term management strategies for increasing SOC sequestration in different areas.

Ecological Modelling

Modeling how to achieve localized areas of reduced white-tailed deer density

Localized management of white-tailed deer ( Odocoileus virginianus ) involves the removal of matriarchal family units with the intent to create areas of reduced deer density. However, application of this approach has not always been successful, possibly because of female dispersal and high deer densities. We developed a spatially explicit, agent-based model to investigate the intensity of deer removal required to locally reduce deer density depending on the surrounding deer density, dispersal behavior, and size and shape of the area of localized reduction. Application of this model is illustrated using the example of abundant deer populations in Pennsylvania, USA. Most scenarios required at least 5 years before substantial deer density reductions occurred. Our model indicated that a localized reduction was successful for scenarios in which the surrounding deer density was lowest (30 deer/mi²), localized antlerless harvest rates were 30%, and the removal area was 5 mi² or larger. When the size of the removal area was < 5 mi2, end population density was highly variable and, in some scenarios, exceeded the initial density. The shape of the area of localized reduction had less influence on the ability to reduce deer density than the size. There were no differences in mean deer density in the same size circle or square removal areas. Similarly, increasing the ratio of sides (length : width) in rectangular removal areas had little influence on the ability to locally reduce deer densities. Situations in which deer density was higher (40 or 50 deer/mi2) required antlerless removal rates to exceed 30% and took more than 5 years to considerably reduce density in the localized area regardless of its size. These results indicate that the size of the area of reduction, surrounding deer density, and antlerless harvest rate are the most influential factors in locally reducing deer density. Therefore, localized management likely can be an effective strategy for lower density herds, especially in larger removal areas. For high density herds, the success of this strategy would depend most on the ability of resource managers to achieve consistently high antlerless harvest rates.

Ecological Modelling

Using integrated population models for insights into monitoring programs: An application using pink-footed geese

Development of integrated population models (IPMs) assume the absence of systematic bias in monitoring programs, yet many potential sources of systematic bias in monitoring data exist (e.g., under-counts of abundance). By integrating multiple sources of data, we can assess whether various sources of monitoring data provide consistent inferences about changes in population size and, thus, whether monitoring programs appear unbiased. For the purposes of understanding how IPMs could provide insights for monitoring programs, we used the Svalbard breeding population of pink-footed goose ( Anser brachyrhynchus ) as a case study. The Svalbard pink-footed goose is a well-studied species, the focus of the first adaptive-harvest-management program in Europe, and the subject of a variety of long-term monitoring programs. We examined two formulations of an IPM, but ultimately relied on the one that provided a satisfactory fit to all the available data as based on Chi-squared goodness of fit tests. Our analyses suggest a negative bias in November counts (-20 %), a negative bias in capture-mark-recapture estimates of survival (-3 %), and a negative bias in indices of productivity (-23 %). We offer possible explanations for these biases, whether the degree of bias seems reasonable considering those explanations, and how bias might be investigated directly and ultimately avoided or corrected. Finally, we discuss implications of our work for developing IPMs and associated monitoring programs for managing pink-footed geese and other waterbird species.

Ecological Modelling

Modeling the impacts of hunting on the population dynamics of red howler monkeys ( Alouatta seniculus )

Overexploitation of wildlife populations occurs across the humid tropics and is a significant threat to the long-term survival of large-bodied primates. To investigate the impacts of hunting on primates and ways to mitigate them, we developed a spatially explicit, individual-based model for a landscape that included hunted and un-hunted areas. We used the large-bodied neotropical red howler monkey ( Alouatta seniculus ) as our case study species because its life history characteristics make it vulnerable to hunting. We modeled the influence of different rates of harvest and proportions of landscape dedicated to un-hunted reserves on population persistence, population size, social dynamics, and hunting yields of red howler monkeys. In most scenarios, the un-hunted populations maintained a constant density regardless of hunting pressure elsewhere, and allowed the overall population to persist. Therefore, the overall population was quite resilient to extinction; only in scenarios without any un-hunted areas did the population go extinct. However, the total and hunted populations did experience large declines over 100 years under moderate and high hunting pressure. In addition, when reserve area decreased, population losses and losses per unit area increased disproportionately. Furthermore, hunting disrupted the social structure of troops. The number of male turnovers and infanticides increased in hunted populations, while birth rates decreased and exacerbated population losses due to hunting. Finally, our results indicated that when more than 55% of the landscape was harvested at high (30%) rates, hunting yields, as measured by kilograms of biomass, were less than those obtained from moderate harvest rates. Additionally, hunting yields, expressed as the number of individuals hunted/year/km 2 , increased in proximity to un-hunted areas, and suggested that dispersal from un-hunted areas may have contributed to hunting sustainability. These results indicate that un-hunted areas serve to enhance hunting yields, population size, and population persistence in hunted landscapes. Therefore, spatial regulation of hunting via a reserve system may be an effective management strategy for sustainable hunting, and we recommend it because it may also be more feasible to implement than harvest quotas or restrictions on season length.

Guárico

Prey patchiness and larval fish growth and survival: inferences from an individual-based model

We used an individual-based simulation model to evaluate how prey patchiness and fish swimming behavior affect larval fish survival and mortality source (predation or starvation). Simulations revealed that cohort survival increased linearly with greater average patch residence times and that patch residence times for individual fish with different fates (survived, starved, eaten) diverged substantially during the first few days of feeding. Further, by examining the interaction of patch spatial distribution (uniform random versus clumped) with three possible swimming behaviors, we found that swimming behavior, via its effect on prey encounter and feeding rates, affected both cohort survival rates and whether fish died from predation or starvation, but that the spatial distribution of patches (fine-scale (100s m)) only influenced whether fish died from predation or starvation. Within a particular patch spatial arrangement, however, patch intensity (division of food between patches and non-patches) had a major effect on survival. Except at high food levels, fish did not survive when there were no patches, suggesting that average, well-mixed prey densities will not support sufficiently rapid growth for survival. As patch intensity increased, survival increased to a maximum and then declined, with peak survival occurring at higher patch intensity as average food concentration declined. Finally, the degree of patchiness also determined the intensity of selection on growth rates. In patchier environments, there was stronger selection for fast growth rates leading to eight-fold differences in average cohort growth rates after only three days of growth. In general, survival was not directly related to fast cohort growth the best survival occurred with high average prey densities and weak patchiness. Prey patchiness, by influencing the average as well as the variance in individual growth rates, can have a substantial impact on survival rates of larval fish cohorts.

Ecological Modelling

Estimating the drivers of species distributions with opportunistic data using mediation analysis

Ecological occupancy modeling has historically relied on high-quality, low-quantity designed-survey data for estimation and prediction. In recent years, there has been a large increase in the amount of high-quantity, unknown-quality opportunistic data. This has motivated research on how best to combine these two data sources in order to optimize inference. Existing methods can be infeasible for large datasets or require opportunistic data to be located where designed-survey data exist. These methods map species occupancies, motivating a need to properly evaluate covariate effects (e.g., land cover proportion) on their distributions. We describe a spatial estimation method for supplementarily including additional opportunistic data using mediation analysis concepts. The opportunistic data mediate the effect of the covariate on the designed-survey data response, decomposing it into a direct and indirect effect. A component of the indirect effect can then be quickly estimated via regressing the mediator on the covariate, while the other components are estimated through a spatial occupancy model. The regression step allows for use of large quantities of opportunistic data that can be collected in locations with no designed-survey data available. Simulation results suggest that the mediated method produces an improvement in relative MSE when the data are of reasonable quality. However, when the simulated opportunistic data are poorly correlated with the true spatial process, the standard, unmediated method is still preferable. A spatiotemporal extension of the method is also developed for analyzing the effect of deciduous forest land cover on red-eyed vireo distribution in the southeastern United States and find that including the opportunistic data do not lead to a substantial improvement. Opportunistic data quality remains an important consideration when employing this method, as with other data integration methods.

eastern United States

Predicting thermally stressful events in rivers with a strategy to evaluate management alternatives

Water temperature is an important factor in river ecology. Numerous models have been developed to predict river temperature. However, many were not designed to predict thermally stressful periods. Because such events are rare, traditionally applied analyses are inappropriate. Here, we developed two logistic regression models to predict thermally stressful events in the Delaware River at the US Geological Survey gage near Lordville, New York. One model predicted the probability of an event >20.0 °C, and a second predicted an event >22.2 °C. Both models were strong (independent test data sensitivity 0.94 and 1.00, specificity 0.96 and 0.96) predicting 63 of 67 events in the >20.0 °C model and all 15 events in the >22.2 °C model. Both showed negative relationships with released volume from the upstream Cannonsville Reservoir and positive relationships with difference between air temperature and previous day's water temperature at Lordville. We further predicted how increasing release volumes from Cannonsville Reservoir affected the probabilities of correctly predicted events. For the >20.0 °C model, an increase of 0.5 to a proportionally adjusted release (that accounts for other sources) resulted in 35.9% of events in the training data falling below cutoffs; increasing this adjustment by 1.0 resulted in 81.7% falling below cutoffs. For the >22.2 °C these adjustments resulted in 71.1% and 100.0% of events falling below cutoffs. Results from these analyses can help managers make informed decisions on alternative release scenarios.

New York

The hydroclimate niche: A tool for predicting and managing riparian plant community responses to streamflow seasonality

Habitat suitability is a consequence of interacting environmental factors. In riparian ecosystems, suitable plant habitat is influenced by interactions between stream hydrology and climate, hereafter referred to as “hydroclimate”. We tested the hypothesis that hydroclimate variables would improve the fit of ecological niche models for a suite of riparian species using occurrence data from the western United States. We focus on the climate conditions (temperature, precipitation and vapor pressure deficit) during the months of lowest and highest streamflow as integrative hydroclimate metrics of resource and stress levels. We found that the inclusion of hydroclimate variables improved model fit for all species in the western USA dataset. We then tested the utility of the improved habitat suitability models by projecting them onto a regulated segment of the Colorado River to assess potential impacts of streamflow seasonality on vegetation metrics of management concern. Species frequency derived from independent survey data in the Colorado River segment was significantly higher for species with predicted suitable habitat than for species without predicted suitable habitat. Under different simulated hydrographs for the Colorado River, overall species richness was predicted to be greatest with peak streamflows during summer, and native-to-non-native species ratios were predicted to be greatest with lowest streamflows in winter. Summer high flows were particularly associated with higher predicted habitat suitability for species that have increased in cover over recent decades (e.g., Pluchea sericea, Baccharis species). We conclude that hydroclimate covariates can be useful tools for predicting how riparian vegetation communities respond to changes in the seasonal timing of low and high streamflows.

River Research and Applications

Abdominally implanted satellite transmitters affect reproduction and survival rather than migration of large shorebirds

Satellite telemetry has become a common technique to investigate avian life-histories, but whether such tagging will affect fitness is a critical unknown. In this study, we evaluate multi-year effects of implanted transmitters on migratory timing and reproductive performance in shorebirds. Shorebirds increasingly are recognized as good models in ecology and evolution. That many of them are of conservation concern adds to the research responsibilities. In May 2009, we captured 56 female Black-tailed Godwits Limosa limosa limosa during late incubation in The Netherlands. Of these, 15 birds were equipped with 26-g satellite transmitters with a percutaneous antenna (7.8 % ± 0.2 SD of body mass), surgically implanted in the coelom. We compared immediate nest survival, timing of migration, subsequent nest site fidelity and reproductive behaviour including egg laying with those of the remaining birds, a comparison group of 41 females. We found no effects on immediate nest survival. Fledging success and subsequent southward and northward migration patterns of the implanted birds conformed to the expectations, and arrival time on the breeding grounds in 2010–2012 did not differ from the comparison group. Compared with the comparison group, in the year after implantation, implanted birds were equally faithful to the nest site and showed equal territorial behaviour, but a paucity of behaviours indicating nests or clutches. In the 3 years after implantation, the yearly apparent survival of implanted birds was 16 % points lower. Despite intense searching, we found only three eggs of two implanted birds; all were deformed. A similarly deformed egg was reported in a similarly implanted Whimbrel Numenius phaeopus returning to breed in central Alaska. The presence in the body cavity of an object slightly smaller than a normal egg may thus lead to egg malformation and, likely, reduced egg viability. That the use of implanted satellite transmitters in these large shorebirds reduced nesting propensity and might also lead to fertility losses argues against the use of implanted transmitters for studies on breeding biology, and for a careful evaluation of the methodology in studies of migration.

Friesl

Top 10 principles for designing healthy coastal ecosystems like the Salish Sea

Like other coastal zones around the world, the inland sea ecosystem of Washington (USA) and British Columbia (Canada), an area known as the Salish Sea, is changing under pressure from a growing human population, conversion of native forest and shoreline habitat to urban development, toxic contamination of sediments and species, and overharvest of resources. While billions of dollars have been spent trying to restore other coastal ecosystems around the world, there still is no successful model for restoring estuarine or marine ecosystems like the Salish Sea. Despite the lack of a guiding model, major ecological principles do exist that should be applied as people work to design the Salish Sea and other large marine ecosystems for the future. We suggest that the following 10 ecological principles serve as a foundation for educating the public and for designing a healthy Salish Sea and other coastal ecosystems for future generations: (1) Think ecosystem: political boundaries are arbitrary; (2) Account for ecosystem connectivity; (3) Understand the food web; (4) Avoid fragmentation; (5) Respect ecosystem integrity; (6) Support nature's resilience; (7) Value nature: it's money in your pocket; (8) Watch wildlife health; (9) Plan for extremes; and (10) Share the knowledge.

British Columbia, Washington

Demography of the Everglade kite: Implications for population management

Simple deterministic and stochastic population modelsi are used to examine the demographic patterns of the Everglade Kite population. These efforts are directed at making inferences about the evolution of the kite life-history pattern, and at providing guidelines for the management of the kite population. The Everglade Kite has apparently evolved high adult survival rates, in partial response to a variable reproductive output. Proper management of this population should include the protection of adults from catastrophic mortality sources, and the provision of adequate water-levels to ensure reproductive success.

Florida

Optimal stochastic control in natural resource management: Framework and examples

A framework is presented for the application of optimal control methods to natural resource problems. An expression of the optimal control problem appropriate for renewable natural resources is given and its application to Markovian systems is presented in some detail. Three general approaches are outlined for determining optimal control of infinite time horizon systems and three examples from the natural resource literature are used for illustration.

Ecological Modelling

Cold-seep habitat mapping: high-resolution spatial characterization of the Blake Ridge Diapir seep field

Relationships among seep community biomass, diversity, and physiographic controls such as underlying geology are not well understood. Previous efforts to constrain these relationships at the Blake Ridge Diapir were limited to observations from piloted deep-submergence vehicles. In August 2012, the autonomous underwater vehicle (AUV) Sentry collected geophysical and photographic data over a 0.131 km2 area at the Blake Ridge Diapir seeps. A nested survey approach was used that began with a regional or reconnaissance-style survey using sub-bottom mapping systems to locate and identify seeps and underlying conduits. This survey was followed by AUV-mounted sidescan sonar and multibeam echosounder systems mapping on a mesoscale to characterize the seabed physiography. At the most detailed survey level, digital photographic imaging was used to resolve sub-meter characteristics of the biology. Four pockmarks (25–70 m diameter) were documented, each supporting chemosynthetic communities. Concentric zonation of mussels and clams suggests the influence of chemical gradients on megafaunal distribution. Data collection and analytical techniques used here yield high-resolution habitat maps that can serve as baselines to constrain temporal evolution of seafloor seeps, and to inform ecological niche modeling and resource management.

Deep-Sea Research Part II: Topical Studies in Ocea