Density-dependent effects of ants on selection for bumble bee pollination in Polemonium viscosum

Mutualisms are commonly exploited by cheater species that usurp rewards without providing reciprocal benefits. Yet most studies of selection between mutualist partners ignore interactions with third species and consequently overlook the impact of cheaters on evolution in the mutualism. Here, we explicitly investigate how the abundance of nectar-thieving ants (cheaters) influences selection in a pollination mutualism between bumble bees and the alpine skypilot, Polemonium viscosum. As suggested in past work with this species, bumble bees accounted for most of the seed production (78% +/- 6% [mean +/- SE]) in our high tundra study population and, in the absence of ants, exerted strong selection for large flowers. We tested for indirect effects of ant abundance on seed set through bumble bee pollination services (pollen delivery and pollen export) and a direct effect through flower damage. Ants reduced seed set per flower by 20% via flower damage. As ant density increased within experimental patches, the rate of flower damage rose, but pollen delivery and export did not vary significantly, showing that indirect effects of increased cheater abundance on pollinator service are negligible in this system. To address how ants affect selection for plant participation in the pollination mutualism we tested the impact of ant abundance on selection for bumble bee-mediated pollination. Results show that the impact of ants on fitness (seed set) accruing under bumble bee pollination is density dependent in P. viscosum. Selection for bumble bee pollination declined with increasing ant abundance in experimental patches, as predicted if cheaters constrain fitness returns of mutualist partner services. We also examined how ant abundance influences selection on flower size, a key component of plant investment in bumble bee pollination. We predicted that direct effects of ants would constrain bumble bee selection for large flowers. However, selection on flower size was significantly positive over a wide range of ant abundance (20-80% of plants visited by ants daily). Although high cheater abundance reduces the fitness returns of bumble bee pollination, it does not completely eliminate selection for bumble bee attraction in P. viscosum.     

Preventing the Spread of Invasive Species: Economic Benefits of Intervention Guided by Ecological Predictions

Abstract:  Preventing the invasion of freshwater aquatic species is the surest way to reduce their impacts, but it is also often expensive. Hence, the most efficient prevention programs will rely on accurate predictions of sites most at risk of becoming invaded and concentrate resources at those sites. Using data from Vilas County, Wisconsin (U.S.A.), collected in the 1970s, we constructed a predictive occurrence model for rusty crayfish ( Orconectes rusticus ) and applied it to an independent data set of 48 Vilas County lakes to predict which of these were most likely to become invaded between 1975 and 2005. We nested this invasion model within an economic framework to determine whether targeted management, derived from our quantitative predictions of likely invasion sites, would increase the economic value of lakes in the independent data set. Although the optimum expenditure on lake protection was high, protecting lakes at this level would have produced net economic benefits of at least $6 million over the last 30 years. We did not attempt to determine the value of nonmarket benefits of protection; thus, our results are likely to underestimate the total benefits from preventing invasions. Our results demonstrate that although few data are available early in an invasion, these data may be sufficient to support targeted, effective, and economically rational management. In addition, our results show that ecological predictions are becoming sufficiently accurate that their application in management can produce net economic benefits.     

Are soil mite assemblages structured by the identity of native and invasive alien grasses?

Associations between plants and animals in aboveground communities are often predictable and specific. This has been exploited for the purposes of estimating the diversity of animal species based on the diversity of plant species. The introduction of invasive alien plants into an ecosystem can result in dramatic changes in both the native plant and animal assemblages. Few data exist at the species level to determine whether belowground animal assemblages share the same degree of association to plants. The hypotheses that soil mites (Acari) form assemblages specifically associated with different native grass species in an unmanipulated natural ecosystem and that invasive alien grasses will impact soil mite assemblage composition in this setting were tested. Soil mites sampled beneath five native and two invasive alien species of grasses at the Konza Prairie Biological Station, Kansas, USA, were similarly abundant, species rich, diverse, and taxonomically distinct. No mite species had affinities for a specific grass species. There was no evidence from analysis of similarity, canonical correspondence analysis, or a nonparametric assemblage analysis that the assemblage composition of soil mites was specific to grass species. Results suggest that soil mite assemblages were more related to characteristics of the plant assemblage as a whole or prevailing soil conditions. The most recent invasive alien grass did not support a successionally younger mite fauna, based on the ratio of mesostigmatid to oribatid mites, and neither of the two invasive grasses influenced mite assemblage structure, possibly because they had not yet substantially altered the soil environment. Our results suggest that extrapolations of soil mite diversity based on assumptions of plant specificity would be invalid.     

Minimizing opportunity costs to aquatic connectivity restoration while controlling an invasive species

Controlling invasive species is critical for conservation but can have unintended consequences for native species and divert resources away from other efforts. This dilemma occurs on a grand scale in the North American Great Lakes, where dams and culverts block tributary access to habitat of desirable fish species and are a lynchpin of long‐standing efforts to limit ecological damage inflicted by the invasive, parasitic sea lamprey (Petromyzon marinus). Habitat restoration and sea‐lamprey control create conflicting goals for managing aging infrastructure. We used optimization to minimize opportunity costs of habitat gains for 37 desirable migratory fishes that arose from restricting sea lamprey access (0–25% increase) when selecting barriers for removal under a limited budget (US$1–105 million). Imposing limits on sea lamprey habitat reduced gains in tributary access for desirable species by 15–50% relative to an unconstrained scenario. Additional investment to offset the effect of limiting sea‐lamprey access resulted in high opportunity costs for 30 of 37 species (e.g., an additional US$20–80 million for lake sturgeon [Acipenser fulvescens]) and often required ≥5% increase in sea‐lamprey access to identify barrier‐removal solutions adhering to the budget and limiting access. Narrowly distributed species exhibited the highest opportunity costs but benefited more at less cost when small increases in sea‐lamprey access were allowed. Our results illustrate the value of optimization in limiting opportunity costs when balancing invasion control against restoration benefits for diverse desirable species. Such trade‐off analyses are essential to the restoration of connectivity within fragmented rivers without unleashing invaders.     

Detecting Insect Pollinator Declines on Regional and Global Scales

Recently there has been considerable concern about declines in bee communities in agricultural and natural habitats. The value of pollination to agriculture, provided primarily by bees, is >$200 billion/year worldwide, and in natural ecosystems it is thought to be even greater. However, no monitoring program exists to accurately detect declines in abundance of insect pollinators; thus, it is difficult to quantify the status of bee communities or estimate the extent of declines. We used data from 11 multiyear studies of bee communities to devise a program to monitor pollinators at regional, national, or international scales. In these studies, 7 different methods for sampling bees were used and bees were sampled on 3 different continents. We estimated that a monitoring program with 200–250 sampling locations each sampled twice over 5 years would provide sufficient power to detect small (2–5%) annual declines in the number of species and in total abundance and would cost U.S.$2,000,000. To detect declines as small as 1% annually over the same period would require >300 sampling locations. Given the role of pollinators in food security and ecosystem function, we recommend establishment of integrated regional and international monitoring programs to detect changes in pollinator communities. Detección de Declinaciones de Insectos Polinizadores a Escalas Regional y Global     

Costs of expanding the network of protected areas as a response to climate change in the Cape Floristic Region

The expansion of protected areas is a critical component of strategies to promote the continued existence of biodiversity (i.e., life at all levels of biological organization) as climate changes, but scientific, social, and economic uncertainties associated with climate change are some of the major obstacles preventing such expansion. New models of climate change and species distribution and new methods of conservation planning now make it possible to explore the uncertainties associated with climate changes and species responses. Yet few reliable estimates of the costs of expanding protected areas and methods for determining these costs exist, largely because of the many (and uncertain) determinants of these costs. We developed a cost-accounting model to estimate the range in costs of various options for expanding protected areas and to explore the variables that drive these costs. Model development was informed by an existing plan to expand protected areas in the Cape Floristic Region of South Africa to address species conservation under a scenario of climate change. The 50-year present value of total costs varied from US$260 million ($1077/ha) for an off-reserve option that involves agreements with landowners and no compensation of forgone production and associated revenue to $1020 million ($4228/ha) for an on-reserve option that involves land acquisition and protection. The costs of acquiring land or compensating landowners for forgone production and development opportunities were the major drivers of the total costs across all options because most of the area identified in the protected-area expansion plan consisted of urban and high-quality agricultural lands. Total costs were also affected by changes in protected area extent and discount rate. Model-generated outputs such as these may be useful for informing implementation strategies and the allocation of future efforts in monitoring, data collection, and model development.     

Environmental pollution and biodiversity: Light pollution and sea turtles in the Caribbean

We examine the impact of pollution on biodiversity by studying the effect of coastal light pollution on the sea turtle population in the Caribbean. To this end we assemble a panel data set of sea turtle nesting activity and satellite-derived measures of nighttime light. Controlling for the surveyor effort, the local economic infrastructure, and spatial spillovers, we find that nighttime light significantly reduces the number of sea turtle nests. According to data on replacement costs for sea turtles raised in captivity, our result suggests that the increase in lighting over the last two decades has resulted in the loss of close to 1800 sea turtles in the Caribbean, worth up to $288 million. Incorporating our empirical estimates into a stage-structured population model, we discover that the dynamic effect of nighttime light on future generations of sea turtles is likely to be much larger, with a cost of approximately $2.8 billion for Guadeloupe alone. More generally, our study provides a new approach to valuing the cost of environmental pollution associated with species extinction.     

Benefits of integrating complementarity into priority threat management

Conservation decision tools based on cost‐effectiveness analysis are used to assess threat management strategies for improving species persistence. These approaches rank alternative strategies by their benefit to cost ratio but may fail to identify the optimal sets of strategies to implement under limited budgets because they do not account for redundancies. We devised a multiobjective optimization approach in which the complementarity principle is applied to identify the sets of threat management strategies that protect the most species for any budget. We used our approach to prioritize threat management strategies for 53 species of conservation concern in the Pilbara, Australia. We followed a structured elicitation approach to collect information on the benefits and costs of implementing 17 different conservation strategies during a 3‐day workshop with 49 stakeholders and experts in the biodiversity, conservation, and management of the Pilbara. We compared the performance of our complementarity priority threat management approach with a current cost‐effectiveness ranking approach. A complementary set of 3 strategies: domestic herbivore management, fire management and research, and sanctuaries provided all species with >50% chance of persistence for $4.7 million/year over 20 years. Achieving the same result cost almost twice as much ($9.71 million/year) when strategies were selected by their cost‐effectiveness ranks alone. Our results show that complementarity of management benefits has the potential to double the impact of priority threat management approaches.     

Rapid assessment of insect pollination services to inform decision-making

Pollinator declines have prompted efforts to assess how land-use change affects insect pollinators and pollination services in agricultural landscapes. Yet many tools to measure insect pollination services require substantial landscape-scale data and technical expertise. In expert workshops, 3 straightforward methods (desk-based method, field survey, and empirical manipulation with exclusion experiments) for rapid insect pollination assessment at site scale were developed to provide an adaptable framework that is accessible to nonspecialist with limited resources. These methods were designed for TESSA (Toolkit for Ecosystem Service Site-Based Assessment) and allow comparative assessment of pollination services at a site of conservation interest and in its most plausible alternative state (e.g., converted to agricultural land). We applied the methods at a nature reserve in the United Kingdom to estimate the value of insect pollination services provided by the reserve. The economic value of pollination services provided by the reserve ranged from US$6163 to US$11,546/year. The conversion of the reserve to arable land would provide no insect pollination services and a net annual benefit from insect-pollinated crop production of approximately $1542/year (US$24∙ha^–1∙year^–1). The methods had wide applicability and were readily adapted to different insect-pollinated crops: rape (Brassica napus) and beans (Vicia faba) crops. All methods were rapidly employed under a low budget. The relatively less robust methods that required fewer resources yielded higher estimates of annual insect pollination benefit.     

Tracing impacts of partner abundance in facultative pollination mutualisms: from individuals to populations

Partner abundance affects costs and benefits in obligate mutualisms, but its role in facultative partnerships is less clear. We address this gap in a pollination web consisting of two clovers (Trifolium) that differ in specialization on a bumble bee pollinator Bombus balteatus. We examine how pollination niche breadth affects plant responses to pollinator abundance, comparing early-flowering (specialized) and late-flowering (generalized) cohorts of T. parryi and early T. parryi to T. dasyphyllum, a pollination generalist. Co-pollinators disrupt the link between B. halteatus visitation and pollination rate for both clovers. Only for early-flowering T. parryi do visitation, pollination, and seed set increase with density of B. balteatus. Bumble bee density also alters timing of seed germination in T. parryi, with seeds from plants receiving augmented B. balteatus germinating sooner than seeds of open-pollinated counterparts. Benefits saturate at intermediate bumble bee densities. Despite strong effects of B. balteatus density on individual plant fitness components, population models suggest little impact of B. balteatus density on lamda in T. parryi or T. dasyphyllum. Findings show that functional redundancy in a pollinator guild mediates host-plant responses to partner density. Unexpected effects of pollinator density on life history schedule have implications for recruitment under pollinator decline.     

On interacting bee/mite populations: a stochastic model with analysis using cumulant truncation

Parasitism by the Varroa mite has had recent drastic impact on both managed and feral bee colonies. This paper proposes a stochastic population dynamics model for interacting African bee colony and Varroa mite populations. Cumulant truncation procedures are used to obtain approximate transient cumulant functions, unconstrained by the usual assumption of bivariate Normality, for an assumed large-scale model. The apparent size of the variance and skewness functions suggest the importance of the proposed truncation procedure which retains some higher-order cumulants, but determining the accuracy of the approximations is problematical. A smaller-scale bee/Varroa mite model is hence proposed and investigated. The accuracy for the means is exceptional, for the second-order cumulants is moderate, and for some third-order cumulants is poor. Notwithstanding the poor accuracy of a skewness approximation, the saddlepoint approximations for the marginal transient population size distributions are excellent. The cumulant truncation methodology is very general, and research is continuing in its application to this new class of host-parasite models.     

Factors Associated with Alien Plants Transitioning from Casual, to Naturalized, to Invasive

Abstract:  To explain current plant invasions, or predict future ones, more knowledge on which factors increase the probability of alien species becoming naturalized and subsequently invasive is needed. We created a database of the alien plants in seminatural habitats in Ireland that included data on taxonomy, invasive status, invasion history, distribution, and biological and ecological plant characteristics. We used information from this database to determine the importance of these factors in increasing the ability of species to become naturalized and invasive. More specifically, we used two multiple logistic regressions to identify factors that distinguish naturalized from casual alien plant species and invasive from noninvasive, naturalized alien species. Clonal growth, moisture-indicator value, nitrogen-indicator value, native range, and date of first record affected (in order of decreasing importance) the probability of naturalization. Factors that distinguished invasive from noninvasive species were ornamental introduction, hermaphrodite flowers, pollination mode, being invasive elsewhere, onset of flowering season, moisture-indicator value, native range, and date of first record. Incorporation of phylogenetic information had little influence on the results, suggesting that the capacity of alien species to naturalize and become invasive evolved largely independently in several phylogenetic lineages. Whereas some of the variables were important for both transitions, others were only important for naturalization or for invasion. This emphasizes the importance of studying different stages of the invasion process when looking for mechanisms of becoming a successful invasive plant, instead of simply comparing invasive with noninvasive alien species. Our results also suggest that a combination of species traits and other variables is likely to produce the most accurate prediction of invasions.     

Land reclamation and strip-mined coal production in appalachia

This study quantifies the short-run impacts of reclamation on strip mining costs, coal prices, production, and employment in Appalachia. A process analysis model is developed and used to estimate short-run strip-mined coal supply functions under conditions of alternative reclamation requirements. Then, an econometric model is developed and used to estimate coal demand relations. Our results show that full reclamation has rather minor impacts. In 1972, full reclamation would have increased strip-mined coal production costs an average of $0.35 per ton, reduced strip-mined coal production by 10 million tons, and cost approximately 1600 jobs in Appalachia.     

An Assessment of Invasion Risk from Assisted Migration

Abstract:  To reduce the risk of extinction due to climate change, some ecologists have suggested human-aided translocation of species, or assisted migration (AM), to areas where climate is projected to become suitable. Such intentional movement, however, may create new invasive species if successful introductions grow out of control and cause ecologic or economic damage. We assessed this risk by surveying invasive species in the United States and categorizing invaders based on origin. Because AM will involve moving species on a regional scale within continents (i.e., range shifts), we used invasive species with an intracontinental origin as a proxy for species that would be moved through AM. We then determined whether intracontinental invasions were more prevalent or harmful than intercontinental invasions. Intracontinental invasions occurred far less frequently than invasions from other continents, but they were just as likely to have had severe effects. Fish and crustaceans pose a particularly high threat of intracontinental invasion. We conclude that the risk of AM to create novel invasive species is small, but assisted species that do become invasive could have large effects. Past experience with species reintroductions may help inform policy regarding AM.     

The ecological impact of invasive cane toads on tropical snakes: field data do not support laboratory-based predictions

Predicting which species will be affected by an invasive taxon is critical to developing conservation priorities, but this is a difficult task. A previous study on the impact of invasive cane toads (Bufo marinus) on Australian snakes attempted to predict vulnerability a priori based on the assumptions that any snake species that eats frogs, and is vulnerable to toad toxins, may be at risk from the toad invasion. We used time-series analyses to evaluate the accuracy of that prediction, based on >3600 standardized nocturnal surveys over a 138-month period on 12 species of snakes and lizards on a floodplain in the Australian wet-dry tropics, bracketing the arrival of cane toads at this site. Contrary to prediction, encounter rates with most species were unaffected by toad arrival, and some taxa predicted to be vulnerable to toads increased rather than declined (e.g., death adder Acanthophis praelongus; Children's python Antaresia childreni). Indirect positive effects of toad invasion (perhaps mediated by toad-induced mortality of predatory varanid lizards) and stochastic weather events outweighed effects of toad invasion for most snake species. Our study casts doubt on the ability of a priori desktop studies, or short-term field surveys, to predict or document the ecological impact of invasive species.     

Local population dynamics of an invasive tree species with a complex life-history cycle: A stochastic matrix model

Biological invasions are widely accepted as having a major impact on ecosystem functioning worldwide, giving urgency to a better understanding of the factors that control their spread. Modelling tools have been developed for this purpose but are often discrete-space, discrete-time spatial-mechanistic models that adopt a computer simulation approach and resist mathematical analysis. We constructed a simple demographic matrix model to explore the local population dynamics of an invasive species with a complex life history and whose invasive success depends on resource availability, which occurs stochastically. As a case study we focused on the American black cherry (Prunus serotina Ehrh.), a gap-dependent tree able both to constitute a long-living seedling bank under unfavourable light conditions and to resprout vigorously once cut-down, which is invading European temperate forests. The model used was a stage-classified matrix population model (i.e., Lefkovitch matrix), integrating environmental stochasticity. Stochastic matrix projection analysis was combined with elasticity analysis and stochastic simulations to search for the species’ ‘Achille heel’. As expected, the population growth rate (i.e., Lyapunov exponent), which measures the risk of P. serotina invasion at the stand scale, increased with light frequency. There was a critical value above which the population of P. serotina explodes and below which it locally goes extinct. The resprouting capacity usually speed up the invasion but appeared to play a minor role. The mean duration of stand invasion was measured and important life stage transitions that mostly contribute to the local stochastic growth rate were identified. Some relevant management implications are discussed and the interest of such models for the understanding of demographic characteristics of invasive species is stressed.     

A simulation model to evaluate the impacts of invasive earthworms on soil carbon dynamics

Recent studies have reported that earthworm invasions alter native communities and impact nutrient cycling in terrestrial ecosystems. We developed a simulation model to evaluate the potential impacts of earthworm invasions on carbon dynamics, taking into consideration earthworm feeding strategies and priming effects on the microorganisms through their casting activities. Responses of carbon stocks (forest litter, soil organic matter, microbial biomass and earthworm populations) and carbon fluxes (litter decomposition, earthworm consumption, and microbial respiration) were used to evaluate an earthworm invasion of a forest ecosystem. Data from a northern temperate forest (Arnot Forest, New York) were adapted for model calibration and evaluation. Simulation results suggest that the impact and outcome of earthworm invasions are affected by pre-invasion resource availability (litter and soil organic matter), invasive earthworm assemblages (particularly feeding strategy), and invasion history (associated with earthworm population dynamics). The abovementioned factors may also determine invasion progress of earthworm species. The accuracy of the model could be improved by the addition of environmental modules (e.g., soil water regimes), precise parameters accounting for individual species attributes under different environmental conditions (e.g. utilization ability of different types of food resources), as well as earthworm population dynamics (size and structure) and interactions with predators and other invasive/indigenous species during the invasion progress. Such an earthworm invasion model could provide valuable evaluation of the complicated responses of carbon dynamics to earthworm invasions in a range of forest ecosystems, particularly under global change scenarios.     

Modeling scale up of anthropogenic impacts from individual pollinator behavior to pollination systems

Understanding how anthropogenic disturbances affect plant–pollinator systems has important implications for the conservation of biodiversity and ecosystem functioning. Previous laboratory studies show that pesticides and pathogens, which have been implicated in the rapid global decline of pollinators over recent years, can impair behavioral processes needed for pollinators to adaptively exploit floral resources and effectively transfer pollen among plants. However, the potential for these sublethal stressor effects on pollinator–plant interactions at the individual level to scale up into changes to the dynamics of wild plant and pollinator populations at the system level remains unclear. We developed an empirically parameterized agent-based model of a bumblebee pollination system called SimBee to test for effects of stressor-induced decreases in the memory capacity and information processing speed of individual foragers on bee abundance (scenario 1), plant diversity (scenario 2), and bee–plant system stability (scenario 3) over 20 virtual seasons. Modeling of a simple pollination network of a bumblebee and four co-flowering bee-pollinated plant species indicated that bee decline and plant species extinction events could occur when only 25% of the forager population showed cognitive impairment. Higher percentages of impairment caused 50% bee loss in just five virtual seasons and system-wide extinction events in less than 20 virtual seasons under some conditions. Plant species extinctions occurred regardless of bee population size, indicating that stressor-induced changes to pollinator behavior alone could drive species loss from plant communities. These findings indicate that sublethal stressor effects on pollinator behavioral mechanisms, although seemingly insignificant at the level of individuals, have the cumulative potential in principle to degrade plant–pollinator species interactions at the system level. Our work highlights the importance of an agent-based modeling approach for the identification and mitigation of anthropogenic impacts on plant–pollinator systems.     

Forecasting the Spread of Invasive Rainbow Smelt in the Laurentian Great Lakes Region of North America

Abstract:  Rainbow smelt ( Osmerus mordax ) have invaded many North American lakes, often resulting in the extirpation of native fish populations. Yet, their invasion is incipient and provides the rationale for identifying ecosystems likely to be invaded and where management and prevention efforts should be focused. To predict smelt presence and absence, we constructed a classification-tree model based on habitat data from 354 lakes in the native range for smelt in southern Maine. Maximum lake depth, lake area, and Secchi depth (surrogate measure of lake productivity) were the most important predictors. We then used our model to identify lakes vulnerable to invasion in three regions outside the smelt's native range: northern Maine (52 of 244 lakes in the non-native range), Ontario (4447 of 8110), and Wisconsin (553 of 5164). We further identified a subset of lakes with a strong potential for impact (potential–impact lakes) based on the presence of fish species that are affected by rainbow smelt. Ninety-four percent of vulnerable lakes in the non-native range in Maine are also potential–impact lakes, as are 94% and 58% of Ontario and Wisconsin's vulnerable lakes, respectively. Our modeling approach can be applied to other invaders and regions to identify invasion-prone ecosystems, thus aiding in the management of invasive species and the efficient allocation of invasive species mitigation and prevention resources.     

Human impacts, plant invasion, and imperiled plant species in California.

Invasive species are one of the fastest growing conservation problems. These species homogenize the world's flora and fauna, threaten rare and endemic species, and impose large economic costs. Here, we examine the distribution of 834 of the more than 1000 exotic plant taxa that have become established in California, USA. Total species richness increases with net primary productivity; however, the exotic flora is richest in low-lying coastal sites that harbor large numbers of imperiled species, while native diversity is highest in areas with high mean elevation. Weedy and invasive exotics are more tightly linked to the distribution of imperiled species than the overall pool of exotic species. Structural equation modeling suggests that while human activities, such as urbanization and agriculture, facilitate the initial invasion by exotic plants, exotics spread ahead of the front of human development into areas with high numbers of threatened native plants. The range sizes of exotic taxa are an order of magnitude smaller than for comparable native taxa. The current small range size of exotic species implies that California has a significant "invasion debt" that will be paid as exotic plants expand their range and spread throughout the state.