Marion E. Wittmann

A Stochastic Model for Annual Reproductive Success

Demographic stochasticity can have large effects on the dynamics of small populations as well as on the persistence of rare genotypes and lineages. Survival is sensibly modeled as a binomial process, but annual reproductive success (ARS) is more complex and general models for demographic stochasticity do not exist. Here we introduce a stochastic model framework for ARS and illustrate some of its properties. We model a sequence of stochastic events: nest completion, the number of eggs or neonates produced, nest predation, and the survival of individual offspring to independence. We also allow multiple nesting attempts within a breeding season. Most of these components can be described by Bernoulli or binomial processes; the exception is the distribution of offspring number. Using clutch and litter size distributions from 53 vertebrate species, we demonstrate that among‐individual variability in offspring number can usually be described by the generalized Poisson distribution. Our model framework allows the demographic variance to be calculated from underlying biological processes and can easily be linked to models of environmental stochasticity or selection because of its parametric structure. In addition, it reveals that the distributions of ARS are often multimodal and skewed, with implications for extinction risk and evolution in small populations.

Forecasting the Impacts of Silver and Bighead Carp on the Lake Erie Food Web

Nonindigenous bigheaded carps (Bighead Carp Hypophthalmichthys nobilis and Silver Carp H. molitrix; hereafter, “Asian carps” [AC]) threaten to invade and disrupt food webs and fisheries in the Laurentian Great Lakes through their high consumption of plankton. To quantify the potential effects of AC on the food web in Lake Erie, we developed an Ecopath with Ecosim (EwE) food web model and simulated four AC diet composition scenarios (high, low, and no detritus and low detritus with Walleye Sander vitreus and Yellow Perch Perca flavescens larvae) and two nutrient load scenarios (the 1999 baseline load and 2£ the baseline [HP]). We quantified the uncertainty of the potential AC effects by coupling the EwE model with estimates of parameter uncertainty in AC production, consumption, and predator diets obtained using structured expert judgment. Our model projected mean § SD AC equilibrium biomass ranging from 52 § 34 to 104 § 75 kg/ha under the different scenarios. Relative to baseline simulations without AC, AC invasion under all detrital diet scenarios decreased the biomass of most fish and zooplankton groups. The effects of AC in the HP scenario were similar to those in the detrital diet scenarios except that the biomasses of most Walleye and Yellow Perch groups were greater under HP because these fishes were buffered from competition with AC by increased productivity at lower trophic levels. Asian carp predation on Walleye and Yellow Perch larvae caused biomass declines among all Walleye and Yellow Perch groups. Large food web impacts of AC occurred in only 2% of the simulations, where AC biomass exceeded 200 kg/ha, resulting in biomass declines of zooplankton and planktivorous fish near the levels observed in the Illinois River. Our findings suggest that AC would affect Lake Erie’s food web by competing with other planktivorous fishes and by providing additional prey for piscivores. Our methods provide a novel approach for including uncertainty into forecasts of invasive species’ impacts on aquatic food webs.

Using structured expert judgment to assess invasive species prevention: Asian carp and the Mississippi-Great Lakes hydrologic connection.

Recently, authors have theorized that invasive species prevention is more cost-effective than control in protecting ecosystem services. However, quantification of the effectiveness of prevention is rare because experiments at field scales are expensive or infeasible. We therefore used structured expert judgment to quantify the efficacy of 17 proposed strategies to prevent Asian carp invasion of the Laurentian Great Lakes via the hydrologic connection between the Mississippi and Great Lakes watersheds. Performance-weighted expert estimates indicated that hydrologic separation would prevent 99% (95,100; median, 5th and 95th percentiles) of Asian carp access, while electric and acoustic-bubble-strobe barriers would prevent 92% (85,95) and 92% (75,95), respectively. For all other strategies, estimated effectiveness was lower, with greater uncertainty. When potential invasions by other taxa are considered, the effectiveness of hydrologic separation increases relative to strategies that are effective primarily for fishes. These results could help guide invasive species management in many waterways globally.

Use of structured expert judgment to forecast invasions by bighead and silver carp in Lake Erie

Identifying which nonindigenous species will become invasive and forecasting the damage they will cause is difficult and presents a significant problem for natural resource management. Often, the data or resources necessary for ecological risk assessment are incomplete or absent, leaving environmental decision makers ill equipped to effectively manage valuable natural resources. Structured expert judgment (SEJ) is a mathematical and performance-based method of eliciting, weighting, and aggregating expert judgments. In contrast to other methods of eliciting and aggregating expert judgments (where, for example, equal weights may be assigned to experts), SEJ weights each expert on the basis of his or her statistical accuracy and informativeness through performance measurement on a set of calibration variables. We used SEJ to forecast impacts of nonindigenous Asian carp (Hypophthalmichthys spp.) in Lake Erie, where it is believed not to be established. Experts quantified Asian carp biomass, production, and consumption and their impact on 4 fish species if Asian carp were to become established. According to experts, in Lake Erie Asian carp have the potential to achieve biomass levels that are similar to the sum of biomasses for several fishes that are harvested commercially or recreationally. However, the impact of Asian carp on the biomass of these fishes was estimated by experts to be small, relative to long term average biomasses, with little uncertainty. Impacts of Asian carp in tributaries and on recreational activities, water quality, or other species were not addressed. SEJ can be used to quantify key uncertainties of invasion biology and also provide a decision-support tool when the necessary information for natural resource management and policy is not available.

Geographic selection bias of occurrence data influences transferability of invasive Hydrilla verticillata distribution models

Due to socioeconomic differences, the accuracy and extent of reporting on the occurrence of native species differs among countries, which can impact the performance of species distribution models. We assessed the importance of geographical biases in occurrence data on model performance using Hydrilla verticillata as a case study. We used Maxent to predict potential North American distribution of the aquatic invasive macrophyte based upon training data from its native range. We produced a model using all available native range occurrence data, then explored the change in model performance produced by omitting subsets of training data based on political boundaries. We also compared those results with models trained on data from which a random sample of occurrence data was omitted from across the native range. Although most models accurately predicted the occurrence of H. verticillata in North America (AUC > 0.7600), data omissions influenced model predictions. Omitting data based on political boundaries resulted in larger shifts in model accuracy than omitting randomly selected occurrence data. For well-documented species like H. verticillata, missing records from single countries or ecoregions may minimally influence model predictions, but for species with fewer documented occurrences or poorly understood ranges, geographic biases could misguide predictions. Regardless of focal species, we recommend that future species distribution modeling efforts begin with a reflection on potential spatial biases of available occurrence data. Improved biodiversity surveillance and reporting will provide benefit not only in invaded ranges but also within under-reported and unexplored native ranges.

Early invasion population structure of quagga mussel and associated benthic invertebrate community composition on soft sediment in a large reservoir

Abstract In 2007 an invasive dreissenid mussel species, Dreissena bugensis (quagga mussel), was discovered in Lake Mead reservoir (AZ–NV). Within 2 years, adult populations have spread throughout the lake and are not only colonizing hard substrates, but also establishing in soft sediments at depths ranging from 1 to >100 m. Dreissena bugensis size class and population density distribution differs between basins; cluster analysis revealed 5 adult cohorts within Boulder Basin and Overton Arm but low densities and low cohort survival in the Las Vegas Basin. Regression analysis suggests depth and temperature are not primary controllers of D. bugensis density in Lake Mead, indicating other factors such as sediment type, food availability or other resource competition may be important. Monthly veliger tows showed at least 2 major spawning events per year, with continuous presence of veligers in the water column. Adult mussels have been found in spawn or post-spawn condition in soft sediments in shallow to deep waters (>80 m) indicating the potential for reproduction at multiple depths. Comparisons to a 1986 benthic survey suggest there have been shifts in nondreissenid macroinvertebrate composition; however, it is unclear if this is due to D. bugensis presence. Current distribution of nondreissenid macroinvertebrates is heterogeneous in all 3 basins, and their biodiversity decreased when D. bugensis density was 2500/m2 or greater.

Tradeoffs among Ecosystem Services Associated with Global Tilapia Introductions

ABSTRACT Previous publications have reached different conclusions about the balance between the socioeconomic benefits of tilapia introduction for aquaculture and capture fisheries, and the potential negative impacts of these species on ecosystem services such as the provisioning of food, habitat, and water quality. This review (1) provides a new estimate of the global scale of tilapia introduction and the reported occurrence of impacts to ecosystem services; (2) assesses whether reported changes to ecosystem services differ among species, regions and type of ecological effect reported; and (3) determine how perceptions of tilapia introduction are related to the reported occurrence of ecological effects and/or the contribution of tilapia to countries’ economies. Feral tilapia populations now exist in at least 114 countries. The majority of research is consistent with the occurrence of ecological change as a result of tilapia introduction, regardless of species or region of study, but there are important regional influences including the economic contributions of tilapia. There is an increasing recognition that introductions provide both significant benefits and considerable but not fully documented harm to ecosystem services. Stakeholders should consider both ecological effects and socioeconomic context, such as reviewed here, in making decisions about the introduction of tilapia.

Confronting species distribution model predictions with species functional traits

Abstract Species distribution models are valuable tools in studies of biogeography, ecology, and climate change and have been used to inform conservation and ecosystem management. However, species distribution models typically incorporate only climatic variables and species presence data. Model development or validation rarely considers functional components of species traits or other types of biological data. We implemented a species distribution model (Maxent) to predict global climate habitat suitability for Grass Carp (Ctenopharyngodon idella). We then tested the relationship between the degree of climate habitat suitability predicted by Maxent and the individual growth rates of both wild (N = 17) and stocked (N = 51) Grass Carp populations using correlation analysis. The Grass Carp Maxent model accurately reflected the global occurrence data (AUC = 0.904). Observations of Grass Carp growth rate covered six continents and ranged from 0.19 to 20.1 g day−1. Species distribution model predictions were correlated (r = 0.5, 95% CI (0.03, 0.79)) with observed growth rates for wild Grass Carp populations but were not correlated (r = −0.26, 95% CI (−0.5, 0.012)) with stocked populations. Further, a review of the literature indicates that the few studies for other species that have previously assessed the relationship between the degree of predicted climate habitat suitability and species functional traits have also discovered significant relationships. Thus, species distribution models may provide inferences beyond just where a species may occur, providing a useful tool to understand the linkage between species distributions and underlying biological mechanisms.

Four decades of change: dramatic loss of zoobenthos in an oligotrophic lake exhibiting gradual eutrophication

Abstract. Native species in lakes commonly are affected by cultural eutrophication and introductions of nonnative species. The effects of these disturbances on benthic communities in large lakes have been understudied, despite the integral role that benthos play in maintaining habitat complexity and ecosystem processes. Lake Tahoe has experienced progressive eutrophication and introductions of nonnative species over the past several decades, but how its unique benthic communities have been affected is unclear. The density of Lake Tahoes benthic fauna was compared between 1960s surveys and our recent (2008–2009) survey, and the association of zoobenthos with macrophytes was examined for contemporary and historical samples. The density of benthic invertebrates and the occurrence of macrophytes in benthic samples have declined dramatically since collections made in the 1960s. Lakewide densities of benthic invertebrate taxa endemic to Lake Tahoe have declined by 80 to 100%, and the community structure of benthic invertebrate assemblages has changed considerably. Several native benthic invertebrate taxa were closely associated with deepwater macrophytes in the 1960s, but contemporary invertebrate association with macrophytes could not be evaluated reliably because of the scarcity of macrophytes in contemporary samples. Declines in native benthic invertebrate density could be related to the loss of habitat and food resources previously provided by abundant deepwater macrophyte assemblages. In addition, establishment and increases in density of nonnative species that occurred after the benthic surveys of the 1960s probably have affected native benthic invertebrate communities. The observed declines in Lake Tahoes native benthic invertebrate and macrophyte communities suggest that they are severely threatened.

A 3D individual-based aquatic transport model for the assessment of the potential dispersal of planktonic larvae of an invasive bivalve

The unwanted impacts of non-indigenous species have become one of the major ecological and economic threats to aquatic ecosystems worldwide. Assessing the potential dispersal and colonization of non-indigenous species is necessary to prevent or reduce deleterious effects that may lead to ecosystem degradation and a range of economic impacts. A three dimensional (3D) numerical model has been developed to evaluate the local dispersal of the planktonic larvae of an invasive bivalve, Asian clam (Corbicula fluminea), by passive hydraulic transport in Lake Tahoe, USA. The probability of dispersal of Asian clam larvae from the existing high density populations to novel habitats is determined by the magnitude and timing of strong wind events. The probability of colonization of new near-shore areas outside the existing beds is low, but sensitive to the larvae settling velocity ws. High larvae mortality was observed due to settling in unsuitable deep habitats. The impact of UV-radiation during the pelagic stages, on the Asian clam mortality was low. This work provides a quantification of the number of propagules that may be successfully transported as a result of natural processes and in function of population size. The knowledge and understanding of the relative contribution of different dispersal pathways, may directly inform decision-making and resource allocation associated with invasive species management.