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Challenges and opportunities for data integration to improve estimation of migratory connectivity

Understanding migratory connectivity, or the linkage of populations between seasons, is critical for effective conservation and management of migratory wildlife. A growing number of tools are available for understanding where migratory individuals and populations occur throughout the annual cycle. Integration of the diverse measures of migratory movements can help elucidate migratory connectivity patterns with methodology that accounts for differences in sampling design, directionality, effort, precision and bias inherent to each data type. The R package MigConnectivity was developed to estimate population-specific connectivity and the range-wide strength of those connections. New functions allow users to integrate intrinsic markers, tracking and long-distance reencounter data, collected from the same or different individuals, to estimate population-specific transition probabilities (estTransition) and the range-wide strength of those transition probabilities (estStrength). We used simulation and real-world case studies to explore the challenges and limitations of data integration based on data from three migratory bird species, Painted Bunting ( Passerina ciris ), Yellow Warbler ( Setophaga petechia ) and Bald Eagle ( Haliaeetus leucocephalus ), two of which had bidirectional data. We found data integration is useful for quantifying migratory connectivity, as single data sources are less likely to be available across the species range. Furthermore, accurate strength estimates can be obtained from either breeding-to-nonbreeding or nonbreeding-to-breeding data. For bidirectional data, integration can lead to more accurate estimates when data are available from all regions in at least one season. The ability to conduct combined analyses that account for the unique limitations and biases of each data type is a promising possibility for overcoming the challenge of range-wide coverage that has been hard to achieve using single data types. The best-case scenario for data integration is to have data from all regions, especially if the question is range-wide or data are bidirectional. Multiple data types on animal movements are becoming increasingly available and integration of these growing datasets will lead to a better understanding of the full annual cycle of migratory animals.

Methods in Ecology and Evolution

Sampling mass mortality events to enable diagnoses: A protocol using freshwater mussels

Many taxa around the globe are threatened by often unexplained mass mortality events (MMEs), which can decimate populations and compromise key ecosystem functions. One example of a highly threatened taxon facing frequent MMEs is freshwater mussels (Unionida). There has been a recent increase in interest in understanding the causes of freshwater mussel MMEs, but standardised methodologies for how best to respond to them to facilitate diagnoses are unavailable. When an MME is observed, swift and appropriate sample collection is imperative owing to the transient nature of these phenomena. Here we provide structured guidance that will facilitate rapid and appropriate sampling of MMEs, using freshwater mussels as an example. We set out standardised procedures for sample collection, preparation and preservation. The procedures we outline will improve our capacity for diagnostic investigations of MMEs and other mortality events, not only in freshwater mussels but also across many other taxa. This, in turn, can inform appropriate management responses.

Methods in Ecology and Evolution

Spatial close-kin mark-recapture models applied to terrestrial species with continuous natal dispersal

Close-kin mark–recapture (CKMR) methods use information on genetic relatedness among individuals to estimate demographic parameters. An individual's genotype can be considered a ‘recapture’ of each of its parent's genotype, and the frequency of kin-pair matches detected in a population sample can directly inform estimates of abundance. CKMR inference procedures require analysts to define kinship probabilities in functional forms, which inevitably involve simplifying assumptions. Among others, population structure can have a strong influence on how kinship probabilities are formulated. Many terrestrial species are philopatric or face barriers to dispersal, and not accounting for dispersal limitation in kinship probabilities, can create substantial bias if sampling is also spatially structured (e.g. via harvest). We present a spatially explicit formulation of CKMR that corrects for incomplete mixing by incorporating natal dispersal distances and spatial distribution of individuals into the kinship probabilities. We used individual-based simulations to evaluate the accuracy of abundance estimates obtained with one spatially naïve and two spatially explicit CKMR models across six scenarios with distinct spatial patterns of relative abundance and sampling probability. Estimates of abundance obtained with a CKMR model naïve to spatial structure were negatively biased when sampling was spatially biased. Incorporating patterns of natal dispersal in the kinship probabilities helped address this bias, but estimates were not always accurate depending on the model used and the scenario considered. Incorporating natal dispersal into spatially structured CKMR models can address the bias created by population structure and heterogeneous sampling but will often require additional assumptions and auxiliary data (e.g. relative abundance indices). The models shown here were designed for terrestrial species with continuous patterns of natal dispersal and high year-to-year site fidelity but could be extended to other species.

Methods in Ecology and Evolution

Simulated soundscapes and transfer learning boost the performance of acoustic classifiers under data scarcity

1. The biodiversity crisis necessitates spatially extensive methods to monitor multiple taxonomic groups for evidence of change in response to evolving environmental conditions. Programs that combine passive acoustic monitoring and machine learning are increasingly used to meet this need. These methods require large, annotated datasets, which are time-consuming and expensive to produce, creating potential barriers to adoption in data- and funding-poor regions. Recently released pre-trained avian acoustic classification models provide opportunities to reduce the need for manual labelling and accelerate the development of new acoustic classification algorithms through transfer learning. Transfer learning is a strategy for developing algorithms under data scarcity that uses pre-trained models from related tasks to adapt to new tasks. 2. Our primary objective was to develop a transfer learning strategy using the feature embeddings of a pre-trained avian classification model to train custom acoustic classification models in data-scarce contexts. We used three annotated avian acoustic datasets to test whether transfer learning and soundscape simulation-based data augmentation could substantially reduce the annotated training data necessary to develop performant custom acoustic classifiers. We also conducted a sensitivity analysis for hyperparameter choice and model architecture. We then assessed the generalizability of our strategy to increasingly novel non-avian classification tasks. 3. With as few as two training examples per class, our soundscape simulation data augmentation approach consistently yielded new classifiers with improved performance relative to the pre-trained classification model and transfer learning classifiers trained with other augmentation approaches. Performance increases were evident for three avian test datasets, including single-class and multi-label contexts. We observed that the relative performance among our data augmentation approaches varied for the avian datasets and nearly converged for one dataset when we included more training examples. 4. We demonstrate an efficient approach to developing new acoustic classifiers leveraging open-source sound repositories and pre-trained networks to reduce manual labelling. With very few examples, our soundscape simulation approach to data augmentation yielded classifiers with performance equivalent to those trained with many more examples, showing it is possible to reduce manual label-ling while still achieving high-performance classifiers and, in turn, expanding the potential for passive acoustic monitoring to address rising biodiversity monitoring needs.

Methods in Ecology and Evolution

Unbiased survival estimates and evidence for skipped breeding opportunities in females

1.  Estimates of demographic parameters for females, in many organisms, are sparse. This is particularly worrisome as more and more species are faced with high extinction probabilities and conservation increasingly depends on actions dictated by complex predictive models that require accurate estimates of demographic parameters for each sex and species. 2.  This study assesses demographic parameters, specifically temporary emigration and survival, for females, a class that has been difficult to investigate historically because of lack of data. Amphibians provide a particularly good example because there is global concern about amphibian decline; yet most demographic parameter estimates are based on data from males, which we show can lead to erroneous conclusions. 3.  We use 10 years of capture–recapture data from boreal toads ( Bufo boreas ) and the multi-state open robust design model to provide evidence for the occurrence of skipped breeding opportunities (i.e. temporary emigration) in females. This is the first time that the open robust design model has been applied to an analysis of an amphibian population that we are aware of. 4.  We determined that the transition from breeder to non-breeder is obligate and the probability of a non-breeder remaining a non-breeder is 64%; thus, temporary emigration is first-order Markovian in nature, where breeding probability is dependent on the previous year’s activity, i.e. if a female did not breed in year one, there is a 36% chance that she will breed in year two. With temporary emigration accounted for, we estimated between-year female survival at 87%. 5.  Establishing the occurrence of temporary emigration not only reduces bias in estimates of survival probabilities but also provides information about expected breeding attempts by females, a critical element in understanding the ecology of an organism and the impacts of outside stressors and conservation actions.

Methods in Ecology and Evolution

Assessing conservation relevance of organism-environment relations using predicted changes in response variables

1. Organism–environment models are used widely in conservation. The degree to which they are useful for informing conservation decisions – the conservation relevance of these relations – is important because lack of relevance may lead to misapplication of scarce conservation resources or failure to resolve important conservation dilemmas. Even when models perform well based on model fit and predictive ability, conservation relevance of associations may not be clear without also knowing the magnitude and variability of predicted changes in response variables. 2. We introduce a method for evaluating the conservation relevance of organism–environment relations that employs confidence intervals for predicted changes in response variables. The confidence intervals are compared to a preselected magnitude of change that marks a threshold (trigger) for conservation action. To demonstrate the approach, we used a case study from the Chihuahuan Desert involving relations between avian richness and broad-scale patterns of shrubland. We considered relations for three winters and two spatial extents (1- and 2-km-radius areas) and compared predicted changes in richness to three thresholds (10%, 20% and 30% change). For each threshold, we examined 48 relations. 3. The method identified seven, four and zero conservation-relevant changes in mean richness for the 10%, 20% and 30% thresholds respectively. These changes were associated with major (20%) changes in shrubland cover, mean patch size, the coefficient of variation for patch size, or edge density but not with major changes in shrubland patch density. The relative rarity of conservation-relevant changes indicated that, overall, the relations had little practical value for informing conservation decisions about avian richness. 4. The approach we illustrate is appropriate for various response and predictor variables measured at any temporal or spatial scale. The method is broadly applicable across ecological environments, conservation objectives, types of statistical predictive models and levels of biological organization. By focusing on magnitudes of change that have practical significance, and by using the span of confidence intervals to incorporate uncertainty of predicted changes, the method can be used to help improve the effectiveness of conservation efforts.

Methods in Ecology and Evolution

Accounting for non-independent detection when estimating abundance of organisms with a Bayesian approach

Summary 1. Binomial mixture models use repeated count data to estimate abundance. They are becoming increasingly popular because they provide a simple and cost‐effective way to account for imperfect detection. However, these models assume that individuals are detected independently of each other. This assumption may often be violated in the field. For instance, manatees ( Trichechus manatus latirostris ) may surface in turbid water (i.e. become available for detection during aerial surveys) in a correlated manner (i.e. in groups). However, correlated behaviour, affecting the non‐independence of individual detections, may also be relevant in other systems (e.g. correlated patterns of singing in birds and amphibians). 2. We extend binomial mixture models to account for correlated behaviour and therefore to account for non‐independent detection of individuals. We simulated correlated behaviour using beta‐binomial random variables. Our approach can be used to simultaneously estimate abundance, detection probability and a correlation parameter. 3. Fitting binomial mixture models to data that followed a beta‐binomial distribution resulted in an overestimation of abundance even for moderate levels of correlation. In contrast, the beta‐binomial mixture model performed considerably better in our simulation scenarios. We also present a goodness‐of‐fit procedure to evaluate the fit of beta‐binomial mixture models. 4. We illustrate our approach by fitting both binomial and beta‐binomial mixture models to aerial survey data of manatees in Florida. We found that the binomial mixture model did not fit the data, whereas there was no evidence of lack of fit for the beta‐binomial mixture model. This example helps illustrate the importance of using simulations and assessing goodness‐of‐fit when analysing ecological data with N‐mixture models. Indeed, both the simulations and the goodness‐of‐fit procedure highlighted the limitations of the standard binomial mixture model for aerial manatee surveys. 5. Overestimation of abundance by binomial mixture models owing to non‐independent detections is problematic for ecological studies, but also for conservation. For example, in the case of endangered species, it could lead to inappropriate management decisions, such as downlisting. These issues will be increasingly relevant as more ecologists apply flexible N‐mixture models to ecological data.

Methods in Ecology and Evolution

Spatial capture-recapture models for search-encounter data

1. Spatial capture–recapture models make use of auxiliary data on capture location to provide density estimates for animal populations. Previously, models have been developed primarily for fixed trap arrays which define the observable locations of individuals by a set of discrete points. 2. Here, we develop a class of models for 'search-encounter' data, i.e. for detections of recognizable individuals in continuous space, not restricted to trap locations. In our hierarchical model, detection probability is related to the average distance between individual location and the survey path. The locations are allowed to change over time owing to movements of individuals, and individual locations are related formally by a model describing individual activity or home range centre which is itself regarded as a latent variable in the model. We provide a Bayesian analysis of the model in WinBUGS, and develop a custom MCMC algorithm in the R language. 3. The model is applied to simulated data and to territory mapping data for the Willow Tit from the Swiss Breeding Bird Survey MHB. While the observed density was 15 territories per nominal 1 km 2 plot of unknown effective sample area, the model produced a density estimate of 21∙12 territories per square km (95% posterior interval: 17–26). 4. Spatial capture–recapture models are relevant to virtually all animal population studies that seek to estimate population size or density, yet existing models have been proposed mainly for conventional sampling using arrays of traps. Our model for search-encounter data, where the spatial pattern of searching can be arbitrary and may change over occasions, greatly expands the scope and utility of spatial capture–recapture models.

Methods in Ecology and Evolution

On thinning of chains in MCMC

1. Markov chain Monte Carlo (MCMC) is a simulation technique that has revolutionised the analysis of ecological data, allowing the fitting of complex models in a Bayesian framework. Since 2001, there have been nearly 200 papers using MCMC in publications of the Ecological Society of America and the British Ecological Society, including more than 75 in the journal Ecology and 35 in the Journal of Applied Ecology . 2. We have noted that many authors routinely 'thin' their simulations, discarding all but every k th sampled value; of the studies we surveyed with details on MCMC implementation, 40% reported thinning. 3. Thinning is often unnecessary and always inefficient, reducing the precision with which features of the Markov chain are summarised. The inefficiency of thinning MCMC output has been known since the early 1990's, long before MCMC appeared in ecological publications. 4. We discuss the background and prevalence of thinning, illustrate its consequences, discuss circumstances when it might be regarded as a reasonable option and recommend against routine thinning of chains unless necessitated by computer memory limitations.

Methods in Ecology and Evolution

Likelihood analysis of species occurrence probability from presence-only data for modelling species distributions

1. Understanding the factors affecting species occurrence is a pre-eminent focus of applied ecological research. However, direct information about species occurrence is lacking for many species. Instead, researchers sometimes have to rely on so-called presence-only data (i.e. when no direct information about absences is available), which often results from opportunistic, unstructured sampling. MAXENT is a widely used software program designed to model and map species distribution using presence-only data. 2. We provide a critical review of MAXENT as applied to species distribution modelling and discuss how it can lead to inferential errors. A chief concern is that MAXENT produces a number of poorly defined indices that are not directly related to the actual parameter of interest – the probability of occurrence ( ψ ). This focus on an index was motivated by the belief that it is not possible to estimate ψ from presence-only data; however, we demonstrate that ψ is identifiable using conventional likelihood methods under the assumptions of random sampling and constant probability of species detection. 3. The model is implemented in a convenient r package which we use to apply the model to simulated data and data from the North American Breeding Bird Survey. We demonstrate that MAXENT produces extreme under-predictions when compared to estimates produced by logistic regression which uses the full (presence/absence) data set. We note that MAXENT predictions are extremely sensitive to specification of the background prevalence, which is not objectively estimated using the MAXENT method. 4. As with MAXENT, formal model-based inference requires a random sample of presence locations. Many presence-only data sets, such as those based on museum records and herbarium collections, may not satisfy this assumption. However, when sampling is random, we believe that inference should be based on formal methods that facilitate inference about interpretable ecological quantities instead of vaguely defined indices.

Methods in Ecology and Evolution

A general theory of multimetric indices and their properties

1. Stewardship of biological and ecological resources requires the ability to make integrative assessments of ecological integrity. One of the emerging methods for making such integrative assessments is multimetric indices (MMIs). These indices synthesize data, often from multiple levels of biological organization, with the goal of deriving a single index that reflects the overall effects of human disturbance. Despite the widespread use of MMIs, there is uncertainty about why this approach can be effective. An understanding of MMIs requires a quantitative theory that illustrates how the properties of candidate metrics relates to MMIs generated from those metrics. 2. We present the initial basis for such a theory by deriving the general mathematical characteristics of MMIs assembled from metrics. We then use the theory to derive quantitative answers to the following questions: Is there an optimal number of metrics to comprise an index? How does covariance among metrics affect the performance of the index derived from those metrics? And what are the criteria to decide whether a given metric will improve the performance of an index? 3. We find that the optimal number of metrics to be included in an index depends on the theoretical distribution of signal of the disturbance gradient contained in each metric. For example, if the rank-ordered parameters of a metric-disturbance regression can be described by a monotonically decreasing function, then an optimum number of metrics exists and can often be derived analytically. We derive the conditions by which adding a given metric can be expected to improve an index. 4. We find that the criterion defining such conditions depends nonlinearly of the signal of the disturbance gradient, the noise (error) of the metric and the correlation of the metric errors. Importantly, we find that correlation among metric errors increases the signal required for the metric to improve the index. 5. The theoretical framework presented in this study provides the basis for understanding the properties of MMIs. It can also be useful throughout the index construction process. Specifically, it can be used to aid understanding of the benefits and limitations of combining metrics into indices; it can inform selection/collection of candidate metrics; and it can be used directly as a decision aid in effective index construction.

Methods in Ecology and Evolution

A two-phase sampling design for increasing detections of rare species in occupancy surveys

1. Occupancy estimation is a commonly used tool in ecological studies owing to the ease at which data can be collected and the large spatial extent that can be covered. One major obstacle to using an occupancy-based approach is the complications associated with designing and implementing an efficient survey. These logistical challenges become magnified when working with rare species when effort can be wasted in areas with none or very few individuals. 2. Here, we develop a two-phase sampling approach that mitigates these problems by using a design that places more effort in areas with higher predicted probability of occurrence. We compare our new sampling design to traditional single-season occupancy estimation under a range of conditions and population characteristics. We develop an intuitive measure of predictive error to compare the two approaches and use simulations to assess the relative accuracy of each approach. 3. Our two-phase approach exhibited lower predictive error rates compared to the traditional single-season approach in highly spatially correlated environments. The difference was greatest when detection probability was high (0·75) regardless of the habitat or sample size. When the true occupancy rate was below 0·4 (0·05-0·4), we found that allocating 25% of the sample to the first phase resulted in the lowest error rates. 4. In the majority of scenarios, the two-phase approach showed lower error rates compared to the traditional single-season approach suggesting our new approach is fairly robust to a broad range of conditions and design factors and merits use under a wide variety of settings. 5. Synthesis and applications. Conservation and management of rare species are a challenging task facing natural resource managers. It is critical for studies involving rare species to efficiently allocate effort and resources as they are usually of a finite nature. We believe our approach provides a framework for optimal allocation of effort while maximizing the information content of the data in an attempt to provide the highest conservation value per unit of effort.

Methods in Ecology and Evolution

Program SPACECAP: software for estimating animal density using spatially explicit capture-recapture models

1. The advent of spatially explicit capture-recapture models is changing the way ecologists analyse capture-recapture data. However, the advantages offered by these new models are not fully exploited because they can be difficult to implement. 2. To address this need, we developed a user-friendly software package, created within the R programming environment, called SPACECAP. This package implements Bayesian spatially explicit hierarchical models to analyse spatial capture-recapture data. 3. Given that a large number of field biologists prefer software with graphical user interfaces for analysing their data, SPACECAP is particularly useful as a tool to increase the adoption of Bayesian spatially explicit capture-recapture methods in practice.

Methods in Ecology and Evolution

Moderating Argos location errors in animal tracking data

1. The Argos System is used worldwide to satellite-track free-ranging animals, but location errors can range from tens of metres to hundreds of kilometres. Low-quality locations (Argos classes A, 0, B and Z) dominate animal tracking data. Standard-quality animal tracking locations (Argos classes 3, 2 and 1) have larger errors than those reported in Argos manuals. 2. The Douglas Argos-filter (DAF) algorithm flags implausible locations based on user-defined thresholds that allow the algorithm's performance to be tuned to species' movement behaviours and study objectives. The algorithm is available in Movebank – a free online infrastructure for storing, managing, sharing and analysing animal movement data. 3. We compared 21,044 temporally paired global positioning system (GPS) locations with Argos location estimates collected from Argos transmitters on free-ranging waterfowl and condors (13 species, 314 individuals, 54,895 animal-tracking days). The 95th error percentiles for unfiltered Argos locations 0, A, B and Z were within 35·8, 59·6, 163·2 and 220·2 km of the true location, respectively. After applying DAF with liberal thresholds, roughly 20% of the class 0 and A locations and 45% of the class B and Z locations were excluded, and the 95th error percentiles were reduced to 17·2, 15·0, 20·9 and 18·6 km for classes 0, A, B and Z, respectively. As thresholds were applied more conservatively, fewer locations were retained, but they possessed higher overall accuracy. 4. Douglas Argos-filter can improve data accuracy by 50–90% and is an effective and flexible tool for preparing Argos data for direct biological interpretation or subsequent modelling.

Methods in Ecology and Evolution

BatTool: Projecting bat populations facing multiple stressors using a demographic model

Bats provide ecologically and agriculturally important ecosystem services but are currently experiencing population declines caused by multiple environmental stressors, including mortality from white-nose syndrome and wind energy development. Analyses of the current and future health and viability of these species may support conservation management decision making. Demographic modeling provides a quantitative tool for decision makers and conservation managers to make more informed decisions, but widespread adoption of these tools can be limited because of the complexity of the mathematical, statistical, and computational components involved in implementing these models. In this work, we provide an exposition of the BatTool R package, detailing the primary components of the matrix projection model, a publicly accessible graphical user interface ( https://rconnect.usgs.gov/battool ) facilitating user-defined scenario analyses, and its intended uses and limitations (Wiens et al., US Geol Surv Data Release 2022; Wiens et al., US Geol Surv Softw Release 2022). We present a case study involving wind energy permitting, weighing the effects of potential mortality caused by a hypothetical wind energy facility on the projected abundance of four imperiled bat species in the Midwestern United States.

Methods in Ecology and Evolution

Coevolution with host fishes shapes parasitic life histories in a group of freshwater mussels (Unionidae: Quadrulini)

Ecological interactions among species often lead to parasitic lineages coevolving with host resources, which is often suggested as the primary driver of parasite diversification. Freshwater mussels are bivalves that possess a parasitic life cycle requiring larval encystment on freshwater vertebrates to complete metamorphosis. The North American freshwater mussel tribe Quadrulini has a suite of life history adaptations including highly specialized patterns of host use, infection strategies, and variable larval morphologies. However, the evolution of life histories has yet to be explored using phylogenetic comparative methods. In this study, we use a holistic approach incorporating biogeographical, ecological, molecular, and morphological datasets to reconstruct the evolution of Quadrulini. Comparative phylogenetic analyses suggested the diversification of Quadrulini has been driven, at least in part, by codiversification with their primary host fishes in Ictaluridae. Major diversification events in both ictalurids and quadrulines were estimated to have occurred in the Mississippi River basin throughout the Miocene. Life history characteristics associated with parasitism were supported to have coevolved with host repertories, supporting the hypothesis that ecological interactions with host fishes have shaped the evolution of highly specialized traits in this group. Our findings demonstrate the importance of ecological interactions with host resources in shaping the evolutionary history of freshwater mussels.

Bulletin of the Society of Systematic Biologists

Positively selected genes in the hoary bat (Lasiurus cinereus) lineage: Prominence of thymus expression, immune and metabolic function, and regions of ancient synteny

Background Bats of the genus Lasiurus occur throughout the Americas and have diversified into at least 20 species among three subgenera. The hoary bat ( Lasiurus cinereus ) is highly migratory and ranges farther across North America than any other wild mammal. Despite the ecological importance of this species as a major insect predator, and the particular susceptibility of lasiurine bats to wind turbine strikes, our understanding of hoary bat ecology, physiology, and behavior remains poor. Methods To better understand adaptive evolution in this lineage, we used whole-genome sequencing to identify protein-coding sequence and explore signatures of positive selection. Gene models were predicted with Maker and compared to seven well-annotated and phylogenetically representative species. Evolutionary rate analysis was performed with PAML. Results Of 9,447 single-copy orthologous groups that met evaluation criteria, 150 genes had a significant excess of nonsynonymous substitutions along the L. cinereus branch ( P < 0.001 after manual review of alignments). Selected genes as a group had biased expression, most strongly in thymus tissue. We identified 23 selected genes with reported immune functions as well as a divergent paralog of Steep1 within suborder Yangochiroptera. Seventeen genes had roles in lipid and glucose metabolic pathways, partially overlapping with 15 mitochondrion-associated genes; these adaptations may reflect the metabolic challenges of hibernation, long-distance migration, and seasonal variation in prey abundance. The genomic distribution of positively selected genes differed significantly from background expectation by discrete Kolmogorov–Smirnov test ( P < 0.001). Remarkably, the top three physical clusters all coincided with islands of conserved synteny predating Mammalia, the largest of which shares synteny with the human cat-eye critical region (CECR) on 22q11. This observation coupled with the expansion of a novel Tbx1 -like gene family may indicate evolutionary innovation during pharyngeal arch development: both the CECR and Tbx1 cause dosage-dependent congenital abnormalities in thymus, heart, and head, and craniodysmorphy is associated with human orthologs of other positively selected genes as well.

PeerJ

Interactions between fire and invasive plants in temperate grasslands of North America

A substantial number of invasive grasses, forbs and woody plants have invaded temperate grasslands in North America. Among the invading species are winter annuals, biennials, cool-season perennials, warm-season perennials, vines, shrubs, and trees. Many of these species have been deliberately introduced and widely planted; some are still used for range improvement, pastures, lawns, and as ornamentals, though many are listed as state or federal noxious weeds. Others have been greatly facilitated by widespread land disturbance. Historically, fire has been a major selective force in the evolution of temperate grasslands. Further, prescribed fire is commonly used as a method of ecological management for native grassland communities as well as in conjunction with restoration efforts. Within this context, it is important to understand how invasive species will interact with natural and prescribed fire regimes. In this paper, we consider what is known about how exotic species that invade temperate grasslands relate to fire. The primary issues addressed for each species are (1) Does fire appear to enhance colonization? (2) To what degree does fire affect the survival of plants? (3) Are plants that are burned able to regrow following fire and, if so, how rapidly can they recover? (4) How important is competition with native species to the response to fire? and (5) What effect does an invasive species have on the characteristics of the fire regime? For many species, results are preliminary, incomplete, or inconsistent among studies. For this reason, many of the conclusions drawn for individual species must be considered preliminary. Based on analyses of individual species, a conceptual framework is presented for considering how invasive plants may interact with fire when they invade an ecosystem. The major categories of influences are the native community, the fire regime, growth conditions for both invasive and native species, and influences that disturbances, human impacts, and landscape characteristics have had in the past and will have in the future. The examples considered in this paper provide support for a few, tentative generalizations. First, among our current worst invaders of temperate grasslands, adaptation to fire is quite variable. Some species are not well adapted to burning and can be easily eliminated; other species are better adapted but can still be eliminated if fire occurs during periods of particular vulnerability and/or at high frequency. There is a set of species that is extremely well adapted to fire and will not be eliminated through burning alone. Second, competitive interactions with native species play a crucial role in the success of nonnative invaders. In cases where differential burn responses between invasive and native species can be exploited, and adequate populations of native dominant species are present, fire can sometimes tip the competitive balance away from invasives. Third, there are a few invasive species that have exceptional attributes and for which there are no easy solutions. The ability of cheatgrass ( Bromus tectorum ) to enhance fire, the ability of Chinese tallow ( Triadica sebifera ) to suppress fire, and the ability of leafy spurge ( Euphorbia esula ) to resprout from repeated injury make these species exceptional threats to native diversity. Finally, the available information for many invasive species is very incomplete, partic- ularly with regard to how fire affects competitive interactions with the native community. There is much more we need to know if we are to consistently predict how invasive species will respond to fire and how burning can best be used to manage for natural diversity.

Conference Paper