Ecological Determinants of Distribution Decline and Risk of Extinction in Moths

Abstract:  For successful conservation of species it is important to identify traits that predispose species to the risk of extinction. By identifying such traits conservation efforts can be directed toward species that are most at risk of becoming threatened. We used data derived from the literature to determine ecological traits that affect distribution, distribution change, and the risk of extinction in Finnish noctuid moths (Lepidoptera, Noctuidae). The ecological traits we examined included body size, larval specificity, length of the flight period, and overwintering stage. In addition, in monophagous species we examined the effects of resource distribution. Larval specificity, length of the flight period, and the overwintering stage each had an independent effect on the risk of extinction when the effects of other traits were controlled by entering all traits into the same regression model. Not a single trait predicted the risk of extinction when analysis was conducted without controlling for the other traits. This discrepancy among the results suggests that a single trait may not be enough to allow prediction of the risk of extinction. Instead, it seems that for successful, predictive conservation science data on several ecological characteristics are needed.     

Can entropy maximization use functional traits to explain species abundances? A comprehensive evaluation

Entropy maximization (EM) is a method that can link functional traits and community composition by predicting relative abundances of each species in a community using limited trait information. We developed a complementary suite of tests to examine the strengths and limitations of EM and the community-aggregated traits (CATs; i.e., weighted averages) on which it depends that can be applied to virtually any plant community data set. We show that suites of CATs can be used to differentiate communities and that EM can address the classic problem of characterizing ecological niches by quantifying constraints (CATs) on complex trait relationships in local communities. EM outperformed null models and comparable regression models in communities with different levels of dominance, diversity, and trait similarity. EM predicted well the abundance of the dominant species that drive community-level traits; it typically identified rarer species as such, although it struggled to predict the abundances of the rarest species in some cases. Predictions were sensitive to choice of traits, were substantially improved by using informative priors based on null models, and were robust to variation in trait measurement due to intraspecific variability or measurement error. We demonstrate how similarity in species' traits confounds predictions and provide guidelines for applying EM.     

Beyond the ecological: biological invasions alter natural selection on a native plant species

Biological invasions can have strong ecological effects on native communities by altering ecosystem functions, species interactions, and community composition. Even though these ecological effects frequently impact the population dynamics and fitness of native species, the evolutionary consequences of biological invasions have received relatively little attention. Here, I show that invasions impose novel selective pressures on a native plant species. By experimentally manipulating community composition, I found that the exotic plant Medicago polymorpha and the exotic herbivore Hypera brunneipennis alter the strength and, in some instances, the direction of natural selection on the competitive ability and anti-herbivore defenses of the native plant Lotus wrangelianus. Furthermore, the community composition of exotics influenced which traits were favored. For example, high densities of the exotic herbivore Hypera selected for increased resistance to herbivores in the native Lotus; however, when Medicago also was present, selection on this defense was eliminated. In contrast, selection on tolerance, another plant defense trait, was highest when both Hypera and Medicago were present at high densities. Thus, multiple exotic species may interact to influence the evolutionary trajectories of native plant populations, and patterns of selection may change as additional exotic species invade the community.     

A trait specific model of competition in a spatially structured plant community

The non-random dispersal of plant propagules is thought to counter competitive exclusion and thus promote the survival of competitively inferior species. We investigated this process by modelling the outcome of interactions between species with competitive ability defined as a function of both life-history traits and the environment with both random and clustered dispersal strategies and in environmentally homogeneous and heterogeneous environments. Four main results emerged: (1) environmental heterogeneity was seen to promote co-existence in conjunction with associated trait variation for tolerance to the environmental variable and where this trait variation was effectively limited by ‘trade-off’ such that no single species had an overall competitive advantage, (2) consistent with theory, random dispersal appeared to enhance the likelihood of competitive exclusion, whereas clustering favoured co-existence, (3) the ecological outcome of interactions between dispersal and competitive relationships varied as a function of the trait determining the competitive advantage within a particular environment, and (4) promotion of co-existence by clustered dispersal was most marked when associated with environmental heterogeneity. It is argued that these results suggest that current ecological models of species interactions may need to be modified to incorporate a more realistic understanding of competitive ability if we are to better understand the factors effecting species co-existence.     

Patterns and biases of climate change threats in the IUCN Red List

International Union for Conservation of Nature (IUCN) Red List assessments rely on published data and expert inputs, and biases can be introduced where underlying definitions and concepts are ambiguous. Consideration of climate change threat is no exception, and recently numerous approaches to assessing the threat of climate change to species have been developed. We explored IUCN Red List assessments of amphibians and birds to determine whether species listed as threatened by climate change display distinct patterns in terms of habitat occupied and additional nonclimatic threats faced. We compared IUCN Red List data with a published data set of species’ biological and ecological traits believed to infer high vulnerability to climate change and determined whether distributions of climate change‐threatened species on the IUCN Red List concur with those of climate change‐threatened species identified with the trait‐based approach and whether species possessing these traits are more likely to have climate change listed as a threat on the IUCN Red List. Species in some ecosystems (e.g., grassland, shrubland) and subject to particular threats (e.g., invasive species) were more likely to have climate change as a listed threat. Geographical patterns of climate change‐threatened amphibians and birds on the IUCN Red List were incongruent with patterns of global species richness and patterns identified using trait‐based approaches. Certain traits were linked to increases or decreases in the likelihood of a species being threatened by climate change. Broad temperature tolerance of a species was consistently related to an increased likelihood of climate change threat, indicating counterintuitive relationships in IUCN assessments. To improve the robustness of species assessments of the vulnerability or extinction risk associated with climate change, we suggest IUCN adopt a more cohesive approach whereby specific traits highlighted by our results are considered in Red List assessments. To achieve this and to strengthen the climate change‐vulnerability assessments approach, it is necessary to identify and implement logical avenues for further research into traits that make species vulnerable to climate change (including population‐level threats).     

Disturbance and resource availability act differently on the same suite of plant traits: revisiting assembly hypotheses

Understanding the mechanisms of trait selection at the scale of plant communities is a crucial step toward predicting community assembly. Although it is commonly assumed that disturbance and resource availability constrain separate suites of traits, representing the regenerative and established phases, respectively, a quantification and test of this accepted hypothesis is still lacking due to limitations of traditional statistical techniques. In this paper we quantify, using structural equation modeling (SEM), the relative contributions of disturbance and resource availability to the selection of suites of traits at the community scale. Our model specifies and reflects previously obtained ecological insights, taking disturbance and nutrient availability as central drivers affecting leaf, allometric, seed, and phenology traits in 156 (semi-) natural plant communities throughout The Netherlands. The common hypothesis positing that disturbance and resource availability each affect a set of mutually independent traits was not consistent with the data. Instead, our final model shows that most traits are strongly affected by both drivers. In addition, trait-trait constraints are more important in community assembly than environmental drivers in half of the cases. Both aspects of trait selection are crucial for correctly predicting ecosystem processes and community assembly, and they provide new insights into hitherto underappreciated ecological interactions.     

A structural equation model to integrate changes in functional strategies during old-field succession

From a functional perspective, changes in abundance, and ultimately species replacement, during succession are a consequence of integrated suites of traits conferring different relative ecological advantages as the environment changes over time. Here we use structural equations to model the interspecific relationships between these integrated functional traits using 34 herbaceous species from a Mediterranean old-field succession and thus quantify the notion of a plant strategy. We measured plant traits related to plant vegetative and reproductive size, leaf functioning, reproductive phenology, seed mass, and production on 15 individuals per species monitored during one growing season. The resulting structural equation model successfully accounts for the pattern of trait covariation during the first 45 years post-abandonment using just two forcing variables: time since site abandonment and seed mass; no association between time since field abandonment and seed mass was observed over these herbaceous stages of secondary succession. All other predicted traits values are determined by these two variables and the cause-effect linkage between them. Adding pre-reproductive vegetative mass as a third forcing variable noticeably increased the predictive power of the model. Increasing the time after abandonment favors species with increasing life span and pre-reproductive biomass and decreasing specific leaf area. Allometric coefficients relating vegetative and reproductive components of plant size were in accordance with allometry theory. The model confirmed the trade-off between seed mass and seed number. Maximum plant height and seed mass were major determinants of reproductive phenology. Our results show that beyond verbal conceptualization, plant ecological strategies can be quantified and modeled.     

Fire reduces morphospace occupation in plant communities

The two main assembly processes claimed to structure plant communities are habitat filtering and competitive interactions. The set of species growing in fire-prone communities has been filtered in such a way that species without fire-persistence traits have not successfully entered the community. Because plant traits are evolutionarily conserved and fire traits are correlated with other plant traits, communities under high fire frequency should not include all possible trait combinations, and thus the morphospace occupation by species in these communities should be lower than expected by chance (underoccupied). In contrast, communities under low fire frequency would lack the filtering factor, and thus their underoccupation of the morphospace is not expected. We test this prediction by comparing the morphospace occupation by species in communities located in the western Mediterranean Basin, five of them subject to high fire frequency (HiFi) and four to low fire frequency (LowFi). We first compile a set of morphological and functional traits for the species growing on the nine sites, then we compute the morphospace occupation of each site as a convex hull volume, and finally, to assert that our results are not a product of a random branching pattern of evolution, we simulate our traits under a null model of neutral evolution and compare the morphospace occupation of the simulated traits with the results from the empirical data. The results suggest that, as predicted, there is a clear differential morphospace occupation between communities under different fire regimes in such a way that the morphospace is underoccupied in HiFi communities only. The simulation of a neutral evolutionary model does not replicate the observed pattern of differential morphospace occupation, and thus it should be attributed to assembly processes. In conclusion, our results suggest that fire is a strong community assembling process, filtering the species that have fire-persistent traits and thus assembling phenotypically and phylogenetically clustered communities with vacant zones in the morphospace.     

Multivariate identification of plant functional response and effect traits in an agricultural landscape

Plant functional traits have been proposed as a linkage between the environmental control of vegetation and ecosystem function. Identification of traits that mediate the response of plant species to the environment is well established, but the identification of effect traits and the linkage between the two sets is less developed. This was attempted for a study of eight contrasting land uses in a marginal agricultural landscape where data on vegetation, management controls of the disturbance regime, and soil characteristics, including nitrogen release, were measured simultaneously with measures of ecosystem function such as litter decomposition rates and primary productivity on 30 sites. Trait data were assembled from databases, and an iterative multivariate approach using the three table (species, trait, environment) method RLQ was employed to identify a parsimonious set of traits that predict plant species responses to the environment and a parsimonious set of traits that link vegetation to ecosystem function. The lists of response and effect traits were similar, and where differences were observed, traits were usually highly correlated with at least one trait in the other list. This approach identified a small number of traits (canopy height, leaf dry matter content, leaf size, and specific leaf area) that provide a means of linking vegetation responses to environmental change with changes in ecosystem function. Other response traits included vegetative spread strategy, start of flowering, and seed terminal velocity, but within the system studied these traits were all significantly correlated to the traits shared between the response and effect lists.     

Butterfly community shifts over two centuries

Environmental changes strongly impact the distribution of species and subsequently the composition of species assemblages. Although most community ecology studies represent temporal snap shots, long‐term observations are rather rare. However, only such time series allow the identification of species composition shifts over several decades or even centuries. We analyzed changes in the species composition of a southeastern German butterfly and burnet moth community over nearly 2 centuries (1840–2013). We classified all species observed over this period according to their ecological tolerance, thereby assessing their degree of habitat specialisation. This classification was based on traits of the butterfly and burnet moth species and on their larval host plants. We collected data on temperature and precipitation for our study area over the same period. The number of species declined substantially from 1840 (117 species) to 2013 (71 species). The proportion of habitat specialists decreased, and most of these are currently endangered. In contrast, the proportion of habitat generalists increased. Species with restricted dispersal behavior and species in need of areas poor in soil nutrients had severe losses. Furthermore, our data indicated a decrease in species composition similarity between different decades over time. These data on species composition changes and the general trends of modifications may reflect effects from climate change and atmospheric nitrogen loads, as indicated by the ecological characteristics of host plant species and local changes in habitat configuration with increasing fragmentation. Our observation of major declines over time of currently threatened and protected species shows the importance of efficient conservation strategies.     

基于数据库的植物功能性状研究现状文献计量学分析

植物功能性状数据库是功能生态学研究的重要工具.了解基于植物功能性状数据库的植物功能性状研究现状,对探讨功能多样性和生物多样性具有重要的指导意义.介绍植物功能性状数据库的类型、影响力的评价方法和使用方法,并整理了17个植物功能性状库(附表).同时,通过Web of Science检索了2008-2018年间基于植物功能性状数据库开展植物功能性状研究的论文,从刊文数量、期刊分布、研究机构、主要作者、研究热点等方面进行分析.结果表明:基于数据库开展植物功能性状研究的文章数量和被引次数呈快速增长的趋势.当前此方向的研究主要以少数学者、机构为核心,大量论文发表在少数期刊上.研究热点主要体现在群落生态学、气候变化和性状进化、恢复和生存策略等方面.未来应加强研究者、研究机构之间的合作,数据共享,建立全球型的数据完善的植物功能性状数据库,同时加强数据标准化处理,解决地下性状和生理性状等难以获取及量化的问题,完善化石记录的植物性状和植物不同生长阶段的植物功能性状数据.(图5表4参60附表1)     

Sensitivity of plant-pollinator-herbivore communities to changes in phenology

For many species in seasonal environments, warmer springs associated with anthropogenic climate change are causing phenological changes. Within ecological communities, the timing of interactions among species may be altered if the species experience asymmetrical phenological shifts. We present a model that examines the consequences of changes in the relative timing of herbivory and pollination in a community of herbivores and pollinators that share a host plant. Our model suggests that phenological shifts can alter the abundances of these species and, in some cases, their population dynamics. If historical patterns of interactions in a community change and herbivores become active before pollinators, the community could see a reduction in pollinators and an increase in herbivores, while if pollinators become active before herbivores, there could be a loss of stable coexistence. Previous studies have warned of the potential for climate change to cause large phenological mismatches whereby species that depend on one another become so separated in time that they can no longer interact. Our results suggest that climate change-induced phenological shifts can have a major impact on communities even in cases where complete phenological mismatches do not occur.     

Ecological Correlates of Extinction Proneness in Tropical Butterflies

Abstract:  Widespread and rapid losses of natural habitats and biodiversity have made the identification of extinction-prone species a major challenge in conservation biology. We assessed the relative importance of biologically relevant species traits (e.g., body size, ecological specialization) obtained from published records to determine the extinction probability of butterflies in a highly disturbed tropical landscape (i.e., Singapore). We also developed a taxon-specific model to estimate the extinction proneness of butterflies in Southeast Asia. Logistic regression analyses showed that adult habitat specialization, larval host plant specificity, geographical distribution, sexual dichromatism, and congenor density were significant and independent determinants of butterfly extinctions in Singapore. Among these traits, specificity of larval host plant and adult habitat specialization were the best correlates of extinction risks. We used this phenomenological extinction-regression model to estimate the relative extinction proneness of 416 butterfly species in Southeast Asia. Our results illustrate the utility of available taxon-specific data for a localized area in estimating the extinction proneness of closely related species on a regional scale. When intensive field studies are not forthcoming, especially in regions suffering from rapid biodiversity losses (e.g., Southeast Asia), similar approaches could be used to estimate extinction threats for other taxonomic groups.     

Trait synergisms and the rarity, extirpation, and extinction risk of desert fishes

Understanding the causes and consequences of species extinctions is a central goal in ecology. Faced with the difficult task of identifying those species with the greatest need for conservation, ecologists have turned to using predictive suites of ecological and life-history traits to provide reasonable estimates of species extinction risk. Previous studies have linked individual traits to extinction risk, yet the nonadditive contribution of multiple traits to the entire extinction process, from species rarity to local extirpation to global extinction, has not been examined. This study asks whether trait synergisms predispose native fishes of the Lower Colorado River Basin (USA) to risk of extinction through their effects on rarity and local extirpation and their vulnerability to different sources of threat. Fish species with "slow" life histories (e.g., large body size, long life, and delayed maturity), minimal parental care to offspring, and specialized feeding behaviors are associated with smaller geographic distribution, greater frequency of local extirpation, and higher perceived extinction risk than that expected by simple additive effects of traits in combination. This supports the notion that trait synergisms increase the susceptibility of native fishes to multiple stages of the extinction process, thus making them prone to the multiple jeopardies resulting from a combination of fewer individuals, narrow environmental tolerances, and long recovery times following environmental change. Given that particular traits, some acting in concert, may differentially predispose native fishes to rarity, extirpation, and extinction, we suggest that management efforts in the Lower Colorado River Basin should be congruent with the life-history requirements of multiple species over large spatial and temporal scales.     

Species' traits predict phenological responses to climate change in butterflies

How do species' traits help identify which species will respond most strongly to future climate change? We examine the relationship between species' traits and phenology in a well-established model system for climate change, the U.K. Butterfly Monitoring Scheme (UKBMS). Most resident U.K. butterfly species have significantly advanced their dates of first appearance during the past 30 years. We show that species with narrower larval diet breadth and more advanced overwintering stages have experienced relatively greater advances in their date of first appearance. In addition, species with smaller range sizes have experienced greater phenological advancement. Our results demonstrate that species' traits can be important predictors of responses to climate change, and they suggest that further investigation of the mechanisms by which these traits influence phenology may aid in understanding species' responses to current and future climate change.     

Life-history strategies of plant pathogens: distribution patterns and phylogenetic analysis

Here we analyze herbarium records, surveys, and studies of fungal plant pathogens in tropical natural systems in order to establish a framework to study plant-pathogen interactions from a life-history perspective. We looked at how life-history traits of pathogens and their host plants affect the distribution of pathogens in different tropical habitats, and the importance of host phylogeny in determining the habitat associations of obligate fungal pathogens. Our study reveals that plant-pathogen interactions are prevalent and widespread in the tropics. Phylogenetic analysis of the distribution of obligate pathogens among plant families did not suggest a strong overall pattern of higher-level host phylogeny in pathogen host range, except that smut pathogens are particularly dominant on Poaceae and Asteraceae, and rusts are most common on Fabaceae, families that dominate disturbed areas in the tropics.     

Identifying robust strategies for assisted migration in a competitive stochastic metacommunity

Assisted migration (AM) is the translocation of species beyond their historical range to locations that are expected to be more suitable under future climate change. However, a relocated population may fail to establish in its donor community if there is high uncertainty in decision-making, climate, and interactions with the recipient ecological community. To quantify the benefit to persistence and risk of establishment failure of AM under different management scenarios (e.g., choosing target species, proportion of population to relocate, and optimal location to relocate), we built a stochastic metacommunity model to simulate several species reproducing, dispersing, and competing on a temperature gradient as temperature increases over time. Without AM, the species were vulnerable to climate change when they had low population sizes, short dispersal, and strong poleward competition. When relocating species that exemplified these traits, AM increased the long-term persistence of the species most when relocating a fraction of the donor population, even if the remaining population was very small or rapidly declining. This suggests that leaving behind a fraction of the population could be a robust approach, allowing managers to repeat AM in case they move the species to the wrong place and at the wrong time, especially when it is difficult to identify a species’ optimal climate. We found that AM most benefitted species with low dispersal ability and least benefited species with narrow thermal tolerances, for which AM increased extinction risk on average. Although relocation did not affect the persistence of nontarget species in our simple competitive model, researchers will need to consider a more complete set of community interactions to comprehensively understand invasion potential.     

Assessing the components of adaptive capacity to improve conservation and management efforts under global change

Natural‐resource managers and other conservation practitioners are under unprecedented pressure to categorize and quantify the vulnerability of natural systems based on assessment of the exposure, sensitivity, and adaptive capacity of species to climate change. Despite the urgent need for these assessments, neither the theoretical basis of adaptive capacity nor the practical issues underlying its quantification has been articulated in a manner that is directly applicable to natural‐resource management. Both are critical for researchers, managers, and other conservation practitioners to develop reliable strategies for assessing adaptive capacity. Drawing from principles of classical and contemporary research and examples from terrestrial, marine, plant, and animal systems, we examined broadly the theory behind the concept of adaptive capacity. We then considered how interdisciplinary, trait‐ and triage‐based approaches encompassing the oft‐overlooked interactions among components of adaptive capacity can be used to identify species and populations likely to have higher (or lower) adaptive capacity. We identified the challenges and value of such endeavors and argue for a concerted interdisciplinary research approach that combines ecology, ecological genetics, and eco‐physiology to reflect the interacting components of adaptive capacity. We aimed to provide a basis for constructive discussion between natural‐resource managers and researchers, discussions urgently needed to identify research directions that will deliver answers to real‐world questions facing resource managers, other conservation practitioners, and policy makers. Directing research to both seek general patterns and identify ways to facilitate adaptive capacity of key species and populations within species, will enable conservation ecologists and resource managers to maximize returns on research and management investment and arrive at novel and dynamic management and policy decisions.     

Importance of species traits for species distribution in fragmented landscapes

Knowledge of the relationship between species traits and species distribution in fragmented landscapes is important for understanding current distribution patterns and as background information for predictive models of the effect of future landscape changes. The existing studies on the topic suffer from several drawbacks. First, they usually consider only traits related to dispersal ability and not growth. Furthermore, they do not apply phylogenetic corrections, and we thus do not know how considerations of phylogenetic relationships can alter the conclusions. Finally, they usually apply only one technique to calculate habitat isolation, and we do not know how other isolation measures would change the results. We studied the issues using 30 species forming congeneric pairs occurring in fragmented dry grasslands. We measured traits related to dispersal, survival, and growth in the species and recorded distribution of the species in 215 grassland fragments. We show many strong relationships between species traits related to both dispersal and growth and species distribution in the landscape, such as the positive relationship between habitat occupancy and anemochory and negative relationships between habitat occupancy and seed dormancy. The directions of these relationships, however, often change after application of phylogenetic correction. For example, more isolated habitats host species with smaller seeds. After phylogenetic correction, however, they turn out to host species with larger seeds. The conclusions also partly change depending on how we calculate habitat isolation. Specifically, habitat isolation calculated from occupied habitats only has the highest predictive power. This indicates slow dynamics of the species. All the results support the expectation that species traits have a high potential to explain patterns of species distribution in the landscape and that they can be used to build predictive models of species distribution. The specific conclusions are, however, dependent on the technique used, and we should carefully consider this when comparing among different studies. Since different techniques answer slightly different questions, we should attempt to use analyses both with and without phylogenetic correction and explore different isolation measures whenever possible and compare the results.