Using avatar species to model the potential distribution of emerging invaders

Aim Anticipating the potential distributions of emerging invasive species is complicated by the tendency for species distribution models to perform better when both native and invasive range data are available for model development. If invasive range data are lacking, species models are liable to under‐estimate distributions for emerging invaders, particularly for species that are not at equilibrium with their native range environment due to historical factors, dispersal limitation and/or ecological interactions. We demonstrate the potential to use well‐quantified niche shifts from established ‘avatar’ (i.e. the remote or virtual manifestation of an entity) invaders to develop plausible distributions for data‐poor emerging invaders contingent on niche shifts of similar magnitude or character. Location Global. Methods Using the globally invasive crayfishes Pacifastacus leniusculus and Procambarus clarkii as our avatar invaders, we quantify how niche position, size and structure differs between native and total ranges using Mahalanobis distance (a measure of multivariate similarity) and the climate predictors of annual minimum and maximum air temperature. We then generalize patterns of niche shift from these species to the emerging crayfish invader Cherax quadricarinatus. Results Some patterns of niche shifts were similar for Pacifastacus leniusculus and Procambarus clarkii, but niche shifts were of considerably greater magnitude for P. clarkii. When a native range model for C. quadricarinatus was modified with generalized niche shifts similar to Pacifastacus leniusculus and Procambarus clarkii, the potential global distribution for this species increased considerably, including many areas not identified by the native range model. Main conclusions We illustrate the potential to use avatar invaders to provide cautionary, niche shift‐assuming species distribution models for emerging invaders. Many theoretical and applied implications of the avatar species concept require additional investigation, including the development of frameworks to select appropriate avatar species and evaluate the performance of avatar‐derived models for emerging invaders. Despite these research needs, we believe this concept will have considerable utility for predicting vulnerability to invasion by data‐poor species; this is a critical management need because shifting pathways of introduction and climate change will produce many novel, emerging invasive species in the future.     

Predicting worldwide invasiveness for four major problematic decapods: an evaluation of using different calibration sets

Recently, there has been much debate whether niche based models (NBM) can predict biological invasions into new areas. These studies have chiefly focused on the type of occurrence data to use for model calibration. Additionally, pseudo‐absences are also known to cause uncertainty in NBM, but are rarely tested for predicting invasiveness. Here we test the implications of using different calibration sets for building worldwide invasiveness models for four major problematic decapods: Cherax destructor, Eriocheir sinensis, Pacifastacus leniusculus and Procambarus clarkii. Using Artificial Neural Networks models we compared predictions containing either native range occurrences (NRO), native and invasive occurrences (NIO) and invasive only (IRO) coupled with three types of pseudo‐absences – based on sampling only 1) the native range (NRA), 2) native and invasive ranges (NIA), and 3) worldwide random (WRA). We further analysed the potential gains in accuracy obtained through averaging across multiple models. Our results showed that NRO and IRO provided the best predictions for native and invaded ranges, respectively. Still, NIO provided the best balance in predicting both ranges. Pseudo‐absences had a large influence on the predictive performance of the models, and were more important for predictiveness than types of occurrences. Specifically, WRA performed the best and NRA and NIA performed poorly. We also found little benefit in combining predictions since best performing single‐models showed consistently higher accuracies. We conclude that NBM can provide useful information in forecasting invasiveness but are largely dependent on the type of initial information used and more efforts should be placed on recognizing its implications. Our results also show extensive areas which are highly suitable for the studied species worldwide. In total these areas reach from three to nine times the species current ranges and large portions of them are contiguous with currently invasive populations.     

Invasion of the shelter snatchers: behavioural plasticity in invasive red swamp crayfish,Procambarus clarkii

1. In many freshwater systems, competition for shelter plays an important role in determining the persistence of both native and alien species. The red swamp crayfish, Procambarus clarkii, is currently invading the native habitat of the signal crayfish, Pacifastacus leniusculus, in southern Oregon, and interspecific competition for shelter may be driving the species replacement in this region. 2. We designed a 2 × 3 factorial mesocosm experiment, with shelter density and species combination as factors, to investigate shelter occupancy and resource competition. Contrary to our predictions, the two crayfish species are equal competitors for shelter. Further, the invasive P. clarkii modified its shelter occupancy behaviour in the presence of the native P. leniusculus and has broader microhabitat preferences. 3. Specifically, we found that P. clarkii alters shelter occupancy and space use patterns when the two species occurred together, such that shelter use was identical between P. clarkii and P. leniusculus in mixed‐species treatments. In such treatments, both species increased their use of shelters when shelter density increased. When P. clarkii was alone, however, individuals did not alter shelter use as a function of shelter density, whereas P. leniusculus exhibited similar density‐dependent behaviour in both mixed‐ and single‐species treatments. 4. In a complementary field survey, we employed an ‘epicentre‐based’ design to sample two field sites. We observed patterns of microhabitat use and breadth for each species similar to those in our mesocosm experiment: the invasive P. clarkii was more abundant across different habitats and used a broader range of microhabitats than the native P. leniusculus. As such, we found that P. clarkii was more abundant across both field sites than the P. leniusculus, occupying microhabitats within and beyond the preferred range of P. leniusculus. Both field sites were affected by urban development and agriculture. 5. The use of microhabitats by both species was similar in the laboratory and the field. This study confirms that P. clarkii individuals can, and do, successfully occupy microhabitats preferred by P. leniusculus in the Willamette Valley. The results from our study may be relevant to other freshwater systems inhabited by P. clarkii and contribute to the understanding of ‘niche opportunity’, a concept which defines the environmental conditions that promote biological invasions.     

Environmental cues during overland dispersal by three freshwater invaders: Eriocheir sinensis,Pacifastacus leniusculus,and Procambarus clarkii (Crustacea,Decapoda)

Freshwater crustaceans are known to disperse overland, aiding the invasion of new environments. To analyze the effect of environmental variables (humidity, temperature, illuminance, vegetation, and slope) on the movement direction of Eriocheir sinensis (H. Milne Edwards, 1853), Pacifastacus leniusculus (Dana, 1852) and Procambarus clarkii (Girard, 1852) in land, the animals were placed at the centre of a metal gutter and given a choice between two values of each variable. Chi-squared tests revealed significant preferences for E. sinensis to move downhill, P. leniusculus to move toward cooler areas and P. clarkii to move downhill and toward cooler areas. To test the ability of P. leniusculus and P. clarkii (E. sinensis was not present at the testing site) to return to water, they were placed in a circular arena near the river bank and their direction of movement was recorded. Chi-squared tests revealed that P. leniusculus, when isolated and in groups of 6, showed a significant tendency to move downstream, but not directly toward the water. P. clarkii showed no significant preference in movement direction. Since movement direction and overland dispersal are influenced by environmental gradients of temperature and elevation, these variables are fundamental for predicting autonomous overland dispersal and colonization patterns.     

Impacts of climate change on the global potential distribution of two notorious invasive crayfishes

1. Invasive alien species and climate change are two of the most serious global environmental threats. In particular, it is of great interest to understand how changing climates could impact the distribution of invaders that pose serious threats to ecosystems and human activities.
2. In this study, we developed ensemble species distribution models for predicting the current and future global distribution of the signal crayfish Pacifastacus leniusculus and the red swamp crayfish Procambarus clarkii, two of the most highly problematic invaders of freshwater ecosystems worldwide. We collected occurrence records of the species, from native and alien established ranges worldwide. These records in combination with averaged observations of current climatic conditions were used to calibrate a set of 10 distinct correlative models for estimating the climatic niche of each species. We next projected the estimated niches into the geographical space for the current climate conditions and for the 2050s and 2070s under representative concentration pathway 2.6 and 8.5 scenarios.
3. Our species distribution models had high predictive abilities and suggest that annual mean temperature is the main driver of the distribution of both species. Model predictions indicated that the two crayfish species have not fully occupied their suitable climates and will respond differently to future climate scenarios in different geographic regions. Suitable climate for P. leniusculus was predicted to shift poleward and to increase in extent in North America and Europe but decrease in Asia. Regions with suitable climate for P. clarkii are predicted to widen in Europe but contract in North America and Asia.
4. This study highlights that invasive species with different thermal preference are likely to respond differently to future climate changes. Our results provide important information for policy makers to design and implement anticipated measures for the prevention and control of these two problematic species.

     

Comparison of trophic function between the globally invasive crayfishes <Emphasis Type="Italic">Pacifastacus leniusculus</Emphasis> and <Emphasis Type="Italic">Procambarus clarkii</Emphasis>

Impacts of invasive species may manifest most strongly if these organisms are highly distinct functionally from the native species they often replace. Yet, should we expect functional differences between native and invasive species of generalist organisms like freshwater crayfish? Some existing evidence has pointed to native and invasive crayfish species as ecologically equivalent. We contribute to this literature by comparing the trophic niches of the globally invasive crayfishes Pacifastacus leniusculus and Procambarus clarkii, by applying carbon and nitrogen stable isotope analyses to replicated allopatric (alone) and sympatric (together) lake populations in western Washington State, USA, where P. clarkii has been recently introduced and P. leniusculus is presumed native. Our study corrected for potential inherent differences in lake food webs as a consequence of lake abiotic or biotic characteristics using random effects in linear mixed effects models. We found that although overall trophic niche size or area of these species was not significantly different, P. leniusculus was significantly higher in trophic position than P. clarkii when also accounting for the effects of body size, sex, and lakes as random effects. This pattern of increased trophic position of P. leniusculus over P. clarkii was conserved over time in one sympatric lake for which we had data over multiple years. Cumulatively, our findings point to trophic differences between the globally cosmopolitan crayfishes P. leniusculus and P. clarkii, particularly when accounting for the ways that ecosystem context can affect food web structure of communities and the trophic resources available to these consumers.     

Geostatistical distribution modelling of two invasive crayfish across dendritic stream networks

Species distribution models combining environmental and spatial components are increasingly used to understand and forecast species invasions. However, modelling distributions of invasive species inhabiting stream networks requires due consideration of their dendritic spatial structure, which may strongly constrain dispersal and colonization pathways. Here we evaluate the application of novel geostatistical tools to species distribution modelling in dendritic networks, using as case study two invasive crayfish (Procambarus clarkii and Pacifastacus leniusculus) in a Mediterranean watershed. Specifically, we used logistic mixed models to relate the probability of occurrence of each crayfish to environmental variables, while specifying three spatial autocorrelation components in random errors. These components described spatial dependencies between sites as a function of (1) straight-line distances (Euclidean model) between sites, (2) hydrologic (along the waterlines) distances between flow-connected sites (tail-up model), and (3) hydrologic distances irrespective of flow connection (tail-down model). We found a positive effect of stream order on P. clarkii, indicating an association with the lower and mid reaches of larger streams, while P. leniusculus was affected by an interaction between stream order and elevation, indicating an association with larger streams at higher altitude. For both species, models including environmental and spatial components far outperformed the pure environmental models, with the tail-up and the Euclidean components being the most important for P. clarkii and P. leniusculus, respectively. Overall, our study highlighted the value of geostatistical tools to model the distribution of riverine and aquatic invasive species, and stress the need to specify spatial dependencies representing the dendritic network structure of stream ecosystems.     

Understanding niche shifts: using current and historical data to model the invasive redlegged earth mite,Halotydeus destructor

Aim Niche conservatism is key to understanding species responses to environmental stress such as climate change or arriving in new geographical space such as biological invasion. Halotydeus destructor is an important agricultural pest in Australia and has been the focus of extensive surveys that suggest this species has undergone a niche shift to expand its invasive range inland to hotter and drier environments. We employ modern correlative modelling methods to examine niche conservatism in H. destructor and highlight ecological differences between historical and current distributions. Location Australia and South Africa. Methods We compile comprehensive distribution data sets for H. destructor, representing the native range in South Africa, its invasive range in Australia in the 1960s (40 yr post‐introduction) and its current range in Australia. Using MAXENT, we build correlative models and reciprocally project them between South Africa and Australia and investigate range expansion with models constructed for historical and current data sets. We use several recently developed model exploration tools to examine the climate similarity between native and invasive ranges and subsequently examine climatic variables that limit distributions. Results The invasive niche of H. destructor in Australia transgresses the native niche in South Africa, and the species has expanded in Australia beyond what is predicted from the native distribution. Our models support the notion that H. destructor has undergone a more recent range shift into hotter and drier inland areas of Australia since establishing a stable distribution in the 1960s. Main conclusions Our use of historical and current data highlights that invasion is an ongoing dynamic process and demonstrates that once a species has reached an established range, it may still expand at a later stage. We also show that model exploration tools help understand factors influencing the range of invasive species. The models generate hypotheses about adaptive shifts in H. destructor.     

Home-field advantage: native signal crayfish (Pacifastacus leniusculus) out consume newly introduced crayfishes for invasive Chinese mystery snail (Bellamya chinensis)

The introduction of non-indigenous plants, animals and pathogens is one of today’s most pressing environmental challenges. Freshwater ecologists are challenged to predict the potential consequences of species invasions because many ecosystems increasingly support novel assemblages of native and non-native species that are likely to interact in complex ways. In this study we evaluated how native signal crayfish (Pacifastacus leniusculus) and non-native red swamp crayfish (Procambarus clarkii) and northern crayfish (Orconectes virilis) utilize a novel prey resource: the non-native Chinese mystery snail (Bellamya chinensis). All species are widespread in the United States, as well as globally, and recent surveys have discovered them co-occurring in lakes of Washington State. A series of mesocosm experiments revealed that crayfish are able to consume B. chinensis, despite the snail’s large size, thick outer shell and trapdoor defense behaviour. Crayfish exhibited size-selective predation whereby consumption levels decreased with increasing snail size; a common pattern among decapod predators. Comparison of prey profitability curves—defined as the yield of food (weight of snail tissue) per second of feeding time (the time taken to crack the shell and consume the contents)—suggests that small and very large snails may represent the most profitable prey choice. By contrast, previous studies have reported the opposite pattern for crayfish consumption on thin-shelled snails. For all snail size classes, we found that native P. leniusculus and invasive O. virilis consumed greater numbers of snails than invasive P. clarkii. Moreover, P. leniusculus consistently handled and consumed snails at a faster pace compared to both invasive crayfishes across the range of snail sizes examined in our study. These results suggest not only that B. chinensis is a suitable food source for crayfish, but also that native P. leniusculus may ultimately out-consume invasive crayfishes for this new prey resource.     

Spatial autocorrelation and the analysis of invasion processes from distribution data: a study with the crayfish <Emphasis Type="Italic">Procambarus clarkii</Emphasis>

Complex spatial dynamics are frequent in invasive species; analyzing distribution patterns can help to understand the mechanisms driving invasions. We used different spatial regression techniques to evaluate processes determining the invasion of the red swamp crayfish Procambarus clarkii. We evaluated four a priori hypotheses on processes that may determine crayfish invasion: landscape alteration, connectivity, wetland suitability for abiotic and biotic features. We assessed the distribution of P. clarkii in 119 waterbodies in a recently invaded area. We used spatially explicit statistical techniques (spatial eigenvector mapping, generalized additive models, Bayesian intrinsic conditional autoregressive models) within an information-theoretic framework to assess the support of hypotheses; we also analyzed the pattern of spatial autocorrelation of data, model residuals, and eigenvectors. We found strong agreement between the results of spatial eigenvector mapping and Bayesian autoregressive models. Procambarus clarkii was significantly associated with the largest, permanent wetlands. Additive models suggested also association with human-dominated landscapes, but tended to overfit data. The results indicate that abiotic wetlands features and landscape alteration are major drivers of the species’ distribution. Species distribution data, residuals of ordinary least squares regression, and spatial eigenvectors all showed positive and significant spatial autocorrelation at distances up to 2,500 m; this may be caused by the dispersal ability of the species. Our analyses help to understand the processes determining the invasion and to identify the areas most at risk where screening and early management efforts can be focused. The comparison of multiple spatial techniques allows a robust assessment of factors determining complex distribution patterns.     

Early assessment of the impact of alien species: differential consequences of an invasive crayfish on adult and larval amphibians

Aim Early assessment of the impact of invasive alien species is crucial to set up timely management, but often the impact is evident when it is too late for action. We evaluated relationships between the alien crayfish, Procambarus clarkii, the distribution of native amphibians, and the abundance of their larvae. We assessed whether considering measures of reproductive success provide a more prompt measure of impact than considering just species distribution. Location One hundred and twenty‐five wetlands in Northern Italy, in an area recently invaded by P. clarkii. Methods We surveyed wetlands to assess the presence of breeding activity of amphibians and the distribution of P. clarkii. We measured the abundance of amphibian larvae before metamorphosis through pipe sampling. We built models analysing the relationships between amphibian and crayfish distribution, while taking into account spatial autocorrelation and environmental features. Analyses were performed at both the species level (generalized linear models and spatial eigenvector mapping) and community level (constrained redundancy analysis). Results In terms of breeding site distribution, only two amphibians (Lissotriton vulgaris and Hyla intermedia) were negatively associated with P. clarkii, while the relationships between other the species and P. clarkii were positive or not significant. However, larval abundance for all amphibian species was negatively associated with the alien crayfish. Analyses performed at community and single species levels yielded consistent results. Main conclusions Procambarus clarkii impacts amphibians through different processes. Newts probably avoid invaded wetlands for breeding. Other species attempt breeding in wetlands with crayfish, but suffer very low success. Considering distribution data alone would not provide a correct picture of the impact of this alien species; measures of reproductive success may allow a more accurate assessment of the impact.     

Indicators of movement and space use for two co-occurring invasive crayfish species

Red swamp crayfish (Procambarus clarkii) and signal crayfish (Pacifastacus leniusculus) are two invasive freshwater species with a worldwide distribution. The objective of this work was to investigate how the two species move and use space in an area of recent coexistence. Simultaneously, we test the use of new tools and indices to describe their movement patterns. To accomplish this we performed a radio-tracking program within a river-type habitat during two different periods (September/October 2010 and June/July 2013). We used spatial analysis tools to map crayfish radio-location data with and without accounting for the curvature of the river. To assess the consistency of the direction of movement and of the distances traveled by crayfish, two indices were developed. To assess the habitat preferences of each species we applied Ivlev's Electivity Index and the Standardized Forage Ratio. Movement of P. clarkii and P. leniusculus differed. The average detected movement was 8.8 m day⁻¹ for P. clarkii and 17.5 m day⁻¹ for P. leniusculus. However, crayfish behavior ranged from almost complete immobility – sometimes during several days – to large movements, in half a day, up to a maximum of 255 m for P. clarkii and 461 m for P. leniusculus. The proportion of upstream or downstream movements was independent of the species and both species displayed no preference for either direction. The indices of consistency of movement showed a large interindividual variation. Species and period (2010 or 2013) affected the mean daily distance traveled, maximum observed distance from location of release and percentage of observations under vegetation cover. The Ivlev's Electivity Index and the Standardized Forage Ratio presented similar results. P. clarkii showed a preference for pool areas with riparian vegetation cover while P. leniusculus preferred riffle and pool areas with riparian vegetation cover. Our work provided new and valuable data for modeling the active dispersal of these two problematic invaders in a context of coexistence.     

Niche modelling to guide conservation actions in France for the endangered crayfish Austropotamobius pallipes in relation to the invasive Pacifastacus leniusculus

1. The white-clawed crayfish (Austropotamobius pallipes) is globally endangered due to the impacts of habitat modification and fragmentation, water pollution, climate change, and invasive species, particularly the signal crayfish (Pacifastacus leniusculus). These pressures have caused the decline of A. pallipes populations in Europe, demonstrating the importance of predicting the species' potential distribution under current and future conditions. Focusing on the watercourses of mainland France, we aimed to identify suitable areas for A. pallipes to guide the conservation of current populations and future introduction actions or protection measures.
2. We applied ecological niche modelling to model the potential distribution of both A. pallipes and P. leniusculus and identified locations suitable for A. pallipes only. We also assessed the potential distribution of the species under two representative concentration pathway (RCP) scenarios: RCP 2.6 and RCP 8.5, respectively describing low-warming and high-warming conditions.
3. We found that A. pallipes and P. leniusculus exploit equivalent niches in France. Despite this, under current conditions, about 5% of the study area simultaneously records a high suitability for A. pallipes and a low suitability for P. leniusculus and is therefore of significant conservation interest. This percentage remains relatively stable under RCP 2.6 for 2050 and 2100, but decreases to 2% under RCP 8.5 for 2100.
4. Ecological niche modelling can supply crucial guidance for conservation actions aimed at protecting endangered species at a national scale by identifying sites most suitable for protection and sites where climate change and invasive species constitute a threat.

     

Geographic expansion of the invasive red crayfish <Emphasis Type="Italic">Procambarus clarkii</Emphasis> (Girard, 1852) (Crustacea: Decapoda) in Mexico

The red crayfish Procambarus clarkii, which is native to southcentral USA and northeastern Mexico, has been successfully introduced into several countries around the world. This study documents the geographic expansion of the exotic red crayfish in Mexico and discusses the consequence of a greater propagation of this species in Mexican inland waters. New state records of this crayfish in the Baja California peninsula and in the states of Durango and Sinaloa indicate its progressive dispersion. The propagation of P. clarkii in Mexico has been caused mainly by human introduction, but it is also facilitated because of the species’ tolerance to an ample range of environmental conditions. Because of the invasive capability of P. clarkii, we suspect that this exotic species is competing for habitat and food with native freshwater shrimp of the genus Macrobrachium in many sites of northern Mexico.     

Niche breadth and trophic diversity: feeding behaviour of the red swamp crayfish (Procambarus clarkii) towards environmental availability of aquatic macroinvertebrates in a rice field (Portugal)

The red swamp crayfish, Procambarus clarkii (Girard 1852), an alien species in Portugal, may have dramatic effects on aquatic communities by depleting all food resources available, just after its introduction. The purpose of this study was to evaluate, through the concept of niche breadth, whether the diet of this species reflected the temporal changes of aquatic macroinvertebrate availability once it is acclimated. Petraitis’ index of niche breadth and Herrera’s trophic diversity index were used to evaluate the trophic behaviour of P. clarkii towards available resources over time and intraspecifically (size classes and sex) in a rice field in Portugal. Results from this study showed that the consumption of aquatic macroinvertebrates by P. clarkii reflected their seasonal availability. The high values of niche breadth and trophic diversity indicated resource use according to trophic availability, diversified diets and different individual exploitation of resources regardless of size or sex. These findings suggest that in habitats where P. clarkii is already acclimated, it adjusted its trophic behaviour to the seasonal availability of aquatic macroinvertebrates. The large niche breadth and high trophic diversity presented by P. clarkii enables it to successfully expand its range to new areas, when other environmental conditions are favourable, as has been observed in Portugal and worldwide.     

Niche conservatism and the potential for the crayfish Procambarus clarkii to invade South America

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Adult neurogenesis in the crayfish brain: Proliferation,migration, and possible origin of precursor cells

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