Characterization of microsatellite loci in the European green crab (Carcinus maenas)

Carcinus maenas (Decapoda: Portunidae) has proven a highly successful invasive marine species whose potential economic and ecological impacts are of great concern worldwide. Here, we characterize 14 polymorphic microsatellite loci in C. maenas and its sister species Carcinus aestuarii. These markers will prove useful for fine‐scale genetic analyses of native and introduced populations, for assessment of the sources and routes of invasion and for evaluation of post‐invasion population dynamics.     

Modelling the biological invasion of Carcinus maenas (the European green crab)

This paper proposes a system of integro-difference equations to model the spread of Carcinus maenas, commonly called the European green crab, that causes severe damage to coastal ecosystems. A model with juvenile and adult classes is first studied. Here, standard theory of monotone operators for integro-difference equations can be applied and yields explicit formulas for the asymptotic spreading speeds of the juvenile and adult crabs. A second model including an infected class is considered by introducing a castrating parasite Sacculina carcini as a biological control agent. The dynamics are complicated and simulations reveal the occurrence of periodic solutions and stacked fronts. In this case, only conjectures can be made for the asymptotic spreading speeds because of the lack of mathematical theory for non-monotone operators. This paper also emphasizes the need for mathematical studies of non-monotone operators in heterogeneous environments and the existence of stacked front solutions in biological invasion models.     

A global invader at home: population structure of the green crab, Carcinus maenas, in Europe

The European green crab, Carcinus maenas, has a native distribution that extends from Norway to Mauritania. It has attracted attention because of its recent invasions of Australia, Tasmania, South Africa, Japan and both coasts of North America. To examine the population structure of this global invader in its native range, we analysed a 502-base-pair fragment of the mitochondrial cytochrome c oxidase I (COI) gene from 217 crabs collected in the North Atlantic and 13 specimens from the Mediterranean. A clear genetic break (11% sequence divergence) occurs between the Mediterranean and Atlantic, supporting the species-level status of these two forms. Populations in the Faeroe Islands and Iceland were genetically distinct from continental populations (F(ST) = 0.264-0.678), with Iceland represented by a single lineage also found in the Faeroes. This break is consistent with a deep-water barrier to dispersal in green crabs. Although there are relatively high levels of gene flow along the Atlantic coast of Europe, slight population structure was found between the central North Sea and populations to the south. Analysis of variance, multidimensional scaling, and the distribution of private haplotypes support this break, located between Bremerhaven, Germany, and Hoek van Holland. Similar biogeographical and genetic associations for other species, such as benthic algae and freshwater eels, suggest that the marine fauna of Europe may be generally subdivided into the areas of Mediterranean, western Europe and northern Europe.     

European green crabs (Carcinus maenas) in the northeastern Pacific: genetic evidence for high population connectivity and current-mediated expansion from a single introduced source population

Aim The European green crab (Carcinus maenas) expanded dramatically after its introduction to the west coast of North America, spreading over 1000 km in < 10 years. We use samples of Carcinus maenas collected over time and space to investigate the genetic patterns underlying the species’ initial establishment and spread, and discuss our findings in the context of the species’ life history characteristics and demography. Location The central west coast of North America, encompassing California, Oregon, and Washington (USA) and British Columbia (Canada). Methods We collected 1040 total samples from 21 sites representing the major episodes of population establishment and expansion along the west coast of North America. Microsatellite markers were used to assess genetic diversity and structure at different time points in the species’ spread, to investigate connectivity between embayments and to estimate both short‐term effective population sizes and the number of original founders. Assignment testing was performed to determine the likely source of the introduction. Results Carcinus maenas in western North America likely derived from a single introduction of a small number of founders to San Francisco Bay, CA from the east coast of North America. Throughout its western North American range, the species experiences periodic migration between embayments, resulting in a minor loss of genetic diversity in more recently established populations versus the populations in the area of initial establishment. Main conclusions Low genetic diversity has not precluded the ability of C. maenas to successfully establish and spread on the west coast of North America. An efficient oceanographic transport mechanism combined with highly conducive life history traits are likely the major drivers of C. maenas spread. Evidence for a single introduction underscores the potential utility of early detection and eradication of high‐risk invasive species.     

Linking ocean conditions to year class strength of the invasive European green crab, Carcinus maenas

Once a non-native species arrives and survives in an area, its long-term persistence depends on its recruitment success. If conditions are not favorable for recruitment it will ultimately disappear. The European green crab (Carcinus maenas) has a 6 year life span and has persisted at low densities in Oregon and Washington coastal estuaries for the past 12 years. We show here that after the arrival of the strong founding year class of 1998, significant self-recruitment to the Oregon and Washington populations occurred only in 2003, 2005 and 2006. Warm winter water temperatures, high Pacific Decadal Oscillation and Multivariate ENSO (El Niño Southern Oscillation) Indices in March, late spring transitions and weak southward shelf currents in March and April are all correlated with the these strong year classes. Cold winter water temperatures, low Pacific Decadal Oscillation Indices, early spring transitions and strong southward (and offshore) currents in March and April are linked to year class failure. Right now, green crabs are still too rare to exert a measurable effect on the native benthic community and on shellfish culture in Oregon and Washington. However, if their numbers were to increase, we would be able to predict the arrival of strong year classes from ocean conditions and alert managers and shellfish growers of possible increases in predation pressure from this invader.     

Prediction and validation of the potential global distribution of a problematic alien invasive species — the American bullfrog

Predicting the probability of successful establishment and invasion of alien species at global scale, by matching climatic and land use data, is a priority for the risk assessment. Both large- and local-scale factors contribute to the outcome of invasions, and should be integrated to improve the predictions. At global scale, we used climatic and land use layers to evaluate the habitat suitability for the American bullfrog Rana catesbeiana , a major invasive species that is among the causes of amphibian decline. Environmental models were built by using Maxent, a machine learning method. Then, we integrated global data with information on richness of native communities and hunting pressure collected at the local scale. Global-scale data allowed us to delineate the areas with the highest suitability for this species. Predicted suitability was significantly related to the invasiveness observed for bullfrog populations historically introduced in Europe, but did not explain a large portion of variability in invasion success. The integration of data at the global and local scales greatly improved the performance of models, and explained > 57% of the variance in introduction success: bullfrogs were more invasive in areas with high suitability and low hunting pressure over frogs. Our study identified the climatic factors entailing the risk of invasion by bullfrogs, and stresses the importance of the integration of biotic and abiotic data collected at different spatial scales, to evaluate the areas where monitoring and management efforts need to be focused.     

提高生态位模型转移能力来模拟入侵物种 的潜在分布

生态位模型利用物种分布点所关联的环境变量去推算物种的生态需求, 模拟物种的分布。在模拟入侵物种分布时, 经典生态位模型包括模型构建于物种本土分布地, 然后将其转移并投射至另一地理区域, 来模拟入侵物种的潜在分布。然而在模型运用时, 出现了模型的转移能力较低、模拟的结果与物种的实际分布不相符的情况, 由此得出了生态位漂移等不恰当的结论。提高生态位模型的转移能力, 可以准确地模拟入侵物种的潜在分布, 为入侵种的风险评估提供参考。作者以入侵种茶翅蝽(Halyomorpha halys)和互花米草(Spartina alterniflora)为例, 从模型的构建材料(即物种分布点和环境变量)入手, 全面阐述提高模型转移能力的策略。在构建模型之前, 需要充分了解入侵物种的生物学特性、种群平衡状态、本土地理分布范围及物种的生物历史地理等方面的知识。在模型构建环节上, 物种分布点不仅要充分覆盖物种的地理分布和生态空间的范围, 同时要降低物种采样点偏差; 环境变量的选择要充分考虑其对物种分布的限制作用、各环境变量之间的空间相关性, 以及不同地理种群间生态空间是否一致, 同时要降低环境变量的空间维度; 模型构建区域要真实地反映物种的地理分布范围, 并考虑种群的平衡状态。作者认为, 在生态位保守的前提下, 如果模型是构建在一个合理方案的基础上, 生态位模型的转移能力是可以保证的, 在以模型转移能力较低的现象来阐述生态位分化时需要引起注意。     

Predicting spread of invasive macrophytes in New Zealand lakes using indirect measures of human accessibility

1. Models predicting invasive macrophyte spread between lakes provide an important tool for focusing proactive management efforts to lakes deemed susceptible to invasion. However, challenges to forecasting macrophyte spread include wide physiological tolerances of invasive macrophytes and a lack of information on the relative importance of the various human vectors (e.g. boating traffic). In New Zealand, three invasive species that reproduce vegetatively, Ceratophyllum demersum, Lagarosiphon major, Egeria densa, and a single species that reproduces sexually, Utricularia gibba, are currently spreading across the lake landscape at a great cost to the local ecology and economy. 2. In this study, we first examined whether variables that indirectly describe weed spread via human access and use, as well as a lake’s position in the landscape, could describe the distribution of these four species using a boosted regression trees (BRT) modelling approach. Then, as these invasive species have not reached their full invasion potential, we examined how giving more influence to infected lakes at the edge of the invasion front, and including all lakes across New Zealand as background samples, simulating ‘absences beyond the invasion front’, influenced our ability to forecast the potential for new lakes to be invaded. 3. The BRT models identified that variables characterising human access and use, as well as lake position, were associated with the occurrence of the three vegetatively reproducing macrophytes. Weed occurrence was more likely when there was a highway in the vicinity, human population density was high and if the lake was large (c. 55 km²). But in the single case of U. gibba, temperature was the variable that best explained occurrence. This is consistent with the suggestion that U. gibba is predominantly dispersed by waterbirds, rather than human activity. 4. But for all four species, the BRT models based on the recorded observations alone predicted observed invasions with low prediction probabilities and did not forecast further spread. By contrast, when observations at the edge of the invasion front were upweighted, and additional background lakes implemented into the model, recorded observations were predicted and additional lakes were forecast to be at risk, suggesting that these models better captured the current and potential distribution of these macrophyte species. 5. The use of variables that characterise weed spread could provide similar insights into other systems where survey information on the nature, strength and direction of invasion vectors is lacking. Furthermore, when weighting the data, many lakes across New Zealand were forecasted to be at risk of invasion. The advantage of weighing the presence data was that insights into the potential for a species to spread were obtained. The probabilistic estimates of risk, as derived from the models, together with other information for prioritising lakes, can be used to focus surveillance and protection efforts.     

Modelling spread of the invasive macrophyte Cabomba caroliniana

1. Predicting spread of non-indigenous species requires an understanding of where propagules are being transported, and whether these propagules can survive in the novel habitat and successfully integrate into the recipient community. In this study, we model potential spread of invading Cabomba caroliniana in Ontario, Canada, using a combination of passive and active dispersal models coupled with an environmental suitability model, thereby considering the first two stages of the invasion process.
2. Measures of propagule pressure incorporated both human-mediated dispersal via trailered boats, and advective flow from invaded to non-invaded systems, while habitat suitability was forecasted by combining native and global data sets and using boosted regression trees.
3. Risk of invasion differed depending on the combination of approaches used and the time period considered. Three lakes appear to be at greatest risk owing to a combination of high boater and water movement from invaded sources, and high environmental suitability. The best predictors of lake suitability were pH, mean lake temperature and dissolved calcium concentration. Hundreds of lakes in Ontario may be suitable for establishment of Cabomba , highlighting the need for vector management.     

Limited filling of the potential range in European tree species

The relative roles of environment and history in controlling large‐scale species distributions are important not only theoretically, but also for forecasting range responses to climatic change. Here, we use atlas data to examine the extent to which 55 tree species fill their climatically determined potential ranges in Europe. Quantifying range filling (R/P) as realized/potential range size ratios using bioclimatic envelope modelling we find mean R/P = 38.3% (±30.3% SD). Many European tree species naturalize extensively outside their native ranges, providing support for interpreting the many low R/Ps as primarily reflecting dispersal limitation. R/P increases strongly with latitudinal range centroid and secondarily with hardiness and decreases weakly with longitudinal range centroid. Hence, European tree species ranges appear strongly controlled by geographical dispersal constraints on post‐glacial expansion as well as climate. Consequently, we expect European tree species to show only limited tracking of near‐future climate changes.     

RAD sequencing reveals genomewide divergence between independent invasions of the European green crab (Carcinus maenas) in the Northwest Atlantic

Genomic studies of invasive species can reveal both invasive pathways and functional differences underpinning patterns of colonization success. The European green crab (Carcinus maenas) was initially introduced to eastern North America nearly 200 years ago where it expanded northwards to eastern Nova Scotia. A subsequent invasion to Nova Scotia from a northern European source allowed further range expansion, providing a unique opportunity to study the invasion genomics of a species with multiple invasions. Here, we use restriction‐site‐associated DNA sequencing‐derived SNPs to explore fine‐scale genomewide differentiation between these two invasions. We identified 9137 loci from green crab sampled from 11 locations along eastern North America and compared spatial variation to mitochondrial COI sequence variation used previously to characterize these invasions. Overall spatial divergence among invasions was high (pairwise F_ST ~0.001 to 0.15) and spread across many loci, with a mean F_ST ~0.052 and 52% of loci examined characterized by F_ST values >0.05. The majority of the most divergent loci (i.e., outliers, ~1.2%) displayed latitudinal clines in allele frequency highlighting extensive genomic divergence among the invasions. Discriminant analysis of principal components (both neutral and outlier loci) clearly resolved the two invasions spatially and was highly correlated with mitochondrial divergence. Our results reveal extensive cryptic intraspecific genomic diversity associated with differing patterns of colonization success and demonstrates clear utility for genomic approaches to delineating the distribution and colonization success of aquatic invasive species.     

Quantifying factors influencing American lobster (Homarus americanus) predation on the invasive green crab (Carcinus maenas) in Newfoundland,Canada

The European green crab (Carcinus maenas) is a newly invasive species in Newfoundland, where it has likely been present for ≤15 years. The green crab has been found in stomach contents of American lobster (Homarus americanus) in New England and Nova Scotia, Canada, but predation on this species has not yet been quantified in Newfoundland. We conducted feeding experiments to determine whether lobsters from Newfoundland were as likely as those from Nova Scotia (which have coexisted with green crabs for >60 years) to recognize and prey upon this new species. We also performed experiments to determine whether green crabs reach a size refuge from predation and whether factors including starvation, availability of alternate food sources, or habitat complexity would influence the probability of lobster attacking or feeding on green crabs. In our trials, lobster origin had no significant effect on crab predation; lobsters, irrespective of origin, were more likely to consume small (<40 mm carapace width [CW]) and medium (40–65 mm CW) crabs than larger (>65 mm CW) ones. Nevertheless, even small lobsters (73–76 mm carapace length, 300 g) were able to kill and consume the largest green crabs (78 mm CW, 100 g). Green crabs were less likely to be attacked or eaten when an alternative food source was present, suggesting that the lobsters were preying on the crabs, rather than simply killing them in a dispute over territory. The addition of a shelter provided a refuge for the green crabs; however, the crabs were only able to avoid being injured or eaten if this shelter was structurally complex. The green crab is slowly spreading westward around the island of Newfoundland, and so its long‐term effects, interactions with other organisms, and contribution to the diet of Newfoundland lobsters remain to be seen.     

Niche shifts during the global invasion of the Asian tiger mosquito, Aedes albopictus Skuse (Culicidae), revealed by reciprocal distribution models

Aim Niche‐based distribution models are often used to predict the spread of invasive species. These models assume niche conservation during invasion, but invasive species can have different requirements from populations in their native range for many reasons, including niche evolution. I used distribution modelling to investigate niche conservatism for the Asian tiger mosquito (Aedes albopictus Skuse) during its invasion of three continents. I also used this approach to predict areas at risk of invasion from propagules originating from invasive populations. Location Models were created for Southeast Asia, North and South America, and Europe. Methods I used maximum entropy (Maxent ) to create distribution models using occurrence data and 18 environmental datasets. One native model was created for Southeast Asia; this model was projected onto North America, South America and Europe. Three models were created independently for the non‐native ranges and projected onto the native range. Niche overlap between native and non‐native predictions was evaluated by comparing probability surfaces between models using real data and random models generated using a permutation approach. Results The native model failed to predict an entire region of occurrences in South America, approximately 20% of occurrences in North America and nearly all Italian occurrences of A. albopictus. Non‐native models poorly predict the native range, but predict additional areas at risk for invasion globally. Niche overlap metrics indicate that non‐native distributions are more similar to the native niche than a random prediction, but they are not equivalent. Multivariate analyses support modelled differences in niche characteristics among continents, and reveal important variables explaining these differences. Main conclusions The niche of A. albopictus has shifted on invaded continents relative to its native range (Southeast Asia). Statistical comparisons reveal that the niche for introduced distributions is not equivalent to the native niche. Furthermore, reciprocal models highlight the importance of controlling bi‐directional dispersal between native and non‐native distributions.     

Organization of a phyllobranchiate gill from the green shore crab Carcinus maenas (Crustacea,Decapoda)

Summary The phyllobranchiate gills of the green shore crab Carcinus maenas have been examined histologically and ultrastructurally. Each gill lamella is bounded by a chitinous cuticle. The apical surface of the branchial epithelium contacts this cuticle, and a basal lamina segregates the epithelium from an intralamellar hemocoel. In animals acclimated to normal sea water, five epithelial cell types can be identified in the lamellae of the posterior gills: chief cells, striated cells, pillar cells, nephrocytes, and glycocytes. Chief cells are the predominant cells in the branchial epithelium. They are squamous or low cuboidal and likely play a role in respiration. Striated cells, which are probably involved in ionoregulation, are also squamous or low cuboidal. Basal folds of the striated cells contain mitochondria and interdigitate with the bodies and processes of adjacent cells. Pillar cells span the hemocoel to link the proximal and distal sides of a lamella. Nephrocytes are large, spherical cells with voluminous vacuoles. They are rimmed by foot processes or pedicels and frequently associate with the pillar cells. Glycocytes are pleomorphic cells packed with glycogen granules and multigranular rosettes. The glycocytes often mingle with the nephrocytes. Inclusion of the nephrocytes and glycocytes as members of the branchial epithelium is justified by their participation in intercellular junctions and their position internal to the epithelial basal lamina.     

Changes in the realized niche of the invasive succulent CAM plant Furcraea foetida

Furcraea foetida (Asparagaceae) is a native plant of Central America and northern South America but there is no information about its country of origin. The species was introduced into Brazil and is now considered invasive, particularly in coastal ecosystems. To date, nothing is known about the environmental factors that constrain its distribution and there is only inconclusive information about its location of origin. We used reciprocal distribution models (RDM) to assess invasion risk of F. foetida across Brazil and to identify source regions in its native range. We also tested the niche conservatism hypothesis using Principal Components Analyses and statistical tests of niche equivalency and similarity between its native and invaded ranges. For RDM analysis, we built two models using maximum entropy, one using records in the native range to predict the invaded distribution (forward‐Ecological Niche Model or forward‐ENM) and one using records in the invaded range to predict the native distribution (reverse‐ENM). Forward‐ENM indicated invasion risk in the Cerrado region and the innermost region of the Atlantic Forest, however, failed to predict the current occurrence in southern Brazil. Reverse‐ENM supported an existing hypothesis that F. foetida originated in the Orinoco river basin, Amazon basin and Caribbean islands. Prediction errors in the RDM and multivariate analysis indicated that the species expanded its realized niche in Brazil. The niche similarity test further suggested that the niche differences are because of differences in habitat availability between the two ranges, not because of evolutionary changes. We hypothesize that physiological pre‐adaptation (especially, the crassulacean acid metabolism), human‐driven propagule pressure and high competitive ability are the main factors determining the current spatial distribution of the species in Brazil. Our study highlights the need to include F. foetida in plant invasion monitoring programs, especially in priority conservation areas where the species has still not been introduced.     

Cryptic niche conservatism among evolutionary lineages of an invasive lizard

Aim There is increasing evidence that the quality and breadth of ecological niches vary among individuals, populations, evolutionary lineages and therefore also across the range of a species. Sufficient knowledge about niche divergence among clades might thus be crucial for predicting the invasion potential of species. We tested for the first time whether evolutionary lineages of an invasive species vary in their climate niches and invasive potential. Furthermore, we tested whether lineage‐specific models show a better performance than combined models. Location Europe. Methods We used species distribution models (SDMs) based on climatic information at native and invasive ranges to test for intra‐specific niche divergence among mitochondrial DNA (mtDNA) clades of the invasive wall lizard Podarcis muralis. Using DNA barcoding, we assigned 77 invasive populations in Central Europe to eight geographically distinct evolutionary lineages. Niche similarity among lineages was assessed and the predictive power of a combination of clade‐specific SDMs was compared with a combined SDM using the pooled records of all lineages. Results We recorded eight different invasive mtDNA clades in Central Europe. The analysed clades had rather similar realized niches in their native and invasive ranges, whereas inter‐clade niche differentiation was comparatively strong. However, we found only a weak correlation between geographic origin (i.e. mtDNA clade) and invasive occurrences. Clades with narrow realized niches still became successful invaders far outside their native range, most probably due to broader fundamental niches. The combined model using data for all invasive lineages achieved a much better prediction of the invasive potential. Conclusions Our results indicate that the observed niche differentiation among evolutionary lineages is mainly driven by niche realization and not by differences in the fundamental niches. Such cryptic niche conservatism might hamper the success of clade‐specific niche modelling. Cryptic niche conservatism may in general explain the invasion success of species in areas with apparently unsuitable climate.     

Positive Darwinian selection on crustacean hyperglycemic hormone (CHH) of the green shore crab, Carcinus maenas

The tissue-specific expression and differential function of the crustacean hyperglycemic hormone (CHH) in Carcinus maenas indicate an interesting evolutionary history. Previous studies have shown that CHH from the sinus gland X-organ (XO-type) has hyperglycemic activity, whereas the CHH from the pericardial organ (PO-type) neither shows hyperglycemic activity nor it inhibits Y-organ ecdysteroid synthesis. Here we examined the types of selective pressures operating on the variants of CHH in Carcinus maenas. Maximum likelihood-based codon substitution analyses revealed that the variants of this neuropeptide in C. maenas have been subjected to positive Darwinian selection indicating adaptive evolution and functional divergence among the CHH variants leading to two unique groups (PO and XO-type). Although the average ratio of nonsynonymous to synonymous substitution (omega) for the entire coding region is 0.5096, few codon sites showed significantly higher omega (10.95). Comparison of models that incorporate positive selection (omega > 1) with models not incorporating positive selection (omega <1) at certain codon sites failed to reject (p=0) evidence of positive Darwinian selection.     

Episodic global dispersal in shallow water marine organisms: the case history of the European shore crabs Carcinus maenas and C. aestuarii

Aim This paper evaluates global collection records, evidence of anthropogenic transport methods, and experimental and distributional data relative to temperature requirements to understand the historical and potential dispersal of a well‐known genus of estuarine crab. Location The records analysed are from temperate and tropical coastal ocean areas. Methods The study is based primarily on literature analysis and examination of museum specimens. Results The human‐mediated successful global dispersal of the European shore crabs Carcinus maenas (Linnaeus, 1758) and C. aestuarii (Nardo, 1847) occurred in three major episodes: around 1800, in the 1850s–70s, and in the 1980s–90s. The nineteenth century introductions occurred through transport by ships (probably in hull fouling or in solid ballast), while the introductions in the 1980s could have occurred through a greater variety of dispersal mechanisms (ships’ hull fouling and seawater system fouling; fouling on semisubmersible drilling platforms; ballast water; transport with fisheries products intended for food or bait; scientific research; releases from aquaria maintained for educational or scientific purposes; or intentional non‐governmental releases for human food production). These introductions have resulted in Carcinus’ establishment in five temperate regions outside of its native Europe in Atlantic North America, Australia, South Africa, Japan and Pacific North America, while releases into tropical regions have not established populations. C. maenas’ range in both its native and introduced regions appears to be regulated by similar temperature parameters, enabling an assessment of its potential distribution. Main conclusions The second episode of Carcinus’ global dispersal, the period from the 1850s to 1870s, may be part of a broader surge of world‐wide invasions caused by an increase in shipping.