Evolutionary history of the little fire ant Wasmannia auropunctata before global invasion: inferring dispersal patterns,niche requirements and past and present distribution within its native range

The evolutionary history of invasive species within their native range may involve key processes that allow them to colonize new habitats. Therefore, phylogeographic studies of invasive species within their native ranges are useful to understand invasion biology in an evolutionary context. Here we integrated classical and Bayesian phylogeographic methods using mitochondrial and nuclear DNA markers with a palaeodistribution modelling approach, to infer the phylogeographic history of the invasive ant Wasmannia auropunctata across its native distribution in South America. We discuss our results in the context of the recent establishment of this mostly tropical species in the Mediterranean region. Our Bayesian phylogeographic analysis suggests that the common ancestor of the two main clades of W. auropunctata occurred in central Brazil during the Pliocene. Clade A would have differentiated northward and clade B southward, followed by a secondary contact beginning about 380 000 years ago in central South America. There were differences in the most suitable habitats among clades when considering three distinct climatic periods, suggesting that genetic differentiation was accompanied by changes in niche requirements, clade A being a tropical lineage and clade B a subtropical and temperate lineage. Only clade B reached more southern latitudes, with a colder climate than that of northern South America. This is concordant with the adaptation of this originally tropical ant species to temperate climates prior to its successful establishment in the Mediterranean region. This study highlights the usefulness of exploring the evolutionary history of invasive species within their native ranges to better understand biological invasions.     

Latitudinal shifts of introduced species: possible causes and implications

This study aims to document shifts in the latitudinal distributions of non-native species relative to their own native distributions and to discuss possible causes and implications of these shifts. We used published and newly compiled data on inter-continentally introduced birds, mammals and plants. We found strong correlations between the latitudinal distributions occupied by species in their native and exotic ranges. However, relatively more non-native species occur at latitudes higher than those in their native ranges, and fewer occur at latitudes lower than those in their native ranges. Only a small fraction of species examined (i.e. <20% of animals and <10% of plants) have expanded their distributions in their exotic range beyond both high- and low-limits of their native latitudes. Birds, mammals and plants tended to shift their exotic ranges in similar ways. In addition, most non-native species (65–85% in different groups) have not reached the distributional extent observed in their native ranges. The possible drivers of latitudinal shifts in the exotic range may include climate change, greater biotic resistance at lower latitudes, historical limitations on ranges in native regions, and the impacts of humans on species distributions. The relatively restricted distribution of most species in their exotic range highlights the great potential of future spread of most introduced species and calls for closely monitoring their directional spread under climate change.     

Latitudinal variation of life-history traits of an exotic and a native impatiens species in Europe

Understanding the responses of invasive and native populations to environmental change is crucial for reliable predictions of invasions in the face of global change. While comparisons of responses across invasive species with different life histories have been performed before, comparing functional traits of congeneric native and invasive species may help to reveal driving factors associated with invasion. Here we compared morphological functional trait patterns of an invasive species (Impatiens parviflora) with its congeneric native species (I. noli-tangere) along an approximately 1600 km European latitudinal gradient from France (49°34′N) to Norway (63°40′N). Soil nitrogen was recorded during six weeks of the growing season, and light, soil moisture, and nutrient availability were estimated for each sampled population using community weighted means of indicator values for co-occurring species. Temperature data were gathered from nearby weather stations.Both the native and invasive species are taller at higher latitudes and this response is strongest in the invasive species. Seed mass and number of seeds per capsule increase in I. noli-tangere but decrease in I. parviflora towards higher latitudes. Surprisingly, plant height in the invasive I. parviflora decreases with increasing soil nitrogen availability. The latitudinal pattern in seed mass is positively related to temperature in I. noli-tangere and negatively in I. parviflora. Leaf area of both species decreases with increasing Ellenberg indicator values for nitrogen and light but increases with increasing soil moisture. Soil nitrogen concentrations and Ellenberg indicator values for nitrogen have significant positive (I. noli-tangere) and negative (I. parviflora) effects on the number of seeds per capsule. Our results show that the native I. noli-tangere has efficient reproduction at its range edge while the invasive I. parviflora shows a marked decrease in seed size and seed number per capsule. These patterns are unrelated to the growth and obtained size of the plants: even low soil nitrogen availability in the north seemed not to limit plant growth and size. Our results suggest that the invasive I. parviflora tends to become more invasive at lower latitudes by producing heavier seeds and more seeds per capsule.     

Do invasive freshwater fish species grow better when they are invasive?

A frequent assumption in invasion ecology is that invasive species have enhanced growth rates in their invasive ranges. However, invasions frequently occur in sub‐tropical and tropical environments where growth could be higher simply due to climatic conditions rather than novel habitat. In this study, a meta‐analysis of growth rates (length‐at‐age data) was completed for six invasive freshwater fish species: common carp Cyprinus carpio, largemouth bass Micropterus salmoides, brown trout Salmo trutta, brown bullhead Ictalurus nebulosus, flathead catfish Pylodictis olivaris and northern snakehead Channa argus. Significant effects of climate on growth were observed for all species except common carp, and following normalization of growth for climate effects, a range of growth responses between native and invasive populations were revealed. Two species (brown trout, flathead catfish) showed significantly increased growth rates in invasive compared to native ranges, but two species (common carp, largemouth bass) showed significantly faster growth in native ranges, and two other species (northern snakehead, brown bullhead) showed no difference in growth rates. No species showed both significantly enhanced growth rates and initial sizes in invasive compared to native ranges. Using the comparative method, countergradient growth variations were apparent for all species within their native ranges and for all but one species in invasive ranges. Invasive populations of freshwater fish do not always grow faster when invasive and future studies need to consider growth covariates (e.g. climate and countergradient growth) prior to comparing life‐history differences between invasive and native populations.     

Does the effect of flowering time on biomass allocation across latitudes differ between invasive and native salt marsh grass Spartina alterniflora?

1. Parallel latitudinal clines in flowering time have been documented in both the invasive and native ranges of plants. Furthermore, flowering time has been found to affect biomass at maturity. Therefore, understanding how these flowering times affect biomass accumulation across latitudes is essential to understanding plant adaptations and distributions.
2. We investigated and compared trends in first flowering day (FFD), aboveground biomass (AGB), belowground biomass (BGB), and BGB:AGB ratio of the salt marsh grass Spartina alterniflora along latitudinal gradients from the invasive (China, 19–40°N) and native range (United States, 27–43°N) in a greenhouse common garden experiment, and tested whether FFD would drive these divergences between invasive and native ranges.
3. The invasive populations produced more (~20%, ~19%) AGB and BGB than native populations, but there were no significant differences in the FFD and BGB:AGB ratio. We found significant parallel latitudinal clines in FFD in both invasive and native ranges. In addition, the BGB:AGB ratio was negatively correlated with the FFD in both the invasive and native ranges but nonsignificant in invasive populations. In contrast, AGB and BGB increased with latitude in the invasive range, but declined with latitude in the native range. Most interestingly, we found AGB and BGB positively correlated with the FFD in the native range, but no significant relationships in the invasive range.
4. Our results indirectly support the evolution of increased competitive ability hypothesis (EICA) that S. alterniflora has evolved to produce greater AGB and BGB in China, but the flowering and allocation pattern of native populations is maintained in the invasive range. Our results also suggest that invasive S. alterniflora in China is not constrained by the trade‐off of earlier flowering with smaller size, and that flowering time has played an important role in biomass allocation across latitudes.

     

Genetic differences in growth of an invasive tree species

Invasive plants are often more vigorous in their introduced ranges than in their native ranges. This may reflect an innate superiority of plants from some habitats or an escape from their enemies. Another hypothesis proposes that invasive plants evolve increased competitive ability in their introduced range. We present the results of a 14-year common garden experiment with the Chinese Tallow Tree ( Sapium sebiferum ) from its native range (Asia), place of introduction to North America (Georgia) and areas colonized a century later (Louisiana and Texas). Invasive genotypes, especially those from recently colonized areas, were larger than native genotypes and more likely to produce seeds but had lower quality, poorly defended leaves. Our results demonstrate significant post-invasion genetic differences in an invasive plant species. Post-introduction adaptation by introduced plants may contribute to their invasive success and make it difficult to predict problem species.     

Different climatic envelopes among invasive populations may lead to underestimations of current and future biological invasions

Aim We explore the impact of calibrating ecological niche models (ENMs) using (1) native range (NR) data versus (2) entire range (ER) data (native and invasive) on projections of current and future distributions of three Hieracium species. Location H. aurantiacum, H. murorum and H. pilosella are native to Europe and invasive in Australia, New Zealand and North America. Methods Differences among the native and invasive realized climatic niches of each species were quantified. Eight ENMs in BIOMOD were calibrated with (1) NR and (2) ER data. Current European, North American and Australian distributions were projected. Future Australian distributions were modelled using four climate change scenarios for 2030. Results The invasive climatic niche of H. murorum is primarily a subset of that expressed in its native range. Invasive populations of H. aurantiacum and H. pilosella occupy different climatic niches to those realized in their native ranges. Furthermore, geographically separate invasive populations of these two species have distinct climatic niches. ENMs calibrated on the realized niche of native regions projected smaller distributions than models incorporating data from species’ entire ranges, and failed to correctly predict many known invasive populations. Under future climate scenarios, projected distributions decreased by similar percentages, regardless of the data used to calibrate ENMs; however, the overall sizes of projected distributions varied substantially. Main conclusions This study provides quantitative evidence that invasive populations of Hieracium species can occur in areas with different climatic conditions than experienced in their native ranges. For these, and similar species, calibration of ENMs based on NR data only will misrepresent their potential invasive distribution. These errors will propagate when estimating climate change impacts. Thus, incorporating data from species’ entire distributions may result in a more thorough assessment of current and future ranges, and provides a closer approximation of the elusive fundamental niche.     

Invasion success of non-indigenous aquatic and semi-aquatic plants in their native and introduced ranges. A comparison between their invasiveness in North America and in France

Aquatic and semi-aquatic plants comprise few species worldwide, yet the introduction of non-indigenous plants represents one of the most severe examples of biological invasions. My goal is to compare the distribution and the biology of aquatic and semi-aquatic plants in their introduced ranges and in their native ranges. The primary objective of this study is to test the hypothesis that invasive species have evolved traits likely to increase their success in the new range. I made two reciprocal comparisons, i.e. I compared European species in France and in North America, and North American species in France and in North America. Twenty-seven species were classified according to their invasiveness in their introduced area. I␣found six invasive macrophyte species in France native to North America and 17 invasive species in North America native to Europe. Four species are invasive in both areas. There is no general tendency for macrophytes to be more vigorous in their introduced ranges. Most non-indigenous aquatic and semi-aquatic species are potentially invasive or widespread and well-established in their introduced country, while few species seem to be restricted in their distribution.     

Prominent genetic structure across native and introduced ranges of Pluchea indica,a mangrove associate,as revealed by microsatellite markers

阔苞菊(Pluchea indica)是一种红树林伴生植物，以其在原产地的药用特性和部分引入地的入侵性而闻名。本研究旨在评估阔苞菊在其分布范围内遗传变异的地理分布，确定影响其遗传结构的因素，并利用这些信息对阔苞菊在原产地和引入地的保护和管理策略提出建议。 我们以来自阔苞菊原产地(亚洲)和引入地(美国)的31个种群共348个个体的15个核微卫星位点数据对阔苞菊的遗传多样性和种群结构进行了评估。在大尺度范围以及局部区域两种空间尺度上对阔苞菊遗传变异的空间格局进行了探讨，并验证了以下假说：地理距离和自然地理屏障将影响种群结构并在空间尺度上产生不同程度的分化。研究结果表明，与所研究区域内的其它红树林物种的遗传多样性参数相比， 我们发现阔苞菊在种群水平上具有相对较高的遗传多样性以及在物种水平上具有明显的遗传分化。大多数阔苞菊种群显示杂合子缺失， 这主要是由于近交和有限的基因流所导致。在较大空间尺度上进行的种群结构分析显示，该物种自然分布范围内存在两个主要遗传谱系，中国的种群与印度尼西亚、马来西亚、新加坡、泰国、柬埔寨和菲律宾的种群分别属于不同的谱系，而美国的种群可能来自于中国的谱系。 此外，在局部区域范围内也同样检测到种群之间的遗传分化。大部分阔苞菊种群所表现出的遗传瓶颈效应强调了其具有本地灭绝的风险。基于上述研究结果，我们建议采用原位保护策略对阔苞菊进行管理，并开展对优先保护种群的保护行动以维持遗传多样性。     

Evidence of ecological niche shift in Rhododendron ponticum (L.) in Britain: Hybridization as a possible cause of rapid niche expansion

Biological invasions threaten global biodiversity and natural resources. Anticipating future invasions is central to strategies for combating the spread of invasive species. Ecological niche models are thus increasingly used to predict potential distribution of invasive species. In this study, we compare ecological niches of Rhododendron ponticum in its native (Iberian Peninsula) and invasive (Britain) ranges. Here, we test the conservation of ecological niche between invasive and native populations of R. ponticum using principal component analysis, niche dynamics analysis, and MaxEnt‐based reciprocal niche modeling. We show that niche overlap between native and invasive populations is very low, leading us to the conclusion that the two niches are not equivalent and are dissimilar. We conclude that R. ponticum occupies novel environmental conditions in Britain. However, the evidence of niche shift presented in this study should be treated with caution because of nonanalogue climatic conditions between native and invasive ranges and a small population size in the native range. We then frame our results in the context of contradicting genetic evidence on possible hybridization of this invasive species in Britain. We argue that the existing contradictory studies on whether hybridization caused niche shift in R. ponticum are not sufficient to prove or disprove this hypothesis. However, we present a series of theoretical arguments which indicate that hybridization is a likely cause of the observed niche expansion of R. ponticum in Britain.     

Native range climate is insufficient to predict anuran invasive potential

Niche conservatism explains biological invasions worldwide. However, a plethora of ecological processes may lead invasive species to occupy environments that are different from those found within native ranges. Here, we assess the potential invadable areas of  the world’s most pervasive invasive amphibians: the cane toad, Rhinella marina?+?R. horribilis, and the North American bullfrog, Lithobates catesbeianus. The uncontrolled spread of such voracious, large-bodied, and disease-tolerant anurans has been documented to impact native faunas worldwide. To disentangle their invasion-related niche dynamics, we compared the predictive ability and distributional forecasts of ecological niche models calibrated with information from native, invaded and pooled (native?+?invaded) ranges. We found that including occurrences from invaded ranges improved model accuracy for both studied species. Non-native occurrences also accounted for 54% and 61% increase in the total area of potential distribution of the cane toad and bullfrog, respectively. Besides, the latter species occupied locations with climatic conditions that are more extreme than those found within its native range. Our results indicate that the occupancy of environments different from those found in native ranges increases the overall potential distribution of the studied invasive anuran species. Therefore, climate information on native ranges alone is insufficient to explain and anticipate the distributional patterns of invasion of cane toads and bullfrogs, underestimating predictions of potential invadable distribution. Moreover, such an observed expansion of realized niches towards occupancy of climates not found within native ranges also has clear implications for invasion risk assessments based on climate modelling worldwide.

     

Niche dynamics of two cryptic <Emphasis Type="Italic">Prosopis</Emphasis> invading South American drylands

Whether or not species track native climatic conditions during invasions (i.e., climate match hypothesis) is fundamental to understand and prevent potential impacts of invasive species. Recent empirical work suggests that climatic mismatches between native and invasive ranges are pervasive. Whether these differences are due to adaptation to new climatic spaces in the invasive range or due to partial filling of the potential climatic space are still subject to debate. Here, we analyze climatic niche dynamics associated with the invasion of the two most common invasive plants in Brazilian semi-arid areas, Prosopis juliflora and Prosopis pallida. These species have been simultaneously introduced in the region, which creates a unique opportunity to compare their niche dynamics during invasion. Given that P. juliflora have a much wider native range size, we expect these species would present different dispersal potentials, which might translate into different unfilling levels. Using an ordination method with kernel smoother and null models, we contrasted climate spaces occupied by each species in both native and invasive ranges. We further used ecological niche models (ENMs) to compare reciprocal predictions of potentially suitable areas. Against our expectation based on differences in native range sizes, climatic niches of P. juliflora and P. pallida overlapped greatly, both in their native and invasive ranges. Our results support niche conservatism during the invasion process. Climatic mismatches among native and invaded ranges were exclusively attributed to unfilling of native climates in the invasive range. Both species showed similar unfilling levels. Likewise, ENMs predicted regions not yet occupied in the invasive range, revealing a potential for further expansion. We discuss colonization time lag and founder effect as potential mechanisms that may have prevented these species to fully occupy their native niches in the invasive range.     

Interactions between invading benthivorous fish and native whitefish in subarctic lakes

相似文献   

Deciphering the worldwide invasion of the Asian long‐horned beetle: A recurrent invasion process from the native area together with a bridgehead effect

Retracing introduction routes is crucial for understanding the evolutionary processes involved in an invasion, as well as for highlighting the invasion history of a species at the global scale. The Asian long‐horned beetle (ALB) Anoplophora glabripennis is a xylophagous pest native to Asia and invasive in North America and Europe. It is responsible for severe losses of urban trees, in both its native and invaded ranges. Based on historical and genetic data, several hypotheses have been formulated concerning its invasion history, including the possibility of multiple introductions from the native zone and secondary dispersal within the invaded areas, but none have been formally tested. In this study, we characterized the genetic structure of ALB in both its native and invaded ranges using microsatellites. In order to test different invasion scenarios, we used an approximate Bayesian “random forest” algorithm together with traditional population genetics approaches. The strong population differentiation observed in the native area was not geographically structured, suggesting complex migration events that were probably human‐mediated. Both native and invasive populations had low genetic diversity, but this characteristic did not prevent the success of the ALB invasions. Our results highlight the complexity of invasion pathways for insect pests. Specifically, our findings indicate that invasive species might be repeatedly introduced from their native range, and they emphasize the importance of multiple, human‐mediated introductions in successful invasions. Finally, our results demonstrate that invasive species can spread across continents following a bridgehead path, in which an invasive population may have acted as a source for another invasion.     

<Emphasis Type="Italic">Acroptilon repens</Emphasis>, an Asian invader,has stronger competitive effects on species from America than species from its native range

The ability to competitively suppress native species is key to successful invasion. Since invasions involve an increase in abundance or dominance of a species in its non-native range, competitive effects might be expected to be stronger in the non-native range of an invader; however, there have been few comparisons of the competitive effects of invasive plants on species from invaded ranges versus species from native ranges. We compared the competitive and allelopathic effects of Acroptilon repens on native North American species to effects on related species from the native range of Acroptilon in Uzbekistan. We also compared the competitive interactions among these North American and Eurasian species, in the absence of Acroptilon, examining the hypothesis that particular regional species pools may show differences in competitive ability. The results showed that Acroptilon had stronger competitive effects against native North American species than against species native to Uzbekistan. There was no difference in the competitive effects among Eurasians and North Americans. The effects of leachates collected from Acroptilon roots were weak but more negative on species from North America than on species from Uzbekistan. Our results suggest that inherently stronger competitive and allelopathic effects of Acroptilon on North American plants than on plants from its native range may contribute to its invasive success.     

Macroecology meets invasion ecology: performance of Australian acacias and eucalypts around the world revealed by features of their native ranges

Native geographical range extent has frequently emerged as a correlate of invasiveness, especially for plant species. We tested whether dimensions of the native range (measured by the area-of-occupancy and its scaling patterns) of 720 Australian eucalypts (genera Angophora, Eucalyptus and Corymbia) could explain introduction and invasion success. We also compared our results with a previous study on 979 Australian acacias (previously grouped in Acacia subgenus Phyllodineae) to investigate whether features of their native ranges explained the much higher invasion success in this group compared to eucalypts. From nine databases we found records that 373 eucalypts have been introduced to areas outside their native ranges; 82 of these have become naturalised, and eight are invasive. A similar proportion of Australian acacias have been introduced, but almost three times as many are invasive (384 species introduced, 71 of which are naturalized and 23 invasive). Eucalypts with large native ranges are more likely to have been introduced and subsequently naturalise, as is the case with acacias. Unlike acacias, however, the native range size of invasive eucalypts was not significantly greater than naturalised (but not invasive) species. Intriguingly, the human preference for introducing species with larger ranges was much greater for acacias than for eucalypts as the geometric mean range sizes of introduced, naturalised and invasive acacias are 2.04, 1.88 and 3.59 times those of eucalypts at the same stage. Moreover, the percolation exponent (i.e. the slope of occupancy scaling) becomes more extreme towards the end of the introduction–naturalization–invasion continuum, decreasing for acacias but increasing for eucalypts, with acacias, except for non-introduced species, having a lower exponent than eucalypts. The selection preference of acacias during introduction is thus for species that can rapidly expand their range; in contrast, slow-spreading eucalypts have been selected for dissemination. In other words, humans appear to have selected for highly invasive acacias but against introducing highly invasive eucalypts.     

Invasion of Lepidium draba (Brassicaceae) in the western United States: distributions and origins of chloroplast DNA haplotypes

Advances in phylogeography are of great value for understanding the population structure and origins of invasive genotypes. Such insights provide constructive information for current or future biological control research efforts. In this study, we investigated a highly variable chloroplast DNA (cpDNA) marker for populations of the weed Lepidium draba (Brassicaceae) in its native Eurasian and invasive US ranges. We sequenced DNA from 684 individuals from Eurasia and the US and found 41 different haplotypes. Our comparative study between the native and invasive ranges showed a 33% reduction in allelic richness (A) and a 7% reduction in haplotype diversity (h) since introduction into the US. Most genetic variation in the native range was observed within geographical regions and populations, not between regions, and this result was similar for the invasive range. Assignment tests indicated the most likely origins of many invasive haplotypes. Some of these occurred in western Europe, supporting an expanded native range that had been proposed for the species. Exact locations were identified for a diverse set of invasive haplotypes which can be used in ongoing host-specificity tests of potential biological control agents.     

Global realized niche divergence in the African clawed frog Xenopus laevis

Although of crucial importance for invasion biology and impact assessments of climate change, it remains widely unknown how species cope with and adapt to environmental conditions beyond their currently realized climatic niches (i.e., those climatic conditions existing populations are exposed to). The African clawed frog Xenopus laevis, native to southern Africa, has established numerous invasive populations on multiple continents making it a pertinent model organism to study environmental niche dynamics. In this study, we assess whether the realized niches of the invasive populations in Europe, South, and North America represent subsets of the species’ realized niche in its native distributional range or if niche shifts are traceable. If shifts are traceable, we ask whether the realized niches of invasive populations still contain signatures of the niche of source populations what could indicate local adaptations. Univariate comparisons among bioclimatic conditions at native and invaded ranges revealed the invasive populations to be nested within the variable range of the native population. However, at the same time, invasive populations are well differentiated in multidimensional niche space as quantified via n‐dimensional hypervolumes. The most deviant invasive population are those from Europe. Our results suggest varying degrees of realized niche shifts, which are mainly driven by temperature related variables. The crosswise projection of the hypervolumes that were trained in invaded ranges revealed the south‐western Cape region as likely area of origin for all invasive populations, which is largely congruent with DNA sequence data and suggests a gradual exploration of novel climate space in invasive populations.     

Evidence of climatic niche shift during biological invasion

Niche-based models calibrated in the native range by relating species observations to climatic variables are commonly used to predict the potential spatial extent of species' invasion. This climate matching approach relies on the assumption that invasive species conserve their climatic niche in the invaded ranges. We test this assumption by analysing the climatic niche spaces of Spotted Knapweed in western North America and Europe. We show with robust cross-continental data that a shift of the observed climatic niche occurred between native and non-native ranges, providing the first empirical evidence that an invasive species can occupy climatically distinct niche spaces following its introduction into a new area. The models fail to predict the current invaded distribution, but correctly predict areas of introduction. Climate matching is thus a useful approach to identify areas at risk of introduction and establishment of newly or not-yet-introduced neophytes, but may not predict the full extent of invasions.     

Niche shift in four non‐native estrildid finches and implications for species distribution models

Non‐native species can have severe impacts on ecosystems. Therefore, predictions of potentially suitable areas that are at risk of the establishment of non‐native populations are desirable. In recent years, species distribution models (SDMs) have been widely applied for this purpose. However, the appropriate selection of species records, whether from the native area alone or also from the introduced range, is still a matter of debate. We combined analyses of native and non‐native realized climate niches to understand differences between models based on all locations, as well as on locations from the native range only. Our approach was applied to four estrildid finch species that have been introduced to many regions around the world. Our results showed that SDMs based on location data from native areas alone may underestimate the potential distribution of a given species. The climatic niches of species in their native ranges differed from those of their non‐native ranges. Niche comparisons resulted in low overlap values, indicating considerable niche shifts, at least in the realized niches of these species. All four species have high potential to spread over many tropical and subtropical areas. However, transferring these results to temperate areas has a high degree of uncertainty, and we urge caution when assessing the potential spread of tropical species that have been introduced to higher latitudes.