Population-specific demography and invasion potential in medfly

Biological invasions are constantly gaining recognition as a significant component of global change. The Mediterranean fruit fly (medfly) constitutes an ideal model species for the study of biological invasions due to its (1) almost cosmopolitan geographic distribution, (2) huge economic importance, and (3) well-documented invasion history. Under a common garden experimental set up, we tested the hypothesis that medfly populations obtained from six global regions [Africa (Kenya), Pacific (Hawaii), Central America (Guatemala), South America (Brazil), Extra-Mediterranean (Portugal), and Mediterranean (Greece)] have diverged in important immature life-history traits such as preadult survival and developmental times. We also tested the hypothesis that medfly populations from the above regions exhibit different population growth rates. For this purpose, data on the life history of immatures were combined with adult survival and reproduction data derived from an earlier study in order to calculate population parameters for the above six populations. Our results clearly show that medfly populations worldwide exhibit significant differences in preadult survival, developmental rates of immatures and important population parameters such as the intrinsic rate of increase. Therefore, geographically isolated medfly populations may share different invasion potential, since population growth rates could influence basic population processes that operate mostly during the last two stages of an invasion event, such as establishment and spread. Our findings provide valuable information for designing population suppression measures and managing invasiveness of medfly populations worldwide.     

An experimental study of the effects of weed invasion on lizard phenotypes

We examined how a weed affected the basking and activity of a diurnal lizard, and the potential cascading effects of these shifts for life history strategies and expression of morphology. Hatchlings of the diurnal lizard Lampropholis delicata were raised to maturity in outdoor enclosures that mimicked high, moderate and low invasion by a sprawling plant (blue periwinkle, Vinca major). Skinks depend on sunlight for growth and maintenance. Periwinkle differs from displaced grassland by being structurally complex and blocking sunlight. Lizards restricted to the enclosure floor achieved preferred body temperatures only when exposure to periwinkle was moderate or low. However, lizards in high invasion enclosures could reach preferred body temperatures by climbing plants and basking on exposed canopy. This shift in basking strategy resulted in lizards growing longer hind limbs compared with animals that rarely (moderate invasion) and never (low invasion) climbed plants. Consequently, lizards reared in high invasion enclosures sprinted faster than conspecifics reared in lower invasion environments. Throughout the study there was no significant variation among treatments in the tendency of animals to be moving when they were not hidden. However, lizards in high invasion treatments hid more often during the day, were lighter in body mass, and females had lighter clutch masses and offspring than did those from moderate and low invasion enclosures. Thus, microhabitat degradation can drive a cascade of changes to an animal’s ecology.     

分子标记技术在昆虫入侵研究中的应用

遗传变异与种群持续性及其进化潜力密切相关,而生物入侵导致种群遗传变异或遗传多样性的改变为研究自然界中各种生态和进化问题提供了理想模式。分子标记技术是调查种群遗传变异的重要工具,揭示了入侵种的入侵过程和结果,并预测未来的发生情况。本综述归纳了分子标记技术在昆虫入侵机制研究中的应用,以典型的研究个案为例,分别综述了分子标记技术在隐蔽入侵的监测应用,分子标记技术在重构入侵历史研究中的推算方式,分子标记技术在探索种群遗传变异与成功入侵机制方面取得的重要进展,并进一步介绍了高分辨率熔解曲线（high-resolution melting, HRM）分析在昆虫入侵研究中的应用前景。     

基于分子生物学方法的外来入侵物种入侵历史重构

生物入侵是一个世界性的问题。全球每年因生物入侵造成的损失超过1万亿美元。探究入侵物种在入侵地的入侵历史对了解生物入侵的生物生态学机制、制定阻截及防除措施有重要意义。分子标记方法的兴起和大规模应用打开了入侵生物入侵历史研究的新天地。采用分子标记的方法可鉴定入侵物种的种类、追溯其来源地、回溯其扩散路径、分析扩散模式及探究物种入侵过程中对入侵种群本身的变化及其对生态系统所造成的各种影响。分子标记的应用使得多个入侵物种的入侵历史得以重现。由于分子标记方法重构的入侵历史受采样范围、采用的分子标记的种类及数量等因素的影响,该方法呈现入侵历史是否是真实发生的入侵过程还存在争议。     

Genomics reveals the history of a complex plant invasion and improves the management of a biological invasion from the South African–Australian biotic exchange

Many plants exchanged in the global redistribution of species in the last 200 years, particularly between South Africa and Australia, have become threatening invasive species in their introduced range. Refining our understanding of the genetic diversity and population structure of native and alien populations, introduction pathways, propagule pressure, naturalization, and initial spread, can transform the effectiveness of management and prevention of further introductions. We used 20,221 single nucleotide polymorphisms to reconstruct the invasion of a coastal shrub, Chrysanthemoides monilifera ssp. rotundata (bitou bush) from South Africa, into eastern Australia (EAU), and Western Australia (WAU). We determined genetic diversity and population structure across the native and introduced ranges and compared hypothesized invasion scenarios using Bayesian modeling. We detected considerable genetic structure in the native range, as well as differentiation between populations in the native and introduced range. Phylogenetic analysis showed the introduced samples to be most closely related to the southern‐most native populations, although Bayesian analysis inferred introduction from a ghost population. We detected strong genetic bottlenecks during the founding of both the EAU and WAU populations. It is likely that the WAU population was introduced from EAU, possibly involving an unsampled ghost population. The number of private alleles and polymorphic SNPs successively decreased from South Africa to EAU to WAU, although heterozygosity remained high. That bitou bush remains an invasion threat in EAU, despite reduced genetic diversity, provides a cautionary biosecurity message regarding the risk of introduction of potentially invasive species via shipping routes.     

Introduction history and invasion success in exotic vines introduced to Australia

The ecological damage caused by invasive vines poses a considerable threat to many natural ecosystems. However, very little data are available for this potentially environmentally destructive functional group in Australia. In order to address this paucity of information, we assembled the first inventory of exotic vines that have become established in natural ecosystems across Australia. The influence that introduction history attributes, variables that relate to the introduction of a species to a new area, may have on the occurrence and distribution of exotic vines was also determined. We asked whether the continent of origin, reason for introduction, and residence time related to the prevalence and distribution of exotic vines across Australia. A total of 179 exotic climbing plant species from 40 different families were found to have become established across continental Australia. However, five families accounted for over 50% of these species. Most exotic vines originated from South America, and were introduced for ornamental purposes. The length of time in which an exotic vine had been present in its new range was significantly related to its distribution, with a positive relationship found between residence time and area of occupancy across the continent. No other introduction history attribute was significantly related to the area of occupancy, or distribution, of a species. This suggests that while the trends found among introduction history attributes are important in explaining the prevalence of exotic vines in Australia, only residence time is currently a useful predictor of their future success.     

A simulation model of plant invasion: long-distance dispersal determines the pattern of spread

Mechanisms and consequences of biological invasions are a global issue. Yet, one of the key aspects, the initial phase of invasion, is rarely observed in detail. Data from aerial photographs covering the spread of Heracleum mantegazzianum (Apiaceae, native to Caucasus) on a local scale of hectares in the Czech Republic from the beginning of invasion were used as an input for an individual-based model (IBM), based on small-scale and short-time data. To capture the population development inferred from the photographs, long-distance seed dispersal, changes in landscape structures and suitability of landscape elements to invasion by H. mantegazzianum were implemented in the model. The model was used to address (1) the role of long-distance dispersal in regional invasion dynamics, and (2) the effect of land-use changes on the progress of the invasion. Simulations showed that already small fractions of seed subjected to long-distance dispersal, as determined by systematic comparison of field data and modelling results, had an over-proportional effect on the spread of this species. The effect of land-use changes on the simulated course of invasion depends on the actual level of habitat saturation; it is larger for populations covering a high proportion of available habitat area than for those in the initial phase of invasion. Our results indicate how empirical field data and model outputs can be linked more closely with each other to improve the understanding of invasion dynamics. The multi-level, but nevertheless simple structure of our model suggests that it can be used for studying the spread of similar species invading in comparable landscapes.     

Human impacts, energy availability and invasion across Southern Ocean Islands

Aim Ongoing biological invasions will enhance the impacts of humans on biodiversity. Nonetheless, the effects of exotic species on diversity are idiosyncratic. Increases in diversity might be a consequence of similar responses by species to available energy, or because of positive relationships between human density, energy and propagule pressure. Here we use data from the Southern Ocean island plants and insects to investigate these issues. Location The Southern Ocean Islands ranging from Tristan da Cunha to Heard Island and South Georgia. Methods Generalized linear models are used to explore the relationships between indigenous and exotic species richness for plants and insects on two different islands. Similar models are used to examine interactions between indigenous and exotic species richness, energy availability and propagule pressure at the regional scale. Results Positive relationships were found between indigenous and exotic species richness at local scales, although for plants, the relationship was partially triangular. Across the Southern Ocean Islands, there was strong positive covariation between indigenous and exotic plant species richness and insect species richness, even taking spatial autocorrelation into account. Both exotic and indigenous plant and insect species richness covaried with energy availability, as did human visitor frequency. When two islands with almost identical numbers of human visits were contrasted, it was clear that energy availability, or perhaps differences in climate‐matching, were responsible for differences in the extent of invasion. Conclusion In plants and insects, there are positive relationships between indigenous and exotic diversity at local and regional scales across the Southern Ocean islands. These relationships are apparently a consequence of similar responses by both groups and by human occupants to available energy. When visitor frequency is held constant, energy availability is the major correlate of exotic species richness, though the exact mechanistic cause of this relationship requires clarification.     

Differences in evolutionary history translate into differences in invasion success of alien mammals in South Africa

Attempts to investigate the drivers of invasion success are generally limited to the biological and evolutionary traits distinguishing native from introduced species. Although alien species introduced to the same recipient environment differ in their invasion intensity – for example, some are “strong invaders”; others are “weak invaders” – the factors underlying the variation in invasion success within alien communities are little explored. In this study, we ask what drives the variation in invasion success of alien mammals in South Africa. First, we tested for taxonomic and phylogenetic signal in invasion intensity. Second, we reconstructed predictive models of the variation in invasion intensity among alien mammals using the generalized linear mixed‐effects models. We found that the family Bovidae and the order Artiodactyla contained more “strong invaders” than expected by chance, and that such taxonomic signal did not translate into phylogenetic selectivity. In addition, our study indicates that latitude, gestation length, social group size, and human population density are only marginal determinant of the variation in invasion success. However, we found that evolutionary distinctiveness – a parameter characterising the uniqueness of each alien species – is the most important predictive variable. Our results indicate that the invasive behavior of alien mammals may have been “fingerprinted” in their evolutionary past, and that evolutionary history might capture beyond ecological, biological and life‐history traits usually prioritized in predictive modeling of invasion success. These findings have applicability to the management of alien mammals in South Africa.     

The role of life history traits in mammalian invasion success

Why some organisms become invasive when introduced into novel regions while others fail to even establish is a fundamental question in ecology. Barriers to success are expected to filter species at each stage along the invasion pathway. No study to date, however, has investigated how species traits associate with success from introduction to spread at a large spatial scale in any group. Using the largest data set of mammalian introductions at the global scale and recently developed phylogenetic comparative methods, we show that human‐mediated introductions considerably bias which species have the opportunity to become invasive, as highly productive mammals with longer reproductive lifespans are far more likely to be introduced. Subsequently, greater reproductive output and higher introduction effort are associated with success at both the establishment and spread stages. High productivity thus supports population growth and invasion success, with barriers at each invasion stage filtering species with progressively greater fecundity.     

Historical land-use legacy and Cortaderia selloana invasion in the Mediterranean region

Two major components of global change: land‐use changes and intentional or accidental species introduction are threatening the conservation of native species worldwide. In particular, Mediterranean coastal areas are highly susceptible to the invasion of alien species and they also have experienced major changes in land use such as agricultural abandonment and urbanization. However, there has been little research done which quantitatively links biological invasions and the components of land‐use changes (i.e. number, trajectory and direction of the changes). We analysed the current distribution and abundance of Cortaderia selloana (Schultes et Schultes fil.) Asch. et Graebner, an alien ornamental species, in 332 fields in Aiguamolls de l'Empordà (Catalonia, NE Spain) and related the patterns of invasion to spatiotemporal data on land‐use changes from 1956 to 2003. Our aim was to determine which land uses had been more susceptible to C. selloana invasion during the last 5 years and to find out which components of land‐use changes triggered invasion. We found that 22.30% of the fields are currently invaded. In the last 5 years, fields have triplicated the total density of C. selloana. The presence of C. selloana decreases with the distance from urban areas. Invasion is over‐represented in pastures and old‐fields, and it has increased with time since abandonment. The presence of C. selloana was also associated to fields that had experienced many changes in land use in the last 46 years. The most heavily invaded fields were those that were pastures in 1956 and are now old fields in 2003. On average, the largest plants are found in agricultural field margins and in fields that had a disturbed land use both in 1956 and in 2003. Furthermore, pastures had the lowest proportion of reproductive plants. Overall, current C. selloana patterns of invasion can be explained by the historical legacy of land‐use changes.     

The invasion of the carnivorous carabid beetle Trechisibus antarcticus on South Georgia (sub-Antarctic) and its effect on the endemic herbivorous beetle Hydromedion spasutum

Recently two species of carabid beetle were accidentally introduced onto the sub-Antarctic island of South Georgia. Both species are carnivorous and flightless. One of the species, Trechisibus antarcticus, is locally very abundant and in the process of invading the coastal lowland area, where the endemic herbivorous beetle Hydromedion sparsutum (Perimylopidae) is common. Field samples showed the abundance of the endemic species to be much lower, and its adult body size to be larger, in carabid-infested locations than in carabid-free locations. The sample data allowed us to estimate the growth rate of the H. sparsutum larvae and to reconstruct the most likely life-cycle of both species. A laboratory experiment showed a high mortality for the first three (out of six) larval instars of H. sparsutum in groups which had been subjected to predation by T. antarcticus. The duration of the period during which the larvae are vulnerable to predation was shown in a growth experiment to depend on food type. Quantitative and qualitative aspects of the interaction between the introduced predator and the endemic prey, and conditions which allowed the former to invade are discussed.     

Fast life history traits promote invasion success in amphibians and reptiles

Competing theoretical models make different predictions on which life history strategies facilitate growth of small populations. While ‘fast’ strategies allow for rapid increase in population size and limit vulnerability to stochastic events, ‘slow’ strategies and bet‐hedging may reduce variance in vital rates in response to stochasticity. We test these predictions using biological invasions since founder alien populations start small, compiling the largest dataset yet of global herpetological introductions and life history traits. Using state‐of‐the‐art phylogenetic comparative methods, we show that successful invaders have fast traits, such as large and frequent clutches, at both establishment and spread stages. These results, together with recent findings in mammals and plants, support ‘fast advantage’ models and the importance of high potential population growth rate. Conversely, successful alien birds are bet‐hedgers. We propose that transient population dynamics and differences in longevity and behavioural flexibility can help reconcile apparently contrasting results across terrestrial vertebrate classes.     

Life history shifts in an exploited African fish following invasion by a castrating parasite

Evolutionary theory predicts that infection by a parasite that reduces future host survival or fecundity should select for increased investment in current reproduction. In this study, we use the cestode Ligula intestinalis and its intermediate fish host Engraulicypris sardella in Wissman Bay, Lake Nyasa (Tanzania), as a model system. Using data about infection of E. sardella fish hosts by L. intestinalis collected for a period of 10 years, we explored whether parasite infection affects the fecundity of the fish host E. sardella, and whether host reproductive investment has increased at the expense of somatic growth. We found that L. intestinalis had a strong negative effect on the fecundity of its intermediate fish host. For the noninfected fish, we observed an increase in relative gonadal weight at maturity over the study period, while size at maturity decreased. These findings suggest that the life history of E. sardella has been shifting toward earlier reproduction. Further studies are warranted to assess whether these changes reflect plastic or evolutionary responses. We also discuss the interaction between parasite and fishery‐mediated selection as a possible explanation for the decline of E. sardella stock in the lake.     

Understanding invasion history: genetic structure and diversity of two globally invasive plants and implications for their management

Aim Resolving the origin of invasive plant species is important for understanding the introduction histories of successful invaders and aiding strategies aimed at their management. This study aimed to infer the number and origin(s) of introduction for the globally invasive species, Macfadyena unguis‐cati and Jatropha gossypiifolia using molecular data. Location Native range: Neotropics; Invaded range: North America, Africa, Europe, Asia, Pacific Islands and Australia. Methods We used chloroplast microsatellites (cpSSRs) to elucidate the origin(s) of introduced populations and calculated the genetic diversity in native and introduced regions. Results Strong genetic structure was found within the native range of M. unguis‐cati, but no genetic structuring was evident in the native range of J. gossypiifolia. Overall, 27 haplotypes were found in the native range of M. unguis‐cati. Only four haplotypes were found in the introduced range, with more than 96% of introduced specimens matching a haplotype from Paraguay. In contrast, 15 haplotypes were found in the introduced range of J. gossypiifolia, with all invasive populations, except New Caledonia, comprising multiple haplotypes. Main conclusions These data show that two invasive plant species from the same native range have had vastly different introduction histories in their non‐native ranges. Invasive populations of M. unguis‐cati probably came from a single or few independent introductions, whereas most invasive J. gossypiifolia populations arose from multiple introductions or alternatively from a representative sample of genetic diversity from a panmictic native range. As introduced M. unguis‐cati populations are dominated by a single haplotype, locally adapted natural enemies should make the best control agents. However, invasive populations of J. gossypiifolia are genetically diverse and the selection of bio‐control agents will be considerably more complex.     

Cold adaptation in the Asian tiger mosquito's native range precedes its invasion success in temperate regions

Adaptation to environmental conditions within the native range of exotic species can condition the invasion success of these species outside their range. The striking success of the Asian tiger mosquito, Aedes albopictus, to invade temperate regions has been attributed to the winter survival of diapause eggs in cold environments. In this study, we evaluate genetic polymorphisms (SNPs) and wing morphometric variation among three biogeographical regions of the native range of A. albopictus. Reconstructed demographic histories of populations show an initial expansion in Southeast Asia and suggest that marine regression during late Pleistocene and climate warming after the last glacial period favored expansion of populations in southern and northern regions, respectively. Searching for genomic signatures of selection, we identified significantly differentiated SNPs among which several are located in or within 20 kb distance from candidate genes for cold adaptation. These genes involve cellular and metabolic processes and several of them have been shown to be differentially expressed under diapausing conditions. The three biogeographical regions also differ for wing size and shape, and wing size increases with latitude supporting Bergmann's rule. Adaptive genetic and morphometric variation observed along the climatic gradient of A. albopictus native range suggests that colonization of northern latitudes promoted adaptation to cold environments prior to its worldwide invasion.     

Resilience of native ant community against invasion of exotic ants after anthropogenic disturbances of forest habitats

The positive association between disturbances and biological invasions is a widely observed ecological pattern in the Anthropocene. Such patterns have been hypothesized to be driven by the superior competitive ability of invaders or by modified environments, as well as by the interaction of these factors. An experimental study that tests these hypotheses is usually less feasible, especially in protected nature areas. An alternative approach is to focus on community resilience over time after the anthropogenic disturbance of habitats. Here, we focused on ant communities within a forest to examine their responses after disturbance over time. We selected the Yanbaru region of northern Okinawa Island, which is a biodiversity hotspot in East Asia. We compared ant communities among roadside environments in forests where the road age differed from 5 to 25 years. We also monitored the ant communities before and after disturbance from forest thinning. We found that the species richness and abundance of exotic ants were higher in recently disturbed environments (roadsides of 5–15 years old roads), where the physical environment was warmer and drier. In contrast, the roadsides of 25‐year‐old roads indicated the potential recovery of the physical environment with cooler and moister conditions, likely owing to regrowth of roadside vegetation. At these sites, there were few exotic ants, except for those immediately adjacent to the road. The population density of the invasive species Technoymex brunneus substantially increased 1–2 years after forest thinning. There was no evidence of the exclusion of native ants by exotic ants that were recorded after disturbance. Our results suggest that local ant communities in the Yanbaru forests have some resilience to disturbance. We suggest that restoration of environmental components is a better strategy for maintaining native ant communities, rather than removing exotic ants after anthropogenic disturbance.     

Global mtDNA genetic structure and hypothesized invasion history of a major pest of citrus,Diaphorina citri (Hemiptera: Liviidae)

The Asian citrus psyllid Diaphorina citri Kuwayama is a key pest of citrus as the vector of the bacterium causing the “huanglongbing” disease (HLB). To assess the global mtDNA population genetic structure, and possible dispersal history of the pest, we investigated genetic variation at the COI gene collating newly collected samples with all previously published data. Our dataset consists of 356 colonies from 106 geographic sites worldwide. High haplotype diversity (H‐mean = 0.702 ± 0.017), low nucleotide diversity (π‐mean = 0.003), and significant positive selection (Ka/Ks = 32.92) were observed. Forty‐four haplotypes (Hap) were identified, clustered into two matrilines: Both occur in southeastern and southern Asia, North and South America, and Africa; lineages A and B also occur in eastern and western Asia, respectively. The most abundant haplotypes were Hap4 in lineage A (35.67%), and Hap9 in lineage B (41.29%). The haplotype network identified them as the ancestral haplotypes within their respective lineages. Analysis of molecular variance showed significant genetic structure (F_ST = 0.62, p < .0001) between the lineages, and population genetic analysis suggests geographic structuring. We hypothesize a southern and/or southeastern Asia origin, three dispersal routes, and parallel expansions of two lineages. The hypothesized first route involved the expansion of lineage B from southern Asia into North America via West Asia. The second, the expansion of some lineage A individuals from Southeast Asia into East Asia, and the third involved both lineages from Southeast Asia spreading westward into Africa and subsequently into South America. To test these hypotheses and gain a deeper understanding of the global history of D. citri, more data‐rich approaches will be necessary from the ample toolkit of next‐generation sequencing (NGS). However, this study may serve to guide such sampling and in the development of biological control programs against the global pest D. citri.