Invasive alien Crustacea: dispersal,establishment, impact and control

The subphylum Crustacea includes the most successful species among aquatic alien invaders. The impacts of invasive alien crustaceans (IAC) are often substantial, due to the complex trophic role of most of these species leading to cascading effects throughout the invaded ecosystems. IAC also have the potential to cause a shift in the ‘keystone’ ecosystem functions, changing energy flux and nutrient cycles which together affect critical ecosystem services such as biodiversity, fisheries yield and water quality. Although no individual trait appears to be a good predictor of invasion success, a combination of some characteristics such as eurytolerance, omnivory and certain r-selected life-history traits results in a high probability of alien crustacean species becoming invasive. Both environmental factors, such as habitat heterogeneity in the invaded ecosystems, and evolutionary factors, such as adaptations to new environmental conditions, also play important roles during establishment. Therefore, individual environmental niche models, including genetic algorithm, have the highest likelihood of providing useful predictive information about invasion success and spread of alien Crustacea. Attempts to control IAC through biocides or mechanical removal have had mixed success in the past but a strategic combination of different methods may lead to some success in the future.     

Commercially Important Trees as Invasive Aliens – Towards Spatially Explicit Risk Assessment at a National Scale

Alien species that are desirable and commercially important in parts of the landscape, but damaging invaders in other parts, present a special challenge for managers, planners, and policy-makers. Objective methods are needed for identifying areas where control measures should be focussed. We analysed the distribution of forestry plantations and invasive (self-sown) stands of Acacia mearnsii and Pinus spp. in South Africa; these two taxa account for 60% of the area under commercial plantations and 54% of the area invaded by alien trees and shrubs. The distribution of commercial forestry plantations and invasive stands of these taxa were mapped and the data was digitised and stored as Geographic Information System (GIS) (Arc/Info) layers. A series of environmental parameters were derived from GIS layers of climate, topography, geology, land use, and natural vegetation. The current distribution of the two taxa was subdivided into three groups according to the degree of invasion, planting history and the precision of the data collection. We used regression-tree analysis to relate, for each taxon, the distribution of invasive stands with environmental variables, and to derive habitat suitability maps for future invasion. The current distribution of invasive stands in South Africa was largely influenced by climatic factors. At a national scale, the distribution of large commercial plantations was a poor predictor of areas invaded by both taxa. Using environmental factors identified by the regression trees, we found that 6.6% and 9.8% of natural habitats currently not invaded and untransformed by urbanisation or agriculture are suitable for invasion by Pinus spp. and A. mearnsii, respectively. We then derived guidelines for policy on alien plant management based on vegetation type, degree of transformation, extent of invasion, and the risk of future alien spread. These factors were used to identify demarcated areas where these alien species can be grown with little risk of invasions, and areas where special measures are needed to manage spread from plantations.     

Restoring habitat for native and endemic plants through the introduction of a fungal pathogen to control the alien invasive tree <Emphasis Type="Italic">Miconia calvescens</Emphasis> in the island of Tahiti

Success of biological control programs is commonly assessed by studying the direct negative impacts of released agents on the target invasive species. Very few quantitative studies have focused on the indirect positive effects on native biodiversity. In this study, we monitored the response of the plant community (both native and alien species) in permanent plots located in four different sites in montane rainforests of the tropical island of Tahiti (South Pacific) severely invaded over decades by the alien invasive tree Miconia calvescens DC (Melastomataceae), after the release of a defoliating fungal pathogen Colletotrichum gloeosporioides f. sp. miconiae Killgore & L. Sugiyama. Results of five years of monitoring showed that total native and endemic species richness and plant cover increased in all sites and plots. Partial defoliation of miconia canopy trees (between 6% and 36%) led to significant recruitment of light-demanding pioneer species, but also to the appearance of some semi-shade and shade tolerant rare endemic species. Native ferns and angiosperms remained dominant (ca. 80%) in the forest understorey during the monitoring period. Colonization by a small number of alien plant species occurred in one permanent plot located at the lower elevation. We conclude that biological control may be considered a tool for partial habitat restoration and recovery of native and endemic species, but long-term monitoring is needed to confirm the stability and resilience of the “novel plant assemblage”.     

Impact assessment revisited: improving the theoretical basis for management of invasive alien species

The theoretical underpinnings of the assessment of invasive alien species impacts need to be improved. At present most approaches are unreliable to quantify impact at regional scales and do not allow for comparison of different invasive species. There are four basic problems that need to be addressed: (1) Some impacted ecosystem traits are spatially not additive; (2) invader effects may increase non-linearly with abundance or there may be effect thresholds impairing estimates of linear impact models; (3) the abundance and impact of alien species will often co-vary with environmental variation; and (4) the total invaded range is an inappropriate measure for quantifying regional impact because the habitat area available for invasion can vary markedly among invasive species. Mathematical models and empirical data using an invasive alien plant species (Heracleum mantegazzianum) indicate that ignoring these issues leads to impact estimates almost an order of magnitude from the real values. Thus, we propose a habitat-sensitive formula for regional impact assessment that is unaffected by non-linearity. Furthermore, we make some statistical suggestions on how to assess invader effects properly and we discuss the quantification of the invaded range. These improvements are crucial for impact assessment with the overall aim of prioritizing management of invasive species.     

Beetle responses to artificial gaps in an oceanic island forest: implications for invasive tree management to conserve endemic species diversity

Natural forests are often replaced by invasive alien trees on isolated oceanic islands. Adequate eradication of invasive trees should be conducted with the goal of biodiversity conservation, because islands support many endemic organisms that depend on native forests. An invasive alien tree, Bischofia javanica Blume (Euphorbiaceae), has invaded and replaced natural forests on the oceanic Ogasawara (Bonin) Islands, Japan, in the northwestern Pacific Ocean. To determine how the removal of B. javanica trees affects insect diversity, we examined flying beetles captured using Malaise traps in B. javanica forests on Hahajima. The abundance, species density, and species composition of wood-boring beetles (Coleoptera: Cerambycidae, Elateridae, Mordellidae, and Scolytidae) were compared between closed-canopy sites and gaps created by girdling B. javanica trees in alien forests during two seasons (June–July and October–November 2005). Of the collected beetles, 75.8, 87.5, 90.0, and 0.0% of cerambycid, elaterid, mordellid, and scolytid beetle species, respectively, were endemic to the Ogasawara Islands. More cerambycid, elaterid, and mordellid individuals were captured in June–July than in October–November; the number of scolytid individuals did not differ between seasons. More cerambycid, elaterid, and scolytid individuals were captured in artificial gaps than on the closed-canopy forest floor. Although fewer mordellid individuals were captured in gaps, more endemic mordellids were captured in gaps. More cerambycid and scolytid species were captured in artificial gaps than in closed-canopy areas. The positive responses of beetles to artificial gaps suggest that the removal of B. javanica increases beetle diversity and the abundance of endemic beetles.     

Contrasting effects of invasive plants in plant–pollinator networks

The structural organization of mutualism networks, typified by interspecific positive interactions, is important to maintain community diversity. However, there is little information available about the effect of introduced species on the structure of such networks. We compared uninvaded and invaded ecological communities, to examine how two species of invasive plants with large and showy flowers (Carpobrotus affine acinaciformis and Opuntia stricta) affect the structure of Mediterranean plant–pollinator networks. To attribute differences in pollination to the direct presence of the invasive species, areas were surveyed that contained similar native plant species cover, diversity and floral composition, with or without the invaders. Both invasive plant species received significantly more pollinator visits than any native species and invaders interacted strongly with pollinators. Overall, the pollinator community richness was similar in invaded and uninvaded plots, and only a few generalist pollinators visited invasive species exclusively. Invasive plants acted as pollination super generalists. The two species studied were visited by 43% and 31% of the total insect taxa in the community, respectively, suggesting they play a central role in the plant–pollinator networks. Carpobrotus and Opuntia had contrasting effects on pollinator visitation rates to native plants: Carpobrotus facilitated the visit of pollinators to native species, whereas Opuntia competed for pollinators with native species, increasing the nestedness of the plant–pollinator network. These results indicate that the introduction of a new species to a community can have important consequences for the structure of the plant–pollinator network.     

Invasion of alien macroalgae in different Mediterranean habitats

The study evaluated different macroalgal invasions in the main Mediterranean coastal habitats on hard bottom. Biodiversity, species composition and structure of macroalgal assemblages were compared among non-invaded areas and areas invaded by the Chlorophyta Caulerpa racemosa var. cylindracea and by the turf-forming Rhodophyta Womersleyella setacea in three different habitats: shallow rocky bottom, deep rocky bottom and dead matte of the seagrass Posidonia oceanica. Results showed that alien macroalgae constituted a relevant component of benthic assemblages in invaded areas of the Mediterranean Sea. Assemblages invaded by Womersleyella setacea and Caulerpa racemosa showed lower values of diversity and large differences in the structure and species composition related to non-nvaded assemblages. The species that mostly suffered from invasion were erect species reproducing sexually; moreover, the dominance of W. setacea led to low abundance of native filamentous algae, while C. racemosa colonization seemed particularly threatening for encrusting algae. All the studied habitats appeared highly invasible by alien macroalgae, even if W. setacea appeared more invasive in deeper habitats, while colonization of C. racemosa seemed more serious in shallower habitats; the dead matte of P. oceanica represented a suitable substrate for the spread of both species. Differences among assemblages in different habitats were reduced in invaded areas.     

重要外来入侵植物随南水北调工程传入京津冀受水区的风险评估

【目的】外来物种在入侵地的扩张蔓延是其造成危害的关键。南水北调工程可为已入侵外来物种的进一步扩张蔓延提供通道和驱动力。风险评估可为预防入侵植物通过该途径的扩散蔓延提供依据。【方法】以国家重点管理外来入侵物种名录中的入侵植物为研究对象,根据生物入侵的发生过程(传入、定殖、扩散、危害)构建了包括4个层次29个指标的外来物种入侵综合风险评估的指标体系,规范了风险指数的计算方法,系统评估了24种入侵植物沿南水北调输水通道向北扩散而入侵京津冀受水区的风险。【结果】紫茎泽兰和互花米草的入侵风险值(R)相对较低(R0.6),其余22种入侵植物的风险值均较高(R0.6),其中,喜旱莲子草、凤眼莲和大薸3种外来水生植物的入侵风险等级最高(R0.8)。【结论】南水北调工程可能会促进入侵植物向北扩张蔓延并最终入侵京津冀受水区。建议尽早开展监测预警工作,以控制入侵物种随南水北调工程的扩张蔓延,进而阻止或降低其对京津冀尤其是华北最大水源地——白洋淀和雄安新区生态环境的威胁。     

Consistent performance of invasive plant species within and among islands of the Mediterranean basin

Since the success of an invasive species depends not only upon its intrinsic traits but also on particular characteristics of the recipient habitat, assessing the performance of an invader across habitats provides a more realistic analysis of risk. Such an analysis will not only provide insights into the traits related to invasiveness, but also the habitat characteristics that underpin vulnerability to invasion that, taken together, will facilitate the selection of management strategies to mitigate the invader’s effect. In the present study, we considered the Mediterranean basin islands as an excellent study region to test how the same invasive species perform in different habitats within a single island, and to scale up differences among islands with similar climate. We tested how the performance of three widespread plant invaders with clonal growth but contrasting life-history traits, a deciduous tree Ailanthus altissima, a succulent subshrub Carpobrotus spp., and an annual geophyte Oxalis pes-caprae, varied depending upon the species identity, habitat, and invaded island. The environmental parameters considered were habitat type, elevation, species diversity in the invaded plot, and several soil traits (% C, % N, C/N, pH, and relative humidity). The study documents that the performance of these three important and widespread plant invaders is dependent mainly on species identity, and less upon the invaded island’s general features. Likewise, differences in performance among habitats were only significant in the case of Ailanthus, whereas Carpobrotus and Oxalis appear to perform equally well in different environments. Ailanthus thus appears to have a broader spectrum of invasiveness, being able to invade a larger number of habitat types. On the contrary, Carpobrotus spp. have not yet invaded habitats different from those where the species have been originally introduced and where they are still commonly spread by humans. Oxalis distribution is mainly related to agricultural activities and disturbed sites, and the total area infested by this geophyte may be more reflection of the extent of suitable habitats than of invasiveness or ecological impact. Our results confirm the potential for these species to significantly alter the functioning of ecosystems in the Mediterranean islands and highlight the risk to other islands not yet invaded.     

An exotic Australian <Emphasis Type="Italic">Acacia</Emphasis> fixes more N than a coexisting indigenous <Emphasis Type="Italic">Acacia</Emphasis> in a South African riparian zone

Acacia mearnsii is an introduced Australian acacia in South Africa and has invaded more than 2.5 million ha, primarily establishing in rangeland and riparian areas. Because acacias have the capability to fix N, A. mearnsii invasions may fundamentally change N dynamics in invaded systems. This study compares biological N₂-fixation in the alien invasive A. mearnsii and the native A. caffra growing in a grassland riparian zone in the Komati Gorge Reserve, Mpumalanga, South Africa. A ¹⁵N natural abundance field survey suggested that both mature alien and native acacias fix N under current conditions in the riparian zone. Significantly depleted δ¹⁵N was observed in both acacias relative to reference species, although variation in δ¹⁵N was not correlated with N concentrations. Calculated contributions of N₂-fixation (%Ndfa) suggest that alien acacias fix significantly more of their N than native acacias (~75 ± 5% SE and 53 ± 9% SE, respectively). There was a larger variation in δ¹⁵N and %Ndfa in the native acacia, suggesting relatively high plasticity in its N₂-fixation contributions. This plasticity was interpreted as a facultative N₂-fixation strategy for the native acacia, while the N₂-fixation strategy of the alien acacia remained unclear. Our results emphasize the importance of potentially elevated N inputs through N₂-fixation by invasive legumes in invaded landscapes. Furthermore, they suggest that N₂-fixation by invasive acacias may not respond to fine-scale patchiness in soil N in the same manner as native acacias, making them potential contributors to N excess in Southern Africa.     

Current distribution and potential extent of the most invasive alien plant species on La Réunion (Indian Ocean,Mascarene islands)

Abstract La Réunion Island has the largest area of intact vegetation of the islands in the Mascarene archipelago. Biological invasions are the primary threat to biodiversity in the intact habitats of the island (those not already transformed by agriculture and urbanization). Our study aimed to identify areas to prioritize in managing invasive alien plants for biodiversity conservation. We used extensive surveys of 238 distinct untransformed areas on La Réunion to define the current distribution patterns of all invasive species. Using expert knowledge, we compiled maps of the current distribution of the 46 most widespread/important invasive plants at the habitat scale (identified according to vegetation structure). Data from 440 botanical relevés for the 20 most threatening invasive alien plant species across the island and climatic envelope models were used to derive climatic suitability surfaces; these were used to map potential distributions for these species. More than 10 species invade 16.7% of the remaining habitat. Five habitat types are invaded by 25 or more species, and eight have fewer than 10 invasive alien plant species. Cluster analysis based on presence/absence of species in the 18 habitat types produced eight groups of species that invade particular habitats. Potential distribution models show that some species have invaded large parts of their potential range (e.g. Fuchsia magellanica, Furcraea foetida, Hiptage benghalensis), whereas others have the potential to increase their range substantially (e.g. Clidemia hirta, Strobilanthes hamiltonianus, Ulex europaeus). Management implications are identified for both groups. Three broad groups of habitats were identified: (i) intact habitats with a low level of invasion (e.g. subalpine shrubland); (ii) moderately invaded habitats with varying levels of intactness (ranging from windward submountain rainforest to the Acacia heterophylla forest); and (iii) habitats with little remaining intact area and high levels of invasion (e.g. lowland rainforest). Different management interventions are appropriate for these three groups.     

Recovery of indigenous butterfly community following control of invasive alien plants in a tropical island’s wet forests

Invasive alien species pose one of the highest threats to biodiversity, especially in isolated oceanic islands where high rates of both endemism and extinction risk also usually prevail. Few studies have investigated the impact of invasive alien plants on butterflies in insular ecosystems, despite butterflies representing a key indicator group for terrestrial arthropod diversity. Using the Pollard Technique, we quantified butterfly species richness and abundance in eight wet lowland forest areas invaded by alien plants, principally the strawberry guava (Psidium cattleianum Sabine) on the tropical volcanic island of Mauritius, and compared the results with paired adjacent forest plots that had been weeded of alien plants between 2 and 12 years previously. Butterfly assemblages in weed-infested and weeded forests were distinctly different with higher species richness and much higher butterfly abundance in the latter. At least some of these differences seemed attributable to weed removal effects on forest structure, but understanding the precise mechanisms involved will require further study. The results suggest that alien plant invasion may have contributed to the extinction of certain endemic taxa and can increase the likelihood of butterfly species extinction by reducing population sizes through reduced habitat quality. Such a shift in a forest’s butterfly assemblage is likely to have negative effects on both their indigenous predators and the plants they pollinate. It is argued that in order to maintain butterfly and other arthropod diversity and function in these forests, alien plant control must be maintained and extended beyond the current 1% of surviving forest remnants.     

苋科(广义)入侵植物墙生藜在中国的新记录

外来物种的归化和入侵对全球环境和社会发展造成了严重影响，已成为当今各国生物多样性管理和生态保护中所面临的全球性问题。我国是遭受外来入侵危害最为严重的国家之一，在外来物种入侵的预警、管理和治理等方面形势严峻。基于野外调查和文献研究，该文报道了苋科(广义)入侵植物墙生藜[Chenopodiastrum murale(L.) S. Fuentes, Uotila&Borsch]在中国的新记录。墙生藜原产于地中海地区，现已扩散到欧洲、美洲、非洲和大洋洲的40多个国家，是一种危害较大的外来入侵植物，同时也是我国海关和检验检疫部门明确规定禁止入境的检疫性有害生物，现于云南省昆明市呈贡区发现该外来入侵植物。该文对其形态特征进行了详细描述，简要介绍了其分类历史，并提供了可供鉴定比对的野外生态照片；此外，对墙生藜可能的传入途径进行了分析，对其危害和风险作了简要评估。该物种的新发现说明我国外来入侵生物的本底调查还存在不足。     

An Ecological Imbalance Induced by a Non-Native Species: The Manila Clam in the Venice Lagoon

Among the 19 non-native species of marine invertebrates which have invaded the Venice Lagoon and have established populations, Ruditapes philippinarum, deliberately introduced in 1983, is surely the most successful species. According to the hypothesis that alien species invasion could be favoured by an altered ecological, chemical or physical state of the system induced by anthropogenic disturbance, R. philippinarum turned out to be ‘the right species at the right moment’. By comparing historical data (1968, 1985, 1990) with 1999 data, changes in macrobenthic community, in particular bivalve molluscs, of the lagoon induced by R. philippinarum introduction and subsequent clam exploiting activity were assessed. It has been possible to describe a sharp reduction, both in terms of distribution area and density, of all other filter feeder bivalves. Moreover, by using the clearance rate of the most abundant bivalve species in 1990 and 1999 (Cerastoderma glaucum and R. philippinarum, respectively), it was possible to estimate that the filtration capacity, expressed as l h⁻¹ m⁻², has more than doubled. This has altered the functioning of the ecosystem, resulting in a stronger benthic–pelagic coupling. In this context, R. philippinarum attains control of the system. Considering all this, it is possible to state that the Venice Lagoon ecosystem has entered into a new state, probably more resistant but less resilient, with implications for future management choices.     

Impacts of Myriophyllum aquaticum invasion in a Mediterranean wetland on plant and macro-arthropod communities

The invasion by alien macrophytes in aquatic ecosystems may produce a strong alteration of the native aquatic vegetation leading to heavy impacts for both plant and faunal native diversity. Myriophyllum aquaticum is an aquatic plant native of Southern America, invasive in several part of the world. We studied the effects of M. aquaticum invasion on plant and macro-arthropod communities in the canals around a protected wetland in the Mediterranean basin. We sampled plant and macro-arthropod communities in 10 transects in invaded and non-invaded tracts of the canals. We assessed the differences in plant and macro-arthropod species richness, diversity, taxonomic diversity and species composition between invaded and non-invaded habitats by means of univariate and multivariate analyses. Our study shows a significant loss of plant diversity between non-invaded to invaded sites, leading to communities numerically and taxonomically impoverished and highly divergent in the species composition. We also detected significant differences in arthropod species composition between invaded and non-invaded transects. Some taxa such as mosquitoes and malacostraca were more frequent in the M. aquaticum-dominated stands. Furthermore, the study shows a positive relation between invaded habitats and juvenile individuals of the invasive alien crayfish Procambarus clarkii.     

Impact of invasions by alien plants on soil seed bank communities: Emerging patterns

Much of our current understanding of the impact of invasive species on plant communities is based on patterns occurring in the above-ground vegetation, while only few studies have examined changes in soil seed banks associated with plant invasions, despite their important role as determinants of vegetation dynamics. Here, we reviewed the literature on the impact of plant invasions on the seed bank and we provide a quantitative synthesis using a meta-analysis approach. Specifically, (1) we quantified the impact of 18 invasive alien plants on (i) species richness and (ii) density of the seed banks of invaded communities, based on 58 pair-wise invaded-uninvaded comparisons (cases); we identified (2) the invasive taxa that are responsible for the largest changes in the seed bank; and (3) the habitats where substantial changes occur. Our study showed three major findings: (1) species richness (68% of cases) and density (58% of cases) were significantly lower in native seed banks invaded by alien plants; (2) species richness and density of native and alien species were remarkably lower in seed banks invaded by large, perennial herbs compared to uninvaded sites; and (3) invaded seed banks were often associated with a larger richness and/or abundance of alien species. This study indicates a need for additional seed bank data in invasion ecology to characterize species-specific and habitat-specific impacts of plant invasions, and to determine whether changes in the seed banks of native and alien species are a symptom of environmental degradation prior to a plant invasion or whether they are its direct result. The findings of this study help improve our capacity to predict the long-term implications of plant invasions, including limitations in the recruitment of native species from the seed bank and the potential for secondary invasions by seeds of other alien species.     

An economic assessment of the contribution of biological control to the management of invasive alien plants and to the protection of ecosystem services in South Africa

This study is a first attempt at a holistic economic evaluation of South African endeavours to manage invasive alien plants using biological control. Our focus was on the delivery of ecosystem services from habitats that are invaded by groups of weeds, rather than by each individual weed species. We established the net present value of the weed biological control efforts, and derived benefit:cost ratios by comparing this value (a cost) to the estimated value of ecosystem services protected by weed biological control. We identified four major functional groupings of invading alien plants, and assessed their impact on water resources, grazing and biodiversity. We estimated the area that remained free of invasions due to all historic control efforts in South Africa, and the proportion that remained free of invasion as a result of biological control (which was initiated in 1913). The estimated value of potential ecosystem services amounted to 152 billion South African rands (ZAR—presently, about US$ 19.7 billion) annually. Although an estimated ZAR 6.5 billion was lost every year due to invading alien plants, this would have amounted to an estimated additional ZAR 41.7 billion had no control been carried out, and 5–75% of this protection was due to biological control. The benefit:cost ratios ranged from 50:1 for invasive sub-tropical shrubs to 3,726:1 for invasive Australian trees. Benefit:cost ratios remained positive and our conclusion, that biological control has brought about a considerable level of protection of ecosystem services, remains robust even when our estimates of the economic impacts of key variables (i.e. sensitivity analyses of indeterminate variables) were substantially reduced.     

Little evidence for negative effects of an invasive alien plant on pollinator services

Many invasive alien plants occur in large populations with abundant flowers which are highly attractive to pollinators, and thus might affect pollination of co-occurring native species. This study focuses on the invasive Heracleum mantegazzianum and distance-dependent effects on pollination of Mimulus guttatus in abandoned grassland over 2 years. First, we examined pollinator abundance in yellow traps at 0, 10, 30 and 60–200 m from H. mantegazzianum. We then placed M. guttatus plants at the same distances to monitor effects of the invasive species on pollinator visitation and seed set of neighbouring plants. Finally, we conducted a garden experiment to test if deposition of H. mantegazzianum pollen reduces seed set in M. guttatus. No distance effect was found for the number of bumblebees in traps, although the invasive species attracted a diverse assemblage of insects, and visitation of M. guttatus was enhanced close to H. mantegazzianum. This positive effect was not reflected by seed set of M. guttatus, and heterospecific pollen decreased seed set in these plants. Overall there is little evidence for negative effects of the invasive species on pollination of neighbouring plants, and flower visitation even increases close to the invaded patches. The functional role of the invader and suitable control strategies need further clarification, since removal of H. mantegazzianum may actually damage local pollinator populations. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Intraguild predation involving Harmonia axyridis: a review of current knowledge and future perspectives

As an effective generalist predator of aphids and other hemipteran pests H. axyridis has been a successful biological control agent. However, the very functional traits that have contributed to its success in this regard also implicate it as an intraguild predator that poses a significant risk not only to the diversity of other natural enemies of Hemiptera (and their associated ecosystem services), but to biodiversity more widely. In this paper we will specifically review the existing data on intraguild predation involving H. axyridis, and consider the strength and symmetry of such interactions both within its native guild and within exotic guilds where it has established as an invasive alien. We will use these studies to interpret the observed population declines in predator diversity in the field, predict species at risk in regions not yet invaded and consider implications for resulting ecosystem services. We will also indicate gaps in our knowledge that require further study in order to identify opportunities for mitigation.     

Different flowering phenology of alien invasive species in Spain: evidence for the use of an empty temporal niche?

Flowering phenology is an important and poorly understood plant trait that may possibly be related to the invasiveness potential of alien species. The present work evaluates whether flowering time of invasive alien species is a key trait to overcome the climatic filters operating in continental Mediterranean ecosystems of Spain (characterised by summer drought and low temperatures in winter). We conducted comparisons between the flowering phenology of the invasive species in their native range and in Spain, and between flowering phenology of 91 coexisting invasive–native species pairs. For the alien species, geographical change from the native to the invaded region did not result in shifts in the start and the length of the flowering period. Overall, climatic conditions in the native range of species selected for a flowering pattern is maintained after translocation of the species to another region. Flowering of tropical and temperate invasive alien species peaked in summer, which contrasts with the spring flowering of native and invasive alien species of Mediterranean climate origin. By exploiting this new temporal niche, these invasive alien species native to tropical and temperate regions benefit from reduced competition with natives for abiotic and biotic resources. We suggest that human-mediated actions have reduced the strength of the summer drought filter in particular microhabitats, permitting the invasion of many summer-flowering aliens.