Population genetics of goldspotted oak borer,Agrilus auroguttatus Schaeffer (Coleoptera: Buprestidae): investigating the origin of an invasive pest of native oaks in California

The goldspotted oak borer, Agrilus auroguttatus Schaeffer, is an invasive woodborer in California USA that is native to oak woodlands across southern Arizona USA. Developing a classical biological control program for this pest in southern California is a high priority due to the continuing ecological and economic damage caused by this insect since its recent introduction into the area. In an attempt to determine the area of origin for this invasive beetle, analyses of the mitochondrial cytochrome oxidase and ribosomal nuclear D2 domain of the 28S gene regions were undertaken and provided insight into the phylogeographic relationship between and within populations of A. auroguttatus in Arizona and California. The area of origin for the invasive population of goldspotted oak borer in California was not determined conclusively, although our molecular data suggests the Dragoon Mountains in Cochise Co., Arizona as a possible source for the California population of A. auroguttatus. Results also confirmed that individuals collected from populations across southern Arizona and California are all A. auroguttatus, and are not part of a cryptic species complex comprised of the morphologically similar A. coxalis. Future surveys for natural enemies of A. auroguttatus will focus on the Dragoon Mountains as a potential source for co-evolved enemies for use in a classical biological control program against this invasive woodborer in southern California.     

Molecular evidence for a single genetic clone of invasive Arundo donax in the United States

Arundo donax (giant reed) is an aggressive invasive weed of riparian habitats throughout the southern half of the United States from California to Maryland. Native to Asia, the species is believed to have been initially introduced into North America from the Mediterranean region although subsequent introductions were from multiple regions. To provide insight into the potential for biological control of A. donax, genetic variation in plants sampled from a wide geographical area in the United States was analyzed using Sequence Related Amplification Polymorphism (SRAP) and transposable element (TE)-based molecular markers. Invasive individuals from 15 states as well as four populations in southern France were genetically fingerprinted using 10 SRAP and 12 TE-based primer combinations. With the exception of simple mutations detected in four plants, A. donax exhibited a single multilocus DNA fingerprint indicating a single genetic clone. The genetic uniformity of invasive A. donax suggests that classical biological control of the species could be successful. A lack of genetic diversity in the invaded range simplifies identification of native source populations to search for natural enemies that could be used as biocontrol agents.     

Loss of microbial (pathogen) infections associated with recent invasions of the red imported fire ant Solenopsis invicta

Loss of natural enemies during colonization is a prominent hypothesis explaining enhanced performance of invasive species in introduced areas. Numerous studies have tested this enemy release hypothesis in a wide range of taxa but few studies have focused on invasive ants. We conducted extensive surveys for the presence of six microbes in recently established populations (California, Australia, New Zealand, Taiwan, and China) of the invasive fire ant Solenopsis invicta. These microbes include Wolbachia, two microsporidia (Kneallhazia solenopsae and Vairimorpha invictae) and three RNA viruses (SINV-1, -2 and -3), all of which previously have been reported in native South American populations of S. invicta. These surveys showed that the total number of enemy species is lower in the recently invaded areas compared with both South American and US populations. Only two microbes were found in any of these recently invaded areas: SINV-1 was detected in all surveyed populations except Australia and New Zealand, and SINV-2 was detected in California and Taiwan only. These results support the general prediction that invasive species lose many of their natural enemies during invasion. Further, the conspicuous absence of some of these microbes in these areas may result from strong selection against founders due to fitness costs associated with harboring detrimental infections rather than the alternative hypothesis that they simply were absent among the original founders. While the successful invasion of S. invicta in these recently invaded areas may be explained partly by the absence of natural enemies, other factors likely have been important as well.     

Tracing the Invasion of the Mediterranean Land Snail Cornu aspersum aspersum Becoming an Agricultural and Garden Pest in Areas Recently Introduced

This study is the first on the genetics of invasive populations of one of the most widely spread land mollusc species known in the world, the “Brown Snail” Cornu aspersum aspersum. Deliberately or accidentally imported, the species has become recently a notorious pest outside its native Mediterranean range. We compared the spatial structure and genetic variability of invasive (America, Oceania, South Africa) versus native populations using five microsatellite loci and mitochondrial (Cyt b and 16S rRNA) genes as a first step towards (i) the detection of potential source populations, and (ii) a better understanding of mechanisms governing evolutionary changes involved in the invasion process. Results based on multivariate analysis (Discriminant Analysis of Principal Components), Bayesian statistical inference (Clustering, Approximate Bayesian Computation) and demographic tests allowed a construction of the introduction pathways of the species over recent centuries. While emigrants originated from only one of the two native lineages, the West one, the most likely scenario involved several introduction events and “source switching” comprising (i) an early stage (around 1660) of simultaneous introductions from Europe (France, Spain) towards Oceania (New Zealand) and California, (ii) from the early 18^th century, a second colonization wave from bridgehead populations successfully established in California, (iii) genetic admixture in invasive areas where highly divergent populations came into contact as in New Zealand. Although these man-made pathways are consistent with historical data, introduction time estimates suggest that the two putative waves of invasion would have occurred long before the first field observations recorded, both in America and in Oceania. A prolonged lag period as the use of an incorrect generation time could explain such 100–150 years discrepancy. Lastly, the contrasting patterns of neutral genetic signal left in invasive populations are discussed in light of possible ways of facing novel environments (standing genetic variation versus new mutation).     

Bacterial Symbiosis in Some New Zealand Plants

A large number of species of Coprasma (Rubiaceae) show bacteria-containingstipular ndules. Hairs which contain bacteria are also present.In other species, for example Myoporum laetum, bacteria arepresent in superficial and in immersed leaf-glands. It is suggestedthat the bacterial symbionts, which can readily be obtainedin culture, may function in the fixation of atmospheric nitrogenand thereby be of importance to the plants when growing in nitrogen-deficientsoils.     

Invasion of the gall mite Aceria genistae (Acari: Eriophyidae), a natural enemy of the invasive weed Cytisus scoparius,into California,U.S.A. and predictions for climate suitability in other regions using ecological niche modelling

Scotch broom (Cytisus scoparius (L.) Link) is a European shrub that has naturalised in several countries worldwide and is recognised as an invasive weed in much of western North America. The mite Aceria genistae (Nalepa) is a coevolved, gall-inducing herbivore associated with Scotch broom in its native range and has been intentionally introduced as a classical weed biological control agent of C. scoparius in Australia and New Zealand. An adventive, never intentionally introduced, population of A. genistae was discovered in Washington and Oregon, U.S.A. in 2005. Surveys for A. genistae in California resulted in the discovery of the gall mite in 11 counties, with a widely scattered distribution. Molecular and morphological assessments confirm the mites collected from galls in California are A. genistae. Whether natural or anthropogenic, the estimated rate of long range dispersal for A. genistae from Washington or Oregon to California ranges from 39 to 62?km/yr. Niche model predictions indicate that A. genistae will continue to expand its distribution throughout much of the Scotch broom-invaded lands of California but areas supporting the weed in the Eastern U.S.A. appear less suitable. Modelling evidence also indicates that portions of Chile and Argentina are suitable for colonisation by A. genistae, also suggesting that expansion of the mite is possible in areas of Tasmania, southeastern Australia, and New Zealand where the mite was released. The environmental safety of A. genistae in relation to non-target plants and the influence of herbivory on Scotch broom fitness are discussed.     

Geographic proximity not a prerequisite for invasion: Hawaii not the source of California invasion by light brown apple moth (Epiphyas postvittana)

Background

The light brown apple moth (LBAM), Epiphyas postvittana (Walker), is native to Australia but invaded England, New Zealand, and Hawaii more than 100 years ago. In temperate climates, LBAM can be a major agricultural pest. In 2006 LBAM was discovered in California, instigating eradication efforts and quarantine against Hawaiian agriculture, the assumption being that Hawaii was the source of the California infestation. Genetic relationships among populations in Hawaii, California, and New Zealand are crucial to understanding LBAM invasion dynamics across the Pacific.

Methodology/Principal Findings

We sequenced mitochondrial DNA (mtDNA) from 1293 LBAM individuals from California (695), Hawaii (448), New Zealand (147), and Australia (3) to examine haplotype diversity and structure among introduced populations, and evaluate the null hypothesis that invasive populations are from a single panmictic source. However, invasive populations in California and New Zealand harbor deep genetic diversity, whereas Hawaii shows low level, shallow diversity.

Conclusions/Significance

LBAM recently has established itself in California, but was in Hawaii and New Zealand for hundreds of generations, yet California and New Zealand show similar levels of genetic diversity relative to Hawaii. Thus, there is no clear relationship between duration of invasion and genetic structure. Demographic statistics suggest rapid expansion occurring in California and past expansions in New Zealand; multiple introductions of diverse, genetically fragmented lineages could contribute to these patterns. Hawaii and California share no haplotypes, therefore, Hawaii is not the source of the California introduction. Paradoxically, Hawaii and California share multiple haplotypes with New Zealand. New Zealand may be the source for the California and Hawaii infestations, but the introductions were independent, and Hawaii was invaded only once. This has significant implications for quarantine, and suggests that probability of invasion is not directly related to geographic distance. Surprisingly, Hawaiian LBAM populations have much lower genetic diversity than California, despite being older.     

Native and introduced range surveys for egg parasitoids of Agrilus auroguttatus,an invasive pest of native oaks in California

The invasive goldspotted oak borer, Agrilus auroguttatus Schaeffer (Coleoptera: Buprestidae), has caused extensive mortality to indigenous oaks in southern California. This woodborer is native to southern Arizona where low population densities may, in part, be due to the presence of co-evolved natural enemies. Surveys were conducted in Arizona and California from June to August 2013 by deploying sentinel egg masses in an attempt to attract, collect and identify potential egg parasitoids of this beetle that could be used for a classical biological control programme in California. In total, 18,052 A. auroguttatus eggs were deployed throughout the native and introduced range. Parasitoids did not attack any eggs deployed in Arizona or California. The inability to detect egg parasitoids could be explained by the following: (1) host-specific egg parasitoids of A. auroguttatus do not exist, (2) surveying time did not coincide with peak activity of egg parasitoids or (3) surveying methods were insufficient at detecting egg parasitoids.     

The enemy release and EICA hypothesis revisited: incorporating the fundamental difference between specialist and generalist herbivores

The success of invasive plants has been attributed to their escape from natural enemies and subsequent evolutionary change in allocation from defence to growth and reproduction. In common garden experiments with Senecio jacobaea, a noxious invasive weed almost worldwide, the invasive populations from North America, Australia, and New Zealand did indeed allocate more resources to vegetative and reproductive biomass. However, invasive plants did not show a complete change in allocation from defence to growth and reproduction. Protection against generalist herbivores increased in invasive populations and pyrrolizidine alkaloids, their main anti‐herbivore compounds, did not decline in invasive populations but were higher overall compared with native populations. In contrast, invasive plants lost additional protection against specialist herbivores adapted to pyrrolizidine alkaloids. Hence, the absence of specialist herbivores in invasive populations resulted in the evolution of lower protection against specialists and increased growth and reproduction, but also allowed a shift towards higher protection against generalist herbivores.     

“Barcoding” and ISSR data illuminate a problematic infraspecific taxonomy for Chrysanthemoides monilifera (Calenduleae; Asteraceae): Lessons for biocontrol of a noxious weed

Chrysanthemoides monilifera Tourn. ex Medik is a noxious weed in Australia and New Zealand. It is a widespread species in southern Africa, where it shows considerable morphological variation that has resulted in a confusing infraspecific taxonomy. We use DNA sequence data from the nuclear Internal Transcribed Spacer (ITS) region from 78 samples of this species from its indigenous distribution range and 10 samples from Australia and New Zealand to determine the regions of origin of the invasive plants. These data are augmented by a smaller study using ISSR markers. Bayesian Inference analysis was somewhat resolved, with many weakly supported nodes. Clades with support tended to correspond to infraspecific taxonomic entities, and were geographically coherent. In contrast, a neighbour-net analysis was not as well resolved and indicated considerable reticulation. All analyses of ITS data retrieved two major clades corresponding to Western and Eastern distributions, with some overlap. Samples from New Zealand and Australia correspond to the taxon C. monilifera subsp. monilifera, and are resolved as most closely related to samples from the greater Cape Town area. Biological control agent populations for C. monilifera subsp. monilifera should be sourced from this region in order to avoid host plant incompatibility problems.     

Pre-release monitoring of Alliaria petiolata (garlic mustard) invasions and the impacts of extant natural enemies in southern Michigan forests

Monitoring of populations of a target weed species prior to releasing natural enemies has the potential to improve the rigor and safety of biological control and to determine the invader’s impacts on native communities while creating a reference point for evaluating the efficacy of subsequent biocontrol agent releases. Eight populations of garlic mustard, Alliaria petiolata (M. Bieb) Cavara and Grande (Brassicaceae), an invasive weed in southern Michigan, were monitored in anticipation of releases of classical biological control agents. The A. petiolata populations were shown to be expanding with 44.4% of initially uninvaded sampling quadrats becoming invaded after four years. While 88.2% of quadrats with A. petiolata showed evidence of foliar damage from pathogens and browsing by mammals, insects and other invertebrates, levels of damage were low and had little impact on rosette or seedling survival. Contrary to expectations, damage was positively correlated with A. petiolata fecundity (P = 0.0465). Given the continued expansion of A. petiolata and the lack of significant herbivore damage by acquired natural enemies, a biological control program should be considered against this invasive plant. If biological control agents are released, the results of this study will provide a benchmark for evaluating their performance.     

空心莲子草响应南方菟丝子寄生的生长-防御权衡

为探讨全寄生植物南方菟丝子(Cuscuta australis)防治入侵植物空心莲子草(Alternanthera philoxeroides)的可行性,以二者野外天然生长的种群为研究对象,分析南方菟丝子寄生对空心莲子草生长及防御的影响,阐明空心莲子草在受到寄生胁迫时如何权衡自身生长与防御的关系,进而发展出一套应对南方菟丝子寄生的生长-防御策略。结果显示:(1)南方菟丝子寄生显著改变空心莲子草茎的形态,茎直径和平均节间长均增加,茎直径变化极显著(P0.01);(2)南方菟丝子寄生显著减少空心莲子草叶片数,但同时显著增加后者茎的分枝数,而茎上的节是潜在的无性繁殖体,故有利于空心莲子草的克隆繁殖;此外,南方菟丝子寄生显著降低了空心莲子草的根、茎、叶生物量和总生物量,抑制空心莲子草的生长;(3)南方菟丝子寄生显著增加空心莲子草茎的单宁、总酚、三萜皂苷含量,增强其防御能力;(4)南方菟丝子寄生的空心莲子草的生物量与茎部木质素、三萜皂苷、单宁和总酚含量均呈现显著负相关性(P0.01),对照组则不存在相关性;且寄生组较对照组相比,生物量的相对百分比显著低于对照组(P0.01),而用于防御的次生代谢产物总含量的相对百分比显著高于对照组(P0.01)。以上结果表明,受到南方菟丝子寄生胁迫后,空心莲子草改变自身的生长-防御策略,减少营养生长投入而将更多的资源投向克隆繁殖,同时增强对"防御"物质的投入,增强其防御能力,以利于后代生存和繁衍。     

Inferring the introduction history of the invasive apomictic grass Cortaderia jubata using microsatellite markers

Aim Reconstructing the introduction history of exotic species is critical to understanding ecological and evolutionary processes that underlie invasive spread and to designing strategies that prevent or manage invasions. The aims of this study were to infer the introduction history of the invasive apomictic bunchgrass Cortaderia jubata and to determine if molecular data support the postulated horticultural origin of invasive populations. Location Invaded areas in the USA (California, Maui) and New Zealand; native areas in Bolivia, Ecuador and Peru. Methods We used nuclear microsatellite markers to genotype 281 plants from invaded areas in California, Maui and New Zealand, and 77 herbarium specimens from native South America, and compared the genotypic and clonal variation of C. jubata from the invaded and native ranges. Clonal diversity was determined from genotypic diversity using two analytical methods. Results Invasive C. jubata from invaded regions in California, Maui and New Zealand consisted of the same single clone that probably originated from a single introduced genotype. In contrast, 14 clones were detected in herbarium specimens from the native areas of Bolivia, Ecuador and Peru. The invasive clone matched the most common clone identified in herbarium specimens from southern Ecuador where horticultural stock is presumed to have originated. Main conclusions The lack of clonal and genotypic diversity in invasive plants, but moderately high diversity detected in native plants, indicates a significant reduction in genetic variation associated with the introduction of C. jubata outside of its native range. Based on historical accounts of the horticultural introduction of C. jubata and the results of this study, a severe founder effect probably occurred during deliberate introduction of C. jubata into cultivation. Our results are consistent with the postulated horticultural origin of invasive C. jubata and point to southern Ecuador as the geographical source of invasive populations.     

A taxonomic,biogeographical and ecological overview of invasive woody plants

Abstract. Invasive plants are considered to be a major threat to the earth's biodiversity, but have not been sufficiently investigated. To address this problem a relational database on invasive woody plants has been set up. It is based on 2045 bibliographical references and contains records on 653 species representing 110 families. The families with the largest number of invasive species are: Rosaceae, Mimosaceae, Papilionaceae and Pinaceae. Out of 1060 recorded invasive events an equal number are reported from continents and oceanic islands. The highest number of invasive woody species are recorded from Europe, followed by the Pacific islands, North America, New Zealand, Australia, Indian Ocean islands and southern Africa. Included in these regions are areas which have fewer highly invasive species, e.g. islands on continental shelves, such as the British Isles. Although most invasions occur in disturbed habitats, most natural communities are susceptible to woody plant invasions. Data on species attributes are only available for a minority of species but indicate that invasive woody plant species may be either insect or wind-pollinated, have a wide array of fruit types, fruit and seed sizes, number of seeds per fruit and dispersal agents. The relative frequency of several attributes varies with the degree of invasiveness. Thus it is not yet possible to determine which set of attributes favours invasiveness and therefore it is difficult to make predictions.     

Latitudinal trends in herbivory and performance of an invasive species, common burdock (Arctium minus)

Plants often lose natural enemies (herbivores and pathogens) while invading new geographic regions, as predicted by the Enemy Release Hypothesis. However, a similar reduction in attack might occur at a more local scale within an invader’s range as plants in marginal areas escape enemies that fail to find them or cannot maintain local populations. In this study, we test the hypothesis that isolated populations near the northern edge of an invader’s range escape the enemies present in more southern populations, using the non-native monocarpic biennial, common burdock (Arctium minus), as a model species. In southern Ontario, this plant is attacked by a wide range of insect herbivores, including generalist leaf chewers as well as leaf-mining flies (Liriomyza arctii, Calycomyza flavinotum) and an abundant lepidopteran seed predator (Metzneria lappella). Surveys over an 815 km transect from temperate southern to boreal northern Ontario indicate that damage by all of these enemies declines sharply with latitude, while plants in more northern areas are slightly larger and more fecund. Critically, seed parasitism drops from more than 85 % in the south to less than 25 % in the north. These results indicate that populations of Arctium near this species’ northern limit escape many of their usual natural enemies, potentially counteracting less favourable environmental conditions. Such escape from enemies near invaders’ range margins may accelerate further spread, including expected migration in response to climate change.     

Increased resistance to generalist herbivores in invasive populations of the California poppy (Eschscholzia californica)

Escape from enemies in the native range is often assumed to contribute to the successful invasion of exotic species. Following optimal defence theory, which assumes a trade‐off between herbivore resistance and plant growth, some have predicted that the success of invasive species could be the result of the evolution of lower resistance to herbivores and increased allocation of resources to growth and reproduction. Lack of evidence for ubiquitous costs of producing plant toxins, and the recognition that invasive species may escape specialist, but not generalist enemies, has led to a new prediction: invasive species may escape ecological trade‐offs associated with specialist herbivores, and evolve increased, rather than decreased, production of defensive compounds that are effective at deterring generalist herbivores in the introduced range. We tested the performance of two generalist lepidopteran herbivores, Trichoplusia ni and Orgyia vetusta, when raised on diets of native and invasive populations of the California poppy, Eschscholzia californica. Pupae of T. ni were significantly larger when reared on native populations. Similarly, caterpillars of O. vetusta performed significantly better when raised on native populations, indicating that invasive populations of the California poppy are more resistant to herbivores than native populations. The chance of successful establishment of some non‐indigenous plant species may be increased by retaining resistance to generalist herbivores, and in some cases, invasive species may be able to escape ecological trade‐offs in their new range and evolve, as we observed, even greater resistance to generalist herbivores than native plants.     

The first record of a terrestrial landhopper (Crustacea: Amphipoda: Talitridae) from Macquarie Island

A terrestrial talitrid crustacean is recorded for the first time from subantarctic Macquarie Island. The species, Puhuruhuru patersoni, is native to the southern part of the South Island of New Zealand. Its distribution on Macquarie Is. is restricted to a single locality near the ANARE station on the Isthmus. All mature females collected in December were carrying eggs; females appeared to be less mobile than males. It seems likely that the species has been transported accidentally from New Zealand, perhaps by the same means that introduced the isopod, Styloniscus otakensis, also from southern New Zealand.     

Can the destruction of California’s oak woodlands be prevented? Potential for biological control of the goldspotted oak borer, <Emphasis Type="Italic">Agrilus auroguttatus</Emphasis>

The goldspotted oak borer (GSOB), Agrilus auroguttatus Schaeffer (Coleoptera: Buprestidae), is an introduced and aggressive phloem/wood borer infesting native oaks in southern California. Elevated levels of oak mortality have occurred continually for the last nine years on three oak species in San Diego Co., California, USA. Biological control is being assessed as an option for long-term and widespread management of the invasive population of GSOB. Foreign exploration in the native ranges of GSOB and a related sibling species (Agrilus coxalis Waterhouse) was conducted to determine life history information, to assess the natural enemy complex, and to collect specimens for molecular analyses that could help to identify the area of origin of California’s introduced population. Two species of parasitoids, Calosota elongata Gibson (Eupelmidae) and Atanycolus simplex Cresson (Braconidae), were discovered with GSOB populations in Arizona and California. No insect natural enemies were found with populations of A. coxalis in southern Mexico. However, Quercus conzatti Trel. and Quercus peduncularis Nee in Oaxaca and Chiapas, respectively, were recorded as the first known hosts of A. coxalis. A comparative analysis of our understanding of the natural enemy complexes for other pestiferous Agrilus with that of GSOB suggests that more effort should be directed at uncovering potential egg parasitoids and microbial pathogens of GSOB. Analyses of mitochondrial and nuclear ribosomal DNA (rDNA) revealed that the California population of GSOB was more similar to the Arizona population. Specimens of A. coxalis from southern Mexico were confirmed as a separate species. Additional surveys and sampling are needed across the complete native range of the GSOB species complex to develop a comprehensive inventory of parasitoid species that could be considered for use in a classical biological control program in California and to delineate the area of origin of California’s population.     

Further evaluation of the southern ladybird (Cleobora mellyi) as a biological control agent of the invasive tomato–potato psyllid (Bactericera cockerelli)

The southern ladybird (Cleobora mellyi Coleoptera: Coccinellidae) is a voracious predator of the invasive tomato–potato psyllid (TPP) (Bactericera cockerelli Hemiptera: Triozidae) in New Zealand. We examined important aspects of the southern ladybird’s ecology to obtain further insight into its potential as a biocontrol agent of TPP in potato crops. We found that the southern ladybird did not prefer TPP over either Myzus persicae Sulzer or Macrosiphum euphorbiae Thomas in choice tests, but avoided consumption of greenhouse whitefly, Trialeurodes vaporariorum (Westwood). Ladybird longevity was tested under the conditions of low prey provision, a floral resource (buckwheat), and a combination of buckwheat and low density of TPP, over a 3 month period. There was no difference in longevity between ladybirds supplied with TPP only or buckwheat only. However, those with access to TPP and buckwheat lived longer than those with only TPP. In a glasshouse microcosm study, the ladybird was able to significantly reduce TPP densities after 3 weeks, and maintain the reduced numbers for 7 weeks. A species-level trophic cascade was found for both number and weight of potato tubers. These results indicate that the southern ladybird has potential as a biological control agent of the invasive tomato–potato psyllid in New Zealand.     

Styela clava Herdman (Urochordata, Ascidiacea), a successful immigrant to North West Europe: ecology, propagation and chronology of spread

Since its first occurrence at Plymouth, southern England, in 1952 the East Asiatic ascidianStyela clava has spread to many localities along the coasts of the south and west British Isles, Ireland, northern France, Belgium, the Netherlands, Denmark and Germany. While some dispersal may occur by natural means, spreading over long distances is probably due to transfer along with oysters when relaid elsewhere. Transport while attached to the hulls of ships or driftingSargassum is also possible.Styela clava is a large, hardy and fast-growing species with a tough, leathery tunic, and has no recorded enemies or native analogues among the NW European ascidian fauna. At many sites it has established dense populations of 500–1000 specimens/m² and in some cases has nearly outcompeted some of the native ascidian species.