Mitochondrial DNA diversity of <Emphasis Type="Italic">Cleruchoides noackae</Emphasis> (Hymenoptera: Mymaridae): a potential biological control agent for <Emphasis Type="Italic">Thaumastocoris peregrinus</Emphasis> (Hemiptera: Thaumastocoridae)

Thaumastocoris peregrinus Carpintero and Dellapé (Hemiptera: Thaumastocoridae) is a native Australian Eucalyptus sap-feeding insect that has become invasive and seriously damaging to commercially grown Eucalyptus in the Southern Hemisphere. Cleruchoides noackae Lin and Huber (Hymenoptera: Mymaridae) was recently discovered as an egg parasitoid of the Thaumastocoridae in Australia. Mitochondrial DNA (mtDNA; cytochrome c oxidase subunit I, COI) sequence diversity amongst 104 individuals from these native C. noackae populations revealed 24 sequence haplotypes. The COI haplotypes of individuals collected from the Sydney and Southeast Queensland clustered in distinct groups, indicating limited spread of the insect between the regions. Individuals collected from Perth in Western Australia were represented by four COI haplotypes. Although this population is geographically more isolated from other populations, two COI haplotypes were identical to haplotypes found in the Sydney region. The results suggest that C. noackae has recently been introduced into Perth, possibly from the Sydney area. The high mtDNA diversity and limited spread that is suggested for C. noackae is in contrast to the lack of geographic associated mtDNA diversity and extensive spread of T. peregrinus. If implemented as a biological control agent, this factor will need to be considered in collecting and releasing C. noackae.     

The potential global distribution of the Bronze bug Thaumastocoris peregrinus Carpintero and Dellapé (Hemiptera: Thaumastocoridae)

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Geographic patterns of genetic diversity from the native range of <Emphasis Type="Italic">Cactoblastis cactorum</Emphasis> (Berg) support the documented history of invasion and multiple introductions for invasive populations

Spread of the invasive cactus-feeding moth Cactoblastis cactorum has been well documented since its export from Argentina to Australia as a biocontrol agent, and records suggest that all non-native populations are derived from a single collection in the moth’s native range. The subsequent global spread of the moth has been complex, and previous research has suggested multiple introductions into North America. There exists the possibility of additional emigrations from the native range in nursery stock during the late twentieth century. Here, we present mitochondrial gene sequence data (COI) from South America (native range) and North America (invasive range) to test the hypothesis that the rapid invasive spread in North America is enhanced by unique genetic combinations from isolated portions of the native range. We found that haplotype richness in the native range of C. cactorum is high and that there was 90% lower richness in Florida than in Argentina. All Florida C. cactorum haplotypes are represented in a single, well-defined clade, which includes collections from the reported region of original export from Argentina. Thus, our data are consistent with the documented history suggesting a single exportation of C. cactorum from the eastern region of the native range. Additionally, the presence of geographic structure in three distinct haplotypes within the same clade across Florida supports the hypothesis of multiple introductions into Florida from a location outside the native range. Because the common haplotypes in Florida are also known to occur in the neighboring Caribbean Islands, the islands are a likely source for independent North American colonization events. Our data show that rapid and successful invasion within North America cannot be attributed to unique genetic combinations. This suggests that successful invasion of the southeastern US is more likely the product of a fortuitous introduction into favorable abiotic conditions and/or defense responses of specific Opuntia hosts, rapid adaptation, or a release from native enemies.     

Geographical distribution modelling of the bronze bug: a worldwide invasion

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High intercontinental migration rates and population admixture in the sapstain fungus Ophiostoma ips

Ophiostoma ips is a common fungal associate of various conifer-infesting bark beetles in their native ranges and has been introduced into non-native pine plantations in the Southern Hemisphere. In this study, we used 10 microsatellite markers to investigate the population biology of O. ips in native (Cuba, France, Morocco and USA) and non-native (Australia, Chile and South Africa) areas to characterize host specificity, reproductive behaviour, and the potential origin as well as patterns of spread of the fungus and its insect vectors. The markers resolved a total of 41 alleles and 75 haplotypes. Higher genetic diversity was found in the native populations than in the introduced populations. Based on the origin of the insect vectors, the populations of O. ips in Australia would be expected to reflect a North American origin, and those in Chile and South Africa to reflect a European origin. However, most alleles observed in the native European population were also found in the native North American population; only the allele frequencies among the populations varied. This admixture made it impossible to confirm the origin of the introduced Southern Hemisphere (SH) populations of O. ips. There was also no evidence for specificity of the fungus to particular bark beetle vectors or hosts. Although O. ips is thought to be mainly self-fertilizing, evidence for recombination was found in the four native populations surveyed. The higher genetic diversity in the North American than in the European population suggests that North America could be the possible source region of O. ips.     

Spread and impact of introduced conifers in South America: Lessons from other southern hemisphere regions

The history of conifers introduced earlier elsewhere in the southern hemisphere suggests that recent invasions in Argentina, Brazil, Chile and Uruguay are likely to increase in number and size. In South Africa, New Zealand and Australia, early ornamental introductions and small forestry plantations did not lead to large‐scale invasions, while subsequent large plantations were followed with a lag of about 20–30 years by troublesome invasions. Large‐scale conifer plantation forestry in South America began about 50–80 years later than in South Africa, Australia and New Zealand, while reports of invasions in South America lagged behind those in the latter nations by a century. Impacts of invading non‐native conifers outside South America are varied and include replacement of grassland and shrubland by conifer forest, alteration of fire and hydrological regimes, modification of soil nutrients, and changes in aboveground and belowground biotic communities. Several of these effects have already been detected in various parts of South America undergoing conifer invasion. The sheer amount of area planted in conifers is already very large in Chile and growing rapidly in Argentina and Brazil. This mass of reproductive trees, in turn, produces an enormous propagule pressure that may accelerate ongoing invasions and spark new ones at an increasing rate. Regulations to control conifer invasions, including measures to mitigate spread, were belatedly implemented in New Zealand and South Africa, as well as in certain Australian states, inspired by observations on invasions in those nations. Regulations in South America are weaker and piecemeal, but the existing research base on conifer invasions elsewhere could be useful in fashioning effective regulations in South America. Pressure from foreign customers in South Africa has led most companies there to seek certification through the Forestry Stewardship Council; a similar programme operates in Australia. Such an approach may be promising in South America.     

Contrasting mitochondrial diversity of European starlings (Sturnus vulgaris) across three invasive continental distributions

European starlings (Sturnus vulgaris) represent one of the most widespread and problematic avian invasive species in the world. Understanding their unique population history and current population dynamics can contribute to conservation efforts and clarify evolutionary processes over short timescales. European starlings were introduced to Central Park, New York in 1890, and from a founding group of about 100 birds, they have expanded across North America with a current population of approximately 200 million. There were also multiple introductions in Australia in the mid‐19th century and at least one introduction in South Africa in the late 19th century. Independent introductions on these three continents provide a robust system to investigate invasion genetics. In this study, we compare mitochondrial diversity in European starlings from North America, Australia, and South Africa, and a portion of the native range in the United Kingdom. Of the three invasive ranges, the North American population shows the highest haplotype diversity and evidence of both sudden demographic and spatial expansion. Comparatively, the Australian population shows the lowest haplotype diversity, but also shows evidence for sudden demographic and spatial expansion. South Africa is intermediate to the other invasive populations in genetic diversity but does not show evidence of demographic expansion. In previous studies, population genetic structure was found in Australia, but not in South Africa. Here we find no evidence of population structure in North America. Although all invasive populations share haplotypes with the native range, only one haplotype is shared between invasive populations. This suggests these three invasive populations represent independent subsamples of the native range. The structure of the haplotype network implies that the native‐range sampling does not comprehensively characterize the genetic diversity there. This study represents the most geographically widespread analysis of European starling population genetics to date.     

Genetic diversity of <Emphasis Type="Italic">Bradysia difformis</Emphasis> (Sciaridae: Diptera) populations reflects movement of an invasive insect between forestry nurseries

The fungus gnat, Bradysia difformis (Sciaridae: Diptera) has recently been recorded for the first time from South Africa where it has been found in forestry nurseries. The presence of this insect in all the major forestry nurseries as the dominant and only sciarid species raises intriguing questions regarding its origin and population genetic structure. A 395 bp portion of the mitochondrial COI gene was analysed from B. difformis individuals collected from four nursery populations in South Africa and three nursery populations in Europe. Shared haplotypes between South African and European populations indicated a historical connection. South African populations showed high genetic diversity and low genetic differentiation. These patterns most likely reflect multiple and/or relatively large introductions of B. difformis into South Africa from its origin combined with subsequent and continued movement of plants between nurseries.     

Molecular relationships within Bostrychia tenuissima (Rhodomelaceae, Rhodophyta)

Bostrychia tenuissima King and Puttock is restricted to southern Australia and New Zealand. Previous studies in Australia have revealed two distinct patterns in the presence of osmotically active polyols. Southern populations only have D-sorbitol whereas northern populations have both D-sorbitol and D-dulcitol. These polyol patterns lead to speculation on the ecotypic differentiation of these two population types. Using single-stranded confirmation polymorphism to rapidly score plastid haplotypes, plus DNA sequencing, a 100% congruence was found between polyol patterns and plastid haplotype. Analysis of 33 B. tenuissima isolates shows that the plastid relationships closely follow biogeo-graphic regions, with south-eastern Australia (southern NSW and Victoria) and South Australia and Tasmania having one haplotype and central and northern NSW having two alternate haplotypes. There is an overlap at the haplotype boundary just south of Sydney, New South Wales.     

Population structure and diversity of an invasive pine needle pathogen reflects anthropogenic activity

Dothistroma septosporum is a haploid fungal pathogen that causes a serious needle blight disease of pines, particularly as an invasive alien species on Pinus radiata in the Southern Hemisphere. During the course of the last two decades, the pathogen has also incited unexpected epidemics on native and non‐native pine hosts in the Northern Hemisphere. Although the biology and ecology of the pathogen has been well documented, there is a distinct lack of knowledge regarding its movement or genetic diversity in many of the countries where it is found. In this study we determined the global population diversity and structure of 458 isolates of D. septosporum from 14 countries on six continents using microsatellite markers. Populations of the pathogen in the Northern Hemisphere, where pines are native, displayed high genetic diversities and included both mating types. Most of the populations from Europe showed evidence for random mating, little population differentiation and gene flow between countries. Populations in North America (USA) and Asia (Bhutan) were genetically distinct but migration between these continents and Europe was evident. In the Southern Hemisphere, the population structure and diversity of D. septosporum reflected the anthropogenic history of the introduction and establishment of plantation forestry, particularly with Pinus radiata. Three introductory lineages in the Southern Hemisphere were observed. Countries in Africa, that have had the longest history of pine introductions, displayed the greatest diversity in the pathogen population, indicating multiple introductions. More recent introductions have occurred separately in South America and Australasia where the pathogen population is currently reproducing clonally due to the presence of only one mating type.     

Attraction of Male Nymphs to Adult Male Volatiles in the Bronze Bug <Emphasis Type="Italic">Thaumastocoris peregrinus</Emphasis> Carpintero &amp; Dellape (Heteroptera: Thaumastocoridae)

The bronze bug, Thaumastocoris peregrinus Carpintero & Dellape (Heteroptera: Thaumastocoridae), is an exotic emerging pest in Eucalyptus commercial forests in South America, Africa and southern Europe. Information on the chemical communication system and reproductive ecology of this insect is scant, and it may be relevant for designing management strategies for eucalypt plantations. Adults and nymphs usually aggregate in the field, possibly by means of chemical signals. Males emit large amounts of 3-methyl-2-butenyl butyrate, which attracts conspecific adult males but not females. The ecological role of this putative male aggregation pheromone remains unknown. Here, we report olfactometer bioassays showing that late-instar male nymphs are also attracted to synthetic 3-methyl-2-butenyl butyrate and to adult male volatile extracts, which contain this compound as the major component. As previously shown for adult females, nymphs that moulted into females were not attracted to either volatile stimulus. The intra-gender attraction of nymphs and adults may be related to the exploitation of food resources, or as a reproductive strategy for newly emerged males. Further studies on the reproductive behaviour and mating system of T. peregrinus will contribute to understanding the ecological significance of male-male, adult-nymph attraction, as well as the practical applications that may result from these findings.     

Variation in mitochondrial cytochrome c oxidase subunit I gene in Nezara viridula (Hemiptera: Pentatomidae) from Argentina

Here, we examine the genetic diversity in the agricultural pest Nezara viridula (Linnaeus, 1758 ) from populations of Argentina using mitochondrial cytochrome c oxidase subunit I (COI) gene sequences. The DNA sequence comparisons of 718 base pairs of the COI gene revealed seven haplotypes. The observed total haplotype diversity (Hd) value was of 0.138, and the nucleotide diversity was of 0.00039 and 0.00135 according to π and θ_W, respectively. Eight out of the 10 populations analysed, mostly from soya bean crops, only presented the more frequent haplotype, while 2 haplotypes were found in a mixed culture and 6 haplotypes in a peanut culture. Factors such as differential insecticide applications, as well as the surrounding habitat, and the host plant preference could be related to the genetic diversity differences observed among samples of N. viridula. The analysis of genetic diversity in samples collected in crops treated and non‐treated with insecticides, as well as in samples collected from different seasons, could help to clarify the role of the factors that led to the pattern of genetic diversity detected in this study. The result of a comparative analysis of COI gene sequences among populations from South America, Africa, Asia and Europe was consistent with the hypothesis of an African origin of N. viridula. On the other hand, the haplotypes of Europe were clustered with haplotypes from South America. In addition, specimens from Madeira (west of Europe) shared ancestry with South America and Europe. It has been suggested that a probable route of colonization of America could have been from Western Europe towards the eastern coasts of South America.     

Genetic diversity and maternal origin of Northeast African and South American donkey populations

To investigate the mtDNA variation and origin of maternal lineages in South American donkeys and to reassess the domestication of donkeys in northeast Africa, we analyzed sequences (489 bp of the D‐loop) from 323 domestic donkeys sampled from Peru, Brazil, Ethiopia and Egypt. Altogether, the 323 sequences displayed 53 different haplotypes (45 in Ethiopia, 14 in Egypt, eight in Peru and six in Brazil). Among the four populations, Egyptian donkeys possessed the highest haplotype diversity (0.910 ± 0.032), followed by Brazilian donkeys (0.879 ± 0.060). The Clade I haplotypes dominated in Peruvian donkeys (65%), whereas Clade II haplotypes dominated in Brazilian donkeys (67%). Estimates of F_ST values showed a high genetic differentiation between Peruvian and Brazilian donkey populations (F_ST = 0.4066), which could be explained by the complex introduction history of South American donkeys. Phylogeographic analysis indicates that northeast Africa could be the most probable domestication center for Clade I donkeys. Analysis of molecular variance confirmed a weak genetic structure in domestic donkey populations among four continents (Europe, Asia, Africa and South America).     

Multiple introductions from multiple sources: invasion patterns for an important Eucalyptus leaf pathogen

Many population studies on invasive plant pathogens are undertaken without knowing the center of origin of the pathogen. Most leaf pathogens of Eucalyptus originate in Australia and consequently with indigenous populations available, and it is possible to study the pathways of invasion. Teratosphaeria suttonii is a commonly occurring leaf pathogen of Eucalyptus species, naturally distributed in tropical and subtropical regions of eastern Australia where it is regarded as a minor pathogen infecting older leaves; however, repeated infections, especially in exotic plantations, can result in severe defoliation and tree deaths. Nine polymorphic microsatellite markers were used to assess the genetic structure of 11 populations of T. suttonii of which four where from within its native range in eastern Australia and the remaining seven from exotic Eucalyptus plantations. Indigenous populations exhibited high allele and haplotype diversity, predominantly clonal reproduction, high population differentiation, and low gene flow. The diversity of the invasive populations varied widely, but in general, the younger the plantation industry in a country or region, the lower the diversity of T. suttonii. Historical gene flow was from Australia, and while self‐recruitment was dominant in all populations, there was evidence for contemporary gene flow, with South Africa being the most common source and Uruguay the most common sink population. This points distinctly to human activities underlying long‐distance spread of this pathogen, and it highlights lessons to be learned regarding quarantine.     

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.     

Recent emergence and worldwide spread of the red tomato spider mite, <Emphasis Type="Italic">Tetranychus evansi</Emphasis>: genetic variation and multiple cryptic invasions

Plant biosecurity is increasingly challenged by emerging crop pests. The spider mite Tetranychus evansi has recently emerged as a new threat to solanaceous crops in Africa and the Mediterranean basin, with invasions characterized by a high reproductive output and an ability to withstand a wide range of temperatures. Mitochondrial (868 bp of COI) and nuclear (1,137 bp of ITS) loci were analyzed in T. evansi samples spanning the current geographical distribution to study the earliest stages of the invasive process. The two sets of markers separate the samples into two main clades that are only present together in South America and Southern Europe. The highest COI diversity was found in South America, consistent with the hypothesis of a South American origin of T. evansi. Among the invaded areas, the Mediterranean region displayed a high level of genetic diversity similar to that present in South America, that is likely the result of multiple colonization events. The invasions of Africa and Asia by T. evansi are characterized by a low genetic variation associated with distinct introductions. Genetic data demonstrate two different patterns of invasions: (1) populations in the Mediterranean basin that are a result of multiple cryptic introductions and (2) emerging invasions of Africa and Asia, each likely the result of propagules from one or limited sources. The recent invasions of T. evansi illustrate not only the importance of human activities in the spread of agricultural pests, but also the limits of international quarantine procedures, particularly for cryptic invasions.     

Mitochondrial DNA reveals multiple Northern Hemisphere introductions of Caprella mutica (Crustacea, Amphipoda)

Caprella mutica (Crustacea, Amphipoda) has been widely introduced to non-native regions in the last 40 years. Its native habitat is sub-boreal northeast Asia, but in the Northern Hemisphere, it is now found on both coasts of North America, and North Atlantic coastlines of Europe. Direct sequencing of mitochondrial DNA (cytochrome c oxidase subunit I gene) was used to compare genetic variation in native and non-native populations of C. mutica . These data were used to investigate the invasion history of C. mutica and to test potential source populations in Japan. High diversity (31 haplotypes from 49 individuals), but no phylogeographical structure, was identified in four populations in the putative native range. In contrast, non-native populations showed reduced genetic diversity (7 haplotypes from 249 individuals) and informative phylogeographical structure. Grouping of C. mutica populations into native, east Pacific, and Atlantic groups explained the most among-region variation (59%). This indicates independent introduction pathways for C. mutica to the Pacific and Atlantic coasts of North America. Two dominant haplotypes were identified in eastern and western Atlantic coastal populations, indicating several dispersal routes within the Atlantic. The analysis indicated that several introductions from multiple sources were likely to be responsible for the observed global distribution of C. mutica , but the pathways were least well defined among the Atlantic populations. The four sampled populations of C. mutica in Japan could not be identified as the direct source of the non-native populations examined in this study. The high diversity within the Japan populations indicates that the native range needs to be assessed at a far greater scale, both within and among populations, to accurately assess the source of the global spread of C. mutica .     

Restricted Genetic Variation in Populations of Achatina (Lissachatina) fulica outside of East Africa and the Indian Ocean Islands Points to the Indian Ocean Islands as the Earliest Known Common Source

The Giant African Land Snail, Achatina ( = Lissachatina) fulica Bowdich, 1822, is a tropical crop pest species with a widespread distribution across East Africa, the Indian subcontinent, Southeast Asia, the Pacific, the Caribbean, and North and South America. Its current distribution is attributed primarily to the introduction of the snail to new areas by Man within the last 200 years. This study determined the extent of genetic diversity in global A. fulica populations using the mitochondrial 16S ribosomal RNA gene. A total of 560 individuals were evaluated from 39 global populations obtained from 26 territories. Results reveal 18 distinct A. fulica haplotypes; 14 are found in East Africa and the Indian Ocean islands, but only two haplotypes from the Indian Ocean islands emerged from this region, the C haplotype, now distributed across the tropics, and the D haplotype in Ecuador and Bolivia. Haplotype E from the Philippines, F from New Caledonia and Barbados, O from India and Q from Ecuador are variants of the emergent C haplotype. For the non-native populations, the lack of genetic variation points to founder effects due to the lack of multiple introductions from the native range. Our current data could only point with certainty to the Indian Ocean islands as the earliest known common source of A. fulica across the globe, which necessitates further sampling in East Africa to determine the source populations of the emergent haplotypes.     

Increasing numbers and intercontinental spread of invasive insects on eucalypts

Native to Australasia, Eucalyptus (sensu lato) is one of the most planted genera of trees in the world. However, the sustainability of Eucalyptus species as plantation trees in non-native areas is increasingly threatened by the introduction and spread of Eucalyptus-feeding insects from Australia. We examine patterns and potential trends with respect to the global spread of Eucalyptus-feeding insects. Likely pathways of introduction and drivers of the rapid distribution of these insects, as well as management options are considered. The rate of introductions is shown to have increased nearly fivefold since the 1980s. As a result, the number of non-native pests of eucalypts outside of Australia has doubled in less than three decades. Furthermore, the rate of secondary spread among continents has also increased. Surprisingly, we found no association between area planted and the number of pests or new introductions. Only a small number of countries have been the points of first detection outside the native range; these countries have acted as bridgeheads to other regions. Quarantine regulations aimed at reducing the spread of invasive organisms appear to be ineffective at a global scale, and pathways allowing these invasions to occur are poorly understood or unknown. An expanded suite of management options are needed to provide resilience against the rapid accrual and homogenization of eucalypt pests, thereby ensuring the sustainability of eucalypt forestry worldwide.