The taxonomic status of Larinus cynarae F. and Larinus latus Herbst (coleoptera: Curculionidae), and its implications for the biological control of Onopordum (asteraceae: Cardueae) in Australia

Analyses were made of the allozyme frequencies of sympatric populations of the capitulum weevils L. latus Herbst and Larinus cyarae F. in Greece. It was found that the two taxa are genetically isolated and that they can therefore be considered as separate species. This complemented ecological data showing separation of the L. latus and L. cynarae populations by their choice of Onopordum illyricum L. and Cynara cardunculus, respectively, as hosts. Onopordum spp. are serious weed problems in Australia while Cynara contains several crop plants. L. latus is a potential biological control agent of Onopordum spp., but its use had been compromised by the uncertainty of the taxonomic distinctness from L. cynarae, biotypes of which are pests of artichoke Cynara scolymus L. Clarification of its taxonomic status lifts these reservations and clears the way for the introduction of L. latus into Australia for the biological control of Onopordum thistles.     

Structure of the phytophagous insect fauna of Onopordum thistles in the northern Mediterranean basin

Thistles of the genus Onopordum (Asteraceae: Cardueae) have become serious weed problems in parts of Australia and North America following introduction from their native Eurasian ranges, and are the target of a biological control programme in Australia. This paper analyses the results of three separate surveys of insects feeding and breeding on thistles of the genus Onopordum in the Mediterranean region. Of the 129 species found feeding, 74 species also bred on these thistles. Endophages comprised 54% of the breeding insects, with species that fed in the capitula and within the rosette crown/root area predominating. The Coleoptera and Hemiptera were the dominant endophage and ectophage taxa, respectively. Differences in the Onopordum fauna were noted between host species and between geographic regions, but this was largely due to the generalist component of the fauna. Onopordum specialists showed little variability across regions or between different species within the host genus. On comparing the fauna of Onopordum with that of other thistle genera, it was suggested that plant-herbivore interactions between Onopordum and its fauna are less highly evolved, in that there is less host specialization and a lesser degree of niche partitioning. This appears to be related to the low rate of speciation and habitat specialization within the host genus itself. The potential of certain genus-specific insects as biological control agents for Onopordum spp. is discussed. It is argued that the absence of a high degree of niche specialization could favour successful biological control, as the agent would be capable of inflicting damage over a wider range of resource gradients.     

Comparative biology of two rosette crown-feeding flies of the genus Botanophila (Dipt., Anthomyiidae) with potential for biological control of their thistle hosts

The comparative biology of two anthomyiid flies with potential for biological control of their host thistles, Botanophila turcica on Carthamus lanatus and Botanophila spinosa on Onopordum acanthium , was studied using field surveys and collections, and by rearing collected eggs and larvae in the laboratory in southern France. The thistle hosts are significant weeds outside their native range, particularly in Australia. Both flies attack the rosette meristems of their hosts prior to flowering. Larval stages are described together with natural attack rates (21–33% of field plants) and the mortality of field-collected larvae reared in the laboratory. A successful rearing protocol for these flies is also described. The results of preliminary host-specificity tests showed that both species are highly specific, being restricted to their host genus in the tests conducted. Furthermore, Botanophila turcica could not complete development on safflower (a congener of its natural host) under natural conditions. Botanophila turcica had an adult activity period that lasted 6 months from late autumn and laid fewer eggs per host plant than B. spinosa , which was active for 2 to 3 months in spring. Only one parasitoid was observed, an endoparasitic eucoilid, Trybliographa sp., which attacked both fly species, causing 18–23% mortality.     

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.     

The lack of genetic bottleneck in invasive Tansy ragwort populations suggests multiple source populations.

Jacobaea vulgaris (Asteraceae) is a species of Eurasian origin that has become a serious non-indigenous weed in Australia, New Zealand, and North America. We used neutral molecular markers to (1) test for genetic bottlenecks in invasive populations and (2) to investigate the invasion pathways. It is for the first time that molecular markers were used to unravel the process of introduction in this species.The genetic variation of 15 native populations from Europe and 16 invasive populations from Australia, New Zealand and North America were compared using the amplified fragment length polymorphisms (AFLP's). An analysis of molecular variance showed that a significant part (10%) of the total genetic variations between all individuals could be explained by native or invasive origin.Significant among-population differentiation was detected only in the native range, whereas populations from the invasive areas did not significantly differ from each other; nor did the Australian, New Zealand and North American regions differ within the invasive range. The result that native populations differed significantly from each other and that the amount of genetic variation, measured as the number of polymorphic bands, did not differ between the native and invasive area, strongly suggests that introductions from multiple source populations have occurred. The lack of differentiation between invasive regions suggests that either introductions may have occurred from the same native sources in all invasive regions or subsequent introductions took place from one into another invasive region and the same mix of genotypes was subsequently introduced into all invasive regions.An assignment test showed that European populations from Ireland, the Netherlands and the United Kingdom most resembled the invasive populations.     

The population genomic basis of geographic differentiation in North American common ragweed (Ambrosia artemisiifolia L.)

Common ragweed (Ambrosia artemisiifolia L.) is an invasive, wind‐pollinated plant nearly ubiquitous in disturbed sites in its eastern North American native range and present across growing portions of Europe, Africa, Asia, and Australia. Phenotypic divergence between European and native‐range populations has been described as rapid evolution. However, a recent study demonstrated major human‐mediated shifts in ragweed genetic structure before introduction to Europe and suggested that native‐range genetic structure and local adaptation might fully explain accelerated growth and other invasive characteristics of introduced populations. Genomic differentiation that potentially influenced this structure has not yet been investigated, and it remains unclear whether substantial admixture during historical disturbance of the native range contributed to the development of invasiveness in introduced European ragweed populations. To investigate fine‐scale population genetic structure across the species' native range, we characterized diallelic SNP loci via a reduced‐representation genotyping‐by‐sequencing (GBS) approach. We corroborate phylogeographic domains previously discovered using traditional sequencing methods, while demonstrating increased power to resolve weak genetic structure in this highly admixed plant species. By identifying exome polymorphisms underlying genetic differentiation, we suggest that geographic differentiation of this important invasive species has occurred more often within pathways that regulate growth and response to defense and stress, which may be associated with survival in North America's diverse climatic regions.     

A taxonomic revision, biology and morphology of immature stages of the Entedon sparetus species group (Hymenoptera: Eulophidae), egg-larval endoparasitoids of weevils (Coleoptera: Curculionidae)

The sparetus-group of Entedon is revised. Six species are recognized for the group, including one new species, E. lucasi Gumovsky sp. n., which is described from Algeria. All species are either diagnosed or described, and illustrated, and new distribution records are provided. An illustrated dichotomous key to all species of the group is given. Biology of two species, E. thomsonianus Erd?s and E. sparetus Walker, is studied in detail: both species proved to be egg-larval parasitoids of stem-boring weevils, Lixus cardui Olivier on the scotch thistle Onopordum acanthium and Gymnetron asellus (Gravenhorst) on mullein Verbascum sp., respectively. The morphology of larval stages and peculiarities of parasitoid-host relationships are described. Possible benefits of egg-larval parasitism for Entedon species are discussed.     

Historical range contraction,and not taxonomy,explains the contemporary genetic structure of the Australian tree <Emphasis Type="Italic">Acacia dealbata</Emphasis> Link

Irrespective of its causes, strong population genetic structure indicates a lack of gene flow. Understanding the processes that underlie such structure, and the spatial patterns it causes, is valuable for conservation efforts such as restoration. On the other hand, when a species is invasive outside its native range, such information can aid management in the non-native range. Here we explored the genetic characteristics of the Australian tree Acacia dealbata in its native range. Two subspecies of A. dealbata have previously been described based on morphology and environmental requirements, but recent phylogeographic data raised questions regarding the validity of this taxonomic subdivision. The species has been widely planted within and outside its native Australian range and is also a highly successful invasive species in many parts of the world. We employed microsatellite markers to investigate the population genetic diversity and structure among 42 A. dealbata populations from across the species’ native range. We also tested whether environmental variables purportedly relevant for the putative separation of subspecies are linked with population genetic differentiation. We found no relationship between population genetic structure of A. dealbata in Australia and these environmental features. Rather, we identified two geographically distinct genetic clusters that corresponded with populations in the northeastern part of mainland Australia, and the southern mainland and Tasmanian range of the species. Our results do not support the taxonomic subdivision of the species into two distinct subspecies based on environmental features. We therefore assume that the observed morphological differences between the putative subspecies are plastic phenotypic responses. This study provides population genetic information that will be useful for the conservation of the species within Australia as well as to better understand the invasion dynamics of A. dealbata.     

Low Genetic Diversity in Wide-Spread Eurasian Liver Fluke Opisthorchis felineus Suggests Special Demographic History of This Trematode Species

Opisthorchis felineus or Siberian liver fluke is a trematode parasite (Opisthorchiidae) that infects the hepato-biliary system of humans and other mammals. Despite its public health significance, this wide-spread Eurasian species is one of the most poorly studied human liver flukes and nothing is known about its population genetic structure and demographic history. In this paper, we attempt to fill this gap for the first time and to explore the genetic diversity in O. felineus populations from Eastern Europe (Ukraine, European part of Russia), Northern Asia (Siberia) and Central Asia (Northern Kazakhstan). Analysis of marker DNA fragments from O. felineus mitochondrial cytochrome c oxidase subunit 1 and 3 (cox1, cox3) and nuclear rDNA internal transcribed spacer 1 (ITS1) sequences revealed that genetic diversity is very low across the large geographic range of this species. Microevolutionary processes in populations of trematodes may well be influenced by their peculiar biology. Nevertheless, we suggest that lack of population genetics structure observed in O. felineus can be primarily explained by the Pleistocene glacial events and subsequent sudden population growth from a very limited group of founders. Rapid range expansion of O. felineus through Asian and European territories after severe bottleneck points to a high dispersal potential of this trematode species.     

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.     

Can an a priori strategy be developed for biological control? The case of Onopordum spp. thistles in Australia

Abstract  Between 1992 and 2000, seven insect agents were released in Australia for the biological control of Onopordum spp. thistles. This paper describes the protocol used for the selection of these agents, starting with the development of a preliminary strategy, based on the ecology and population dynamics of the target weed. The strategy informed the surveys for natural enemies in the native range of Onopordum , targeting insects that attacked key transitional stages of the weed's life cycle. Ongoing studies of Onopordum populations in both Australia and Europe, plus experimental studies on the ecology, potential impact and preliminary host specificity of the agents, led to the refinement of the strategy and the selection and prioritisation of the agents. It is argued that development of an explicit strategy prior to release should be encouraged, as it forces researchers to revisit the rationale for and aims of particular biological control projects, ensuring that the process of agent selection remains focused. It also provides a tool to improve the process of agent selection, as subsequent results can be measured against the strategy and agent success or failure evaluated against the a priori expectations.     

Cultivation shapes genetic novelty in a globally important invader

Acacia saligna is a species complex that has become invasive in a number of countries worldwide where it has caused substantial environmental and economic impacts. Understanding genetic and other factors contributing to its success may allow managers to limit future invasions of closely related species. We used three molecular markers to compare the introduced range (South Africa) to the native range (Western Australia). Nuclear markers showed that invasive populations are divergent from native populations and most closely related to a cultivated population in Western Australia. We also found incongruence between nuclear and chloroplast data that, together with the long history of cultivation of the species, suggest that introgressive hybridization (coupled with chloroplast capture) may have occurred within A. saligna. While we could not definitively prove introgression, the genetic distance between cultivated and native A. saligna populations was comparable to known interspecific divergences among other Acacia species. Therefore, cultivation, multiple large‐scale introductions and possibly introgressive hybridization have rapidly given rise to the divergent genetic entity present in South Africa. This may explain the known global variation in invasiveness and inaccuracy of native bioclimatic models in predicting potential distributions.     

Widespread hybridization in the introduced hog deer population of Victoria,Australia, and its implications for conservation

In Australia, many species have been introduced that have since undergone drastic declines in their native range. One species of note is the hog deer (Axis porcinus) which was introduced in the 1860s to Victoria, Australia, and has since become endangered in its native range throughout South‐East Asia. There is increased interest in using non‐native populations as a source for genetic rescue; however, considerations need to be made of the genetic suitability of the non‐native population. Three mitochondrial markers and two nuclear markers were sequenced to assess the genetic variation of the Victorian population of hog deer, which identified that the Victorian population has hybrid origins with the closely related chital (Axis axis), a species that is no longer present in the wild in Victoria. In addition, the mitochondrial D‐loop region within the Victorian hog deer is monomorphic, demonstrating that mitochondrial genetic diversity is very low within this population. This study is the first to report of long‐term persistence of hog deer and chital hybrids in a wild setting, and the continual survival of this population suggests that hybrids of these two species are fertile. Despite the newly discovered hybrid status in Victorian hog deer, this population may still be beneficial for future translocations within the native range. However, more in‐depth analysis of genetic diversity within the Victorian hog deer population and investigation of hybridization rates within the native range are necessary before translocations are attempted.     

Locating the sources of an invasive pest,grape phylloxera,using a mitochondrial DNA gene genealogy

Range expansions through human introductions have increased with global commerce and have led to the extinction of native species, alterations in community structure and pest status of the invasive species. Inferring the evolutionary history of invasive species can help to build a firmer footing for management tactics. This study used mitochondrial DNA (mtDNA) sequence comparisons of samples collected from the native and introduced ranges of a pest herbivore of cultivated grapes, grape phylloxera (Daktulosphaira vitifoliae Fitch, Phylloxeridae) to infer the sources and pattern of introductions into worldwide viticulture. Introductions into viticulture from its native North American range first occurred in the mid-19th century. The pattern of spread has suggested a focus of introduction into France, but independent introductions may have occurred elsewhere. The results show that the introduced population represents a limited subsample of the native genetic diversity. The data suggest that most grape phylloxera in viticulture, including all European, have originated in the northeastern USA where the grape species Vitis riparia dominates. There was evidence for independent introductions into South Africa and California. Most California haplotypes were most closely related to native grape phylloxera from the Atlantic Coast on V. vulpina. It is likely that subsequent spread from California into Australia, New Zealand and Peru has occurred.     

Within-Range Translocations and Their Consequences in European Larch

In contrast to biological invasions, translocations of individuals within a species range are understudied, due to difficulties in systematically detecting them. This results in limited knowledge about the corresponding processes and uncertainties regarding the status of extant populations. European larch, a forest tree whose fragmented native distribution is restricted to the Alps and to other Central European mountains, has been massively planted for at least 300 years. Here we focus on the genetic characterization of translocations having taken place within its native range. Microsatellite variation at 13 nuclear loci and sequence data of two mitochondrial DNA fragments were analyzed on the basis of a comprehensive range-wide population sample. Two complementary methods (Geneclass and Structure) were used to infer translocation events based on nuclear data whereas mitochondrial data were used for validation of these inferences. Using Geneclass, we found translocation events in a majority of populations. Additional cases of translocation and many instances of admixture were identified using Structure, thanks to the clear-cut ancestral genetic structure detected in this species. In particular, a strong divide between Alpine and Central European populations, also apparent at mitochondrial markers, helped uncover details on translocation events and related processes. Translocations and associated admixture events were found to be heterogeneously distributed across the species range, with a particularly high frequency in Central Europe. Furthermore, translocations frequently involved multiple geographic sources, some of which were over-represented. Our study illustrates the importance of range-wide investigations for tracing translocations back to their origins and for revealing some of their consequences. It provides some first clues for developing suitable conservation and management strategies.     

The effect of habitat fragmentation on the bee visitor assemblages of three Australian tropical rainforest tree species

Tropical forest loss and fragmentation can change bee community dynamics and potentially interrupt plant–pollinator relationships. While bee community responses to forest fragmentation have been investigated in a number of tropical regions, no studies have focused on this topic in Australia. In this study, we examine taxonomic and functional diversity of bees visiting flowers of three tree species across small and large rainforest fragments in Australian tropical landscapes. We found lower taxonomic diversity of bees visiting flowers of trees in small rainforest fragments compared with large forest fragments and show that bee species in small fragments were subsets of species in larger fragments. Bees visiting trees in small fragments also had higher mean body sizes than those in larger fragments, suggesting that small‐sized bees may be less likely to persist in small fragments. Lastly, we found reductions in the abundance of eusocial stingless bees visiting flowers in small fragments compared to large fragments. These results suggest that pollinator visits to native trees living in small tropical forest remnants may be reduced, which may in turn impact on a range of processes, potentially including forest regeneration and diversity maintenance in small forest remnants in Australian tropical countryside landscapes.     

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

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

Evidence for multiple sources of invasion and intraspecific hybridization in Brachypodium sylvaticum (Hudson) Beauv. in North America

We compared the levels and distribution of genetic diversity in Eurasian and North American populations of Brachypodium sylvaticum (Huds.) Beauv. (false brome), a newly invasive perennial bunchgrass in western North America. Our goals were to identify source regions for invasive populations, determine the number of independent invasion events, and assess the possibility that postinvasion bottlenecks and hybridization have affected patterns of genetic diversity in the invaded range. We tested the hypothesis that this Eurasian grass was accidentally introduced into two areas in Oregon and one site in California by examining nuclear microsatellites and chloroplast haplotype variation in 23 introduced and 25 native populations. In the invaded range, there was significantly lower allelic richness (R(S)), observed heterozygosity (H(O)) and within-population gene diversity (H(S)), although a formal test failed to detect a significant genetic bottleneck. Most of the genetic variation existed among populations in the native range but within populations in the invaded range. All of the allelic variation in the invaded range could be explained based on alleles found in western European populations. The distribution of identified genetic clusters in the North American populations and the unique alleles associated with them is consistent with two historical introductions in Oregon and a separate introduction to California. Further analyses of population structure indicate that intraspecific hybridization among genotypes from geographically distinct regions of western Europe occurred following colonization in Oregon. The California populations, however, are more likely to be derived from one or perhaps several genetically similar regions in the native range. The emergence and spread of novel recombinant genotypes may be facilitating the rapid spread of this invasive species in Oregon.     

Historical range expansion determines the phylogenetic diversity introduced during contemporary species invasion

For a species rapidly expanding its geographic range, such as during biological invasion, most alleles in the introduced range will have their evolutionary origins in the native range. Yet, the way in which historical processes occurring over evolutionary time in the native range contribute to the diversity sampled during contemporary invasion is largely unknown. We used chloroplast DNA (cpDNA) gene genealogies and coalescent methods to study two congeneric plants, Silene latifolia and S. vulgaris. We examined how phylogenetic diversity was shaped by demographic growth and historical range expansions in the native European range, and how this history affected the diversity sampled during their recent invasion of North America. Genealogies from both species depart from neutrality, likely as a result of demographic expansion in the ancestral range, the timing of which corresponds to shortly after each species originated. However, the species differ in the spatial distribution of cpDNA lineages across the native range. Silene latifolia shows a highly significant phylogeographic structure that most likely reflects different avenues of the post-glacial expansion into northern Europe from Mediterranean refugia. By contrast, cpDNA lineages in S. vulgaris have been widely scattered across Europe during, or since, the most recent post-glacial expansion. These different evolutionary histories resulted in dramatic differences in how phylogenetic diversity was sampled during invasion of North America. In S. latifolia, relatively few, discrete invasion events from a structured native range resulted in a rather severe genetic bottleneck, but also opportunities for admixture among previously isolated lineages. In S. vulgaris, lack of genetic structure was accompanied by more representative sampling of phylogenetic diversity during invasion, and reduced potential for admixture. Our results provide clear insights into how historical processes may feed forward to influence the phylogenetic diversity of species invading new geographic ranges.