Bactrocera dorsalis (Hendel) (Diptera: Tephritidae) is not invasive through Asia: It's been there all along

Oriental fruit fly, Bactrocera dorsalis (Hendel), is a highly polyphagous fruit fly which, in the last 15 years, has invaded (with or without establishment) Africa, Europe and North America. As a direct result of these invasions, there is increasing research interest in the invasion history and spread patterns of this fly. A statement being repeatedly used in the B. dorsalis invasion literature is that the species was first identified from Taiwan in 1912 and that it subsequently spread through South‐East and South Asia during the 20th century. This assumption is incorrect and stems from: (a) an incomplete knowledge of B. dorsalis taxonomic history; and (b) a confounding of first taxonomic record with first invasion record. Rather than being first detected in Taiwan in 1912, the first record of oriental fruit fly was from “East India” (India orientali) under the synonymous name of Musca ferruginea by Fabricius in 1794, and in the 1910s, it was known not only from Taiwan, but widely across tropical Asia with records from India, Burma, Bengal, Sri Lanka (as Ceylon), Singapore and Indonesia (multiple islands). The taxonomic literature is very clear that oriental fruit fly has not invaded the rest of Asia from Taiwan since 1912, and this error should not continue to be repeated in the literature.     

Phylogenetics,delimitation and historical biogeography of the pantropical tree genus Thespesia (Malvaceae,Gossypieae)

Thespesia consists of 16 species of trees and shrubs from Southeast Asia–Oceania, Africa and America, the most well known being T. populnea, a small tree of tropical coastal areas around the world. Phylogenetic relationships in the genus and among its allies in tribe Gossypieae were inferred using three plastid and two nuclear regions to ascertain its generic delimitation and explore its biogeographical history. Maximum‐likelihood and Bayesian analyses confirmed that Thespesia is not monophyletic and, based on these results, Azanza is reinstated to accommodate the two species previously placed in Thespesia section Lampas. Dating analyses and ancestral range estimation indicated that Thespesia s.s. most likely originated in Southeast Asia–Oceania c. 30 Mya, but extant species did not begin to differentiate until the late Miocene. Two dispersal events, one into Africa c. 11 Mya and another into America (Antilles) c. 9 Mya, gave rise to the African and the Greater Antillean endemics, respectively. The two most widespread hydrochorous species, T. populnea and T. populneoides, originated in Southeast Asia–Oceania from where they spread to other parts of the world. Our analysis also indicated a much earlier origin than previously reported for the eumalvoid clade and its tribes Gossypieae, Malveae and Hibisceae suggesting that vicariance might have had an important role early in the history of these groups.     

Resistance to three thrips species in Capsicum spp. depends on site conditions and geographic regions

Capsicum species are commercially grown for pepper production. This crop suffers severely from thrips damage and the identification of natural sources of thrips resistance is essential for the development of resistant cultivars. It is unclear whether resistance to Frankliniella occidentalis as assessed in a specific environment holds under different conditions. Additionally, other thrips species may respond differently to the plant genotypes. Screening for robust and general resistance to thrips encompasses testing different Capsicum accessions under various conditions and with different thrips species. We screened 11 Capsicum accessions (C. annuum and C. chinense) for resistance to F. occidentalis at three different locations in the Netherlands. Next, the same 11 accessions were screened for resistance to Thrips palmi and Scirtothrips dorsalis at two locations in Asia. This resulted in a unique analysis of thrips resistance in Capsicum at five different locations around the world. Finally, all accessions were also screened for resistance to F. occidentalis in the Netherlands using a leaf disc choice assay, allowing direct comparison of whole plant and leaf disc assays. Resistance to F. occidentalis was only partially consistent among the three sites in the Netherlands. The most susceptible accessions were consistently susceptible, but which accession was the most resistant differed among sites. In Asia, one C. chinense accession was particularly resistant to S. dorsalis and T. palmi, but this was not the most resistant accession to F. occidentalis. Overall, resistance to F. occidentalis correlated with S. dorsalis but not with T. palmi resistance in the C. annuum accessions. Damage inflicted on leaf discs reflected damage on the whole plant level. Our study showed that identifying broad spectrum resistance to thrips in Capsicum may prove to be challenging. Breeding programmes should focus on developing cultivars suitable for growing in defined geographic regions with specific thrips species and abiotic conditions.     

Global genetic diversity of Aedes aegypti

Mosquitoes, especially Aedes aegypti, are becoming important models for studying invasion biology. We characterized genetic variation at 12 microsatellite loci in 79 populations of Ae. aegypti from 30 countries in six continents, and used them to infer historical and modern patterns of invasion. Our results support the two subspecies Ae. aegypti formosus and Ae. aegypti aegypti as genetically distinct units. Ae. aegypti aegypti populations outside Africa are derived from ancestral African populations and are monophyletic. The two subspecies co‐occur in both East Africa (Kenya) and West Africa (Senegal). In rural/forest settings (Rabai District of Kenya), the two subspecies remain genetically distinct, whereas in urban settings, they introgress freely. Populations outside Africa are highly genetically structured likely due to a combination of recent founder effects, discrete discontinuous habitats and low migration rates. Ancestral populations in sub‐Saharan Africa are less genetically structured, as are the populations in Asia. Introduction of Ae. aegypti to the New World coinciding with trans‐Atlantic shipping in the 16th to 18th centuries was followed by its introduction to Asia in the late 19th century from the New World or from now extinct populations in the Mediterranean Basin. Aedes mascarensis is a genetically distinct sister species to Ae. aegypti s.l. This study provides a reference database of genetic diversity that can be used to determine the likely origin of new introductions that occur regularly for this invasive species. The genetic uniqueness of many populations and regions has important implications for attempts to control Ae. aegypti, especially for the methods using genetic modification of populations.     

Allopatric speciation in ticks: genetic and reproductive divergence between geographic strains of Rhipicephalus (Boophilus) microplus

Background  

The cattle tick, Rhipicephalus (Boophilus) microplus, economically impact cattle industry in tropical and subtropical regions of the world. The morphological and genetic differences among R. microplus strains have been documented in the literature, suggesting that biogeographical and ecological separation may have resulted in boophilid ticks from America/Africa and those from Australia being different species. To test the hypothesis of the presence of different boophilid species, herein we performed a series of experiments to characterize the reproductive performance of crosses between R. microplus from Australia, Africa and America and the genetic diversity of strains from Australia, Asia, Africa and America.     

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

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

The photokinesis of oriental fruit flies,Bactrocera dorsalis,to LED lights of various wavelengths

The oriental fruit fly, Bactrocera dorsalis Hendel (Diptera: Tephritidae), is an invasive pest of orchards around the world, particularly in Asian countries such as China. Light traps offer a potential means for pest monitoring and management. This study aimed to evaluate the sensitivity of the fly to light and investigate the impact of monochromatic light in the sensitivity spectrum on B. dorsalis. Six light wavelengths in LEDs – green (522 nm), yellow (596 nm), blue (450 nm), red (633 nm), purple (440 nm), and white (compound light) – were adapted to test responses of 5‐, 10‐, and 20‐day‐old B. dorsalis adults kept in laboratory conditions. We also tested the effects of green and red lights on pupal development and adults’ life activities. The results indicated a phototaxis preference rank in B. dorsalis adults to monochromatic LEDs with, in decreasing order, green, yellow, purple, blue, and red. Moreover, positive phototaxis significantly increased with age. Male adults are more sensitive than female adults to test lights, mainly at the age of 10 and 20 days. Emergence rates of pupae exposed to 12 and 24 h green light daily were 42 and 67%, respectively, whereas controls held in red light emerged at 33 and 37%, respectively. Furthermore, body weight, female fecundity, and mortality of B. dorsalis in night‐time exposure of green light (from 21:00 to 09:00 hours; during daytime flies were illuminated by white LED light) were significantly higher than in red‐light test groups and dark controls. In conclusion, B. dorsalis displayed preference toward green light, and fly age and gender seemed to significantly impact the phototactic behavior. Green LED light exposure during nighttime remarkably improved the emergence rates of B. dorsalis, and it enhanced growth, development, and ovipositing peak period, but decreased adult lifespan. This research lays a foundation for the development of new trap models, e.g., with green sticky cards or green light, for monitoring and control of B. dorsalis in the field.     

Cryptic genetic variation in an inbreeding and cosmopolitan pest,Xylosandrus crassiusculus,revealed using ddRADseq

Each year new exotic species are transported across the world through global commerce, causing considerable economic and ecological damage. An important component of managing invasion pathways is to identify source populations. Some of the most widespread exotic species are haplodiploid ambrosia beetles. The ability to mate with siblings (inbreed) and their transportable food source (symbiotic fungus) have enabled them to colonize most of the world and become pests of plant nurseries, lumber, and forests. One of the fastest spreading ambrosia beetles is Xylosandrus crassiusculus. In order to discover the source populations of this globally invasive species, track its movement around the world, and test biogeographical scenarios, we combined restriction site‐associated DNA sequencing (RADseq) with comprehensive sampling across the species native and introduced range. From 1,365 genotyped SNP loci across 198 individuals, we determined that in its native range, X. crassiusculus is comprised of a population in Southeast Asia that includes mainland China, Thailand, and Taiwan, and a second island population in Japan. North America and Central America were colonized from the island populations, while Africa and Oceania were colonized from the mainland Asia, and Hawaii was colonized by both populations. Populations of X. crassiusculus in North America were genetically diverse and highly structured, suggesting (1) numerous, repeated introductions; (2) introduction of a large founding population; or (3) both scenarios with higher than expected outcrossing. X. crassiusculus, other wood‐boring insects, and indeed many other pests with unusual genetic structure continue to spread around the world. We show that contemporary genetic methods offer a powerful tool for understanding and preventing pathways of future biosecurity threats.     

The global spread of <Emphasis Type="Italic">Harmonia axyridis</Emphasis> (Coleoptera: Coccinellidae): distribution,dispersal and routes of invasion

Released as a biological control agent of aphids and coccids, Harmonia axyridis (Coleoptera: Coccinellidae) has spread from Asia to four additional continents. Since 1988 H. axyridis has established in at least 38 countries in its introduced range: three countries in North America, six in South America, 26 in Europe and three in Africa. In different continents the species has spread at rates estimated between 100 and 500 km year⁻¹. Here, the global spread of H. axyridis is thoroughly reviewed. Mechanisms of short- and long-distance dispersal in coccinellids are discussed, as are the reasons for them, with particular emphasis on H. axyridis. Dispersal via anthropogenic means has been particularly important in the case of H. axyridis. Preliminary studies investigating the invasion routes of H. axyridis using genetic analyses (involving both microsatellite and mitochondrial DNA) are outlined.     

基于MaxEnt模型的入侵植物白花鬼针草的分布预测及适生性分析

【背景】白花鬼针草为农区恶性杂草,原产于美洲,现已广泛分布于世界热带及亚热带地区,但其在全球和中国的适生区域及适生等级还不明确。【方法】利用MaxEnt生态位模型对白花鬼针草在全球以及中国的潜在适生区进行预测。【结果】白花鬼针草在全球的分布更多受到温度因素的影响。白花鬼针草的适生区主要集中在北半球和南半球15°~30°之间的热带和亚热带地区。其中,北美南部、南美中南部、非洲南部、东南亚北部以及大洋洲中南部沿海地区为白花鬼针草中、高度适生区。白花鬼针草在中国的适生区主要位于广东、广西、海南、云南、福建、台湾。到2070年,白花鬼针草在全球的适生区面积与当前相似,但在中国的适生区有所增大。【结论】白花鬼针草在我国有进一步扩张的风险。     

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.     

One and the same: integrative taxonomic evidence that Bactrocera invadens (Diptera: Tephritidae) is the same species as the Oriental fruit fly Bactrocera dorsalis

The invasive fruit fly Bactrocera invadens Drew, Tsuruta & White, and the Oriental fruit fly Bactrocera dorsalis (Hendel) are highly destructive horticultural pests of global significance. Bactrocera invadens originates from the Indian subcontinent and has recently invaded all of sub‐Saharan Africa, while B. dorsalis principally occurs from the Indian subcontinent towards southern China and South‐east Asia. High morphological and genetic similarity has cast doubt over whether B. invadens is a distinct species from B. dorsalis. Addressing this issue within an integrative taxonomic framework, we sampled from across the geographic distribution of both taxa and: (i) analysed morphological variation, including those characters considered diagnostic (scutum colour, length of aedeagus, width of postsutural lateral vittae, wing size, and wing shape); (ii) sequenced four loci (ITS1, ITS2, cox1 and nad4) for phylogenetic inference; and (iii) generated a cox1 haplotype network to examine population structure. Molecular analyses included the closely related species, Bactrocera kandiensis Drew & Hancock. Scutum colour varies from red‐brown to fully black for individuals from Africa and the Indian subcontinent. All individuals east of the Indian subcontinent are black except for a few red‐brown individuals from China. The postsutural lateral vittae width of B. invadens is narrower than B. dorsalis from eastern Asia, but the variation is clinal, with subcontinent B. dorsalis populations intermediate in size. Aedeagus length, wing shape and wing size cannot discriminate between the two taxa. Phylogenetic analyses failed to resolve B. invadens from B. dorsalis, but did resolve B. kandiensis. Bactrocera dorsalis and B. invadens shared cox1 haplotypes, yet the haplotype network pattern does not reflect current taxonomy or patterns in thoracic colour. Some individuals of B. dorsalis/B. invadens possessed haplotypes more closely related to B. kandiensis than to conspecifics, suggestive of mitochondrial introgression between these species. The combined evidence fails to support the delimitation of B. dorsalis and B. invadens as separate biological species. Consequently, existing biological data for B. dorsalis may be applied to the invasive population in Africa. Our recommendation, in line with other recent publications, is that B. invadens be synonymized with B. dorsalis.     

Mutation in the ace‐1 gene of the tomato leaf miner (Tuta absoluta) associated with organophosphates resistance

The tomato leaf miner, Tuta absoluta (Lepidoptera: Gelechiidae), is a major invasive pest that has spread throughout many countries in the Mediterranean basin and parts of Asia over the last decade. The control of T. absoluta has relied heavily on the use of chemical insecticides, a strategy that has led to the evolution of resistance. In this study, biological and molecular methods were used to determine the susceptibility of five strains of T. absoluta to the organophosphate chlorpyrifos and to investigate the molecular mechanisms underlying resistance to this class of insecticides. High levels of resistance to chlorpyrifos were observed in all five strains tested. Cloning and sequencing of the gene encoding the organophosphate target site, ace‐1, of T. absoluta revealed the presence of an alanine to serine substitution at a position that has been previously linked with organophosphate resistance across a range of different insect and mite species. The presence of this mutation at high frequency in T. absoluta populations originating from various countries further supports the suggestion that the rapid expansion of this species is, in part, mediated by the resistance of this pest to chemical insecticides.     

Demographic expansion and genetic load of the halophyte model plant Eutrema salsugineum

The halophyte model plant Eutrema salsugineum (Brassicaceae) disjunctly occurs in temperate to subarctic Asia and North America. This vast, yet extremely discontinuous distribution constitutes an ideal system to examine long‐distance dispersal and the ensuing accumulation of deleterious mutations as expected in expanding populations of selfing plants. In this study, we resequenced individuals from 23 populations across the range of E. salsugineum. Our population genomic data indicate that E. salsugineum migrated “out of the Altai region” at least three times to colonize northern China, northeast Russia and western China. It then expanded its distribution into North America independently from northeast Russia and northern China, respectively. The species colonized northern China around 33.7 thousand years ago (kya) and underwent a considerable expansion in range size approximately 7–8 kya. The western China lineage is likely a hybrid derivative of the northern China and Altai lineages, originating approximately 25–30 kya. Deleterious alleles accumulated in a stepwise manner from (a) Altai to northern China and North America and (b) Altai to northeast Russia and North America. In summary, E. salsugineum dispersed from Asia to North America and deleterious mutations accumulated in a stepwise manner during the expansion of the species’ distribution.     

Population structure and gene flow in the global pest,Helicoverpa armigera

Helicoverpa armigera is a major agricultural pest that is distributed across Europe, Asia, Africa and Australasia. This species is hypothesized to have spread to the Americas 1.5 million years ago, founding a population that is at present, a distinct species, Helicoverpa zea. In 2013, H. armigera was confirmed to have re‐entered South America via Brazil and subsequently spread. The source of the recent incursion is unknown and population structure in H. armigera is poorly resolved, but a basic understanding would highlight potential biosecurity failures and determine the recent evolutionary history of region‐specific lineages. Here, we integrate several end points derived from high‐throughput sequencing to assess gene flow in H. armigera and H. zea from populations across six continents. We first assemble mitochondrial genomes to demonstrate the phylogenetic relationship of H. armigera with other Heliothine species and the lack of distinction between populations. We subsequently use de novo genotyping‐by‐sequencing and whole‐genome sequences aligned to bacterial artificial chromosomes, to assess levels of admixture. Primarily, we find that Brazilian H. armigera are derived from diverse source populations, with strong signals of gene flow from European populations, as well as prevalent signals of Asian and African ancestry. We also demonstrate a potential field‐caught hybrid between H. armigera and H. zea, and are able to provide genomic support for the presence of the H. armigera conferta subspecies in Australasia. While structure among the bulk of populations remains unresolved, we present distinctions that are pertinent to future investigations as well as to the biosecurity threat posed by H. armigera.     

Efficacy of biorational insecticides against chilli thrips,Scirtothrips dorsalis (Thysanoptera: Thripidae), infesting roses under nursery conditions

We evaluated biopesticides based on entomopathogenic fungi, azadirachtin and horticultural oils for the management of the chilli thrips, Scirtothrips dorsalis Hood, an economically significant invasive pest in the United States. Insecticides were applied four times at 7‐ to 10‐day intervals against established S. dorsalis infestations on shrub roses KnockOut^®, Rosa x ‘Radrazz’ under simulated nursery conditions. When applied as stand‐alone treatments, Beauveria bassiana GHA (BotaniGard^® ES), Metarhizium brunneum F52 (Met‐52 EC), a horticultural oil (SuffOil‐X^®) and azadirachtin (Molt‐X^®) at label rates provided significant control, reducing populations of S. dorsalis by 48–71% compared with control over 4–6 weeks. Similar results were observed when the biopesticides were applied in rotation with each other. A conventional standard, spinosad (Conserve^® SC), was consistently the most effective treatment in these studies, reducing thrips populations by >95% overall. In another study, more effective control (87%–92%) was achieved in biopesticide rotation programmes that included spinosad, when compared with those that did not. Results also showed that these biopesticides can be tank‐mixed. However, there was no evidence that B. bassiana or M. brunneum combined with azadirachtin resulted in additive or synergistic control, as neither tank‐mix treatment improved control compared with azadirachtin alone. These findings highlight the potential use of biopesticides in rotation programmes with conventional insecticides to manage S. dorsalis on roses. Biopesticides evaluated in this study can be incorporated into an IPM programme for roses.     

Genetic diversity analysis of French goat populations reveals selective sweeps involved in their differentiation

After domestication 11 000 years ago in Asia Minor, the goat followed human migration to Europe and Asia. It was then introduced in Africa and is now raised all over the world. In this study, we exploited a dataset composed of 54 000 SNPs (Illumina goat DNA chip) to analyze the genetic diversity of 223 individuals belonging to eight French breeds (Alpine, Angora, Corse, Fossés, Poitevine, Provençale, Pyrénées and Saanen). Analyses carried out included individual‐based approaches (principal component analysis and population structure) and population‐based approaches (phylogenetic tree constructions). The results of the genetic diversity analyses revealed that French breeds are clearly differentiated, in particular, the Angora breed that originates from south west Asia. The Provençale breed shows a very original genetic pattern that could be the result of ancient admixture. Then, selection signatures were detected by identifying regions of outlying genetic differentiation between populations. Five genomic regions were detected under selection on chromosomes 5, 6, 11, 13 and 20, revealing mainly soft selective sweeps and a few hard selective sweeps and highlighting candidate genes that had been selected for during the evolutionary history of these breeds. Among them, two coat coloration genes (ADAMTS20 and ASIP) and one gene related to milk composition (CSN1S1) were involved.