Fly Community Research Articles

Diversity and species composition of fruit flies (Diptera: Tephritidae) in Lombok Island, Indonesia

Abstract. Hudiwaku S, Himawan T, Rizali A. 2021. Diversity and species composition of fruit flies (Diptera: Tephritidae) in Lombok Island, Indonesia. Biodiversitas 22: 4608-4616. Fruit flies (Diptera: Tephritidae) are pests of several horticultural crops that can reduce the quality and quantity of fruit production. Information on fruit flies in Lombok Island, Indonesia, is still limited. However, it is predicted to have a high diversity of fruit flies because this island belongs to the Wallacea region. The aims of this research was to study the diversity and species composition of fruit flies in different habitat types in Lombok Island. The research was carried out on two habitat types, i.e., tropical rainforest and orchard with each habitat type consisted of three different sites that located spread across Lombok Island as replication. The research was carried out on two habitat types, i.e., tropical rainforest and orchard, with each habitat type consisted of three different sites spread across Lombok Island as replication. A sampling of fruit flies was conducted using parapheromone traps from March to June 2020. Twenty-two species and 210,267 individual fruit flies were collected from all locations during the study period. The most dominant species were Bactrocera carambolae, Bactrocera limbifera, Zeugodacus caudatus, and Bactrocera dorsalis. Based on the ANOVA, habitat types significantly affected the abundance but not the species richness of fruit flies. The visualization results obtained from the NMDS ordination indicated a difference in the species composition of fruit flies between the two habitats. In conclusion, habitat types are an essential factor in shaping the community of fruit flies in Lombok Island.

The importance of environmental microbes for Drosophila melanogaster during seasonal macronutrient variability

Experiments manipulating the nutritional environment and the associated microbiome of animals have demonstrated their importance for key fitness components. However, there is little information on how macronutrient composition and bacterial communities in natural food sources vary across seasons in nature and on how these factors affect the fitness components of insects. In this study, diet samples from an orchard compost heap, which is a natural habitat for many Drosophila species and other arthropods, were collected over 9 months covering all seasons in a temperate climate. We developed D. melanogaster on diet samples and investigated stress resistance and life-history traits as well as the microbial community of flies and compost. Nutrient and microbial community analysis of the diet samples showed marked differences in macronutrient composition and microbial community across seasons. However, except for the duration of development on these diet samples and Critical Thermal maximum, fly stress resistance and life-history traits were unaffected. The resulting differences in the fly microbial community were also more stable and less diverse than the microbial community of the diet samples. Our study suggests that when D. melanogaster are exposed to a vastly varying nutritional environment with a rich, diverse microbial community, the detrimental consequences of an unfavourable macronutrient composition are offset by the complex interactions between microbes and nutrients.

Analysis on the relationship between winter precipitation and the annual variation of horse stomach fly community in arid desert steppe, Northwest China (2007-2019).

Gasterophilus spp. have been found to be widespread in reintroduced Przewalski's horses in the Kalamaili Nature Reserve (Northwest China). However, data on the annual variation in Gasterophilus infections are lacking. To analyze the epidemiological features and determine the cause of the annual variation in Gasterophilus infections, we treated 110 Przewalski's horses with ivermectin and collected Gasterophilus larvae from fecal samples each winter from 2007 to 2019. All 110 Przewalski's horses studied were found to be infected by Gasterophilus spp., and a total of 141 379 larvae were collected. Six species of Gasterophilus were identified with the following prevalence: G. pecorum (100%), G. nasalis (96.36%), G. nigricornis (94.55%), G. haemorrhoidalis (56.36%), G. intestinalis (59.09%), and G. inermis (3.64%). The mean infection intensity of Gasterophilus spp. larvae in Przewalski's horses was 1285 ± 653. G. pecorum (92.96% ± 6.71%) was the most abundant species. The intensity of Gasterophilus spp. (r = –0.561, P < 0.046) was significantly correlated with winter precipitation. Our findings confirmed that, in the Kalamaili Nature Reserve, gasterophilosis is a severe parasitic disease in Przewalski's horses. Winter precipitation at the beginning of the year can indirectly affect the intensity and composition of Gasterophilus spp. in Przewalski's horses at the end of the year. Therefore, the water‐related ecological regulation should be carried out to help reduce the parasite infection of Przewalski's horses.

Ecological determinants of black fly assemblages of relict mountain streams in northeastern Algeria, plus new records (Diptera: Simuliidae)

Despite their ecological and socio-economic significance, the black fly communities of Algerian hydrosystems have been understudied. The simuliid fauna of relict mountain streams of northeastern Algeria was sampled at 24 localities in two distinct and relatively uncharted watersheds. Species richness in the El Kala Highlands varied from eight taxa in the most species-rich localities to the virtual absence of any taxon in some localities. Species richness also mirrored the hydroperiod of most localities and reached its peak in late winter/early spring. Co-inertia analysis indicated a significant co-structure between environmental descriptors and black fly assemblages, suggesting a strong influence of the habitat templet on aquatic insect community structure. A comparison with an additional survey of 23 localities spread across northern Algeria suggests distinct black fly assemblages characterizing the El Kala Highlands.

Culturable bacteria in adults of a Southeast Asian black fly, Simulium tani (Diptera:Simuliidae)

Although the microbiome of blood-feeding insects serves an integral role in host physiology, both beneficial and pathogenic, little is known of the microbial community of black flies. An investigation, therefore, was undertaken to identify culturable bacteria from one of Malaysia's most common black flies, Simulium tani Takaoka and Davies, using 16S rDNA sequencing, and then evaluate the isolates for antibiotic resistance and virulence genes. A total of 20 isolates representing 11 bacterial species in four genera were found. Five isolates showed β-hemolysis on Columbia agar, and virulence genes were found in three of these isolates. Some degree of resistance to six of the 12 tested antibiotics was found among the isolates. The baseline data from this study suggest rich opportunities for comparative studies exploring the diversity and roles of the microbiome of S. tani and other Southeast Asian black flies.

A Superfamily-wide Activity Atlas of Serine Hydrolases in Drosophila melanogaster.

The serine hydrolase (SH) superfamily is, perhaps, one of the largest functional enzyme classes in all forms of life and consists of proteases, peptidases, lipases, and carboxylesterases as representative members. Consistent with the name of this superfamily, all members, without any exception to date, use a nucleophilic serine residue in the enzyme active site to perform hydrolytic-type reactions via a two-step ping-pong mechanism involving a covalent enzyme intermediate. Given the highly conserved catalytic mechanism, this superfamily has served as a classical prototype in the development of several platforms of chemical proteomics techniques, activity-based protein profiling (ABPP), to globally interrogate the functions of its different members in various native, yet complex, biological settings. While ABPP-based proteome-wide activity atlases for SH activities are available in numerous organisms, including humans, to the best of our knowledge, such an analysis for this superfamily is lacking in any insect model. To address this, we initially report a bioinformatics analysis toward the identification and categorization of nonredundant SHs in Drosophila melanogaster. Following up on this in silico analysis, leveraging discovery chemoproteomics, we identify and globally map the full complement of SH activities during various developmental stages and in different adult tissues of Drosophila. Finally, as a proof of concept of the utility of this activity atlas, we highlight sexual dimorphism in SH activities across different tissues in adult D. melanogaster, and we propose new research directions, resources, and tools that this study can provide to the fly community.

Interactions between plants and pollinators across urban and rural farming landscapes

Arthropods are responsible for pollinating the majority of food and fuel crops worldwide. However, global declines in bee populations threaten the delivery of pollination services in both managed and natural ecosystems. Alternative pollinators such as flies, butterflies, beetles, and wasps may provide a buffer that protects agriculture from bee population declines. Flower-visiting flies are abundant in both rural and urban agricultural settings, and in some cases may pollinate plants avoided by bees. Here we explored the structure of plant-pollinator networks for both bee and non-bee pollinator communities across rural and urban farms in western Washington State, USA. We identified several broad groups of pollinators comprising 13 morphogroups of insects and arthropods. While bees were the pollinators observed on reproductive parts of flowers most frequently, syrphid flies contributed to a high number of arthropod-flower visits. We also found that syrphid flies visited four plant types not visited by bees, and that urban and rural farm settings were characterized by different syrphid fly communities. When comparing the networks of plant-arthropod interactions, urban farm settings had more diverse networks overall, with a greater richness of plant-arthropod interactions for both bee and non-bee pollinators. These findings underscore the need for greater focus on alternative pollinators as contributors to pollination services, particularly in urban settings where pollination services may be more impacted by bee declines and habitat loss. Our work will inform specific pollinator conservation efforts in the Pacific Northwest region and more broadly across the United States.

Anastrepha spp. (Diptera: Tephritidae) in Ximenia americana and other tropical fruits in a Brazilian savannah

The aim of this work was to evaluate the infestation of Anastrepha species in several fruit tree species and A. alveata in wild plum Ximenia americana L. (Olacaceae), as well as the fruit fly community in a Savannah area using faunal indexes and population fluctuation. For this, we installed McPhail traps and carried out collections at Palmeiras home garden, municipality of São Pedro of Piauí, state of Piaui, northeastern Brazil. We noted high infestation of A. alveata in wild plum, with 144.2 puparia/kg, 216.1 puparia/kg and 321.6 puparia/kg for the years 2014, 2015 and 2016. The infestation indexes of A. obliqua in red mombin (Spondias purpurea) were 146.1 puparia/kg in 2016 and 479.2 puparia/kg in 2017, and in umbu-hog plum (Spondias sp.) 607.3 puparia/kg in 2017, both from the Anacardiaceae family. The species that composed the fruit fly community were A. alveata, A. ethalea (predominant), A. obliqua, A. zenildae (dominant and accessory), A. sororcula, A. fraterculus and Anastrepha sp. (accidental). The population fluctuation of A. ethalea occurred from March to October, while A. alveata presented population peaks in the fruiting of wild plum, between October and December

Effects of Bacillus thuringiensis var. israelensison the Black Fly Communities (Diptera, Simuliidae) in Tropical Streams.

Bacillus thuringiensis var. israelensis (Bti) Berliner, 1911 is widely used in the biological control of black fly and mosquito populations. The objective of this study was to evaluate the effects of Bti on the black fly communities in streams in the Atlantic Forest domain. The study was carried out in eight streams of Serra do Mar, in the municipality of Ubatuba, São Paulo. Some parts of the streams in this locality have received applications of Bti fortnightly for more than 25years by the sanitary agency of the region. In each stream, two sections were sampled, with and without application of Bti (June 2015 and 2016). Black flies were collected and identified to the species level in the laboratory with 1382 larvae being identified, distributed in six species. Of this total, 73% of the individuals were found in sections where Bti was not applied. There was a difference in the abundance of simulids between the sections with and without Bti application, being more abundant in the latter. We measured the head capsule of individuals of the most abundant species, Simulium pertinax Kollar 1832, using a stereoscopic microscope with millimeter lens. These measurements demonstrated that there was a difference between the sections regarding the age structure of Simuliidae populations. In the sections without Bti application, there was a higher proportion of larvae in the last instar, while in the Bti-treated sections, smaller instars were predominant, possibly due to constant recolonization processes.

Exploring the influence of different habitats and their volatile chemistry in modulating sand fly population structure in a leishmaniasis endemic foci, Kenya.

Phlebotomine sand flies transmit many viral protozoan and bacterial pathogens of public health importance. Knowledge of the ecologic factors influencing their distribution at local scale can provide insights into disease epidemiology and avenues for targeted control. Animal sheds, termite mounds and houses are important peri-domestic and domestic habitats utilized by different sand flies as resting or breeding habitats. However, our knowledge for selection of these habitats by sand flies remains poor. Here, we tested the hypothesis that these habitat types harbor different composition of sand fly species and differ in their volatile chemistry that could influence sand fly selection. To achieve this, we employed CDC light traps following a cross-sectional survey to investigate the distribution of sand flies in the three habitats in an endemic site for leishmaniasis in Kenya. The study was carried out during the dry season, when sand flies are optimally abundant in 2018 and 2020. Sand fly abundance did not vary between the habitats, but species-specific differences in abundance was evident. Measures of sand fly community structure (Shannon diversity and richness) were highest in animal shed, followed by termite mound and lowest inside human dwelling (house). This finding indicates broader attraction of both sexes of sand flies and females of varying physiological states to animal sheds potentially used as breeding or resting sites, but also as a signal for host presence for a blood meal. Furthermore, gas chromatography-mass spectrometric analysis of volatiles collected from represented substrates associated with these habitats viz: human foot odor on worn socks (houses indoors), cow dung (animal sheds) and termite mounds (enclosed vent), revealed a total of 47 volatile organic compounds. Of these, 26, 35 and 16 were detected in human socks, cow dung and enclosed termite vent, respectively. Of these volatiles, 1-octen-3-ol, 6-methyl-5-hepten-2-one, α-pinene, benzyl alcohol, m-cresol, p-cresol and decanal, previously known as attractants for sandflies and other blood-feeding insects, were common to the habitats. Our results suggest that habitat volatiles may contribute to the composition of sand flies and highlight their potential for use in monitoring sand fly populations.

Scarless engineering of the Drosophila genome near any site-specific integration site.

We describe a simple and efficient technique that allows scarless engineering of Drosophila genomic sequences near any landing site containing an inverted attP cassette, such as a MiMIC insertion. This two-step method combines phiC31 integrase-mediated site-specific integration and homing nuclease-mediated resolution of local duplications, efficiently converting the original landing site allele to modified alleles that only have the desired change(s). Dominant markers incorporated into this method allow correct individual flies to be efficiently identified at each step. In principle, single attP sites and FRT sites are also valid landing sites. Given the large and increasing number of landing site lines available in the fly community, this method provides an easy and fast way to efficiently edit the majority of the Drosophila genome in a scarless manner. This technique should also be applicable to other species.

The Effect of Urbanization on Bats and Communities of Bat Flies (Diptera: Nycteribiidae and Streblidae) in the Amazon, Northern Brazil

Bats and bat flies are intimately associated and are relevant models to study the effects of anthropogenic impacts when bats are highly distributed in urban environments. The lack of knowledge on bat's responses to the anthropogenic effects as related to their bat flies leads to ineffective conservation strategies. Thus, our goal is to describe the association between bats and bat flies in urban and non-urban environments and measuring how much urbanization can alter the parasite-host associations. We collected bats and bat flies in 11 sites (six urban areas and five non-urban areas) distributed among five municipalities of the state of Para. To identify a possible pattern on communities, we conducted a Composition Principal Component Analysis and performed a permutational multivariate analysis of variance using distance matrices — PERMANOVA. We captured 513 bats and examined 867 bat flies. Our findings show that bat and bat fly communities of rural environments in the Amazon are more diverse than those from urban environments. Urbanization seems to play an essential role as an environmental filter of bats and bat fly species that are exclusively rural.

Assemblages of fruit flies (Diptera: Tephritidae) along an elevational gradient in the rainforests of Papua New Guinea

Abstract Papua New Guinea (PNG) has the greatest diversity of dacine fruit fly species (Diptera: Tephritidae: Dacinae) in the world, many of which are significant agricultural pests. Although their taxonomy is relatively well known, there is limited research on their geographical and elevational distribution. We undertook a survey of PNG's fruit fly fauna along a complete elevational gradient on Mt Wilhelm (175–3700 m a.s.l.) to determine the elevational species distribution and turnover of fruit fly communities. Fruit flies were sampled using a selection of male parapheromone lures, including Zingerone, a new and promising male attractant. In total, we collected 10 700 fruit flies representing 77 species. The total abundance and species richness of fruit flies decreased with increasing elevation. Species richness was similarly high at 175 and 200 m a.s.l. (30 and 27 species) and at 700 and 1200 m a.s.l. (16 and 20 species, respectively) but dropped suddenly to only two species at 1700 m a.s.l. and to zero at all four sites above this elevation. There were no flies attracted exclusively to Zingerone. One species (Bactrocera recurrens) exhibited dramatic, temporal changes in abundance during the study period. Fruit fly assemblages were significantly influenced by both the lure type and elevation. Similarity of fruit fly communities decayed linearly with increasing elevational distance. We concluded that the upper limit of fruit fly distribution in PNG occurs between 1700 and 2200 m a.s.l. and the centre of diversity occurs between 0 and 700 m a.s.l.

Diversity of robber flies (Diptera: Asilidae) in a tropical deciduous forest of central Mexico

Robber flies are an important group in ecosystems, because of their ecological role as predators, thus contributing to maintain a balance in insect populations. Nevertheless, little is known about their diversity and ecological interactions in Mexico. In this study, we analyzed the structure of a community of robber flies collected during three years in a tropical deciduous forest of central Mexico. In this type of vegetation, the dry and rainy seasons are contrasting and generate drastic biotic and abiotic changes that could impact on robber fly populations. Therefore, we compared the structure of the robber fly community across three seasons: cold, dry, and rainy. We collected a total of 191 individuals which included eight species and 21 morphospecies. The rainy season presented higher species richness and diversity, as well as a species composition that differed from that of the cold and dry seasons. However, abundance did not vary significantly across seasons. We also record 14 genera and ten species previously unregistered in the state of Puebla.

Evaluation of environmental and local factors on urban sarcosaprophagous Diptera: a case study with Sarcophagidae.

The urbanization process affects the structure of insect communities, favouring biotic homogenization processes by promoting adequate conditions for the establishment of synanthropic species. Considering the scarce knowledge about the ecology of Neotropical sarcophagids, we compared the flesh fly community from two different urban zones in two contrasting seasons and examined the response of the Sarcophagidae assemblage to landscape and microhabitat variables. Adult flies were collected in 13 sites with two traps (total number of traps 26) baited with 200 g of hydrated bone meal. A total of 2312 adult sarcophagids were collected belonging to 27 species; Oxysarcodexia paulistanensis (Mattos, 1919) represented >48% of the total sample. The local factor of insolation explained both the abundance and species richness of sarcophagids, collections being higher and richer in traps exposed to sunlight compared with traps placed in the shade regardless of the season. Richness was also affected by the environment, being higher in traps exposed in suburban rather than urban sites. The combination of these factors (sub-urban environment and availability of open areas) could explain the Sarcophagidae assemblage. The average number of sarcophagid species collected was related to the season, providing a good opportunity to assess the temporal variability of this family in the region.

First Study of Diptera Seasonal Dynamics and their Community Structure in Moroccan Vineyards

The purpose of this study was to describe the structure and composition of Diptera in Moroccan vineyard, using wet-traps sampling technique throughout 2017. The study focused on five varieties within Loudaya vineyard (west region of Marrakech, Morocco). Eleven different fly species were identified belonging to 10 families, Ceratitis capitata, Drosophilla suzukii and Thaumatiya notata were the most abundant species and first time behaviourally dominant flies species. In addition, more flies were captured in the local varieties (10 579 individuals) than in the introduced ones (966). Moreover, a significant difference was observed in fly richness, evenness and diversity between varieties. Fly community structure in Loudaya showed a clear seasonal dynamics, which richness, abundance, diversity and evenness were significantly (P < 0.001) dissimilar among months. Furthermore, Diptera species were positively influenced by humidity and rainfall. Richness (expected by sample-based rarefaction) and diversity peaked in the middle of vineyard phenology, corresponding to mid-spring to late-summer in all varieties. Fly species occurrence and seasonal climatic variation in Loudaya vineyard might be among various ecological factors such as habitat distribution, climate and the availability of resources. Taking into account the periods of fly abundance, the survey could give relevant indications on new approaches to integrated pest management (IPM) in Moroccan vineyards, particularly in the Loudaya region.

Soil salinity determines the diversity of snail-killing flies (Diptera: Sciomyzidae) in brackish marshes

Brackish marshes resulting from embankments are a rare habitat in Europe and for which there is little knowledge of the entomofauna. There is an urgent need to document the insect diversity and its unique properties in the light of ongoing habitat losses. The current study focuses on snail-killing flies in the remaining Belgian brackish marshes, in particular the role of soil salinity in determining their diversity and how it compares with that in freshwater marshes. Snail-killing fly communities in brackish marshland were relatively species poor and clearly different from those in freshwater, yet species richness and abundance increased with decreasing soil salinity. We recorded no habitat-specific species, but Tetanocera arrogans was more abundant in brackish marshes. Malaise traps proved to be a successful and repeatable method of sampling sciomyzid communities, the species composition of which was strongly site-specific. We conclude that extant Belgian brackish marshes have poor to moderately rich snail-killing fly communities that lack unique species. Richness and abundance, however, increased as soil salinity decreased.

Characterizing the microbiome of ectoparasitic louse flies feeding on migratory raptors.

Louse flies (Diptera: Hippoboscidae) are obligate ectoparasites that often cause behavioral, pathogenic, and evolutionary effects on their hosts. Interactions between ectoparasites and avian hosts, especially migrating taxa, may influence avian pathogen spread in tropical and temperate ecosystems and affect long-term survival, fitness and reproductive success. The purpose of this study was to characterize the vector-associated microbiome of ectoparasitic louse flies feeding on migrating raptors over the fall migration period. Surveys for louse flies occurred during fall migration (2015–2016) at a banding station in Pennsylvania, United States; flies were collected from seven species of migrating raptors, and we sequenced their microbial (bacteria and archaea) composition using high-throughput targeted amplicon sequencing of the 16S rRNA gene (V4 region). All louse flies collected belonged to the same species, Icosta americana. Our analysis revealed no difference in bacterial communities of louse flies retrieved from different avian host species. The louse fly microbiome was dominated by a primary endosymbiont, suggesting that louse flies maintain a core microbial structure despite receiving blood meals from different host species. Thus, our findings highlight the importance of characterizing both beneficial and potentially pathogenic endosymbionts when interpreting how vector-associated microbiomes may impact insect vectors and their avian hosts.

Effects of forest degradation on the sand fly communities of northeast Brazil.

The richness and abundance of sand fly species were studied in northeastern Brazil in areas of leishmaniasis transmission. The study was carried out in two forest areas with different deforestation times for agricultural and livestock activities: one modified by long-term settlement (more than 50 years of occupation) and another less impacted by short-term settlement (10 years). The sand flies were captured with CDC light traps from 18:00 to 06:00 for three consecutive nights, once a month, from May, 2012 to April, 2014. The study captured 21,708 specimens and also 33 species of Lutzomyia and two of Brumptomyia. Species richness and abundance were higher in the more conserved area of short-term occupation (31 species; 61.7%) than in the more degraded area with long-term occupation (17 species; 38.3%). In the most conserved area, the species richness was higher in the forest fragment than in the rural settlement, whereas in the degraded area the richness was higher in the peri-domicile than in the forest. The diversity was higher in the degraded area forest. There were significant statistical differences when comparing the means of total abundance with the intra-domicile, peri-domicile, and forest environments. The average abundance was statistically higher in the peri-domicile compared to the forest (p = 0.009), but there were no statistically significant differences between intra-domicile-peri-domicile (p = 0.11) and forest-intra-domicile (p = 0.87). In conclusion, a change in vegetation cover negatively affects the richness and abundance of sand flies in the natural environment.

The puzzling mitochondrial phylogeography of the black soldier fly (Hermetia illucens), the commercially most important insect protein species

BackgroundThe black soldier fly (Diptera: Stratiomyidae, Hermetia illucens) is renowned for its bioconversion ability of organic matter, and is the worldwide most widely used source of insect protein. Despite varying extensively in morphology, it is widely assumed that all black soldier flies belong to the same species, Hermetia illucens. We here screened about 600 field-collected and cultured flies from 39 countries and six biogeographic regions to test this assumption based on data for three genes (mitochondrial COI, nuclear ITS2 & 28S rDNA) and in order to gain insights into the phylogeography of the species.ResultsOur study reveals a surprisingly high level of intraspecific genetic diversity for the mitochondrial barcoding gene COI (divergences up to 4.9%). This level of variability is often associated with the presence of multiple species, but tested nuclear markers (ITS2 and 28S rDNA) were invariant and fly strain hybridization experiments under laboratory conditions revealed reproductive compatibility. COI haplotype diversity is not only very high in all biogeographic regions (56 distinct haplotypes in total), but also in breeding facilities and research centers from six continents (10 haplotypes: divergences up to 4.3%). The high genetic diversity in fly-breeding facilities is mostly likely due to many independent acquisitions of cultures via sharing and/or establishing new colonies from field-collected flies. However, explaining some of the observed diversity in several biogeographic regions is difficult given that the origin of the species is considered to be New World (32 distinct haplotypes) and one would expect severely reduced genetic diversity in the putatively non-native populations in the remaining biogeographic regions. However, distinct, private haplotypes are known from the Australasian (N = 1), Oriental (N = 4), and the Eastern Palearctic (N = 4) populations. We reviewed museum specimen records and conclude that the evidence for introductions is strong for the Western Palearctic and Afrotropical regions which lack distinct, private haplotypes.ConclusionsBased on the results of this paper, we urge the black soldier fly community to apply molecular characterization (genotyping) of the fly strains used in artificial fly-breeding and share these data in research publications as well as when sharing cultures. In addition, fast-evolving nuclear markers should be used to reconstruct the recent invasion history of the species.