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In recent years, the oak lace bug, Corythucha arcuata, has emerged as a significant threat to European oak forests. This species, native to North America, has in the last two decades rapidly extended its range in Europe, raising concerns about its potential impact on the continent's invaluable oak populations. To address this growing concern, we conducted an extensive study to assess the distribution, colonization patterns, and potential ecological niche of the oak lace bug in Europe. We gathered 1792 unique presence coordinates from 21 Eurasian countries, utilizing diverse sources such as research observations, citizen science initiatives, GBIF database, and social media reports. To delineate the realized niche and future distribution, we employed an ensemble species distribution modelling (SDM) framework. Two future greenhouse gas scenarios (RCP 4.5 and RCP 8.5) were considered across three-time intervals (2021-2040, 2061-2080, and 2081-2100) to project and evaluate the species' potential distribution in the future. Our analysis revealed that significant hotspots rich in host species occurrence for this invasive insect remain uninvaded so far, even within its suitable habitat. Furthermore, the native ranges of Turkey oak (Quercus cerris L.) and Hungarian oak (Quercus frainetto L.) species offer entirely suitable environments for the oak lace bug. In contrast, the pedunculate oak and sessile oak distribution ranges currently show only 40 % and 50 % suitability for colonization, respectively. However, our predictive models indicate a significant transformation in the habitat suitability of the oak lace bug, with suitability for these two oak species increasing by up to 90 %. This shift underlines an evolving landscape where the oak lace bug may exploit more of its available habitats than initially expected. It emphasises the pressing need for proactive measures to manage and stop its expanding presence, which may lead to a harmful impact on the oak population across the European landscape.
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Cambio Climático , Especies Introducidas , Quercus , Europa (Continente) , Animales , Ecosistema , Heterópteros , Distribución AnimalRESUMEN
The diameter distribution of a given species of deciduous trees is well approximated by a Gamma distribution. Here we give new experimental evidence for this conjecture by analyzing deciduous tree size data in mature semi-natural forest and ancient, traditionally managed wood-pasture from Central Europe. These distribution functions collapse on a universal shape if the tree sizes are normalized to the mean value in the considered sample. A new evolutionary master equation is used to model the observed distribution. The model incorporates four ecological processes: tree growth, mortality, recruitment, and diversification. Utilizing simple and realistic kernel functions describing the first three, along with an assumed multiplicative dilution due to diversification, the stationary solution of the master equation yields the experimentally observed Gamma distribution. The model as it is formulated allows an analytically compact solution and has only two fitting parameters whose values are consistent with the experimental data related to these processes. We found that the equilibrium size distribution of tree species with different ecology, originating from two contrastingly different semi-natural ecosystem types can be accurately described by a single dynamical mean-field model.
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The North American gall mite Aceria fraxiniflora was first recorded in Europe in southeast Hungary in 2017. Since then, it has shown a remarkably rapid spread on its host, the also North American green ash (Fraxinus pennsylvanica). By the beginning of 2023 it has been recorded in eight Central-Eastern European countries. In 2022 it was recorded on the other North American ash (Fraxinus Americana) in Zagreb (Croatia) and in Szarvas Arboretum (SE Hungary). Possible reasons and outcomes of this spread are discussed.
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Escarabajos , Fraxinus , Ácaros , Animales , Europa (Continente) , Europa OrientalRESUMEN
Non-native pests, climate change, and their interactions are likely to alter relationships between trees and tree-associated organisms with consequences for forest health. To understand and predict such changes, factors structuring tree-associated communities need to be determined. Here, we analysed the data consisting of records of insects and fungi collected from dormant twigs from 155 tree species at 51 botanical gardens or arboreta in 32 countries. Generalized dissimilarity models revealed similar relative importance of studied climatic, host-related and geographic factors on differences in tree-associated communities. Mean annual temperature, phylogenetic distance between hosts and geographic distance between locations were the major drivers of dissimilarities. The increasing importance of high temperatures on differences in studied communities indicate that climate change could affect tree-associated organisms directly and indirectly through host range shifts. Insect and fungal communities were more similar between closely related vs. distant hosts suggesting that host range shifts may facilitate the emergence of new pests. Moreover, dissimilarities among tree-associated communities increased with geographic distance indicating that human-mediated transport may serve as a pathway of the introductions of new pests. The results of this study highlight the need to limit the establishment of tree pests and increase the resilience of forest ecosystems to changes in climate.
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Ecosistema , Micobioma , Animales , Humanos , Filogenia , Bosques , Geografía , Cambio Climático , InsectosRESUMEN
International trade in plants and climate change are two of the main factors causing damaging tree pests (i.e. fungi and insects) to spread into new areas. To mitigate these risks, a large-scale assessment of tree-associated fungi and insects is needed. We present records of endophytic fungi and insects in twigs of 17 angiosperm and gymnosperm genera, from 51 locations in 32 countries worldwide. Endophytic fungi were characterized by high-throughput sequencing of 352 samples from 145 tree species in 28 countries. Insects were reared from 227 samples of 109 tree species in 18 countries and sorted into taxonomic orders and feeding guilds. Herbivorous insects were grouped into morphospecies and were identified using molecular and morphological approaches. This dataset reveals the diversity of tree-associated taxa, as it contains 12,721 fungal Amplicon Sequence Variants and 208 herbivorous insect morphospecies, sampled across broad geographic and climatic gradients and for many tree species. This dataset will facilitate applied and fundamental studies on the distribution of fungal endophytes and insects in trees.
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Endófitos , Hongos , Insectos , Animales , Biodiversidad , ÁrbolesRESUMEN
Novel interactions between introduced oaks and their natural enemies across different continents provide an opportunity to test the enemy release hypothesis (ERH) at local and global scales. Based on the ERH, we assessed the impacts of native seed-feeding insects on introduced and native oaks within and among continents. We combined a common-garden experiment in China and biogeographic literature surveys to measure seed predation by insects and the proportion of acorn embryos surviving after insect infestation among 4 oak species with different geographical origins: Quercus mongolica origin from China, Q. robur and Q. petraea from Europe, and Q. rubra from North America. Mostly supporting the ERH, oaks in introduced continents escaped seed predation compared to those in native continents and compared to other native oaks in introduced continents. Common-garden comparisons showed that total acorn infestation rate of introduced Q. rubra (section Lobatae) was considerably lower than that of native oaks (section Quercus) in China and Europe, likely because of the differences in seed traits associated with different oak sections. Literature surveys showed that seed predation of introduced oaks was lower in the introduced continent than in the native continent. Embryo survival was higher in introduced Q. rubra than native oaks in China and Poland. However, insect seed predation of recently introduced Q. rubra in China was similar to that in Europe, which is not consistent with the ERH. Our results suggest that reduced acorn attack by native insects and higher embryo survival after acorn damage could increase the establishment success or invasion risk of introduced oaks in non-native continents.
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Quercus , Animales , China , Insectos , Conducta Predatoria , SemillasRESUMEN
(1) Background: The principal aim of our work was to identify pheromone components for Plagionotus detritus (L.) (Coleoptera: Cerambycidae), which could be exploited for developing a pheromone-based monitoring system for the complementary purposes of plant protection in areas where it might become a pest, and natural conservation in areas where it is rare or endangered. (2) Methods: Collection and analysis of headspace volatiles were carried out with field-collected beetles. Bioactive volatile compounds identified [(R)-3-hydroxyhexan-2-one and (S)-2-hydroxyoctan-3-one] from extracts of males were purchased [(±)-3-hydroxyhexan-2-one], and synthesized [(S)-2-hydroxyoctan-3-one] and field-tested. Electroantennogram assays showed that antennae of the predatory beetle Clerus mutillarius F. (Coleoptera: Cleridae) also responded to the synthetic compounds. (3) Results: A two-component aggregation-sex pheromone consisting of (R)-3-hydroxyhexan-2-one and (S)-2-hydroxyoctan-3-one was identified for P. detritus. (±)-3-hydroxyhexan-2-one and (S)-2-hydroxyoctan-3-one attracted adults of P. detritus in field bioassays. Adults of the clerid C. mutillarius also were attracted to both compounds. The cerambycid Xylotrechus antilope Schönh was significantly attracted to traps baited with (S)-2-hydroxyoctan-3-one alone or the blend containing this compound. (4) Conclusions: Our data confirmed that 3-hydroxyhexan-2-one and 2-hydroxyoctan-3-one are male-produced pheromone components for P. detritus. These results show that both intraspecific and interspecific communication may play key roles in longhorn beetle life history and ecology, with closely and more distantly related species eavesdropping on each other's signals.
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Sirex noctilio is an invasive Eurasian woodwasp that can kill pine (Pinus spp.) trees and has been introduced to areas of the Southern Hemisphere where plantations of introduced pines are grown. The main method of control of this invasive pest has been introduction and augmentation of a parasitic nematode, Deladenus siricidicola. The strain of D. siricidicola used for biological control of S. noctilio in the Southern Hemisphere originated in Sopron, Hungary. The genotype of D. siricidicola used for biological control sterilizes females of the strain of S. noctilio present in Australia. However, different strains of S. noctilio have been introduced to different geographic areas that have been invaded and different combinations of D. siricidicola and S. noctilio genotypes vary in whether these nematodes sterilize female S. noctilio. Moreover, even in the event of sterilization, partial sterilization can occur, where not all woodwasp eggs are compromised. Sirex noctilio has now invaded North America accidentally, putatively accompanying D. siricidicola, but these host/parasite pairings do not result in female sterilization. More information is needed about the genetic diversity of D. siricidicola both where it is native and introduced. In addition, the host range of these nematodes is necessary to understand to evaluate their potential use in areas where pine communities are native. We collected and evaluated Deladenus parasitizing S. noctilio, S. juvencus, and associated insects in Hungary, Denmark, Spain, and Italy, as well as in the United States. Phylogenetic analyses were unable to fully provide fine resolution, although some community structure was evident. Many D. siricidicola samples from Hungary had identical COI and ITS sequences to the strain of D. siricidicola accidentally introduced to North America putatively when S. noctilio invaded. The same or similar strains of D. siricidicola parasitize two different Sirex species that utilize pines as well as a Sirex parasitoid and a pine-boring beetle, demonstrating some limited variability in host specificity of this species. These results highlight the genetic diversity of Deladenus siricidicola in its native range in Europe.
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Variación Genética , Control de Insectos , Control Biológico de Vectores , Tylenchida/genética , Animales , Europa (Continente) , Femenino , Interacciones Huésped-Parásitos , Himenópteros , Especies Introducidas , América del NorteRESUMEN
We evaluated the presence and impact of Entomophaga maimaiga on both target and non-target phytophagous larvae. All six study plots, with low gypsy moth population density, were situated in Central and Southeastern European oak forests and E. maimaiga had previously been reported from these plots. Totally, 45 of 4,045 (1.13%) collected non-target larvae died due to fungal infections. No non-target insect specimen was infected by E.maimaiga, although the presence of the pathogen could not be fully excluded in three cadavers. Out of 1,780L.dispar larvae collected, 15individuals (0.84%) were infected by E.maimaiga.
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Entomophthorales , Larva/parasitología , Mariposas Nocturnas/parasitología , Control Biológico de Vectores/métodos , Animales , Europa (Continente) , Bosques , Quercus/parasitologíaRESUMEN
Studies of transient population dynamics have largely focused on temporal changes in dynamical behaviour, such as the transition between periods of stability and instability. This study explores a related dynamic pattern, namely transient synchrony during a 49-year period among populations of five sympatric species of forest insects that share host tree resources. The long time series allows a more comprehensive exploration of transient synchrony patterns than most previous studies. Considerable variation existed in the dynamics of individual species, ranging from periodic to aperiodic. We used time-averaged methods to investigate long-term patterns of synchrony and time-localized methods to detect transient synchrony. We investigated transient patterns of synchrony between species and related these to the species' varying density dependence structures; even species with very different density dependence exhibited at least temporary periods of synchrony. Observed periods of interspecific synchrony may arise from interactions with host trees (e.g., induced host defences), interactions with shared natural enemies or shared impacts of environmental stochasticity. The transient nature of synchrony observed here raises questions both about the identity of synchronizing mechanisms and how these mechanisms interact with the endogenous dynamics of each species. We conclude that these patterns are the result of interspecific interactions that act only temporarily to synchronize populations, after which differences in the endogenous population dynamics among the species acts to desynchronize their dynamics.
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Bosques , Mariposas Nocturnas/fisiología , Animales , Hungría , Dinámica Poblacional , Estaciones del AñoRESUMEN
Approximate Bayesian computation (ABC) is a powerful and widely used approach in inference of population history. However, the computational effort required to discriminate among alternative historical scenarios often limits the set that is compared to those considered more likely a priori. While often justifiable, this approach will fail to consider unexpected but well-supported population histories. We used a hierarchical tournament approach, in which subsets of scenarios are compared in a first round of ABC analyses and the winners are compared in a second analysis, to reconstruct the population history of an oak gall wasp, Synergus umbraculus (Hymenoptera, Cynipidae) across the Western Palaearctic. We used 4,233 bp of sequence data across seven loci to explore the relationships between four putative Pleistocene refuge populations in Iberia, Italy, the Balkans and Western Asia. We compared support for 148 alternative scenarios in eight pools, each pool comprising all possible rearrangements of four populations over a given topology of relationships, with or without founding of one population by admixture and with or without an unsampled "ghost" population. We found very little support for the directional "out of the east" scenario previously inferred for other gall wasp community members. Instead, the best-supported models identified Iberia as the first-regional population to diverge from the others in the late Pleistocene, followed by divergence between the Balkans and Western Asia, and founding of the Italian population through late Pleistocene admixture from Iberia and the Balkans. We compare these results with what is known for other members of the oak gall community, and consider the strengths and weaknesses of using a tournament approach to explore phylogeographic model space.
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Teorema de Bayes , Genética de Población , Modelos Genéticos , Avispas/genética , Animales , Asia , Europa (Continente) , Marcadores Genéticos , Variación Genética , Medio Oriente , Tasa de Mutación , Filogenia , Filogeografía , Quercus , Refugio de FaunaRESUMEN
As global biodiversity continues to decline steeply, it is becoming increasingly important to understand diversity patterns at local and regional scales. Changes in land use and climate, nitrogen deposition and invasive species are the most important threats to global biodiversity. Because land use changes tend to benefit a few species but impede many, the expected outcome is generally decreasing population sizes, decreasing species richness at local and regional scales, and increasing similarity of species compositions across sites (biotic homogenization). Homogenization can be also driven by invasive species or effects of soil eutrophication propagating to higher trophic levels. In contrast, in the absence of increasing aridity, climate warming is predicted to generally increase abundances and species richness of poikilotherms at local and regional scales. We tested these predictions with data from one of the few existing monitoring programmes on biodiversity in the world dating to the 1960s, where the abundance of 878 species of macro-moths have been measured daily at seven sites across Hungary. Our analyses revealed a dramatic rate of regional species loss and homogenization of community compositions across sites. Species with restricted distribution range, specialized diet or dry grassland habitat were more likely than others to disappear from the community. In global context, the contrasting effects of climate change and land use changes could explain why the predicted enriching effects from climate warming are not always realized.
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Biodiversidad , Cambio Climático , Mariposas Nocturnas , Animales , Ecosistema , Europa (Continente) , PlantasRESUMEN
Spectral mating preferences were examined in male Agrilus angustulus (Buprestidae: Coleoptera), a member of a taxon known for its high species diversity and striking metallic coloration. The spectral emission profile of a typical A. angustulus female displays low chroma, broadly overlapping that of the green oak leaves they feed and rest upon, while also including longer wavelengths. To pinpoint behaviorally significant spectral regions for A. angustulus males during mate selection, we observed their field approaches to females of five Agrilus planipennis color morphs that have greater chroma than the normal conspecific female targets. Agrilus angustulus males would initially fly equally frequently toward any of the three longest wavelength morphs (green, copper and red) whose spectral emission profiles all overlap that of typical A. angustulus females. However, they usually only completed approaches toward the two longest wavelength morphs, but not the green morphs. Thus, spectral preference influenced mate selection by A. angustulus males, and their discrimination of suitable targets became greater as these targets were approached. This increasing spectral discrimination when approaching targets may have evolved to allow female emissions to remain somewhat cryptic, while also being visible to conspecifics as distinct from the background vegetation and heterospecific competitors.
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Sirex noctilio is a woodwasp of Eurasian origin that was inadvertently introduced to the southern hemisphere in the 1900s and to North America over a decade ago. Its larvae bore in Pinus spp. and can cause significant mortality in pine plantations. S noctilio is associated with a symbiotic white rot fungus, Amylostereum areolatum, which females inject into trees when they oviposit and which is required for survival of developing larvae. We compared the genetic diversity of A. areolatum isolated from S. noctilio and other woodwasps collected from Europe and from northeastern North America to determine the origin of introduction(s) into the United States. Multilocus genotyping of nuclear ribosomal regions and protein coding genes revealed two widespread multilocus genotypes (MLGs) among the European samples, one of which is present in the US. The other two MLGs associated with S. noctilio in the US represented unique haplotypes. These latter two haplotypes were likely from unrepresented source populations, and together with the introduced widespread haplotype reveal multiple A. areolatum MLGs introduced by S. noctilio and indicate possible multiple S. noctilio introductions to North America from Europe. Our results also showed a lack of fidelity between woodwasp hosts and Amylostereum species.
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Basidiomycota/aislamiento & purificación , Himenópteros/microbiología , Pinus/parasitología , Animales , Basidiomycota/clasificación , Basidiomycota/genética , Europa (Continente) , Femenino , Variación Genética , Genotipo , Himenópteros/crecimiento & desarrollo , Especies Introducidas , Masculino , Datos de Secuencia Molecular , Tipificación de Secuencias Multilocus , América del Norte , Filogenia , Enfermedades de las Plantas/parasitología , Estados UnidosRESUMEN
Long-term data sets, covering several decades, could help to reveal the effects of observed climate change on herbivore damage to plants. However, sufficiently long time series in ecology are scarce. The research presented here analyzes a long-term data set collected by the Hungarian Forest Research Institute over the period 1961-2009. The number of hectares with visible defoliation was estimated and documented for several forest insect pest species. This resulted in a unique time series that provides us with the opportunity to compare insect damage trends with trends in weather patterns. Data were analyzed for six lepidopteran species: Thaumetopoea processionea, Tortrix viridana, Rhyacionia buoliana, Malacosoma neustria, Euproctis chrysorrhoea, and Lymantria dispar. All these species exhibit outbreak dynamics in Hungary. Five of these species prefer deciduous tree species as their host plants, whereas R. buoliana is a specialist on Pinus spp. The data were analyzed using general linear models and generalized least squares regression in relation to mean monthly temperature and precipitation. Temperature increased considerably, especially over the last 25 years (+1.6°C), whereas precipitation exhibited no trend over the period. No change in weather variability over time was observed. There was increased damage caused by two species on deciduous trees. The area of damage attributed to R. buoliana decreased over the study period. There was no evidence of increased variability in damage. We conclude that species exhibiting a trend toward outbreak-level damage over a greater geographical area may be positively affected by changes in weather conditions coinciding with important life stages. Strong associations between the geographical extent of severe damage and monthly temperature and precipitation are difficult to confirm, studying the life-history traits of species could help to increase understanding of responses to climate change.
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We show that the population ecology of the 9- to 10-year cyclic, broadleaf-defoliating winter moth (Operophtera brumata) and other early-season geometrids cannot be fully understood on a local scale unless population behaviour is known on a European scale. Qualitative and quantitative data on O. brumata outbreaks were obtained from published sources and previously unpublished material provided by authors of this article. Data cover six decades from the 1950s to the first decade of twenty-first century and most European countries, giving new information fundamental for the understanding of the population ecology of O. brumata. Analyses on epicentral, regional and continental scales show that in each decade, a wave of O. brumata outbreaks travelled across Europe. On average, the waves moved unidirectionally ESE-WNW, that is, toward the Scandes and the Atlantic. When one wave reached the Atlantic coast after 9-10 years, the next one started in East Europe to travel the same c. 3000 km distance. The average wave speed and wavelength was 330 km year(-1) and 3135 km, respectively, the high speed being incongruous with sedentary geometrid populations. A mapping of the wave of the 1990s revealed that this wave travelled in a straight E-W direction. It therefore passed the Scandes diagonally first in the north on its way westward. Within the frame of the Scandes, this caused the illusion that the wave moved N-S. In analogy, outbreaks described previously as moving S-N or occurring contemporaneously along the Scandes were probably the result of continental-scale waves meeting the Scandes obliquely from the south or in parallel. In the steppe zone of eastern-most and south-east Europe, outbreaks of the winter moth did not participate in the waves. Here, broadleaved stands are small and widely separated. This makes the zone hostile to short-distance dispersal between O. brumata subpopulations and prevents synchronization within meta-populations. We hypothesize that hostile boundary models, involving reciprocal host-herbivore-enemy reactions at the transition between the steppe and the broadleaved forest zones, offer the best explanation to the origin of outbreak waves. These results have theoretical and practical implications and indicate that multidisciplinary, continentally coordinated studies are essential for an understanding of the spatio-temporal behaviour of cyclic animal populations.
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Mariposas Nocturnas/fisiología , Animales , Monitoreo del Ambiente , Europa (Continente) , Dinámica Poblacional , Factores de TiempoRESUMEN
How geographically widespread biological communities assemble remains a major question in ecology. Do parallel population histories allow sustained interactions (such as host-parasite or plant-pollinator) among species, or do discordant histories necessarily interrupt them? Though few empirical data exist, these issues are central to our understanding of multispecies evolutionary dynamics. Here we use hierarchical approximate Bayesian analysis of DNA sequence data for 12 herbivores and 19 parasitoids to reconstruct the assembly of an insect community spanning the Western Palearctic and assess the support for alternative host tracking and ecological sorting hypotheses. We show that assembly occurred primarily by delayed host tracking from a shared eastern origin. Herbivores escaped their enemies for millennia before parasitoid pursuit restored initial associations, with generalist parasitoids no better able to track their hosts than specialists. In contrast, ecological sorting played only a minor role. Substantial turnover in host-parasitoid associations means that coevolution must have been diffuse, probably contributing to the parasitoid generalism seen in this and similar systems. Reintegration of parasitoids after host escape shows these communities to have been unsaturated throughout their history, arguing against major roles for parasitoid niche evolution or competition during community assembly.
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Ecosistema , Interacciones Huésped-Parásitos/fisiología , Quercus/parasitología , Avispas/parasitología , Animales , Regiones Árticas , Teorema de Bayes , Evolución Biológica , ADN Mitocondrial/genética , Modelos Biológicos , Datos de Secuencia Molecular , Enfermedades de las Plantas/parasitología , Factores de TiempoRESUMEN
BACKGROUND: Biological invasions provide a window on the process of community assembly. In particular, tracking natural enemy recruitment to invading hosts can reveal the relative roles of co-evolution (including local adaptation) and ecological sorting. We use molecular data to examine colonisation of northern Europe by the parasitoid Megastigmus stigmatizans following invasions of its herbivorous oak gallwasp hosts from the Balkans. Local host adaptation predicts that invading gallwasp populations will have been tracked primarily by sympatric Balkan populations of M. stigmatizans (Host Pursuit Hypothesis). Alternatively, ecological sorting allows parasitoid recruitment from geographically distinct populations with no recent experience of the invading hosts (Host Shift Hypothesis). Finally, we test for long-term persistence of parasitoids introduced via human trade of their hosts' galls (Introduction Hypothesis). RESULTS: Polymorphism diagnostic of different southern refugial regions was present in both mitochondrial and nuclear microsatellite markers, allowing us to identify the origins of northern European invaded range M. stigmatizans populations. As with their hosts, some invaded range populations showed genetic variation diagnostic of Balkan sources, supporting the Host Pursuit Hypothesis. In contrast, other invading populations had an Iberian origin, unlike their hosts in northern Europe, supporting the Host Shift Hypothesis. Finally, both British and Italian M. stigmatizans populations show signatures compatible with the Introduction Hypothesis from eastern Mediterranean sources. CONCLUSIONS: These data reveal the continental scale of multi-trophic impacts of anthropogenic disturbance and highlight the fact that herbivores and their natural enemies may face very different constraints on range expansion. The ability of natural enemies to exploit ecologically-similar hosts with which they have had no historical association supports a major role for ecological sorting processes in the recent assembly of these communities. The multitude of origins of invading natural enemy populations in this study emphasises the diversity of mechanisms requiring consideration when predicting consequences of other biological invasions or biological control introductions.
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Interacciones Huésped-Parásitos/fisiología , Avispas/genética , Avispas/patogenicidad , Animales , ADN Mitocondrial/genética , Europa (Continente) , Haplotipos , Repeticiones de Microsatélite/genética , Filogenia , Quercus/parasitología , Avispas/clasificaciónRESUMEN
Little is known about the evolutionary history of most complex multi-trophic insect communities. Widespread species from different trophic levels might evolve in parallel, showing similar spatial patterns and either congruent temporal patterns (Contemporary Host-tracking) or later divergence in higher trophic levels (Delayed Host-tracking). Alternatively, host shifts by natural enemies among communities centred on different host resources could disrupt any common community phylogeographic pattern. We examined these alternative models using two Megastigmus parasitoid morphospecies associated with oak cynipid galls sampled throughout their Western Palaearctic distributions. Based on existing host cynipid data, a parallel evolution model predicts that eastern regions of the Western Palaearctic should contain ancestral populations with range expansions across Europe about 1.6 million years ago and deeper species-level divergence at both 8-9 and 4-5 million years ago. Sequence data from mitochondrial cytochrome b and multiple nuclear genes showed similar phylogenetic patterns and revealed cryptic genetic species within both morphospecies, indicating greater diversity in these communities than previously thought. Phylogeographic divergence was apparent in most cryptic species between relatively stable, diverse, putatively ancestral populations in Asia Minor and the Middle East, and genetically depauperate, rapidly expanding populations in Europe, paralleling patterns in host gallwasp species. Mitochondrial and nuclear data also suggested that Europe may have been colonized multiple times from eastern source populations since the late Miocene. Temporal patterns of lineage divergence were congruent within and across trophic levels, supporting the Contemporary Host-tracking Hypothesis for community evolution.
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Evolución Molecular , Especiación Genética , Filogenia , Quercus/parasitología , Avispas/genética , Animales , Núcleo Celular/genética , ADN Mitocondrial/genética , Europa (Continente) , Geografía , Modelos Genéticos , Análisis de Secuencia de ADN , Avispas/clasificaciónRESUMEN
We examine phylogenetic relationships within the Synergus complex of herbivorous inquiline gallwasps (Hymenoptera; Cynipidae; Synergini) associated with cynipid host galls on oak, a biologically diverse group whose genus-level morphological taxonomy has long been considered stable but whose species level taxonomy is problematic. We incorporate data for over 70% of recognised Western Palaearctic species in five morphology-based genera (Ceroptres, Saphonecrus, Synergus, Synophrus, Ufo), comprising sequence for two mitochondrial loci (coxI, cytb) and one nuclear locus (28S D2). In particular, we assess the evidence for monophyly of two long-established, morphology-defined sections within the genus Synergus that differ in a range of biological traits. To aid analyses of ecological interactions within oak cynipid communities, we also consider the utility of cytochrome oxidase I (coxI) DNA barcodes in the oak inquilines. In this assessment, we do not assume that species are delineated at a single threshold value of sequence divergence for a single gene, but examine concordance in the composition of molecular operational Taxonomic units (MOTUs) across a range of sequence divergences in each gene and across genes. We also assess the impact of sampling effort on MOTU stability. Phylogenetic reconstructions for all three loci support monophyly for Synergus and Synophrus, but reject monophyly for Saphonecrus and for the two sections within Synergus. The suites of traits associated with the two sections of the genus Synergus are thus homoplasious. All three loci also reject monophyly for three Synergus species (S. hayneanus, S. pallipes, S. umbraculus). Sequences for each locus identify robust MOTUs that are largely concordant across loci for a range of cut-off values. Though many MOTU's correspond to recognised Linnean species, there is significant, multigene disagreement between groupings supported by morphology and sequence data, with both allocation of different morphospecies to the same MOTU and allocation of the same morphospecies to multiple MOTUs, regardless of cut-off value. Our results imply that while DNA barcoding has considerable utility within this group, morphology-based identification needs major revision at both genus and species levels. Further, lifehistory traits currently attributed to single morphospecies probably confound attributes of multiple lineages. Revealing patterns of character state evolution in Synergus requires collection of new host association and life history data explicitly linked to DNA barcode data for the specimens concerned.