Host Plant Shifts Research Articles

Adapting to change: Exploring the consequences of climate-induced host plant shifts in two specialist Lepidoptera species.

Asynchronous migration of insect herbivores and their host plants towards higher elevations following climate warming is expected to generate novel plant-insect interactions. While the disassociation of specialised interactions can challenge species' persistence, consequences for specialised low-elevation insect herbivores encountering novel high-elevation plants under climate change remain largely unknown. To explore the ability of two low-elevation Lepidoptera species, Melitaea celadussa and Zygaena filipendulae, to undergo shifts from low- to high-elevation host plants, we combined a translocation experiment performed at two elevations in the Swiss Alps with experiments conducted under controlled conditions. Specifically, we exposed M. celadussa and Z. filipendulae to current low- and congeneric high-elevation host plants, to test how shifts in host plant use impact oviposition probability, number of eggs clutches laid, caterpillar feeding preference and growth, pupation rate and wing size. While our study shows that both M. celadussa and Z. filipendulae can oviposit and feed on novel high-elevation host plants, we reveal strong preferences towards ovipositing and feeding on current low-elevation host plants. In addition, shifts from current low- to novel high-elevation host plants reduced pupation rates as well as wing size for M. celadussa, while caterpillar growth was unaffected by host plant identity for both species. Our study suggests that populations of M. celadussa and Z. filipendulae have the ability to undergo host plant shifts under climate change. However, these shifts may impact the ability of populations to respond to rapid climate change by altering developmental processes and morphology. Our study highlights the importance of considering altered biotic interactions when predicting consequences for natural populations facing novel abiotic and biotic environments.

Evolution of chemosensory genes in Colorado potato beetle, Leptinotarsa decemlineata.

Associating with plant hosts is thought to have elevated the diversification of insect herbivores, which comprise the majority of global species diversity. In particular, there is considerable interest in understanding the genetic changes that allow host-plant shifts to occur in pest insects and in determining what aspects of functional genomic diversity impact host-plant breadth. Insect chemoreceptors play a central role in mediating insect-plant interactions, as they directly influence plant detection and sensory stimuli during feeding. Although chemosensory genes evolve rapidly, it is unclear how they evolve in response to host shifts and host specialization. We investigate whether selection at chemosensory genes is linked to host-plant expansion from the buffalo burr, Solanum rostratum, to potato, Solanum tuberosum, in the super-pest Colorado potato beetle (CPB), Leptinotarsa decemlineata (Coleoptera: Chrysomelidae). First, to refine our knowledge of CPB chemosensory genes, we developed novel gene expression data for the antennae and maxillary-labial palps. We then examine patterns of selection at these loci within CPB, as well as compare whether rates of selection vary with respect to 9 closely related, non-pest Leptinotarsa species that vary in diet breadth. We find that rates of positive selection on olfactory receptors are higher in host-plant generalists, and this signal is particularly strong in CPB. These results provide strong candidates for further research on the genetic basis of variation in insect chemosensory performance and novel targets for pest control of a notorious super-pest.

Evolutionary host shifts across plant orders despite high host specificity in tree stem surface-living Stomaphis aphids inferred from molecular phylogeny

We reconstructed Stomaphis phylogeny and analyzed evolutionary host-plant shifts. The molecular phylogeny revealed 23 well-supported lineages, each specialized to use specific host plant family, whilst host plants of Stomaphis aphids ranged across 6 orders, 11 families, 21 genera, and 28 species. This combination of high host specificity with evolutionarily distant host shifts is exceptional in herbivores. To explain this pattern, we propose one hypothesis among several possibilities: Stomaphis aphids are generalists with respect to the defensive chemicals produced by the plant, but specialists with respect to the stem surface structure of the host tree. This hypothesis predicts that tree taxa having stem surface structures preferred by Stomaphis would be used again and again by genetically distinct Stomaphis lineages. Consistent with this prediction, we found that different (occasionally phylogenetically distant) Stomaphis lineages shared the same host plant genera such as Alnus and Betula. This result suggests that, in the course of their evolutionary history, Japanese Stomaphis aphids have repeatedly colonized a limited number of host plant genera.

Differential olfactory responses associated with host plant shift by the fruit-piercing moth, Eudocima phalonia, in the Pacific islands

ABSTRACT The fruit-piercing moth (FPM) Eudocima phalonia is widely present in the tropics, causing major damages to the citrus industry. In its ancestral distribution range, e.g. Australia, FPM develops on vines from the Menispermaceae family. But in more recently colonized islands, e.g. New Caledonia, FPM populations have shifted to new host plants of the Fabaceae family, Erythrina spp.. To understand this host shift, we studied the chemical ecology of FPM as a mechanism driving host plant acceptance. We collected volatile headspace samples of Erythrina species and Menispermaceae and compared their chemical spectra. We assessed the electrophysiological responses of FPM populations from the two countries to the plant chemical extracts and identified bioactive compounds. The volatile profiles from each species were quite different between and within each plant family. However, five compounds common across the two families triggered electrophysiological responses in both FPM populations. Those common bioactive compounds could have facilitated the host shift to completely different plant family. Furthermore, the diverging history between the two FPM populations may explain differences in electrophysiological sensitivity to other specific compounds. These results are discussed in the context of the production of a generic lure that could be attractive to all FPM populations.

Phylogenomics resolves major relationships of Catocala underwing moths

AbstractUnderwing moths in the genus Catocala Schrank are among the most charismatic of Lepidoptera. Catocala is also one of the most diverse genera worldwide in the speciose family Erebidae, but a phylogenetic framework for the genus is lacking. Here we reconstruct the first comprehensive molecular phylogeny for the genus based on 685 anchored hybrid enrichment loci sampled from 161 Catocala species (99 Nearctic, 62 Palearctic), four species of Ulotrichopus Wallengren and 33 outgroups. Phylogenetic analysis unambiguously recovers Catocala and Catocala + Ulotrichopus as monophyletic with strong support and resolves many backbone relationships within Catocala. Our results confirm the classification of previously proposed taxonomic subgroups of Catocala, including seven based on recent molecular/morphological evidence, and ten based on early twentieth‐century morphological research. Mapping of larval host plant use onto the tree shows Fabaceae to be the likely ancestral host plant family for Catocala and Catocala + Ulotrichopus. There appear to have been at least 18 independent larval host plant shifts to nine plant families, the most common shift being from Fabaceae to Fagaceae. Larval host plant use has likely played an important role in the evolutionary history of Catocala, with several rapid diversification events propelled by shifts to novel larval host plants, particularly in the North American Catocala fauna.

Molecular phylogeny, divergence time, biogeography and trends in host plant usage in the agriculturally important tortricid tribe Grapholitini (Lepidoptera: Tortricidae: Olethreutinae).

The leaf-roller moth tribe Grapholitini comprises about 1200 described species and contains numerous notorious pests of fruits and seeds. The phylogeny of the tribe has been little studied using contemporary methods, and the monophyly of several genera remains questionable. In order to provide a more robust phylogenetic framework for the group, we conducted a multiple-gene phylogenetic analysis of 104 species representing 27 genera of Grapholitini and 29 outgroup species. Divergence time, ancestral area, and host plant usage were also inferred to explore evolutionary trends in the tribe. Our analyses indicate that Larisa and Corticivora, traditionally assigned to Grapholitini, are best excluded from the tribe. After removal of these two genera, the tribe is found to be monophyletic, represented by two major lineages-a Dichrorampha clade and a Cydia clade, the latter of which can be divided into seven generic groups. The genus Grapholita was found to be polyphyletic, comprising three different clades, and we propose three genera to accommodate these groups: Grapholita (sensu stricto), Aspila (formerly a subgenus of Grapholita) and Ephippiphora (formerly considered a synonym of Grapholita). We summarize each generic group, including related genera not included in our analysis, providing morphological, pheromone and food plant characters that support particular branches within the molecular hypotheses. Biogeographical analyses indicate that Grapholitini probably originated in the Nearctic, Afrotropical and Neotropical regions in the Lutetian of the middle Eocene (ca. 44.3 Ma). Our results also indicate that most groups in Grapholitini originated from Fabaceae-feeding monophagous or oligophagous ancestors, and that host plant shifts probably promoted species diversification within the tribe.

Alpine butterflies want to fly high: Species and communities shift upwards faster than their host plants.

Despite sometimes strong codependencies of insect herbivores and plants, the responses of individual taxa to accelerating climate change are typically studied in isolation. For this reason, biotic interactions that potentially limit species in tracking their preferred climatic niches are ignored. Here, we chose butterflies as a prominent representative of herbivorous insects to investigate the impacts of temperature changes and their larval host plant distributions along a 1.4-km elevational gradient in the German Alps. Following a sampling protocol of 2009, we revisited 33 grassland plots in 2019 over an entire growing season. We quantified changes in butterfly abundance and richness by repeated transect walks on each plot and disentangled the direct and indirect effects of locally assessed temperature, site management, and larval and adult food resource availability on these patterns. Additionally, we determined elevational range shifts of butterflies and host plants at both the community and species level. Comparing the two sampled years (2009 and 2019), we found a severe decline in butterfly abundance and a clear upward shift of butterflies along the elevational gradient. We detected shifts in the peak of species richness, community composition, and at the species level, whereby mountainous species shifted particularly strongly. In contrast, host plants showed barely any change, neither in connection with species richness nor individual species shifts. Further, temperature and host plant richness were the main drivers of butterfly richness, with change in temperature best explaining the change in richness over time. We concluded that host plants were not yet hindering butterfly species and communities from shifting upwards. However, the mismatch between butterfly and host plant shifts might become a problem for this very close plant-herbivore relationship, especially toward higher elevations, if butterflies fail to adapt to new host plants. Further, our results support the value of conserving traditional extensive pasture use as a promoter of host plant and, hence, butterfly richness.

Phylogenomics provides insights into the evolution of cactophily and host plant shifts in Drosophila

Cactophilic species of the Drosophila buzzatii cluster (repleta group) comprise an excellent model group to investigate genomic changes underlying adaptation to extreme climate conditions and host plants. In particular, these species form a tractable system to study the transition from chemically simpler breeding sites (like prickly pears of the genus Opuntia) to chemically more complex hosts (columnar cacti). Here, we report four highly contiguous genome assemblies of three species of the buzzatii cluster. Based on this genomic data and inferred phylogenetic relationships, we identified candidate taxonomically restricted genes (TRGs) likely involved in the evolution of cactophily and cactus host specialization. Functional enrichment analyses of TRGs within the buzzatii cluster identified genes involved in detoxification, water preservation, immune system response, anatomical structure development, and morphogenesis. In contrast, processes that regulate responses to stress, as well as the metabolism of nitrogen compounds, transport, and secretion were found in the set of species that are columnar cacti dwellers. These findings are in line with the hypothesis that those genomic changes brought about key mechanisms underlying the adaptation of the buzzatii cluster species to arid regions in South America.

Transcriptional responses are oriented towards different components of the rearing environment in two Drosophila sibling species

BackgroundThe chance to compare patterns of differential gene expression in related ecologically distinct species can be particularly fruitful to investigate the genetics of adaptation and phenotypic plasticity. In this regard, a powerful technique such as RNA-Seq applied to ecologically amenable taxa allows to address issues that are not possible in classic model species. Here, we study gene expression profiles and larval performance of the cactophilic siblings Drosophila buzzatii and D. koepferae reared in media that approximate natural conditions and evaluate both chemical and nutritional components of the diet. These closely related species are complementary in terms of host-plant use since the primary host of one is the secondary of the other. D. koepferae is mainly a columnar cactus dweller while D. buzzatii prefers Opuntia hosts.ResultsOur comparative study shows that D. buzzatii and D. koepferae have different transcriptional strategies to face the challenges posed by their natural resources. The former has greater transcriptional plasticity, and its response is mainly modulated by alkaloids of its secondary host, while the latter has a more canalized genetic response, and its transcriptional plasticity is associated with the cactus species.ConclusionsOur study unveils a complex pleiotropic genetic landscape in both species, with functional links that relate detox responses and redox mechanisms with developmental and neurobiological processes. These results contribute to deepen our understanding of the role of host plant shifts and natural stress driving ecological specialization.

Non-parallel morphological divergence following colonization of a new host plant

Adaptation to new ecological niches is known to spur population diversification and may lead to speciation if gene flow is ceased. While adaptation to the same ecological niche is expected to be parallel, it is more difficult to predict whether selection against maladaptive hybridization in secondary sympatry results in parallel divergence also in traits that are not directly related to the ecological niches. Such parallelisms in response to selection for reproductive isolation can be identified through estimating parallelism in reproductive character displacement across different zones of secondary contact. Here, we use a host shift in the phytophagous peacock fly Tephritis conura, with both host races represented in two geographically separate areas East and West of the Baltic Sea to investigate convergence in morphological adaptations. We asked (i) if there are consistent morphological adaptations to a host plant shift and (ii) if the response to secondary sympatry with the alternate host race is parallel across contact zones. We found surprisingly low and variable, albeit significant, divergence between host races. Only one trait, the length of the female ovipositor, which serves an important function in the interaction with the hosts, was consistently different between host races. Instead, co-existence with the other host race significantly affected the degree of morphological divergence, but the divergence was largely driven by different traits in different contact zones. Thus, local stochastic fixation or reinforcement could generate trait divergence, and additional evidence is needed to conclude whether divergence is locally adaptive.

Use of an exotic host plant shifts immunity, chemical defense, and viral burden in wild populations of a specialist insect herbivore.

Defense against natural enemies constitutes an important driver of herbivore host range evolution in the wild. Populations of the Baltimore checkerspot butterfly, Euphydryas phaeton (Nymphalidae), have recently incorporated an exotic plant, Plantago lanceolata (Plantaginaceae), into their dietary range. To understand the tritrophic consequences of utilizing this exotic host plant, we examined immune performance, chemical defense, and interactions with a natural entomopathogen (Junonia coenia densovirus, Parvoviridae) across wild populations of this specialist herbivore. We measured three immune parameters, sequestration of defensive iridoid glycosides (IGs), and viral infection load in field‐collected caterpillars using either P. lanceolata or a native plant, Chelone glabra (Plantaginaceae). We found that larvae using the exotic plant exhibited reduced immunocompetence, compositional differences in IG sequestration, and higher in situ viral burdens compared to those using the native plant. On both host plants, high IG sequestration was associated with reduced hemocyte concentration in the larval hemolymph, providing the first evidence of incompatibility between sequestered chemical defenses and the immune response (i.e., the “vulnerable host” hypothesis) from a field‐based study. However, despite this negative relationship between IG sequestration and cellular immunity, caterpillars with greater sequestration harbored lower viral loads. While survival of virus‐infected individuals decreased with increasing viral burden, it ultimately did not differ between the exotic and native plants. These results provide evidence that: (1) phytochemical sequestration may contribute to defense against pathogens even when immunity is compromised and (2) herbivore persistence on exotic plant species may be facilitated by sequestration and its role in defense against natural enemies.

Host specialization is ancestral in Torymus (Hymenoptera, Chalcidoidea) cynipid gall parasitoids

AbstractThe host range of parasitoids varies greatly among species: Some only parasitize one to a few hosts (specialists), while others parasitize multiple species or a variety of host types (generalists). The direction of most host range shifts in parasitoid groups, that is from generalist to specialist or, alternatively, from specialist to generalist, is unknown. To explore the origin of host range shifts, we studied a clade within the genus Torymus (Hymenoptera, Chalcidoidea) that includes both generalist and specialist parasitoids of Cynipidae (Hymenoptera). We analysed the phylogenetic relationships of the species of Torymus on the basis of two gene fragments (cox1 and ITS2) of 246 specimens and performed an ancestral state reconstruction of the specialist/generalist trait. Our results revealed the following: (a) The ancestral state of this group of Torymus is specialist, with the generalist state evolving through a loss of specialization. (b) The species Torymus cyaneus and Torymus flavipes both have a strong genetic structure, suggesting the existence of different biological identities. (c) There has been a host plant shift in the lineage(s) leading to Torymus rubi and Torymus bedeguaris from galls on Quercus to those on Rosaceae. (d) The alien species Torymus sinensis and the native European species Torymus notatus are phylogenetically closely related. (e) Speciation within Torymus was likely associated with the diversification of their cynipid hosts, which itself was driven by the dramatic changes in climate and vegetation that occurred during the Miocene.

Host plant shift differentially alters olfactory sensitivity in female and male Drosophila mojavensis

Animals may vary in their utilization of plants depending on plant availability, and also on the sex of the animal. Evolutionary adaptations may arise, particularly in specialist animals to the chemistry of the host plants, and these adaptations may differ between the sexes due to differences in their interactions with the plants. Drosophila mojavensis uses different host cacti across its range, and volatile chemicals emitted by the host are the primary cue for host plant identification. In this study, we measured responses of individual olfactory sensory neurons to a large suite of odorants across males and females of the two southern D. mojavensis populations. We show that a switch in host plant is accompanied by changes in the olfactory system, but the effect of this switch is minor compared to that of sex. That is, we observe differences in olfactory receptor neuron specificity and sensitivity to odorants between sexes, and to a lesser extent between populations. The majority of sensory differences are restricted to only three of the 17 sensory neurons measured. Further, we found numerous differences between sexes that only occur within one population, i.e., sex-by-population interactions.

Punctuational ecological changes rather than global factors drive species diversification and the evolution of wing phenotypes inMorphobutterflies

Assessing the relative importance of geographical and ecological drivers of evolution is paramount to understand the diversification of species and traits at the macroevolutionary scale. Here, we use an integrative approach, combining phylogenetics, biogeography, ecology and quantified phenotypes to investigate the drivers of both species and phenotypic diversification of the iconic Neotropical butterfly genus Morpho. We generated a time-calibrated phylogeny for all known species and inferred historical biogeography. We fitted models of time-dependent (accounting for rate heterogeneity across the phylogeny) and paleoenvironment-dependent diversification (accounting for global effect on the phylogeny). We used geometric morphometrics to assess variation of wing size and shape across the tree and investigated their dynamics of evolution. We found that the diversification of Morpho is best explained when considering variable diversification rates across the tree, possibly associated with lineages occupying different microhabitat conditions. First, a shift from understory to canopy was characterized by an increased speciation rate partially coupled with an increasing rate of wing shape evolution. Second, the occupation of dense bamboo thickets accompanying a major host-plant shift from dicotyledons towards monocotyledons was associated with a simultaneous diversification rate shift and an evolutionary 'jump' of wing size. Our study points to a diversification pattern driven by punctuational ecological changes instead of a global driver or biogeographic history.

Recursive adaptation in action: allochronic isolation and divergence of host‐associated populations of the apple maggot fly, Rhagoletis pomonella, following its recent introduction to the western USA

AbstractAn outstanding issue in the study of insect host races concerns the idea of ‘recursive adaptive divergence’, whereby adaptation can occur repeatedly across space and/or time, and the most recent adaptive episode is defined by one or more previously similar cases. The host plant shift of the apple maggot fly, Rhagoletis pomonella (Walsh) (Diptera: Tephritidae, Carpomyini), from ancestral downy hawthorn [Crataegus mollis (Torr. & A. Gray) Scheele] to introduced, domesticated apple (Malus domestica Borkh.) in the eastern USA has long served as a model system for investigating ecologically driven host race formation in phytophagous insect specialists. Here, we report results from an annual geography survey of eclosion time demonstrating a similar ecological pattern among nascent host‐associated populations of the fly recently introduced ca. 40 years ago from its native range in the east into the Pacific Northwest (PNW) region of the USA. Specifically, using data collected from 25 locations across 5 years, we show that apple‐infesting fly populations in the PNW have rapidly and repeatedly shifted (and maintained differences in) their adult eclosion life‐history timing to infest two novel hawthorn hosts with different fruiting phenologies – a native species (Crataegus douglasii Lindl.) and an introduced species (Crataegus monogyna Jacq.) – generating partial allochronic reproductive isolation in the process. The shifts in the PNW parallel the classic case of host race formation in the eastern USA, but have occurred bi‐directionally to two hawthorn species with phenologies slightly earlier (black hawthorn) and significantly later (ornamental hawthorn) than apple. Our results imply that R. pomonella can both possess and retain extensive‐standing variation (i.e., ‘adaptive memory’) in diapause traits, even following introductions, to rapidly and temporally track novel phenological host opportunities when they arise. Thus, ‘specialized’ host races may not constitute evolutionary dead ends. Rather, adaptive phenotypic and genetic memory may carry over from one host shift to the next, recursively facilitating host race formation in phytophagous insects.

Host range determination in a novel outbreak pest of sugarcane, Cacosceles newmannii (Coleoptera: Cerambycidae, Prioninae), inferred from stable isotopes

AbstractAn outbreak of Cacosceles newmannii (Coleoptera: Cerambycidae) was detected for the first time on sugarcane (Saccharum spp.) in 2015 in KwaZulu‐Natal, South Africa. Although primary host plants of this native species remain unknown, these are central to testing hypotheses concerning the outbreak. We hypothesized that this species has undergone a host plant shift (i.e. a feeding association with a novel host plant). We compared δ13C and δ15N ratios of adult beetles retrieved from South African museum collections, collected between 1891 and 2016 (n = 23; ‘pre‐outbreak’), with samples from infested fields in 2017 (n = 9, ‘post‐outbreak’) and in 2019 (n = 23, ‘post‐outbreak’), as well as diverse, plausible host plants (n = 42 samples across 10 species) from infested fields and surrounding patches of indigenous and commercial forest vegetation. We used Bayesian isotope mixing models to infer the relative contribution of the different plants to the diet of C. newmannii. Pre‐outbreak, C3 plants contributed strongly to the larval diet, whereas post‐outbreak, C4 plants were the largest component of their diet. There was some indication of C4 plants contributing to their diet pre‐outbreak. Our results suggest that the outbreak of this polyphagous beetle was not a dramatic host shift but rather a rapid increase in the proportion of C4 plants already in their diet. We concluded that plants from the families Fabaceae and Poaceae are the most likely host plants of this species. Nevertheless, the drivers of this rapid outbreak on sugarcane remain poorly determined and should be the focus of future research.

A reversal in sensory processing accompanies ongoing ecological divergence and speciation in Rhagoletis pomonella.

Changes in behaviour often drive rapid adaptive evolution and speciation. However, the mechanistic basis for behavioural shifts is largely unknown. The tephritid fruit fly Rhagoletis pomonella is an example of ecological specialization and speciation in action via a recent host plant shift from hawthorn to apple. These flies primarily use specific odours to locate fruit, and because they mate only on or near host fruit, changes in odour preference for apples versus hawthorns translate directly to prezygotic reproductive isolation, initiating speciation. Using a variety of techniques, we found a reversal between apple and hawthorn flies in the sensory processing of key odours associated with host fruit preference at the first olfactory synapse, linking changes in the antennal lobe of the brain with ongoing ecological divergence. Indeed, changes to specific neural pathways of any sensory modality may be a broad mechanism for changes in animal behaviour, catalysing the genesis of new biodiversity.

Genome-wide macroevolutionary signatures of key innovations in butterflies colonizing new host plants

The mega-diversity of herbivorous insects is attributed to their co-evolutionary associations with plants. Despite abundant studies on insect-plant interactions, we do not know whether host-plant shifts have impacted both genomic adaptation and species diversification over geological times. We show that the antagonistic insect-plant interaction between swallowtail butterflies and the highly toxic birthworts began 55 million years ago in Beringia, followed by several major ancient host-plant shifts. This evolutionary framework provides a valuable opportunity for repeated tests of genomic signatures of macroevolutionary changes and estimation of diversification rates across their phylogeny. We find that host-plant shifts in butterflies are associated with both genome-wide adaptive molecular evolution (more genes under positive selection) and repeated bursts of speciation rates, contributing to an increase in global diversification through time. Our study links ecological changes, genome-wide adaptations and macroevolutionary consequences, lending support to the importance of ecological interactions as evolutionary drivers over long time periods.

Short term transcriptional responses of P450s to phytochemicals in insects and mites.

Cytochrome P450 monooxygenases (P450s) play a key role in the detoxification of phytochemicals in arthropod herbivores. We present here an overview of recent progress in understanding the breadth and specificity of gene expression plasticity of P450s in response to phytochemicals. We discuss experimental setups and new findings in mechanisms of P450 regulation. Whole genome transcriptomic analysis of arthropod herbivores, either after direct administration of phytochemicals or after host plant shifts, allowed to integrate various levels of chemical complexity and lead to the unbiased identification of responsive P450 genes. However, despite progress in identification of inducible P450s, the link between induction and metabolism is still largely unexplored, and to what extent the overall response is biologically functional should be further investigated. In the near future, such studies will be more straightforward as forward and reverse genetic tools become more readily available.

The Role of Sympatric and Allopatric Speciation in the Origin of Biodiversity of Herbivorous Insects, with Palaearctic Species of the Genus Macropsis Lewis, 1836 Taken as an Example (Homoptera, Auchenorrhyncha, Cicadellidae, Eurymelinae, Macropsini)

Using 60 Palaearctic species of the leafhopper genus Macropsis as an example, the contribution of sympatric and allopatric speciation to the origin of herbivorous insect biodiversity was studied. According to the previously reconstructed evolutionary scenarios, in the temperate zone of the Palaearctic Region the genus Macropsis probably dispersed from east to west and formed 10 natural groups, with species within each group being similar both in morphological traits and in host specialization. The hypothetical speciation modes for each group can be largely inferred from geography. If related species replace each other in different zoogeographic regions, they are likely to have arisen as the result of allopatric speciation. If the species show overlapping distributions and live on the same host plant, they must have originated in allopatry and become secondarily sympatric due to subsequent dispersal. Finally, if related species have strongly overlapping ranges and populate similar habitats but differ in host specializations, this may indicate their origin as the result of sympatric speciation due to host plant shifts. For 31 species (51.7%), allopatric speciation seems to be the most likely or even the only possible scenario, whereas for 20 species (33.3%), the hypothesis of sympatric speciation is either convincingly substantiated or at least quite feasible. The origins of the remaining 9 species (15%) remain unknown. Therefore, sympatric speciation is not a rare phenomenon in Macropsis, but it certainly does not predominate over the allopatric mode. Thus, even within the same genus of herbivorous insects, speciation can occur in different ways: in the allopatric mode, when a host plant shift is possible but not essential, and in the sympatric mode, when divergence is triggered by the host shift. Cases of intraspecific differentiation, which can be considered as initial stages of speciation, are also discussed. These are allopatric color forms associated with the same host plant, allopatric color forms associated with different plants and, finally, forms living on different sympatric or allopatric hosts but showing no difference in coloration. It is shown that divergence is not always caused by transition to a new host plant even in specialized herbivores.