Higher-order species interactions cause time-dependent niche and fitness differences: Experimental evidence in plant-feeding arthropods.

Species interact in different ways, including competition, facilitation and predation. These interactions can be non-linear or higher order and may depend on time or species densities. Although these higher-order interactions are virtually ubiquitous, they remain poorly understood, as they are challenging both theoretically and empirically. We propose to adapt niche and fitness differences from modern coexistence theory and apply them to species interactions over time. As such, they may not merely inform about coexistence, but provide a deeper understanding of how species interactions change. Here, we investigated how the exploitation of a biotic resource (plant) by phytophagous arthropods affects their interactions. We performed monoculture and competition experiments to fit a generalized additive mixed model to the empirical data, which allowed us to calculate niche and fitness differences. We found that species switch between different types of interactions over time, including intra- and interspecific facilitation, and strong and weak competition.

Mites inhabiting nests of wood warbler, Phylloscopus sibilatrix (Aves: Passeriformes), in the Wielkopolska National Park in western Poland.

The wood warbler, Phylloscopus sibilatrix (Aves: Passeriformes), is a well-known model organism for studying bird migration, breeding habitat selection and nest predation. The nest acarofauna of this bird species has not been extensively studied so far. To provide a comprehensive report on mite species inhabiting wood warbler nests and to assess infestation parameters (prevalence, intensity, and abundance) for mite species and orders, we collected 45 nests of this bird species in the Wielkopolska National Park in western Poland. Analyses revealed a huge diversity (198 species) of mites inhabiting wood warbler nests. We found individuals belonging to the Mesostigmata, Trombidiformes and Sarcoptiformes. The Trombidiformes, represented in our study only by the Prostigmata, achieved statistically significantly lower intensity and abundance, compared to representatives of other orders. However, the number of recorded prostigmatid species was high (65). The most common were: Stigmaeus sphagneti (22 nests), Stigmaeus longipilis (16), Eupodes voxencollinus (15), Cunaxa setirostris (14), Stigmaeus pilatus (11), and Linopodes sp. 2 (10). The prevalence of Mesostigmata and Sarcoptiformes was equal, reaching 91.1%. Most of Gamasina (Mesostigmata) species found in this study were more characteristic of the soil environment and forest litter than bird nests, but there was also a typical bird parasite, viz. Ornithonyssus sylviarum. None of the observed species of Uropodina (Mesostigmata) or Oribatida (Sarcoptiformes) was typical for bird nests. Among the Uropodina, the highest parameters of nest infestation were achieved by Oodinychus ovalis, whereas among the Oribatida, they were achieved by Metabelba pulverosa. We discuss the importance of wood warbler nests for mite dispersal, survival and reproduction.

A sink host allows a specialist herbivore to persist in a seasonal source.

In seasonal environments, sinks that are more persistent than sources may serve as temporal stepping stones for specialists. However, this possibility has to our knowledge, not been demonstrated to date, as such environments are thought to select for generalists, and the role of sinks, both in the field and in the laboratory, is difficult to document. Here, we used laboratory experiments to show that herbivorous arthropods associated with seasonally absent main (source) habitats can endure on a suboptimal (sink) host for several generations, albeit with a negative growth rate. Additionally, they dispersed towards this host less often than towards the main host and accepted it less often than the main host. Finally, repeated experimental evolution attempts revealed no adaptation to the suboptimal host. Nevertheless, field observations showed that arthropods are found in suboptimal habitats when the main habitat is unavailable. Together, these results show that evolutionary rescue in the suboptimal habitat is not possible. Instead, the sink habitat functions as a temporal stepping stone, allowing for the persistence of a specialist when the source habitat is gone.

Hitchhiking or hang gliding? Dispersal strategies of two cereal-feeding eriophyoid mite species.

Dispersal shapes the dynamics of populations, their genetic structure and species distribution; therefore, knowledge of an organisms' dispersal abilities is crucial, especially in economically important and invasive species. In this study, we investigated dispersal strategies of two phytophagous eriophyoid mite species: Aceria tosichella (wheat curl mite, WCM) and Abacarus hystrix (cereal rust mite, CRM). Both species are obligatory plant parasites that infest cereals and are of economic significance. We investigated their dispersal success using different dispersal agents: wind and vectors. We hypothesised that in both mite species the main mode of dispersal is moving via wind, whereas phoretic dispersal is rather accidental, as the majority of eriophyoid mite species do not possess clear morphological or behavioural adaptations for phoresy. Results confirmed our predictions that both species dispersed mainly with wind currents. Additionally, WCM was found to have a higher dispersal success than CRM. Thus, this study contributes to our understanding of the high invasive potential of WCM.

Temperature-dependent development and survival of an invasive genotype of wheat curl mite, Aceria tosichella.

Quantifying basic biological data, such as the effects of variable temperatures on development and survival, is crucial to predicting and monitoring population growth rates of pest species, many of which are highly invasive. One of the most globally important pests of cereals is the eriophyoid wheat curl mite (WCM), Aceria tosichella, which is the primary vector of several plant viruses. The aim of this study was to evaluate temperature-dependent development and survival of WCM at a wide range of constant temperatures in the laboratory (17-33 °C). The development time of each stage depended significantly on temperature and it was negatively correlated with temperature increase. At high temperatures (27-33 °C), individuals had shorter developmental times, with the shortest (6 days) at 33 °C, whereas at the lowest tested temperatures (17-19 °C), developmental time was almost 3× longer. Moreover, temperature had a clear effect on survival: the higher the temperature, the lower the survival rate. These data provide information promoting more efficient and effective manipulation of WCM laboratory colonies, and further our understanding of the ramifications of temperature change on WCM physiology and implications for the growth and spread of this globally invasive pest.

A comprehensive and cost-effective approach for investigating passive dispersal in minute invertebrates with case studies of phytophagous eriophyid mites.

Dispersal is a fundamental biological process that operates at different temporal and spatial scales with consequences for individual fitness, population dynamics, population genetics, and species distributions. Studying this process is particularly challenging when the focus is on microscopic organisms that disperse passively, whilst controlling neither the transience nor the settlement phase of their movement. In this work we propose a comprehensive approach for studying passive dispersal of microscopic invertebrates and demonstrate it using wind and phoretic vectors. The protocol includes the construction of versatile, modifiable dispersal tunnels as well as a theoretical framework quantifying the movement of species via wind or vectors, and a hierarchical Bayesian approach appropriate to the structure of the dispersal data. The tunnels were used to investigate the three stages of dispersal (viz., departure, transience, and settlement) of two species of minute, phytophagous eriophyid mites Aceria tosichella and Abacarus hystrix. The proposed devices are inexpensive and easy to construct from readily sourced materials. Possible modifications enable studies of a wide range of mite species and facilitate manipulation of dispersal factors, thus opening a new important area of ecological study for many heretofore understudied species.

Integrative taxonomy of Abacarus mites (Eriophyidae) associated with hybrid sugarcane plants, including description of a new species.

Phytophagous mites belonging to the Eriophyoidea are extremely diverse and highly host-specific. Their accurate morphological identification is hampered by their reduced size and simplified bodies and by the existence of cryptic species complexes. Previous studies have demonstrated the urgency of applying multisource methods to accurate taxonomic identification of eriophyoid mites, especially species belonging to the genus Abacarus. This genus comprises 65 species, of which 37 are associated with grasses and four with sugarcane Saccharum (Poaceae). Recently, Abacarus specimens very similar to Abacarus sacchari were collected from the sugarcane crop in Brazil; however, their taxonomic placement was uncertain. In this study, we used an integrative approach to determine whether A. aff. sacchari specimens belong to A. sacchari or constitute a cryptic species. Morphological data were combined with molecular phylogeny based on the nucleotide sequences of three markers, one mitochondrial (COI) and two nuclear (D2 region of 28S and ITS). Morphological differences were observed between A. aff. sacchari, A. sacchari and A. doctus. The phylogenetic relationships among these three taxa and the genetic distances separating them revealed an interspecific divergence. The results of the morphological and molecular methods were congruent and supported the existence of a new species: Abacarus neosacchari n. sp. Duarte and Navia, herein described. This species belongs to the Abacarus cryptic species complex associated with sugarcane in the Americas. The results of this study, presenting the occurrence of multiple Abacarus species associated with sugarcane, contribute to the knowledge on plants and mites diversity by adding up one more clue highlighting that plant hybridization can be an important mechanism contributing to the speciation of plant-feeding arthropods.

Genetics of lineage diversification and the evolution of host usage in the economically important wheat curl mite, Aceria tosichella Keifer, 1969.

BACKGROUND: Understanding the mechanisms that underlie the diversification of herbivores through interactions with their hosts is important for their diversity assessment and identification of expansion events, particularly in a human-altered world where evolutionary processes can be exacerbated. We studied patterns of host usage and genetic structure in the wheat curl mite complex (WCM), Aceria tosichella, a major pest of the world's grain industry, to identify the factors behind its extensive diversification. RESULTS: We expanded on previous phylogenetic research, demonstrating deep lineage diversification within the taxon, a complex of distinctive host specialist and generalist lineages more diverse than previously assumed. Time-calibrated phylogenetic reconstruction inferred from mitochondrial DNA sequence data suggests that lineage diversification pre-dates the influence of agricultural practices, and lineages started to radiate in the mid Miocene when major radiations of C4 grasses is known to have occurred. Furthermore, we demonstrated that host specificity is not phylogenetically constrained, while host generalization appears to be a more derived trait coinciding with the expansion of the world's grasslands. Demographic history of specialist lineages have been more stable when compared to generalists, and their expansion pre-dated all generalist lineages. The lack of host-associated genetic structure of generalists indicates gene flow between mite populations from different hosts. CONCLUSIONS: Our analyses demonstrated that WCM is an unexpectedly diverse complex of genetic lineages and its differentiation is likely associated with the time of diversification and expansion of its hosts. Signatures of demographic histories and expansion of generalists are consistent with the observed proliferation of the globally most common lineages. The apparent lack of constrains on host use, coupled with a high colonization potential, hinders mite management, which may be further compromised by host range expansion. This study provides a significant contribution to the growing literature on host-association and diversification in herbivorous invertebrates.

Competition-driven niche segregation on a landscape scale: Evidence for escaping from syntopy towards allotopy in two coexisting sibling passerine species.

The role of interspecific competition for generating patterns in species' distribution is hotly debated and studies taking into account processes occurring at both large and small spatial scales are almost missing. Theoretically, competition between species with overlapping niches should result in divergence of their niches in sympatry to reduce the costs of competition. Many species show a mosaic distribution within sympatric zones, with the syntopic sites occupied by both species, and allotopic sites where only one species occurs. It is unclear whether such mosaics arise as a consequence of competition-driven niche segregation or due to the decline of their abundances towards range edges driven by environmental gradients. If the interspecific competition matters, we should observe (1) a shift in habitat preferences of one or both species between syntopy and allotopy, and (2) between allopatry and allotopy. Moreover, (3) species should show greater divergence in their habitat preferences in allotopy than in allopatry where (4) no differences in habitat preferences may occur. Finally, (5) shifts should be generally greater in the competitively subordinate species than in the dominant species. We used a unique dataset on abundance of two closely related passerine species, the Common Nightingale (Luscinia megarhynchos) and the Thrush Nightingale (Luscinia luscinia), collected across their syntopy, allotopy and allopatry. The predictions were tested within a generalized mixed-effects modelling framework. After accounting for environmental gradients perpendicular to the species' contact zone, we found a strong support for all but one prediction. Habitat preferences of both species shifted markedly between syntopy and allotopy, as well as between allopatry and allotopy. Whereas the species preferred the same habitats in allopatry, their preferences became strikingly different in allotopy where the abundance of the Common Nightingale increased towards dry and warm sites with low coverage of pastures, while the abundance of the Thrush Nightingale showed exactly opposite trends. Fifth prediction was not supported. Our results indicate that the competition between closely related species can result in considerable changes in habitat use across their geographic ranges accompanied with divergence in their habitat preferences in sympatry. Here, the species "escape" from competition to allotopic sites covered by habitats avoided by the competitor. Therefore, we argue that the interspecific competition is an important driver of species' distribution at both large and small spatial scales.

Cryptic diversity within grass-associated Abacarus species complex (Acariformes: Eriophyidae), with the description of a new species, Abacarus plumiger n. sp.

Accurate estimation of species richness is often complex as genetic divergence is not always accompanied by appreciable morphological differentiation. In consequence, cryptic lineages or species evolve. Cryptic speciation is common especially in taxa characterized by small and simplified bodies, what makes their proper identification challenging. The cereal rust mite, Abacarus hystrix, was regarded for a long time as a species associated with a wide range of grass hosts, whereas wide host ranges are rather rare in eriophyoid mites. Therefore, the generalist status of A. hystrix was questioned. In this paper we demonstrate that the diversity within Abacarus species associated with grasses is more complex than it was previously thought. The 78 Abacarus mtDNA COI sequences used in this study formed 10 highly supported clades (bootstrap value 99%) and four more distinct genetic lineages were represented by unique sequences. The genetic distances between them ranged from 6.6 to 26.5%. Moreover, morphological study and genetic approach based on the combination of the Poisson Tree Processes model for species delimitation (PTP) and a Bayesian implementation of PTP (bPTP), and Neighbour Joining analyses led to delimitation of a new species within the Abacarus complex: Abacarus plumiger, specialized on smooth brome (Bromus inermis). Furthermore, our analyses demonstrated a pattern of host-associated differentiation within the complex. Overall, our study indicates that cryptic speciation occurs in the grass-associated Abacarus genus, and suggests the need for more extensive sampling using integrative methods.

Is body size important? Seasonal changes in morphology in two grass-feeding Abacarus mites.

Overwintering strategies in herbivorous mites (Acariformes: Eriophyoidea) are poorly understood. A study of two Abacarus spp. was conducted to compare body size parameters of adult females in different seasons. Mites of Abacarus n. sp. (under description) and A. lolli were sampled from Bromopsis inermis and Lolium perenne, respectively, in April, September and December of 2001 in Poznan, Poland; 21 morphological traits were measured for each specimen. A principal component analysis revealed significant differences in body size parameters between collection dates, with larger females collected in December in both species. Larger body size in winter is consistent with the hypothesis that mites of these species, for which deutogyny has not been observed, undergo physiological changes such as accumulation of nutritional reserves, that enable them to withstand adverse environmental conditions. Larger body size has also been shown in other invertebrates to reduce heat loss in cold conditions. Filling gaps in the current knowledge of eriophyoid overwintering strategies, whether in the presence or absence of deutogyny, will contribute to both basic and applied future studies of this important arthropod group.

Population growth rate of dry bulb mite, Aceria tulipae (Acariformes: Eriophyidae), on agriculturally important plants and implications for its taxonomic status.

Dry bulb mite (DBM), Aceria tulipae, is an economically important mite with a worldwide distribution and a broad host range. As a generalist, it is the most important eriophyoid mite attacking bulbous plants such as garlic, onion and tulip. To date, DBM has been recorded on host plants belonging to the families Liliaceae, Amaryllidaceae, Melanthiaceae and Asparagaceae. However, a precise understanding of DBM host range is lacking as it is largely based on casual records of mites on plants, some of which may include accidental hosts. Moreover, the possible existence of cryptic species has not been considered. In this study the hypothesis that DBM may be a complex of distinct genetic lineages or cryptic species was tested by comparing the common barcode sequence marker mtDNA COI of specimens from several populations originating from the Netherlands and Poland. The population growth rate of DBM on seven agriculturally important plant species and on various parts of the garlic plant was also experimentally assessed in the laboratory. The results did not support the first hypothesis, and indicated that DBM populations originating from Poland and the Netherlands shared essentially the same genome. In addition, they indicated that DBM reached the highest population growth rate on leek and also displayed high growth rates on garlic, chive and red onion, whereas white onion and wheat were not colonized by the mites. Answering the question of whether DBM is a single polyphagous species rather than a complex of cryptic lineages is of particular importance since the misidentification of pests may lead to ineffective control strategies. Moreover, improved knowledge of DBM host range is essential for assessing risk to crops.

Change in abundance of three phytophagous mite species (Acari: Eriophyidae, Tetranychidae) on quackgrass in the presence of choke disease.

Phytophagous mites and endophytic fungi may interact when sharing a host plant, potentially influencing one another's growth or population dynamics; however, interactions between them are poorly known and remain largely unexplored. In this study, quantitative associations between three species of phytophagous mites and the endophytic fungus Epichloë bromicola Leuchtm. & Schardl (Clavicipitaceae, Ascomycotina) on quackgrass, Elymus repens (L.) Gould are reported. The mites' abundance was assessed on field-collected grass shoots that were either exhibiting choke disease symptoms or without the fungus. Overall, the abundance of Tetranychus urticae and Aculodes mckenziei was significantly lower on quackgrass plants infected by E. bromicola compared to plants without the fungus. Conversely, populations of Abacarus hystrix were significantly larger on plants colonised by the fungus than on uninfected plants. Thus, the presence of this endophytic fungus may have divergent effects on different phytophagous mite species although the basis of these effects is not yet known.

Cryptic speciation in the Acari: a function of species lifestyles or our ability to separate species?

There are approximately 55,000 described Acari species, accounting for almost half of all known Arachnida species, but total estimated Acari diversity is reckoned to be far greater. One important source of currently hidden Acari diversity is cryptic speciation, which poses challenges to taxonomists documenting biodiversity assessment as well as to researchers in medicine and agriculture. In this review, we revisit the subject of biodiversity in the Acari and investigate what is currently known about cryptic species within this group. Based on a thorough literature search, we show that the probability of occurrence of cryptic species is mainly related to the number of attempts made to detect them. The use of, both, DNA tools and bioassays significantly increased the probability of cryptic species detection. We did not confirm the generally-accepted idea that species lifestyle (i.e. free-living vs. symbiotic) affects the number of cryptic species. To increase detection of cryptic lineages and to understand the processes leading to cryptic speciation in Acari, integrative approaches including multivariate morphometrics, molecular tools, crossing, ecological assays, intensive sampling, and experimental evolution are recommended. We conclude that there is a demonstrable need for future investigations focusing on potentially hidden mite and tick species and addressing evolutionary mechanisms behind cryptic speciation within Acari.

Genetic and morphological diversity of Trisetacus species (Eriophyoidea: Phytoptidae) associated with coniferous trees in Poland: phylogeny, barcoding, host and habitat specialization.

Eriophyoid species belonging to the genus Trisetacus are economically important as pests of conifers. A narrow host specialization to conifers and some unique morphological characteristics have made these mites interesting subjects for scientific inquiry. In this study, we assessed morphological and genetic variation of seven Trisetacus species originating from six coniferous hosts in Poland by morphometric analysis and molecular sequencing of the mitochondrial cytochrome oxidase subunit I gene and the nuclear D2 region of 28S rDNA. The results confirmed the monophyly of the genus Trisetacus as well as the monophyly of five of the seven species studied. Both DNA sequences were effective in discriminating between six of the seven species tested. Host-dependent genetic and morphological variation in T. silvestris and T. relocatus, and habitat-dependent genetic and morphological variation in T. juniperinus were detected, suggesting the existence of races or even distinct species within these Trisetacus taxa. This is the first molecular phylogenetic analysis of the Trisetacus species. The findings presented here will stimulate further investigations on the evolutionary relationships of Trisetacus as well as the entire Phytoptidae family.

Understanding the joint evolution of dispersal and host specialisation using phytophagous arthropods as a model group.

Theory generally predicts that host specialisation and dispersal should evolve jointly. Indeed, many models predict that specialists should be poor dispersers to avoid landing on unsuitable hosts while generalists will have high dispersal abilities. Phytophagous arthropods are an excellent group to test this prediction, given extensive variation in their host range and dispersal abilities. Here, we explore the degree to which the empirical literature on this group is in accordance with theoretical predictions. We first briefly outline the theoretical reasons to expect such a correlation. We then report empirical studies that measured both dispersal and the degree of specialisation in phytophagous arthropods. We find a correlation between dispersal and levels of specialisation in some studies, but with wide variation in this result. We then review theoretical attributes of species and environment that may blur this correlation, namely environmental grain, temporal heterogeneity, habitat selection, genetic architecture, and coevolution between plants and herbivores. We argue that theoretical models fail to account for important aspects, such as phenotypic plasticity and the impact of selective forces stemming from other biotic interactions, on both dispersal and specialisation. Next, we review empirical caveats in the study of this interplay. We find that studies use different measures of both dispersal and specialisation, hampering comparisons. Moreover, several studies do not provide independent measures of these two traits. Finally, variation in these traits may occur at scales that are not being considered. We conclude that this correlation is likely not to be expected from large-scale comparative analyses as it is highly context dependent and should not be considered in isolation from the factors that modulate it, such as environmental scale and heterogeneity, intrinsic traits or biotic interactions. A stronger crosstalk between theoretical and empirical studies is needed to understand better the prevalence and basis of the correlation between dispersal and specialisation.

Phylogenetic analyses reveal extensive cryptic speciation and host specialization in an economically important mite taxon.

The wheat curl mite (WCM) is a major pest in cereal crops around the world and the vector of at least four known pathogens capable of reducing yields in crops such as wheat, corn, barley, oats, millet and rye. Current taxonomy recognizes WCM as a single species, Aceriatosichella; however, recent genetic, physiological and ecological studies have shown that WCM is likely to be a species complex. In this study we assessed genetic variation and phylogenetic relationships among WCM from four continents and a wide range of host plants using DNA sequence data from one mitochondrial gene, one nuclear gene and a single nuclear intergenic spacer region. Phylogenetic analyses revealed 11 unique mite lineages associated with specific plant hosts including wheat and barley. Host associations were consistent across continents, often with a single haplotype dominating a host plant regardless of geographic origin. The genetic and ecological differences identified in this study support the notion that WCM is a species complex in need of major taxonomic revision. These findings have implications for control of WCM globally, particularly within the context of identifying plants that form 'green bridge' refuges, assessing disease transmission risk, and identifying resistance in cereal genotypes to WCM and associated pathogens.

Wheat curl mite, Aceria tosichella, and transmitted viruses: an expanding pest complex affecting cereal crops.

The wheat curl mite (WCM), Aceria tosichella, and the plant viruses it transmits represent an invasive mite-virus complex that has affected cereal crops worldwide. The main damage caused by WCM comes from its ability to transmit and spread multiple damaging viruses to cereal crops, with Wheat streak mosaic virus (WSMV) and Wheat mosaic virus (WMoV) being the most important. Although WCM and transmitted viruses have been of concern to cereal growers and researchers for at least six decades, they continue to represent a challenge. In older affected areas, for example in North America, this mite-virus complex still has significant economic impact. In Australia and South America, where this problem has only emerged in the last decade, it represents a new threat to winter cereal production. The difficulties encountered in making progress towards managing WCM and its transmitted viruses stem from the complexity of the pathosystem. The most effective methods for minimizing losses from WCM transmitted viruses in cereal crops have previously focused on cultural and plant resistance methods. This paper brings together information on biological and ecological aspects of WCM, including its taxonomic status, occurrence, host plant range, damage symptoms and economic impact. Information about the main viruses transmitted by WCM is also included and the epidemiological relationships involved in this vectored complex of viruses are also addressed. Management strategies that have been directed at this mite-virus complex are presented, including plant resistance, its history, difficulties and advances. Current research perspectives to address this invasive mite-virus complex and minimize cereal crop losses worldwide are also discussed.

Mechanisms of dispersal and colonisation in a wind-borne cereal pest, the haplodiploid wheat curl mite.

Dispersal and colonisation determine the survival and success of organisms, and influence the structure and dynamics of communities and ecosystems in space and time. Both affect the gene flow between populations, ensuring sufficient level of genetic variation and improving adaptation abilities. In haplodiploids, such as Aceria tosichella (wheat curl mite, WCM), a population may be founded even by a single unfertilised female, so there is a risk of heterozygosity loss (i.e. founder effect). It may lead to adverse outcomes, such as inbreeding depression. Yet, the strength of the founder effect partly depends on the genetic variation of the parental population. WCM is an economically important pest with a great invasive potential, but its dispersal and colonisation mechanisms were poorly studied before. Therefore, here we assessed WCM dispersal and colonisation potential in relation to the genetic variation of the parental population. We checked whether this potential may be linked to specific pre-dispersal actions (e.g. mating before dispersal and collective behaviour). Our study confirms that dispersal strategies of WCM are not dependent on heterozygosity in the parental population, and the efficient dispersal of this species depends on collective movement of fertilised females.

Effective specialist or jack of all trades? Experimental evolution of a crop pest in fluctuating and stable environments.

Understanding pest evolution in agricultural systems is crucial for developing effective and innovative pest control strategies. Types of cultivation, such as crop monocultures versus polycultures or crop rotation, may act as a selective pressure on pests' capability to exploit the host's resources. In this study, we examined the herbivorous mite Aceria tosichella (commonly known as wheat curl mite), a widespread wheat pest, to understand how fluctuating versus stable environments influence its niche breadth and ability to utilize different host plant species. We subjected a wheat-bred mite population to replicated experimental evolution in a single-host environment (either wheat or barley), or in an alternation between these two plant species every three mite generations. Next, we tested the fitness of these evolving populations on wheat, barley, and on two other plant species not encountered during experimental evolution, namely rye and smooth brome. Our results revealed that the niche breadth of A. tosichella evolved in response to the level of environmental variability. The fluctuating environment expanded the niche breadth by increasing the mite's ability to utilize different plant species, including novel ones. Such an environment may thus promote flexible host-use generalist phenotypes. However, the niche expansion resulted in some costs expressed as reduced performances on both wheat and barley as compared to specialists. Stable host environments led to specialized phenotypes. The population that evolved in a constant environment consisting of barley increased its fitness on barley without the cost of utilizing wheat. However, the population evolving on wheat did not significantly increase its fitness on wheat, but decreased its performance on barley. Altogether, our results indicated that, depending on the degree of environmental heterogeneity, agricultural systems create different conditions that influence pests' niche breadth evolution, which may in turn affect the ability of pests to persist in such systems.