6Pgdh polymorphism in wild bulb mite populations: prevalence, environmental correlates and life history trade-offs.

Genetic polymorphism in key metabolic genes plays a pivotal role in shaping phenotypes and adapting to varying environments. Polymorphism in the metabolic gene 6-phosphogluconate dehydrogenase (6Pgdh) in bulb mites, Rhizoglyphus robini is characterized by two alleles, S and F, that differ by a single amino acid substitution and correlate with male reproductive fitness. The S-bearing males demonstrate a reproductive advantage. Although the S allele rapidly fixes in laboratory settings, the persistence of polymorphic populations in the wild is noteworthy. This study examines the prevalence and stability of 6Pgdh polymorphism in natural populations across Poland, investigating potential environmental influences and seasonal variations. We found widespread 6Pgdh polymorphism in natural populations, with allele frequencies varying across locations and sampling dates but without clear geographical or seasonal clines. This widespread polymorphism and spatio-temporal variability may be attributed to population demography and gene flow between local populations. We found some correlation between soil properties, particularly cation content (Na, K, Ca, and Mg) and 6Pgdh allele frequencies, showcasing the connection between mite physiology and soil characteristics and highlighting the presence of environment-dependent balancing selection. We conducted experimental fitness assays to determine whether the allele providing the advantage in male-male competition has antagonistic effects on life-history traits and if these effects are temperature-dependent. We found that temperature does not differentially influence development time or juvenile survival in different 6Pgdh genotypes. This study reveals the relationship between genetic variation, environmental factors, and reproductive fitness in natural bulb mite populations, shedding light on the dynamic mechanisms governing 6Pgdh polymorphism.

Transitional chelal digit patterns in saprophagous astigmatan mites.

Changes in the functional shape of astigmatan mite moveable digit profiles are examined to test if Tyrophagus putrescentiae (Acaridae) is a trophic intermediate between a typical micro-saprophagous carpoglyphid (Carpoglyphus lactis) and a common macro-saprophagous glycyphagid (Glycyphagus domesticus). Digit tip elongation in these mites is decoupled from the basic physics of optimising moveable digit inertia. Investment in the basal ramus/coronoid process compared to that for the moveable digit mastication length varies with feeding style. A differentiated ascending ramus is indicated in C. lactis and in T. putrescentiae for different trophic reasons. Culturing affects relative investments in C. lactis. A markedly different style of feeding is inferred for the carpoglyphid. The micro-saprophagous acarid does not have an intermediate pattern of trophic functional form between the other two species. Mastication surface shape complexity confirms the acarid to be heterodontous. T. putrescentiae is a particularly variably formed species trophically. A plausible evolutionary path for the gradation of forms is illustrated. Digit form and strengthening to resist bending under occlusive loads is explored in detail. Extensions to the analytical approach are suggested to confirm the decoupling of moveable digit pattern from cheliceral and chelal adaptations. Caution is expressed when interpreting ordinations of multidimensional data in mites.

Rhizoglyphus robini, a pest mite of saffron, is unable to resist extracellular ice formation.

The saffron mite, Rhizoglyphus robini Claparède (Acari, Astigmata: Acaridae), is one of the most important pests of saffron-producing regions in Iran. It causes yellowing and decreases saffron growth, and finally it destroys the bulbs. In this research, the cold tolerance and supercooling point (SCP) of the saffron mite were measured in three populations and two temperature regimes. Our results showed that the mean SCP of the saffron mite was approximately -14.6 °C without significant difference among the populations. On the contrary, acclimation of the mites significantly decreased their SCP to a mean of approximately -16.5 °C. Exposure of the mites for 24 h to 0 and -2.5 °C had no significant effect on the survival of the mites but when the mites were exposed to -5.0 °C for 24 h, survival of the three populations reached the lowest level of roughly 60%. By 24-h exposure to -7.5 °C, survival of the mites was almost negligible. As a large proportion of mortality was observed above the SCP, and LT50 > SCP, it can be inferred that the saffron mite is likely a chill-susceptible species. This suggests that the saffron mite lacks the ability to withstand extracellular ice formation. Overall, the results of the current study suggest no significant physiological differences between populations of the saffron mite.

Hidden biodiversity in microarthropods (Acari, Oribatida, Eremaeoidea, Caleremaeus).

A challenge for taxonomists all over the world and across all taxonomic groups is recognizing and delimiting species, and cryptic species are even more challenging. However, an accurate identification is fundamental for all biological studies from ecology to conversation biology. We used a multidisciplinary approach including genetics as well as morphological and ecological data to assess if an easily recognizable, widely distributed and euryoecious mite taxon represents one and the same species. According to phylogenetic (based on mitochondrial and nuclear genes) and species delimitation analyses, five distinct putative species were detected and supported by high genetic distances. These genetic lineages correlate well with ecological data, and each species could be associated to its own (micro)habitat. Subsequently, slight morphological differences were found and provide additional evidence that five different species occur in Central and Southern Europe. The minuteness and the characteristic habitus of Caleremaeus monilipes tempted to neglect potential higher species diversity. This problem might concern several other "well-known" euryoecious microarthropods. Five new species of the genus Caleremaeus are described, namely Caleremaeus mentobellus sp. nov., C. lignophilus sp. nov., C. alpinus sp. nov., C. elevatus sp. nov., and C. hispanicus sp. nov. Additionally, a morphological evaluation of C. monilipes is presented.

Deformed wings of small hive beetle independent of virus infections and mites.

Insect wing deformities can be caused by viruses, mites and other environmental stressors during development. Here we conducted differential diagnostics of deformed wings in small hive beetles, Aethina tumida (SHB). Adult SHB with and without deformed wings from individual and mass reared scenarios were evaluated for the mite Tyrophagus putrescentiae and for deformed wing virus. Viral load and mite number were similar for SHB with deformed wings and unaffected beetles. Because deformed wings were only observed in individually pupating SHB, a humidity challenge most likely explains the observed clinical symptoms. Our observations support the importance of differential diagnostics.

Juvenile diet quality and intensity of sexual conflict in the mite Sancassania berlesei.

BACKGROUND: Differing evolutionary interests of males and females may result in sexual conflict, whereby traits or behaviours that are beneficial for male reproductive success (e.g., traits related to male-male competition) are costly for females. Since sexual conflict may play an important role in areas such as speciation, population persistence or evolution of life history traits, understanding what factors modulate the intensity of sexual conflict is important. This study aims to examine juvenile diet quality as one of the underestimated ecological factors that may affect the intensity of sexual conflict via individual conditions. I used food manipulation during the development of the mite Sancassania berlesei to investigate the effects on male reproductive behaviour and competitiveness, male-induced harm to female fitness and female resistance to this harm. RESULTS: Males that were exposed to low-quality food started mating later than the control males, and number of their mating attempts were lower compared to those of control males. Moreover, males from the low-quality diet treatment sired fewer offspring under competition than males from the control treatment. However, the fitness of females exposed to males reared on a poor diet did not differ from that of females mated with control males. Furthermore, female diet quality did not alter their resistance to male-induced harm. CONCLUSION: Overall, diet quality manipulation affected male reproductive behaviour and mating success. However, I found no evidence that the intensity of sexual conflict in S. berlesei depends on male or female conditions. Investigating a broader range of environmental factors will provide a better understanding of sexual conflict dynamics and its feedback into associated evolutionary mechanisms.

Storage Mite Infestation of Dry-Stored Food Products and Its Relation to Human Intestinal Acariasis in the City of Minia, Egypt.

Initial data on the prevalence of storage mites in dry-stored food products and estimates of the presence of mites in human stool in the city of Minia, Egypt are provided. In total, 847 samples were collected randomly from houses and retail stores between March 2017 and February 2018. In addition, 1,000 human stool samples were collected for the detection of the presence of mites. Mites were extracted from 285 of 840 (33.9%) samples, and mite contamination was found to be most prevalent in wheat flour (73.3%). In total, 11 mite species belonging to six families were identified, with the pest species Tyrophagus putrescentiae (Schrank) (Acari: Acaridae) (TP) being the most prevalent (91.2% of samples). The seasonal density distribution showed the highest storage mite density in March-April, followed by October, and the lowest in January. In addition, mites were detected in 87 (8.7%) human stool samples, with significant associations between certain occupations and some personal characteristics. Therefore, more attention needs to be paid to intestinal acariasis arising from mite infestation of dry-stored food products.

Oral delivery of water-soluble compounds to the phytoseiid mite Neoseiulus californicus (Acari: Phytoseiidae).

Phytoseiids are predatory mites that prey on other mites and small arthropods, and several species are used in commercial agriculture for biological control of pests. To optimize phytoseiid mites' use in biocontrol, an efficient method for oral delivery of test compounds is required to assess their sensitivities to pesticides, RNAi for gene functional analysis and artificial diets. Here we developed four methods for oral delivery of a solution of xenobiotics to different life stages of the commercially available generalist predatory mite Neoseiulus californicus: (i) soaking mites in the solution, or allowing them to feed on (ii) spider mites soaked in the solution, (iii) a solution droplet, or (iv) solution-saturated filter paper. As measured by ingestion of a tracer dye, the droplet-based feeding system was most efficient; the dye was observed in the alimentary canal of >90% test mites of all life stages, with no mortality. The droplet-based feeding system was also effective for the commercially available specialist predatory mite Phytoseiulus persimilis, with >80% delivery efficiency. This study paves the way for development of methods for high-throughput RNAi and for toxicological or nutritional assays in phytoseiid mites.

Cross-level considerations for explaining selection pressures and the maintenance of genetic variation in condition-dependent male morphs.

Condition-dependent expression of alternative male morphologies (AMMs) exists in many arthropods. Understanding their coexistence requires answering (at least) two questions: (i) what are the ecological selection pressures that maintain condition-dependent plasticity of AMM expression, and (ii) what maintains the associated genetic variation? Focusing on acarid mites, we show that the questions should not be conflated. We argue how, instead, answers should be sought by testing phenotype-level (question 1) or genotype-level (question 2) hypotheses. We illustrate that energy allocation restrictions and physiological trade-offs are likely to play a crucial role in AMM expression in acarid mites. We thus conclude that these aspects require specific attention in identifying selection pressures maintaining condition-dependent plasticity, and evolutionary processes that maintain genetic variation in condition-dependent phenotypic plasticity.

Population growth of the stored product pest Tyrophagus putrescentiae (Acari: Acaridae) on environmentally and medically important fungi.

The stored food mite Tyrophagus putrescentiae (Schrank) (Acari: Acaridae) has been associated with the presence of several fungal species. The aims of this work were to evaluate T. putrescentiae population growth associated to environmental and medically important fungal species to determine on which fungal species populations of T. putrescentiae performs best, and to evaluate their ability to disperse each fungal species. First, 24 fungal species were inoculated separately in Petri dishes containing Sabouraud agar medium. One week after inoculation, 50 mites were added to each plate. On the 28th evaluation day, mites and eggs were counted in each plate, and 50 mites randomly collected from each replicate were transferred to new plates containing only Sabouraud agar medium. Then, mites, eggs, and fungal population were evaluated in each plate on day 28 again. The highest population increases were on Trichophyton mentagrophytes, Alternaria sp., Microsporum gypseum, and Aspergillus chevalieri. With Fusarium guttiforme and the medically important fungi Microsporum canis, M. gypseum, T. mentagrophytes, and Sporothrix sp., mites were observed to feed on whole mycelium. Only eight fungal species were dispersed by T. putrescentiae to the new Petri dishes: Aspergillus clavatus, Candida tropicalis, Candida albicans, Fusarium guttiforme, Hyphopichia burtonii, Penicillium citrinum, Rhizophus azygosporus, and Trichophyton mentagrophytes. The best performance of T. putrescentiae was found feeding on F. guttiforme, P. citrinum, and T. mentagrophytes. In conclusion, T. putrescentiae successfully used fungi as a food source, and it proved to be an important tool for disseminating both environmental and medically important fungi.

Storage mite concentrations are underestimated compared to house dust mite concentrations.

Dwellings are increasingly well insulated to save energy and this leads to higher humidity and temperature, which improves conditions for mites. Dermatophagoides antigens are the main allergens involved and tested in atopic asthma. We developed three new species-specific quantitative PCR (qPCR) methods for house dust mites (Dermatophagoides pteronyssinus and D. farinae) and storages mites (Acarus siro, Glycyphagus domesticus, Lepidoglyphus destructor). We sampled dust with electrostatic dust collectors, in the bedrooms, under beds and in the kitchens of patients with allergies (n = 24) and healthy controls (n = 18). Mite quantification was carried out with the three new qPCRs and the qPCR previously described for the Dermatophagoides genus. The qPCRs were highly specific and efficient for house dust mite species and the storage mites. Storage mite concentrations were higher than house dust mite concentrations and were higher in dwellings of patients with allergies. Consequently, allergists should test more often patients against the storage mite antigens by prick tests or IgE serology. Dampness is a major factor in storage mite development and the presence of effective mechanical ventilation can reduce storage mite concentrations four-fold. In addition, to limit exposure to dust mites, treatments should be used throughout dwellings and not only in patients' bedrooms.

Risk of exposure of a selected rural population in South Poland to allergenic mites. Part II: acarofauna of farm buildings.

Exposure to mite allergens, especially from storage and dust mites, has been recognized as a risk factor for sensitization and allergy symptoms that could develop into asthma. The aim of this study was to investigate the occurrence of mites in debris and litter from selected farm buildings of the Malopolskie province, South Poland, with particular reference to allergenic and/or parasitic species as a potential risk factor of diseases among farmers. Sixty samples of various materials (organic dust, litter, debris and residues) from farm buildings (cowsheds, barns, chaff-cutter buildings, pigsties and poultry houses) were subjected to acarological examination. The samples were collected in Lachowice and Kurów (Suski district, Malopolskie). A total of 16,719 mites were isolated including specimens from the cohort Astigmatina (27 species) which comprised species considered as allergenic (e.g., Acarus siro complex, Tyrophagus putrescentiae, Lepidoglyphus destructor, Glycyphagus domesticus, Chortoglyphus arcuatus and Gymnoglyphus longior). Species of the families Acaridae (A. siro, A. farris and A. immobilis), Glycyphagidae (G. domesticus, L. destructor and L. michaeli) and Chortoglyphidae (C. arcuatus) have been found as numerically dominant among astigmatid mites. The majority of mites were found in cowsheds (approx. 32%) and in pigsties (25.9%). The remaining mites were found in barns (19.6%), chaff-cutter buildings (13.9%) and poultry houses (8.8%). The results suggest that the allergenic mites may constitute an occupational hazard for agricultural workers in all farming environments examined.

The Mite Tyrophagus putrescentiae Hosts Population-Specific Microbiomes That Respond Weakly to Starvation.

The effect of short-term nutrient deprivation was studied in five populations of the mite Tyrophagus putrescentiae with different microbiomes. The fresh weight, nutrient status, respiration, and population growth of the mites were observed for the five mite population-scale samples. The starvation caused the larvae and nymphs to be eliminated, resulting in a significant increase in the fresh weight of starved adult specimens. Three populations were negatively influenced by starvation, and the starved specimens were characterized by a decrease in nutrient status, respiration, and population growth. One population was not influenced or was slightly influenced by starvation, which had no effect on population growth or nutrient contents but caused a significant decrease in respiration. One population was positively influenced by starvation; the population growth increased in starved specimens, and starvation had no effect on respiration. Although starvation altered the bacterial profiles of the microbiomes, these differences were much smaller than those between the populations. The bacterial profiles of Staphylococcus, Bacillus, Kocuria, Brevibacterium, and unidentified Micrococcaceae and Enterobacteriaceae increased in starved specimens, whereas those of Bartonella and Solitalea-like genera were reduced in the starved mite populations. The profiles of the intracellular symbiont Cardinium decreased in the starved specimens, and the Wolbachia profile changes were dependent on the mite population. In mite populations, when the symbionts were rare, their profiles varied stochastically. Correlations between changes in the profiles of the bacterial taxa and mite fitness parameters, including nutrient status (lipids, proteins, saccharides, and glycogen contents), mite population growth, and respiration, were observed. Although the microbiomes were resistant to the perturbations caused by nutrition deficiency, the responses of the mites differed in terms of their population growth, respiration, and nutrient status.

Costs of weaponry: Unarmed males sire more offspring than armed males in a male-dimorphic mite.

Morphological structures used as weapons in male-male competition are not only costly to develop but are also probably costly to maintain during adulthood. Therefore, having weapons could reduce the energy available for other fitness-enhancing actions, such as post-copulatory investment. We tested the hypothesis that armed males make lower post-copulatory investments than unarmed males, and that this difference will be most pronounced under food-limited conditions. We performed two experiments using the male-dimorphic bulb mite Rhizoglyphus robini, in which males are either armed "fighters" or unarmed "scramblers." Firstly, we tested whether fighters and scramblers differed in their reproductive output after being starved or fed for 1 or 2 weeks. Secondly, we measured the reproductive output of scramblers and fighters (starved or fed) after one, two or three consecutive matings. Scramblers sired more offspring than fighters after 1 week, but scramblers and fighters only sired a few offspring after 2 weeks. Scramblers also sired more offspring than fighters at the first mating, and males rarely sired offspring after consecutive matings. Contrary to our hypothesis, the fecundity of starved and fed males did not differ. The higher reproductive output of scramblers suggests that, regardless of nutritional state, scramblers make larger post-copulatory investments than fighters. Alternatively, (cryptic) female choice generally favours scramblers. Why the morphs differed in their reproductive output is unclear. Neither morph performed well relatively late in life or after multiple matings. It remains to be investigated to what extent the apparent scrambler advantage contributes to the maintenance and evolution of male morph expression.

Mite color alteration and acaricidal activity of 3,7-dimethyl-2,6-octadienal and its structural analogues against the stored food pest mite Tyrophagus putrescentiae.

Acaricidal activities of the active component isolated from Melissa officinalis oil and its structural analogues against Tyrophagus putrescentiae were evaluated using fumigant and contact bioassays. The structure of 3,7-dimethyl-2,6-octadienal purified from M. officinalis oil was elucidated with EI-MS, 1H- and 13C-NMR, 1H-1H COSY, and DEPT-NMR. Based on the LD50 values of 3,7-dimethyl-2,6-octadienal analogues in fumigant and contact bioassays, respectively, 2,4-octadienal showed the highest activity (LD50 = 2.09 µg/cm3 and 11.08 µg/cm2), followed by 3,7-dimethyl-6-octenal (3.60 µg/cm3 and 29.34 µg/cm2), 3,7-dimethyl-2,6-octadienal (6.18 µg/cm3 and 36.17 µg/cm2), 2-octenal (7.45 µg/cm3 and 47.36 µg/cm2) and M. officinalis oil (8.89 µg/cm3 and 23.83 µg/cm2). Comparing the acaricidal activities of the aldehyde group based on the degree of unsaturation, 2,4-octadienal containing two double bonds was more potent than 2-octenal with a single double bond. Based on the acaricidal activities of the methyl group, on the other hand, 3,7-dimethyl-6-octenal containing a single double bond was more acaricidal than 3,7-dimethyl-2,6-octadienal with two double bonds. These results indicate that 3,7-dimethyl-2,6-octadienal analogues are useful to control food mites.

Response of the predatory mite Cydnoseius negevi (Acari: Phytoseiidae) to webbing of the date palm mite, Oligonychus afrasiaticus (Acari: Tetranychidae), on date palm fruits and leaves.

The behavioral response of a generalist phytoseiid predator, Cydnoseius negevi (Swirskii & Amitai) (Acari: Phytoseiidae) to the complicated webbing of the date palm mite (DPM) Oligonychus afrasiaticus (McGregor) (Acari: Tetranychidae) was evaluated for the first time on date palm leaves and fruits, both in the laboratory and on field-collected samples. Cydnoseius negevi used its first pair of legs in a swimming-like to-and-fro movement to penetrate the complicated webbing of DPM and demonstrated three attack behaviors, i.e., patrolling (PG), web invasion (WI), and web penetration (WP), against the webbing of DPM on date fruits and leaves. The time spent by the predator on attack behaviors was significantly longer in the laboratory for treatments where either more prey females or immature stages along with females were present. The time spent by C. negevi on response behaviors such as searching, grooming, and resting increased with increased number of DPM females and with the addition of immature stages along with DPM females. Cydnoseius negevi, even though being a generalist phytoseiid predator, showed its potential towards penetrating the complex dense webs of DPM on date palm fruits and leaves. If released in suitable numbers prior to establishment of DPM colonies, C. negevi may prove to be an effective biological control agent.

A Gustatory Receptor Used for Rapid Detection of Tyrophagus putrescentiae in Fungi Hosts.

The storage mite, Tyrophagus putrescentiae, found worldwide in many habitats, is an important pest of edible fungi in China. Storage mites are tiny and difficult to observe, especially when they occur in fungi composts. In this study, one gustatory receptor protein (TputGR1) was identified from the transcriptome of T. putrescentiae. Phylogenetic analysis of GRs families from 10 arthropod species revealed that TputGR1 had high homology with the SccaGR1 of Sarcoptes scabiei and TurtGR1-2 of Tetranychus urticae, but low homology with other insect species, Drosophila melanogaster, Anopheles gambiae, Bombyx mori, Aedes aegypti, Culex quinquefasciatus, and Pediculus humanus. We developed a detection system for the mite on fungi hosts using the GR protein and the loop-mediated isothermal amplification (LAMP). This procedure was rapid (60 min from sampling to result) and had high sensitivity (0.5 ng/mL). LAMP provided rapid and reliable detection of T. putrescentiae. It has good specificity for single samples and for large-scale surveys.

Fitness consequences of threshold trait expression subject to environmental cues.

Most cases of alternative reproductive tactics (ARTs) are thought to represent conditional strategies, whereby high-status males express highly competitive phenotypes, whereas males below a certain status threshold resort to sneaky tactics. The underlying evolutionarily stable strategy (ESS) model assumes that males of high competitive ability achieve higher fitness when expressing the territorial phenotype, whereas the less competitive males are more fit as sneakers, caused by fitness functions for the ARTs having different slopes and intersecting at a threshold value of competitive ability. The model, however, is notoriously difficult to test as it requires access to low-status territorials and high-status sneakers, that rarely occur in nature. Here, we test the conditional ESS in the androdimorphic acarid mite Sancassania berlesei, where large males tend to develop into an armoured, aggressive 'fighter' morph, while small males become unarmoured, non-aggressive 'scramblers'. In addition to body size, male morph is affected by pheromones produced by big populations, with fighters being suppressed in dense colonies. By manipulating pheromone concentration, we obtained high-status scramblers and low-status fighters. We also estimated status- and size-dependent fitness functions for male morphs across a range of population sizes. Fighters had the highest fitness in small populations and their fitness declined with increasing density, whereas the reverse was true for scramblers, providing support for condition-dependent ESS with respect to demography. However, whereas male fitness increased with body size, the fitness functions did not differ significantly between morphs. Thus, although we found evidence for the intersection of morph fitness functions with respect to demography, we did not find such an intersection in relation to male body size. Our results highlight how demography can exert selection pressures shaping the evolution of the conditional strategy in species with ARTs.

Trait-based predictions and responses from laboratory mite populations to harvesting in stochastic environments.

Predictions on population responses to perturbations are often derived from trait-based approaches like integral projection models (IPMs), but are rarely tested. IPMs are constructed from functions that describe survival, growth and reproduction in relation to the traits of individuals and their environment. Although these functions comprise biologically non-informative statistical coefficients within standard IPMs, model parameters of the recently developed dynamic energy budget IPM (DEB-IPM) are life-history traits like "length at maturation" and "maximum reproduction rate". Testing predictions from mechanistic IPMs against empirical observations can therefore provide functional insights into the links between individual life history, the environment and population dynamics. Here, we compared the population dynamics of the bulb mite (Rhizoglyphus robini) predicted by a DEB-IPM with those observed in an experiment where populations experienced daily food rations that were either positively correlated over time (red noise), negatively (blue noise) or uncorrelated (white noise). We also selectively harvested large adults in half of these populations. The model failed to generate detailed predictions of population structure as juvenile numbers were overestimated; likely because juvenile-adult interference competition was underestimated. The model performed well at the population level as, for both harvested and unharvested populations, simulations matched the observed, long-term stochastic growth rate λs . We next generalised the model to investigate how stochastic change affects mite λs , which correlated well with the frequency f of experiencing periods of good environment, but, due to the relationship between f and noise colour ρ, did not correlate well with shifts in ρ. The sensitivity of λs to perturbations in life-history parameters depended on the type of stochastic change, as well as population growth. Our findings show that responses to differential mortality depend on individual life-history traits, environmental characteristics and population growth. As long-term climate change causes ever greater environmental fluctuations, trait-based approaches will be increasingly important in predicting population responses to change. We therefore conclude by illustrating what questions can be examined with mechanistic trait-based models like the DEB-IPM, the answers to which will advance our knowledge of the functional links between individual traits, the environment and population dynamics.

Morph-specific artificial selection reveals a constraint on the evolution of polyphenisms.

Theory predicts that the evolution of polyphenic variation is facilitated where morphs are genetically uncoupled and free to evolve towards their phenotypic optima. However, the assumption that developmentally plastic morphs can evolve independently has not been tested directly. Using morph-specific artificial selection, we investigated correlated evolution between the sexes and male morphs of the bulb mite Rhizoglyphus echinopus Large 'fighter' males have a thick and sharply terminating pair of legs used to kill rival males, while small 'scrambler' males have unmodified legs, and search for unguarded females, avoiding fights. We selected on the relative leg width of only the fighter male morph, tracked the evolutionary responses in fighters and the correlated evolutionary responses in scramblers and females that were untouched by direct selection. Fighters diverged in relative leg thickness after six generations; assaying scramblers and females at the ninth generation we observed correlated responses in relative leg width in both. Our results represent strong evidence for the evolution of intraspecific phenotypic diversity despite correlated evolution between morphs and sexes, challenging the idea that male morphs are genetically uncoupled and free to independently respond to selection. We therefore question the perceived necessity for genetic independence in traits with extreme phenotypic plasticity.