Toward an understanding of the chemical ecology of alternative reproductive tactics in the bulb mite (Rhizoglyphus robini)

Background Under strong sexual selection, certain species evolve distinct intrasexual, alternative reproductive tactics (ARTs). In many cases, ARTs can be viewed as environmentally-cued threshold traits, such that ARTs coexist if their relative fitness alternates over the environmental cue gradient. Surprisingly, the chemical ecology of ARTs has been underexplored in this context. To our knowledge, no prior study has directly quantified pheromone production for ARTs in a male-polymorphic species. Here, we used the bulb mite—in which males are either armed fighters that kill conspecifics, or unarmed scramblers (which have occasionally been observed to induce mating behavior in other males)—as a model system to gain insight into the role of pheromones in the evolutionary maintenance of ARTs. Given that scramblers forgo investment into weaponry, we tested whether scramblers produce higher quantities of the putative female sex-pheromone α-acaridial than fighters, which would improve the fitness of the scrambler phenotype through female mimicry by allowing avoidance of aggression from competitors. To this end, we sampled mites from a rich and a poor nutritional environment and quantified their production of α-acaridial through gas chromatography analysis. Results We found a positive relationship between pheromone production and body size, but males exhibited a steeper slope in pheromone production with increasing size than females. Females exhibited a higher average pheromone production than males. We found no significant difference in slope of pheromone production over body size between fighters and scramblers. However, scramblers reached larger body sizes and higher pheromone production than fighters, providing some evidence for a potential female mimic strategy adopted by large scramblers. Pheromone production was significantly higher in mites from the rich nutritional environment than the poor environment. Conclusion Further elucidation of pheromone functionality in bulb mites, and additional inter- and intrasexual comparisons of pheromone profiles are needed to determine if the observed intersexual and intrasexual differences in pheromone production are adaptive, if they are a by-product of allometric scaling, or diet-mediated pheromone production under weak selection. We argue chemical ecology offers a novel perspective for research on ARTs and other complex life-history traits.

Size-structured mating niche differentiation in the male-polyphenic mite Rhizoglyphus robini

Condition-dependent expression of alternative male morphs evolves when males of different sizes experience different mating niches, requiring different morphologies. Such mating niche differentiation can be due to competitive asymmetry between large and small males in contests over mating opportunities. Here, we tested the hypothesis that aggressive interactions among males cause size-structured mating niches in an acarid mite with condition-dependent male polyphenism: the bulb mite Rhizoglyphus robini. In this species, large males mature as armed fighters with enlarged legs, and small males as scramblers without modified legs. We staged experimental dyadic contests over a mating opportunity between either a pair of fighter males, or a fighter and a scrambler male. We predicted that the larger male would have a higher likelihood of mating first in contests among fighters, that the fighter male would have a higher likelihood of mating first in fighter-scrambler contests, that fighters would have a higher likelihood of interrupting ongoing copulations if they are larger than their mating rival, and that copulations in the presence of a fighter rival therefore last shorter than copulations with a scrambler rival present. We found that in contests among fighters, the larger male had a higher probability of mating first. In contests among a fighter and scrambler, the fighter male was more likely to mate first regardless of the body size difference between the contestants. Ongoing copulations were only rarely interrupted by the rival male (always by a fighter), and the probability of interruption did not depend on the body size difference between the mating male and its rival. Copulations lasted shorter in the presence of a rival fighter, but this effect was not attributable to interruption of copulations. We conclude that the fighting niche is particularly accessible for large males, as larger males have a higher probability of winning pre-copulatory contests. Such mating niche differentiation likely contributes to evolutionary maintenance of condition-dependent male polyphenism, where small males are forced to adopt an alternative mating tactic and hence develop a dedicated morphology.

Toward an understanding of the chemical ecology of alternative reproductive tactics in the bulb mite (Rhizoglyphus robini)

BackgroundUnder strong sexual selection, certain species evolve distinct intrasexual, alternative reproductive tactics (ARTs). In many cases, ARTs can be viewed as environmentally-cued threshold traits, such that ARTs coexist if their relative fitness alternates over the environmental cue gradient. Surprisingly, the chemical ecology of ARTs has been underexplored in this context. To our knowledge, no prior study has directly quantified pheromone production for ARTs in a male-polymorphic species. Here, we used the bulb mite—in which males are either armed fighters that kill conspecifics, or unarmed scramblers—as a model system to gain insight into the role of pheromones in the evolutionary maintenance of ARTs. Given that scramblers forgo investment into weaponry, we tested whether scramblers produce higher pheromone quantities than fighters, which would improve the fitness of the scrambler phenotype, e.g. through female mimicry to avoid aggression from competitors. To this end, we sampled mites from a rich and a poor nutritional environment and quantified their production of the female sex pheromone α-acaridial through gas chromatography analysis. ResultsWe found a positive relationship between pheromone production and body size, but males exhibited a steeper slope in pheromone production with increasing size than females. Females exhibited a higher average pheromone production than males. We found no significant difference in slope of pheromone production over body size between fighters and scramblers. However, scramblers reached larger body sizes and higher pheromone production than fighters, providing some evidence for a potential female mimic strategy adopted by large scramblers. Pheromone production was significantly higher in mites from the rich nutritional environment than the poor environment. ConclusionFurther elucidation of pheromone functionality in bulb mites, and additional inter- and intrasexual comparisons of pheromone profiles are needed to determine if the observed intersexual and intrasexual differences in pheromone production are adaptive, if they are a by-product of allometric scaling, or diet-mediated pheromone production under weak selection. We argue chemical ecology offers a novel perspective for research on ARTs and other complex life-history traits.

Toward an understanding of the chemical ecology of alternative reproductive tactics in the bulb mite (Rhizoglyphus robini)

Background Under strong sexual selection, certain species evolve distinct intrasexual, alternative reproductive tactics (ARTs). In many cases, ARTs can be viewed as environmentally cued threshold traits, such that ARTs coexist if their relative fitness alternates over the environmental cue gradient. Surprisingly, the chemical ecology of ARTs has been underexplored in this context. To our knowledge, no prior study has directly quantified pheromone production for ARTs in a male polymorphic species. Here, we used the bulb mite — in which males are either armed fighters that kill conspecifics, or unarmed scramblers — as a model system to gain insight into the role of pheromones in the evolutionary maintenance of ARTs. Given that scramblers forgo investment into weaponry, we tested whether scramblers produce higher pheromone quantities than fighters, which would improve the fitness of the scrambler phenotype, e.g. through female mimicry to avoid aggression from competitors. To this end, we sampled mites from a rich and a poor nutritional environment and quantified their production of the female sex pheromone α acaridial through gas chromatography analysis. Results We found a positive relationship between pheromone production and body size, but males exhibited a steeper slope in pheromone production with increasing size than females. Females exhibited a higher average pheromone production than males. We found no significant difference in slope of pheromone production over body size between fighters and scramblers. However, scramblers reached larger body sizes and higher pheromone production than fighters, providing some evidence for a potential female mimic strategy adopted by large scramblers. Pheromone production was significantly higher in mites from the rich nutritional environment than the poor environment. Conclusion Further elucidation of pheromone functionality in bulb mites, and additional inter and intrasexual comparisons of pheromone profiles are needed to determine if the observed intersexual and intrasexual differences in pheromone production are adaptive, if they are a byproduct of allometric scaling, or diet mediated pheromone production under weak selection. We argue chemical ecology offers a novel perspective for research on ARTs and other complex life-history traits.

Efficacy of soil isolates of entomopathogenic fungi against the bulb mite, Rhizoglyphus robini (Acari: Acaridae)

The bulb mite, Rhizoglyphus robini, is a serious pest of garlic, onion and other crops. The mite is usually found in association with dangerous fungal pathogens such as Fusarium spp. Control of this pest has relied upon the use of synthetic acaricides but chemical control of the bulb mite is difficult because it is able to develop resistance quickly. Thus, alternative control methods, e.g. biological control, need to be developed and implemented. The aim of this study was to assess efficacy of selected strains of entomopathogenic fungi (EPF) against adult females of R. robini under laboratory conditions. New EPF strains were isolated from soil samples collected in onion and garlic fields in the Czech Republic and Israel using soil elution and cultivation on selective media. Fungal species were determined using macroscopic, microscopic and molecular markers. The efficacy against R. robini females was tested in 17 isolated and 3 reference strains of EPF. Results revealed high variability among species and strains. The highest efficacy against R. robini mites was found in strains of Metarhizium anisopliae isolated from soil samples collected in the Czech Republic which caused mortality up to 99.3%, and a Metarhizium indigoticum strain from Israel causing 98.3% mortality after four days of bioassay. Isaria fumosorosea strains did not caused mortality higher than 40%. The lowest virulence was found in Beauveria spp. strains causing mortality of mites between 5 and 25%. Median lethal time (LT50) and median lethal concentration (LC50) in the three most virulent strains ranged between 2 and 4 days and between 1.01×104 and 2.36×105 spores/ml, respectively. The concentration-response models indicated that the M. indigoticum strain is more lethal than M. anisopliae strains. The present study showed that some strains of entomopathogenic fungi, especially from the genus Metarhizium, could be perspective biocontrol agents against R. robini.

Acute toxicity and sublethal effects of metal oxide nanoparticles against the bulb mite

Crocus sativus L. (Iridaceae) known as saffron is the most commercially important medicinal food product in Iran and worldwide because of its significance for exporting commercial. However, the health of this plant is adversely threatened by abiotic and biotic stress. Bulb mite Rhizoglyphus robini Claparede (Acari: Acaridae) as abiotic stress is oligophagous and the main pest of tuber plants, commonly found in all the cultivated saffron producing areas could decrease saffron economic value. The mites tunnel through corms allowing the entry of fungal and bacterial pathogens. The widespread use of commercial pesticides in modern agriculture to protect crops from pests has caused significant public concern because of their side impacts on the environment and non-target species. Nanotechnology has been used in the production, processing, storage, packaging, and transport of agricultural products. Newly, oxide nanoparticles have been introduced as novel pesticides against pests and fertilizer. This study was conducted as an effective and environmentally-friendly method to control the growth of Rhizoglyphus robini. The lethal and sublethal effects of metal oxide nanoparticles (MNPs) including nano copper oxides (nano-CuO), nano magnesium oxide (nano-MgO), nano zinc oxide (nano-ZnO), nano titanium dioxide (nano-TiO2), and nano iron oxide (nano-Fe2O3) which their lethal concentrations ranged between 159.2 and 1036.4 mg L-1 were studied on Rhizoglyphus robini adult females. The contact effect of the tested MNPs was evaluated using the corm dip bioassay method. Assays showed that the contact toxicity of the nano-CuO treatment (229.1 mg L-1) was higher than other MNPs. To assess the sublethal effects of the MNPs, adult females were exposed to the LC25 concentration of MNPs, and life history and life-table parameters of the surviving Rhizoglyphus robini were investigated. The exposure to sublethal concentrations of the tested MNPs prolonged the egg incubation, larval period, and nymphal period significantly when compared to the control mites. Compared to other MNPs or control mite, nano-MgO and nano-TiO2 led to the longest duration of the total immature stages (11.32 and 11.32, respectively). The net reproductive rate (R0), the intrinsic rate of increase (rm), the finite rate of increase (λ), and the generation time (T) were significantly different between control and MNP treatments. The population exposed to ZnO-, MgO-, and TiO2-nanoformulations had a much lower rm value (0.164, 0.164, and 0.171 d-1, respectively) compared to the control population (0.231 d-1) and those exposed to Fe3O4- and CuO-nanoformulations (1.19 and 1.19 d-1, respectively). According to our findings, all tested MNPs, especially CuO-, ZnO-, and MgO-nano formulations, have a potential application for use in the management of Rhizoglyphus robini.

Interactions of Metarhizium brunneum-7 with Phytophagous Mites Following Different Application Strategies

Metarhizium brunneum is a generalist entomopathogenic fungus known to be virulent against Acari. We investigated Metarhizium brunneum-7 (Mb7) interactions in three systems of phytophagous mites and their respective plant hosts: Volkamer lemon (Citrus volkameriana) and the citrus rust mite Phyllocoptruta oleivora; common bean (Phaseolus vulgaris) and the two-spotted spider mite Tetranychus urticae; and spring onion (Allium cepa) and the bulb mite Rhizoglyphus robini. All three mite species were susceptible to directly applied Mb7 conidia. Results obtained using the standard method for studying endophytic colonization vs. live confocal imaging of plant tissues using the green fluorescent protein (GFP)-transformed fungus differed markedly, demonstrating that microscopy validation was more definite than the standard process of recovery from plant tissue. Endophytic colonization was observed in conidium-infiltrated citrus leaves and in roots of onion plants treated with soil-drenched conidia, but not in common bean treated by either spray or drench of conidia. Endophytic colonization of citrus leaves did not affect the citrus mite population. Drench application in common bean reduced two-spotted mite population. Similarly, drench application in onion reduced bulb mite population. This study emphasizes the importance of the host plant effects on Mb7 control efficacy of mite pests, and the merits of live-imaging techniques in studying endophytic interaction.