Community structure,trophic position and reproductive mode of soil and bark-living oribatid mites in an alpine grassland ecosystem

The community structure, stable isotope ratios (¹⁵N/¹⁴N, ¹³C/¹²C) and reproductive mode of oribatid mites (Acari, Oribatida) were investigated in four habitats (upper tree bark, lower tree bark, dry grassland soil, forest soil) at two sites in the Central Alps (Tyrol, Austria). We hypothesized that community structure and trophic position of oribatid mites of dry grassland soils and bark of trees are similar since these habitats have similar abiotic characteristics (open, dry) compared with forest soil. Further, we hypothesized that derived taxa of oribatid mites reproducing sexually dominate on the bark of trees since species in this habitat consume living resources such as lichens. In contrast to our hypothesis, the community structure of oribatid mites differed among grassland, forest and bark indicating the existence of niche differentiation in the respective oribatid mite species. In agreement with our hypothesis, sexually reproducing taxa of oribatid mites dominated on the bark of trees whereas parthenogenetic species were more frequent in soil. Several species of bark-living oribatid mites had stable isotope signatures that were similar to lichens indicating that they feed on lichens. However, nine species that frequently occurred on tree bark did not feed on lichens according to their stable isotope signatures. No oribatid mite species could be ascribed to moss feeding. We conclude that sexual reproduction served as preadaptation for oribatid mites allowing them to exploit new habitats and new resources on the bark of trees. Abiotic factors likely are of limited importance for bark-living oribatid mites since harsh abiotic conditions are assumed to favor parthenogenesis.     

Parthenogenetic vs. sexual reproduction in oribatid mite communities

The dominance of sex in Metazoa is enigmatic. Sexual species allocate resources to the production of males, while potentially facing negative effects such as the loss of well‐adapted genotypes due to recombination, and exposure to diseases and predators during mating. Two major hypotheses have been put forward to explain the advantages of parthenogenetic versus sexual reproduction in animals, that is, the Red Queen hypothesis and the Tangled Bank/Structured Resource Theory of Sex. The Red Queen hypothesis assumes that antagonistic predator—prey/ parasite–host interactions favor sex. The Structured Resource Theory of Sex predicts sexual reproduction to be favored if resources are in short supply and aggregated in space. In soil, a remarkable number of invertebrates reproduce by parthenogenesis, and this pattern is most pronounced in oribatid mites (Oribatida, Acari). Oribatid mites are abundant in virtually any soil across very different habitats, and include many sexual and parthenogenetic (thelytokous) species. Thereby, they represent an ideal model group to investigate the role of sexual versus parthenogenetic reproduction across different ecosystems and habitats. Here, we compiled data on oribatid mite communities from different ecosystems and habitats across biomes, including tropical rainforests, temperate forests, grasslands, arable fields, salt marshes, bogs, caves, and deadwood. Based on the compiled dataset, we analyzed if the percentage of parthenogenetic species and the percentage of individuals of parthenogenetic species are related to total oribatid mite density, species number, and other potential driving factors of the reproductive mode including altitude and latitude. We then interpret the results in support of either the Red Queen hypothesis or the Structured Resource Theory of Sex. Overall, the data showed that low density of oribatid mites due to harsh environmental conditions is associated with high frequency of parthenogenesis supporting predictions of the Structured Resource Theory of Sex rather than the Red Queen hypothesis.     

No evidence for the 'Meselson effect' in parthenogenetic oribatid mites (Oribatida, Acari)

It has been hypothesized that in ancient apomictic, nonrecombining lineages the two alleles of a single copy gene will become highly divergent as a result of the independent accumulation of mutations (Meselson effect). We used a partial sequence of the elongation factor-1alpha (ef-1alpha) and the heat shock protein 82 (hsp82) genes to test this hypothesis for putative ancient parthenogenetic oribatid mite lineages. In addition, we tested if the hsp82 gene is fully transcribed by sequencing the cDNA and we also tested if there is evidence for recombination and gene conversion in sexual and parthenogenetic oribatid mite species. The average maximum intra-specific divergence in the ef-1alpha was 2.7% in three parthenogenetic species and 8.6% in three sexual species; the average maximum intra-individual genetic divergence was 0.9% in the parthenogenetic and 6.0% in the sexual species. In the hsp82 gene the average maximum intra-individual genetic divergence in the sexual species Steganacarus magnus and in the parthenogenetic species Platynothrus peltifer was 1.1% and 1.2%, respectively. None of the differences were statistically significant. The cDNA data indicated that the hsp82 sequence is transcribed and intron-free. Likelihood permutation tests indicate that ef-1alpha has undergone recombination in all three studied sexual species and gene conversion in two of the sexual species, but neither process has occurred in any of the parthenogenetic species. No evidence for recombination or gene conversion was found for sexual or parthenogenetic oribatid mite species in the hsp 82 gene. There appears to be no Meselson effect in parthenogenetic oribatid mite species. Presumably, their low genetic divergence is due to automixis, other homogenizing mechanisms or strong selection to keep both the ef-1alpha and the hsp82 gene functioning.     

Radiation in sexual and parthenogenetic oribatid mites (Oribatida, Acari) as indicated by genetic divergence of closely related species

The D3 domain and its flanking regions of 28S rRNA of four pairs of closely related sexual species (Eupelops hirtus and E. torulosus; Oribatella calcarata and O. quadricornuta; Chamobates voigtsi and Ch. borealis; Liacarus coracinus and L. subterraneus) and four pairs of closely related parthenogenetic species (Nanhermannia nana and Na. coronata; Nothrus silvestris and No. palustris; Tectocepheus sarekensis and T. minor; Camisia spinifer and Ca. segnis) of oribatid mites were sequenced to investigate (1) if the D3 region can be used as a species marker and (2) if there is genetic variation among closely related species pairs and if its magnitude is related to reproductive mode. Furthermore, we investigated the world-wide genetic variation of the D3 region from the oribatid mite species Platynothrus peltifer. There was no intraspecific genetic variation in the D3 region in any of the species studied; it was even identical in two closely related parthenogenetic species (Na. nana and Na. coronata) and two closely related sexual species (E. hirtus and E. torulosus). The genetic differences of the other species pairs indicated that both parthenogenetic and sexual lineages have various ages. On average, however, the differences between the closely related parthenogenetic species were larger than those between closely related sexual species, indicating that parthenogenetic lineages exist historically and may radiate slower than sexual species. The findings of this study support the hypothesis that some of the parthenogenetic oribatid mite taxa (Tectocepheus, Nothrus) are ‘ancient asexuals’. The absence of intraspecific or intra-individual variation in the D3 region of parthenogenetic species is consistent with the presence of concerted evolution in the 28S rRNA gene. From this we infer the existence of a meiotic process, which is consistent with the automixy known from several other parthenogenetic oribatid species. This revised version was published online in July 2006 with corrections to the Cover Date.     

Resource Availability as Driving Factor of the Reproductive Mode in Soil Microarthropods (Acari,Oribatida)

The availability of high quality resources is an important factor driving community structure and reproductive mode of animals. Parthenogenetic reproduction prevails when resources are available in excess, whereas sexuality correlates with resource shortage. We investigated the effect of resource availability on the community structure of oribatid mites in a laboratory experiment. Availability of food resources was increased by addition of glucose to leaf litter and reduced by leaching of nutrients from leaf litter. Experimental systems were incubated at three different temperatures to establish different regimes of resource exploitation. Community structure of oribatids and numbers of eggs per female were measured over a period of ten months. We expected the density of oribatid mites to decline in the reduced litter quality treatment but to increase in the glucose treatment. Both effects were assumed to be more pronounced at higher temperatures. We hypothesized sexual species to be less affected than parthenogenetic species by reduced resource quality due to higher genetic diversity allowing more efficient exploitation of limited resources, but to be outnumbered by parthenogenetic species in case of resource addition due to faster reproduction. In contrast to our hypotheses, both sexual and parthenogenetic oribatid mite species responded similarly with their densities declining uniformly during incubation. The parthenogenetic Brachychthoniidae and Tectocepheus dominated early in the experiment but were replaced later by parthenogenetic Desmonomata and Rhysotritia. In parthenogenetic species the number of eggs per female increased during the experiment while the number of eggs in sexual females remained constant or decreased slightly; in general, egg numbers were higher in sexual than in parthenogenetic species. The results indicate that for sustaining oribatid mite populations other resources than litter and associated saprotrophic microorganisms are needed. They also indicate that there are two groups of parthenogenetically reproducing species: exploiters of easily available resources and consumers of leaf litter associated resources.     

Molecular phylogeny of oribatid mites (Oribatida, Acari): evidence for multiple radiations of parthenogenetic lineages

Nucleotide sequences of the D3 expansion segment and its flanking regions of the 28S rDNA gene were used to evaluate phylogenetic relationships among representative sexual and asexual oribatid mites (Oribatida, Acariformes). The aim of this study was to investigate the hypothesis that oribatid mites consist of species-rich clusters of asexual species that may have radiated while being parthenogenetic. Furthermore, the systematic position of the astigmate mites (Astigmata, Acariformes) which have been hypothesised to represent a paedomorphic lineage within the oribatid mites, is investigated. This is the first phylogenetic tree for oribatid mites s.l. (incl. Astigmata) based on nucleotide sequences. Intraspecific genetic variation in the D3 region was very low, confirming the hypothesis that this region is a good species marker. Results from neighbour joining (NJ) and maximum parsimony (MP) algorithms indicate that several species-rich parthenogenetic groups like Camisiidae, Nanhermanniidae and Malaconothridae are monophyletic, consistent with the hypothesis that some oribatid mite groups diversified despite being parthenogenetic. The MP and maximum likelihood (ML) method indicated that the D3 region is a good tool for elucidating the relationship of oribatid mite species on a small scale(genera, families) but is not reliable for large-scale taxonomy, because branches from the NJ algorithm collapsed in the MP and ML tree. In all trees calculated by different algorithms the Astigmata clustered within the oribatid mites, as proposed earlier.     

Positive correlation of trophic level and proportion of sexual taxa of oribatid mites (Acari: Oribatida) in alpine soil systems

We investigated community structure, trophic ecology (using stable isotope ratios; ¹⁵N/¹⁴N, ¹³C/¹²C) and reproductive mode of oribatid mites (Acari, Oribatida) along an altitudinal gradient (2,050–2,900 m) in the Central Alps (Obergurgl, Austria). We hypothesized that (1) the community structure changes with altitude, (2) oribatid mites span over four trophic levels, (3) the proportion of sexual taxa increases with altitude, and (4) the proportion of sexual taxa increases with trophic level, i.e. is positively correlated with the δ¹⁵N signatures. Oribatid mite community structure changed with altitude indicating that oribatid mites occupy different niches at different altitudes. Oribatid mites spanned over 12 δ¹⁵N units, i.e. about four trophic levels, which is similar to lowland forest ecosystems. The proportion of sexually reproducing taxa increased from 2,050 to 2,900 m suggesting that limited resource availability at high altitudes favors sexual reproduction. Sexual taxa more frequently occurred higher in the food web indicating that the reproductive mode is related to nutrition of oribatid mites. Generally, oribatid mite community structure changed from being decomposer dominated at lower altitude to being dominated by fungal and lichen feeders, and predators at higher altitude. This supports the view that resources from dead organic material become less available with increasing altitude forcing species to feed on living resources such as fungi, lichens and nematodes. Our findings support the hypothesis that limited resource accessibility (at high altitudes) favors sexually reproducing species whereas ample resource supply (at lower altitudes) favors parthenogenetic species.     

Population genetic structure of sexual and parthenogenetic damselflies inferred from mitochondrial and nuclear markers

It has been postulated that obligate asexual lineages may persist in the long term if they escape from negative interactions with either sexual lineages or biological enemies; and thus, parthenogenetic populations will be more likely to occur in places that are difficult for sexuals to colonize, or those in which biological interactions are rare, such as islands or island-like habitats. Ischnura hastata is the only known example of natural parthenogenesis within the insect order Odonata, and it represents also a typical example of geographic parthenogenesis, as sexual populations are widely distributed in North America, whereas parthenogenetic populations of this species have only been found at the Azores archipelago. In order to gain insight in the origin and distribution of parthenogenetic I. hastata lineages, we have used microsatellites, mitochondrial and nuclear DNA sequence data, to examine the population genetic structure of this species over a wide geographic area. Our results suggest that sexual populations of I. hastata in North America conform to a large subdivided population that has gone through a recent spatial expansion. A recent single long distance dispersal event, followed by a demographic expansion, is the most parsimonious hypothesis explaining the origin of the parthenogenetic population of this species in the Azores islands.     

The ecological distribution of reproductive mode in oribatid mites, as related to biological complexity

The high incidence of asexuality in oribatid mites presents an unusual opportunity for examining hypotheses for the maintenance of sex. There is a presumed range in age of asexual species: many oribatid species are phylogenetically clustered, occurring in speciose early-derivative families or genera without sexual species, while others are phylogenetically isolated from other asexual species, occurring in later derivative taxa with sexual congeners. We examined the distribution of oribatid mite reproductive mode in soil of corn fields, grassy and shrub fields, and forests in central New York State (three replicate plots of each type, with 25 samples per plot), to test three ecological predictions from current theory. (1) If overall biotic uncertainty, as generated by competitors and predators, mediates the ecological distribution of oribatid mites, then the proportion of asexual oribatid mites should be negatively correlated with biological diversity; we examine this prediction using literature data as well. (2) If Muller’s Ratchet (the stochastic loss of best genotypes, which is independent of environment), mediates the success of asexuality, then no ecological pattern should exist. (3) If general purpose genotypes are characteristic of asexual oribatid mites, their habitat distribution should be broader than that of sexual species. For each plot the level of asexuality was compared to indices of overall biotic diversity, as calculated from the pooled oribatid mite (competitors) and mesostigmatid mite (predator) communities. We found no negative correlation in this relationship in our own data or in the literature analysis of 290 faunal surveys from 50 literature sources, so we reject biotic uncertainty as an important determinant of reproductive mode distribution. When only data on phylogenetically clustered asexuals are considered, there is instead a positive correlation between asexuality and diversity that is not explained. Because of the latter pattern we tentatively reject Muller’s Ratchet as the primary factor maintaining reproductive mode in these mites, but cannot reject it for isolated asexual species. Niche breadth in sexual and asexual oribatid mites provides no support for widespread general purpose genotypes but broad patterns in the literature suggest that the idea needs further investigation. Possible complicating or unknown factors that are discussed include historical disturbance in the study area, the relationship between parasitism and general biotic diversity, and the level and source of genetic diversity in asexual oribatid mites.     

High genetic divergences indicate ancient separation of parthenogenetic lineages of the oribatid mite Platynothrus peltifer (Acari, Oribatida)

Theories on the evolution and maintenance of sex are challenged by the existence of ancient parthenogenetic lineages such as bdelloid rotifers and darwinulid ostracods. It has been proposed that several parthenogenetic and speciose taxa of oribatid mites (Acari) also have an ancient origin. We used nucleotide sequences of the mitochondrial gene cytochrome oxidase I to estimate the age of the parthenogenetic oribatid mite species Platynothrus peltifer. Sixty-five specimens from 16 sites in North America, Europe and Asia were analysed. Seven major clades were identified. Within-clade genetic distances were below 2 % similar to the total intraspecific genetic diversity of most organisms. However, distances between clades averaged 56 % with a maximum of 125 %. We conclude that P. peltifer, as it is currently conceived, has existed for perhaps 100 million years, has an extant distribution that results from continental drift rather than dispersal and was subject to several cryptic speciations.     

Cryptic diversification in ancient asexuals: evidence from the bdelloid rotifer Philodina flaviceps

Bdelloid rotifers, darwinulid ostracods and some oribatid mites have been called 'ancient asexuals' as they speciated and survived over long-term evolutionary timescale without sexual recombination. Data on their genetic diversification are contrasting: within-species diversification is present mostly at a continental scale in a parthenogenetic oribatid mite, whereas almost no genetic diversification at all seems to occur within darwinulid ostracod species. Strangely enough, no clear data for bdelloid rotifers are available so far. In this paper, we analyse partial COI mtDNA sequences to show that a bdelloid rotifer, Philodina flaviceps, so far considered a single traditional morphological species, has actually been able to diversify into at least nine distinct evolutionary entities, with genetic distances between lineages comparable with those between different traditional species within the same genus. We discovered that local coexistence of such different independent lineages is very common: up to four lineages were found in a same stream, and up to three in a single moss sample of 5 cm(2). In contrast to the large-scale geographic pattern that has recently been reported in the oribatid mite, the spatial distribution of the bdelloid lineages provided evidence of micro-phylogeographic patterns. If the mtDNA diversity indicates that the lineages are independent and represent sympatric cryptic species within P. flaviceps, then the actual bdelloid diversity can be expected to be much greater than that recognized today.     

The Persistence of Facultative Parthenogenesis in Drosophila albomicans

Parthenogenesis has evolved independently in more than 10 Drosophila species. Most cases are tychoparthenogenesis, which is occasional or accidental parthenogenesis in normally bisexual species with a low hatching rate of eggs produced by virgin females; this form is presumed to be an early stage of parthenogenesis. To address how parthenogenesis and sexual reproduction coexist in Drosophila populations, we investigated several reproductive traits, including the fertility, parthenogenetic capability, diploidization mechanisms, and mating propensity of parthenogenetic D. albomicans. The fertility of mated parthenogenetic females was significantly higher than that of virgin females. The mated females could still produce parthenogenetic offspring but predominantly produced offspring by sexual reproduction. Both mated parthenogenetic females and their parthenogenetic-sexual descendants were capable of parthenogenesis. The alleles responsible for parthenogenesis can be propagated through both parthenogenesis and sexual reproduction. As diploidy is restored predominantly by gamete duplication, heterozygosity would be very low in parthenogenetic individuals. Hence, genetic variation in parthenogenetic genomes would result from sexual reproduction. The mating propensity of females after more than 20 years of isolation from males was decreased. If mutations reducing mating propensities could occur under male-limited conditions in natural populations, decreased mating propensity might accelerate tychoparthenogenesis through a positive feedback mechanism. This process provides an opportunity for the evolution of obligate parthenogenesis. Therefore, the persistence of facultative parthenogenesis may be an adaptive reproductive strategy in Drosophila when a few founders colonize a new niche or when small populations are distributed at the edge of a species'' range, consistent with models of geographical parthenogenesis.     

Patterns and mechanisms in instances of endosymbiont‐induced parthenogenesis

Female‐producing parthenogenesis can be induced by endosymbionts that increase their transmission by manipulating host reproduction. Our literature survey indicates that such endosymbiont‐induced parthenogenesis is known or suspected in 124 host species from seven different arthropod taxa, with Wolbachia as the most frequent endosymbiont (in 56–75% of host species). Most host species (81%, 100 out of 124) are characterized by haplo‐diploid sex determination, but a strong ascertainment bias likely underestimates the frequency of endosymbiont‐induced parthenogenesis in hosts with other sex determination systems. In at least one taxon, hymenopterans, endosymbionts are a significant driver of transitions from sexual to parthenogenetic reproduction, with one‐third of lineages being parthenogenetic as a consequence of endosymbiont infection. Endosymbiont‐induced parthenogenesis appears to facilitate the maintenance of reproductive polymorphism: at least 50% of species comprise both sexual (uninfected) and parthenogenetic (infected) strains. These strains feature distribution differences similar to the ones documented for lineages with genetically determined parthenogenesis, with endosymbiont‐induced parthenogens occurring at higher latitudes than their sexual relatives. Finally, although gamete duplication is often considered as the main mechanism for endosymbiont‐induced parthenogenesis, it underlies parthenogenesis in only half of the host species studied thus far. We point out caveats in the methods used to test for endosymbiont‐induced parthenogenesis and suggest specific approaches that allow for firm conclusions about the involvement of endosymbionts in the origin of parthenogenesis.     

Repeated convergent evolution of parthenogenesis in Acariformes (Acari)

The existence of old species‐rich parthenogenetic taxa is a conundrum in evolutionary biology. Such taxa point to ancient parthenogenetic radiations resulting in morphologically distinct species. Ancient parthenogenetic taxa have been proposed to exist in bdelloid rotifers, darwinulid ostracods, and in several taxa of acariform mites (Acariformes, Acari), especially in oribatid mites (Oribatida, Acari). Here, we investigate the diversification of Acariformes and their ancestral mode of reproduction using 18S rRNA. Because parthenogenetic taxa tend to be more frequent in phylogenetically old taxa of Acariformes, we sequenced a wide range of members of this taxon, including early‐derivative taxa of Prostigmata, Astigmata, Endeostigmata, and Oribatida. Ancestral character state reconstruction indicated that (a) Acariformes as well as Oribatida evolved from a sexual ancestor, (b) the primary mode of reproduction during evolution of Acariformes was sexual; however, species‐rich parthenogenetic taxa radiated independently at least four times (in Brachychthonioidea (Oribatida), Enarthronota (Oribatida), and twice in Nothrina (Oribatida), (c) parthenogenesis additionally evolved frequently in species‐poor taxa, for example, Tectocepheus, Oppiella, Rostrozetes, Limnozetes, and Atropacarus, and (d) sexual reproduction likely re‐evolved at least three times from species‐rich parthenogenetic clusters, in Crotonia (Nothrina), in Mesoplophora/Apoplophora (Mesoplophoridae, Enarthronota), and in Sphaerochthonius/Prototritia (Protoplophoridae, Enarthronota). We discuss possible reasons that favored the frequent diversification of parthenogenetic taxa including the continuous long‐term availability of dead organic matter resources as well as generalist feeding of species as indicated by natural variations in stable isotope ratios.     

Origin and Genetic Diversity of Diploid Parthenogenetic Artemia in Eurasia

There is wide interest in understanding how genetic diversity is generated and maintained in parthenogenetic lineages, as it will help clarify the debate of the evolution and maintenance of sexual reproduction. There are three mechanisms that can be responsible for the generation of genetic diversity of parthenogenetic lineages: contagious parthenogenesis, repeated hybridization and microorganism infections (e.g. Wolbachia). Brine shrimps of the genus Artemia (Crustacea, Branchiopoda, Anostraca) are a good model system to investigate evolutionary transitions between reproductive systems as they include sexual species and lineages of obligate parthenogenetic populations of different ploidy level, which often co-occur. Diploid parthenogenetic lineages produce occasional fully functional rare males, interspecific hybridization is known to occur, but the mechanisms of origin of asexual lineages are not completely understood. Here we sequenced and analysed fragments of one mitochondrial and two nuclear genes from an extensive set of populations of diploid parthenogenetic Artemia and sexual species from Central and East Asia to investigate the evolutionary origin of diploid parthenogenetic Artemia, and geographic origin of the parental taxa. Our results indicate that there are at least two, possibly three independent and recent maternal origins of parthenogenetic lineages, related to A. urmiana and Artemia sp. from Kazakhstan, but that the nuclear genes are very closely related in all the sexual species and parthenogegetic lineages except for A. sinica, who presumable took no part on the origin of diploid parthenogenetic strains. Our data cannot rule out either hybridization between any of the very closely related Asiatic sexual species or rare events of contagious parthenogenesis via rare males as the contributing mechanisms to the generation of genetic diversity in diploid parthenogenetic Artemia lineages.     

Vertical stratification of oribatid (Acari: Oribatida) communities in relation to their morphological and life-history traits and tree structures in a subtropical forest in Japan

To clarify the effect of tree structure on the diversity of oribatid mites (Acari: Oribatida), we collected a total of 16,325 oribatids in 181 morphospecies from the leaves, branches, and trunk bark of trees and from the forest-floor soil and litter in a subtropical forest, Okinawa, Japan, and tested three predictions: (1) moisture stress in arboreal habitats would lead to larger body size (not supported); (2) morphological traits related to gripping a surface (number and size of claws) would be more developed in arboreal species (supported); and (3) advantages in colonization (no cost for searching mates and doubled population growth) would favor parthenogenesis in the arboreal oribatid communities (not supported). We observed vertical stratification among the five habitats in terms of mite density, species diversity, and species composition, but found no difference in the body length of oribatid species between the arboreal and forest-floor habitats. However, (homo-)tridactylous species predominated in the arboreal habitat, suggesting that this claw morphology facilitates adherence to and movement on arboreal substrates. Sexual species were common in the arboreal oribatid communities, whereas about half of the dominant species collected from the forest floor were likely to be parthenogenetic. It is unclear how these different reproductive systems may be advantageous for oribatid mites in arboreal and forest-floor habitats. Nevertheless, the structural complexity provided by trees appears to enhance and maintain the diversity of oribatid communities through vertical stratification in this subtropical forest.     

Phenology dictates the impact of climate change on geographic distributions of six co‐occurring North American grasshoppers

Throughout the last century, climate change has altered the geographic distributions of many species. Insects, in particular, vary in their ability to track changing climates, and it is likely that phenology is an important determinant of how well insects can either expand or shift their geographic distributions in response to climate change. Grasshoppers are an ideal group to test the hypothesis that phenology correlates with range expansion, given that co‐occurring confamilial, and even congeneric, species can differ in phenology. Here, I tested the hypothesis that early‐ and late‐season species should possess different range expansion potentials, as estimated by habitat suitability from ecological niche models. I used nine different modeling techniques to estimate habitat suitability of six grasshopper species of varying phenology under two climate scenarios for the year 2050. My results suggest that, of the six species examined here, early‐season species were more sensitive to climate change than late‐season species. The three early‐season species examined here might shift northward during the spring, while the modeled geographic distributions of the three late‐season species were generally constant under climate change, likely because they were pre‐adapted to hot and dry conditions. Phenology might therefore be a good predictor of how insect distributions might change in the future, but this hypothesis remains to be tested at a broader scale.     

Sexual selection,feather wear,and time constraints on the pre‐basic molt explain the acquisition of the pre‐alternate molt in European passerines

Avian feathers need to be replaced periodically to fulfill their functions, with natural, social, and sexual selection presumably driving the evolution of molting strategies. In temperate birds, a common pattern is to molt feathers immediately after the breeding season, the pre‐basic molt. However, some species undergo another molt in winter‐spring, the pre‐alternate molt. Using a sample of 188 European passerine species, Bayesian phylogenetic mixed models, and correlated evolution analyses, we tested whether the occurrence of the pre‐alternate molt was positively associated with proxies for sexual selection (sexual selection hypothesis) and nonsexual social selection (social selection hypothesis), and with factors related to feather wear (feather wear hypothesis) and time constraints on the pre‐basic molt (time constraints hypothesis). We found that the pre‐alternate molt was more frequent in migratory and less gregarious species inhabiting open/xeric habitats and feeding on the wing, and marginally more frequent in species with strong sexual selection and those showing a winter territorial behavior. Moreover, an increase in migratory behavior and sexual selection intensity preceded the acquisition of the pre‐alternate molt. These results provide support for the feather wear hypothesis, partial support for the sexual selection and time constraints hypotheses, and no support for the social selection hypothesis.     

Evidence for Frozen-Niche Variation in a Cosmopolitan Parthenogenetic Soil Mite Species (Acari,Oribatida)

Parthenogenetic lineages may colonize marginal areas of the range of related sexual species or coexist with sexual species in the same habitat. Frozen-Niche-Variation and General-Purpose-Genotype are two hypotheses suggesting that competition and interclonal selection result in parthenogenetic populations being either genetically diverse or rather homogeneous. The cosmopolitan parthenogenetic oribatid mite Oppiella nova has a broad ecological phenotype and is omnipresent in a variety of habitats. Morphological variation in body size is prominent in this species and suggests adaptation to distinct environmental conditions. We investigated genetic variance and body size of five independent forest - grassland ecotones. Forests and grasslands were inhabited by distinct genetic lineages with transitional habitats being colonized by both genetic lineages from forest and grassland. Notably, individuals of grasslands were significantly larger than individuals in forests. These differences indicate the presence of specialized genetic lineages specifically adapted to either forests or grasslands which coexist in transitional habitats. Molecular clock estimates suggest that forest and grassland lineages separated 16-6 million years ago, indicating long-term persistence of these lineages in their respective habitat. Long-term persistence, and morphological and genetic divergence imply that drift and environmental factors result in the evolution of distinct parthenogenetic lineages resembling evolution in sexual species. This suggests that parthenogenetic reproduction is not an evolutionary dead end.     

The trophic structure of bark-living oribatid mite communities analysed with stable isotopes (15N, 13C) indicates strong niche differentiation

The aim of the present study was to identify food sources of bark-living oribatid mites to investigate if trophic niche differentiation contributes to the diversity of bark living Oribatida. We measured the natural variation in stable isotope ratios (¹⁵N/¹⁴N, ¹³C/¹²C) in oribatid mites from the bark of oak (Quercus robur), beech (Fagus sylvatica), spruce (Picea abies) and pine (Pinus sylvestris) trees and their potential food sources, i.e., the covering vegetation of the bark (bryophytes, lichens, algae, fungi). As a baseline for calibration the stable isotope signatures of the bark of the four tree species were measured and set to zero. Oribatid mite stable isotope ratios spanned over a range of about 13 δ units for ¹⁵N and about 7 δ units for ¹³C suggesting that they span over about three trophic levels. Different stable isotope signatures indicate that bark living oribatid mites feed on different food sources, i.e., occupy distinct trophic niches. After calibration stable isotope signatures of respective oribatid mite species of the four tree species were similar indicating close association of oribatid mites with the corticolous cover as food source. Overall, the results support the hypothesis that trophic niche differentiation of bark living oribatid mites contributes to the high diversity of the group.