“Jack‐of‐all‐trades” is parthenogenetic

Sex is evolutionarily more costly than parthenogenesis, evolutionary ecologists therefore wonder why sex is much more frequent than parthenogenesis in the majority of animal lineages. Intriguingly, parthenogenetic individuals and species are as common as or even more common than sexuals in some major and putative ancient animal lineages such as oribatid mites and rotifers. Here, we analyzed oribatid mites (Acari: Oribatida) as a model group because these mites are ancient (early Paleozoic), widely distributed around the globe, and include a high number of parthenogenetic species, which often co‐exist with sexual oribatid mite species. There is evidence that the reproductive mode is phylogenetically conserved in oribatid mites, which makes them an ideal model to test hypotheses on the relationship between reproductive mode and species'' ecological strategies. We used oribatid mites to test the frozen niche variation hypothesis; we hypothesized that parthenogenetic oribatid mites occupy narrow specialized ecological niches. We used the geographic range of species as a proxy for specialization as specialized species typically do have narrower geographic ranges than generalistic species. After correcting for phylogenetic signal in reproductive mode and demonstrating that geographic range size has no phylogenetic signal, we found that parthenogenetic lineages have a higher probability to have broader geographic ranges than sexual species arguing against the frozen niche variation hypothesis. Rather, the results suggest that parthenogenetic oribatid mite species are more generalistic than sexual species supporting the general‐purpose genotype hypothesis. The reason why parthenogenetic oribatid mite species are generalists with wide geographic range sizes might be that they are of ancient origin reflecting that they adapted to varying environmental conditions during evolutionary history. Overall, our findings indicate that parthenogenetic oribatid mite species possess a widely adapted general‐purpose genotype and therefore might be viewed as “Jack‐of‐all‐trades.”     

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.     

The phylogenetic relationship between Astigmata and Oribatida (Acari) as indicated by molecular markers

Astigmata comprise a diverse group of acariform mite species with a remarkable range of life histories, most of which involve parasitic or commensal relationships with other organisms. Several authors have suggested that Astigmata evolved as a paedomorphic clade from within Oribatida, and both morphology and gland-chemistry strongly suggest that their sister-clade is within the oribatid subgroup Desmonomata. The biologies of these groups contrast greatly, since oribatid mites are mostly soil-living detritivores and fungivores, and have life cycles that are much longer than those in Astigmata. We tested the hypothesis that Astigmata evolved from within Desmonomata using two molecular markers, the ribosomal 18S region (18S) and the nuclear elongation factor 1 alpha (ef1α) gene. Representative acariform mites included 28 species of Oribatida, eight of Astigmata, two of Prostigmata and two of Endeostigmata; outgroups included members of Opilioacariformes, Parasitiformes and Ricinulei. To minimize the possibility of long-branch attraction artifacts, we limited highly variable sites by removing gaps (18S) and third codon positions (ef1α) from the sequences. Maximum parsimony, neighbor-joining and Bayesian algorithms formed trees that consistently placed Astigmata outside monophyletic Oribatida, usually as sister-group of the endeostigmatid mite Alicorhagia sp. Analyses with and without outgroups resulted in similar topologies, showing no evidence for long-branch artifacts and leaving the conflict with morphological and biochemical data unexplained.     

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.     

Ecdysial cleavage lines of acariform mites (Arachnida, Acari)

Among mites the ancestral ecdysial cleavage line, or line of dehiscence (a), is inferred by outgroup comparison to be prodehiscent: U-shaped, passing around the front of the mite just above the insertions of the appendages, such that the mite ecloses anteriorly. From preserved and living individuals and exuviae, we found prodehiscence (or its slight variations) to be widespread in Acariformes. It appears to be pervasive in endeostigmatic mites, eupodine Prostigmata, and basal taxa in the Oribatida (Enarthronota, Palaeosomata); it is dominant in cleutherengone Prostigmata and is present in at least one anystine family (Caeculidae). Three general modes of dehiscence are considered to be derived within acariform mites. (1) Merodehiscence is a transverse splitting of the dorsal cuticle at or near the juncture of proterosoma and hysterosoma; it evolved separately in thrcc groups of Prostigmata (Tetranychidae, an undefined subgroup within Cheylctidae, and active instars of Parasitengona) and in a genus of Astigmata (Histiogaster). (2) Trarnsdehiscence is a transverse splitting of the dorsal hysterosomal cuticle anterior to the opisthosomal glands; it occurs in middle-derivative oribatid mites (the paraphyletic Desmonomata), and new observations show it to be widespread in Astigmata, lending support to the hypothesis that the latter group evolved from within Desmonomata. (3) Circumdehiscence is a circumferential splitting of the hysterosomal cuticle that may be incomplete anteriorly; it has long been known to characterize the monophyletic oribatid taxon Brachypylina, but it is convergent with a similar dehiscence in an unrelated family, Lohmanniidae. Transdehiscent and circumdehiscent mites eclose posteriorly. Astigmata exhibit the greatest variety of modes of dehiscence, including the three derived modes and a probable reversal to prodehiscence in Algophagidae. Furthermore, heteromorphic deutonymphs (hypopi) may ecdyse differently from other immature instars of the same species.     

Morphological evidence for the evolutionary origin of Astigmata (Acari: Acariformes)

A century ago, Antonio Berlese first discussed the close phylogenetic relationship between the large mite groups Oribatida and Astigmata. Since then, information having phylogenetic value has greatly increased and the paradigms within which we interpret it have changed. Herein I refine the general hypothesis that Astigmata originated within oribatid mites and suggest Malaconothridae as a possible sister group. Among the 14 apomorphies used to support the origin of Astigmata within oribatid mites are possession of lateral opisthosomal glands, regression of hysterosomal setal pair f1, paired prelarval denticles, partially internalized chelicerae with incomplete adaxial walls, an atelobasic rutellum, pretarsal condylophores that articulate posteriorly with the tarsus, a dorsally fused palp tibia and tarsus and transdehiscent ecdysis. A further 13 apomorphies support the origin of Astigmata at some level within Malaconothroidea. These include absence of an oblique labiogenal articulation, presence of a distal rutellar lamella, shortening of the palp tarsus, larval regression of hysterosomal seta f2, loss of the bothridial seta in all instars, and several losses and modifications of leg setae. The hypothesis brings to light evolutionary questions that were previously obscured by incorrect or inappropriate classifications. The nomenclatural problems that arise from it are best solved by considering Astigmata as a subgroup within Oribatida.     

亚热带不同树种组成森林中土壤甲螨群落结构特征: 以江西新岗山为例

在全球环境变化的大背景下, 生物多样性丧失日益加剧。土壤动物作为生物多样性重要组成之一, 受到广泛的关注。位于我国江西省新岗山的亚热带森林生物多样性与生态系统功能实验样地(BEF-China)是全世界25个森林生物多样性控制实验样地之一。本研究自2019年9月至2022年4月在BEF-China两个不同树种组成的样地(A样地和B样地)内采样, 共获得甲螨23,704头, 隶属于34科50属61种。本文分析和对比了两个样地内甲螨群落结构的差异, 及其多度、物种丰富度、Shannon多样性指数的季节性差异; 通过Pearson检验探讨了甲螨多度与环境因子的关系。结果表明: 在A、B两个不同树种组成的森林生态系统内, 土壤甲螨群落结构及其季节动态具有显著差异。具体表现在: A样地奥甲螨科、罗甲螨科、若甲螨科和尖棱甲螨科的相对多度高于B样地; B样地菌甲螨科、盖头甲螨科和礼服甲螨科的相对多度高于A样地。A样地中夏季和秋季甲螨多度、物种丰富度和Shannon多样性指数显著低于春季和冬季; 而B样地中秋季甲螨多度和物种丰富度与春季差异不显著。Pearson检验结果显示, 凋落物木质素含量与单翼甲螨科和菌甲螨科多度呈负相关关系, 而与奥甲螨科多度呈正相关关系。菌甲螨科多度与土壤和凋落物同一理化因子的相关性基本相同(碳氮比除外), 但与凋落物碳氮比呈正相关关系而与土壤碳氮比呈负相关关系。     

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.     

Multiple convergent evolution of arboreal life in oribatid mites indicates the primacy of ecology

Frequent convergent evolution in phylogenetically unrelated taxa points to the importance of ecological factors during evolution, whereas convergent evolution in closely related taxa indicates the importance of favourable pre-existing characters (pre-adaptations). We investigated the transitions to arboreal life in oribatid mites (Oribatida, Acari), a group of mostly soil-living arthropods. We evaluated which general force—ecological factors, historical constraints or chance—was dominant in the evolution of arboreal life in oribatid mites. A phylogenetic study of 51 oribatid mite species and four outgroup taxa, using the ribosomal 18S rDNA region, indicates that arboreal life evolved at least 15 times independently. Arboreal oribatid mite species are not randomly distributed in the phylogenetic tree, but are concentrated among strongly sclerotized, sexual and evolutionary younger taxa. They convergently evolved a capitate sensillus, an anemoreceptor that either precludes overstimulation in the exposed bark habitat or functions as a gravity receptor. Sexual reproduction and strong sclerotization were important pre-adaptations for colonizing the bark of trees that facilitated the exploitation of living resources (e.g. lichens) and served as predator defence, respectively. Overall, our results indicate that ecological factors are most important for the observed pattern of convergent evolution of arboreal life in oribatid mites, supporting an adaptationist view of evolution.     

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.     

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.     

Sperm structure and phylogeny of Astigmata

The Astigmata, a large and variable group, is still a subject of taxonomic dispute. Particularly, their origin from ancestors of the lower oribatid mites (e.g., Malaconothroidea) seems well documented by many lines of evidence. The structure of spermatozoa has been successfully applied to phylogenetic investigations in many animal groups. The aim of our study was to provide new data on spermatozoon structure in Astigmata and to consider its appropriateness in phylogenetic studies. The study reveals information on spermatozoa in 17 species of Astigmata (11 species studied for the first time) extending our knowledge to 18 species (one species known only from the literature) representing 12 families and 7 superfamilies. Spermatozoa have the same basic structure in all species: cells are multiform and the chromatin forms thin threads embedded directly in the cytoplasm; the acrosome is absent. The cytoplasm in most species contains electron-dense lamellae, varying in both number and arrangement within the cell. In Sarcoptoidea, electron-dense tubules in contact with lamellae margins were also observed in Psoroptidae (Psoroptes equi), whereas in two representatives of Sarcoptidae (Notoedres cati and Sarcoptes scabiei), only electron-dense tubules were found. In two species, Canestrinia sellnicki (Canestrinioidea: Canestriniidae) and Scutulanyssus obscurus (Analgoidea: Pteronyssidae), neither lamellae nor tubules were present. The mitochondria in a spermatozoon are usually gathered at the cell periphery and their structure is usually modified to form so-called mitochondrial derivatives. The chromatin threads are an autapomorphy strongly supporting the monophyly of Astigmata. As spermatozoa vary considerably between species in Astigmata, we deduce that sperm structure may be useful for phylogenetic analyses within the group. Several conclusions concerning the affinities within Astigmata are presented. Spermatology seems to be unhelpful, however, in questions on the origin of Astigmata (particularly for Astigmata-Oribatida relationships), since their sperm do not possess synapomorphies with sperm of the remaining groups of Acariformes, i.e., Endeostigmata, Prostigmata, and Oribatida.     

Tree hollows harbour a specialised oribatid mite fauna

Tree hollows are known to harbour a species-rich and specialized beetle fauna, while other invertebrates, such as for instance mites, have been much less studied. The importance of hollows in oak trees (Quercus robur) for local oribatid mite diversity was studied at three sites in south eastern Sweden. The qualitative and quantitative composition of the oribatid mite fauna was studied in hollows of fifteen 240–420 years old oak trees and compared to that in the surrounding soil. A total of 5,530 specimen of adult oribatid mites were determined belonging to 63 taxa. Taxonomic composition and community structure of the tree hollow communities differed markedly from the soil communities. The most dominant and frequent component of the tree hollow communities comprised a Carabodes species new to Sweden that accounted on average for 44 % of all Oribatida. This species, that closely resembles both Carabodes oenipontanus and Carabodes granulatus, was specific to the tree hollows. Dominance patterns in the soil communities were more even, with the most common taxa also occurring in the tree hollows but exhibiting a significant preference for the soil. Overall, there was little taxonomic overlap between the communities, suggesting that tree hollows harbour an independent mite community from the soil and therefore significantly contribute to the overall mite diversity on the landscape level. The present study therefore strongly supports the use of hollow trees as biodiversity indicators and also their conservation, which will preserve specialised invertebrate communities, including mites.     

Efficiencies of different genes and different tree-building methods in recovering a known vertebrate phylogeny

The relative efficiencies of different protein-coding genes of the mitochondrial genome and different tree-building methods in recovering a known vertebrate phylogeny (two whale species, cow, rat, mouse, opossum, chicken, frog, and three bony fish species) was evaluated. The tree-building methods examined were the neighbor joining (NJ), minimum evolution (ME), maximum parsimony (MP), and maximum likelihood (ML), and both nucleotide sequences and deduced amino acid sequences were analyzed. Generally speaking, amino acid sequences were better than nucleotide sequences in obtaining the true tree (topology) or trees close to the true tree. However, when only first and second codon positions data were used, nucleotide sequences produced reasonably good trees. Among the 13 genes examined, Nd5 produced the true tree in all tree-building methods or algorithms for both amino acid and nucleotide sequence data. Genes Cytb and Nd4 also produced the correct tree in most tree-building algorithms when amino acid sequence data were used. By contrast, Co2, Nd1, and Nd41 showed a poor performance. In general, large genes produced better results, and when the entire set of genes was used, all tree-building methods generated the true tree. In each tree-building method, several distance measures or algorithms were used, but all these distance measures or algorithms produced essentially the same results. The ME method, in which many different topologies are examined, was no better than the NJ method, which generates a single final tree. Similarly, an ML method, in which many topologies are examined, was no better than the ML star decomposition algorithm that generates a single final tree. In ML the best substitution model chosen by using the Akaike information criterion produced no better results than simpler substitution models. These results question the utility of the currently used optimization principles in phylogenetic construction. Relatively simple methods such as the NJ and ML star decomposition algorithms seem to produce as good results as those obtained by more sophisticated methods. The efficiencies of the NJ, ME, MP, and ML methods in obtaining the correct tree were nearly the same when amino acid sequence data were used. The most important factor in constructing reliable phylogenetic trees seems to be the number of amino acids or nucleotides used.      

分子系统进化关系分析的一种新方法--贝叶斯法在硬蜱属中的应用

距离矩阵邻接法、最大简约法和最大似然法是重建生物系统关系的3种主要方法。普遍认为最大似然法在原理上优于前二种方法，但其计算复杂费时。由于现行计算机的能力尚达不到其要求而实用性差，特别是在处理大数据集样本(即大于25个分类单元)时，用此方法几乎不可能。新近提出的贝叶斯法(Bayesianmethod)既保留了最大似然法的基本原理，又引进了马尔科夫链的蒙特卡洛方法，并使计算时间大大缩短。本文用贝叶斯法对硬蜱属(Ixodes)19个种的线粒体16S rDNA片段进行了系统进化分析。从总体上看，分析结果与现有的基于形态学的分类体系基本吻合。但与现存的假说相反，莱姆病的主要宿主蓖籽硬蜱复合种组并非单系。通过比较贝叶斯法与其它三种方法的结果，我们认为贝叶斯法是一种系统进化分析的好方法，它既能根据分子进化的现有理论和各种模型用概率重建系统进化关系，又克服了最大似然法计算速度慢、不适用于大数据集样本的缺陷。贝叶斯法根据后验概率直观地表示系统进化关系的分析结果，不需要用自引导法进行检验。可以预料，贝叶斯法将会被广泛地应用到系统进化分析上[动物学报49(3)：380—388，2003]。     

Comparative Genomics Reveals Insights into the Divergent Evolution of Astigmatic Mites and Household Pest Adaptations

Highly diversified astigmatic mites comprise many medically important human household pests such as house dust mites causing ∼1–2% of all allergic diseases globally; however, their evolutionary origin and diverse lifestyles including reversible parasitism have not been illustrated at the genomic level, which hampers allergy prevention and our exploration of these household pests. Using six high-quality assembled and annotated genomes, this study not only refuted the monophyly of mites and ticks, but also thoroughly explored the divergence of Acariformes and the diversification of astigmatic mites. In monophyletic Acariformes, Prostigmata known as notorious plant pests first evolved, and then rapidly evolving Astigmata diverged from soil oribatid mites. Within astigmatic mites, a wide range of gene families rapidly expanded via tandem gene duplications, including ionotropic glutamate receptors, triacylglycerol lipases, serine proteases and UDP glucuronosyltransferases. Gene diversification after tandem duplications provides many genetic resources for adaptation to sensing environmental signals, digestion, and detoxification in rapidly changing household environments. Many gene decay events only occurred in the skin-burrowing parasitic mite Sarcoptes scabiei. Throughout the evolution of Acariformes, massive horizontal gene transfer events occurred in gene families such as UDP glucuronosyltransferases and several important fungal cell wall lytic enzymes, which enable detoxification and digestive functions and provide perfect drug targets for pest control. This comparative study sheds light on the divergent evolution and quick adaptation to human household environments of astigmatic mites and provides insights into the genetic adaptations and even control of human household pests.     

基于Cyt b基因探讨羚羊亚科原羚属系统发生关系

原羚属物种在羚羊亚科中的分类地位尚存在很多争议。本文测定了原羚属的黄羊和藏原羚细胞色素b基因全序列(1140bp),并与牛科其它属31个种的同源序列进行比较,对其碱基组成变异情况及核苷酸序列差异进行了分析。基于细胞色素b基因全序列,用简约法(MP)、邻接法(NJ)和似然法(ML)构建了系统进化树。结果表明：黄羊和藏原羚的序列差异为3．78％,颠换数目近乎为0,其突变远未饱和;原羚属内黄羊和藏原羚为不同种,单系发生;原羚属与赛加羚羊属、犬羚属及跳羚属等并系发生,原羚属隶属于羚羊亚科,应为独立属;羚羊亚科组成属间多为并系起源。根据序列差异值2％／百万年的细胞色素6分子钟,推测黄羊和藏原羚分歧时间大约为1～2百万年;原羚属与羚羊亚科其它属分歧时间大约在5．7～8百万年。     

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.     

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.     

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.