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.     

Taxonomic distribution of defensive alkaloids in Nearctic oribatid mites (Acari,Oribatida)

The opisthonotal (oil) glands of oribatid mites are the source of a wide diversity of taxon-specific defensive chemicals, and are likely the location for the more than 90 alkaloids recently identified in oribatids. Although originally recognized in temperate oribatid species, alkaloids have also been detected in related lineages of tropical oribatids. Many of these alkaloids are also present in a worldwide radiation of poison frogs, which are known to sequester these defensive chemicals from dietary arthropods, including oribatid mites. To date, most alkaloid records involve members of the superfamily Oripodoidea (Brachypylina), although few species have been examined and sampling of other taxonomic groups has been highly limited. Herein, we examined adults of more than 60 species of Nearctic oribatid mites, representing 46 genera and 33 families, for the presence of alkaloids. GC–MS analyses of whole body extracts led to the detection of 15 alkaloids, but collectively they occur only in members of the genera Scheloribates (Scheloribatidae) and Protokalumma (Parakalummidae). Most of these alkaloids have also been detected previously in the skin of poison frogs. All examined members of the oripodoid families Haplozetidae and Oribatulidae were alkaloid-free, and no mites outside the Oripodoidea contained alkaloids. Including previous studies, all sampled species of the cosmopolitan oripodoid families Scheloribatidae and Parakalummidae, and the related, mostly tropical families Mochlozetidae and Drymobatidae contain alkaloids. Our findings are consistent with a generalization that alkaloid presence is widespread, but not universal in Oripodoidea. Alkaloid presence in tropical, but not temperate members of some non-oripodoid taxa (in particular Galumnidae) deserves further study.     

Expanding the ‘enemy-free space’ for oribatid mites: evidence for chemical defense of juvenile Archegozetes longisetosus against the rove beetle Stenus juno

Adult oribatid mites are thought to live functionally in ‘enemy-free space’ due to numerous morphological and chemical defensive strategies. Most juvenile oribatid mites, however, lack hardened cuticles and are thus thought to be under stronger predation pressure. On the other hand, the majority of oribatids have exocrine oil glands in all developmental stages, possibly rendering chemical defense the crucial survival strategy in juvenile Oribatida. We manipulated tritonymphs of the model oribatid mite Archegozetes longisetosus to completely discharge their oil glands and offered these chemically disarmed specimens to the polyphagous rove beetle Stenus juno. Disarmed specimens were easily consumed. By contrast, specimens with filled oil glands were significantly protected, being rejected by the beetles. This is the first direct evidence that oil gland secretions provide soft-bodied juvenile oribatids with chemical protection against large arthropod predators.     

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.     

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.     

Oribatid Mites as Potential Vectors for Soil Microfungi: Study of Mite-Associated Fungal Species

The ability of soil-living oribatid mites to disperse fungal propagules on their bodies was investigated. Classical plating methods were applied to cultivate these fungi and to study their morphology. Molecular markers were used for further determination. The nuclear ribosomal large subunit and the nuclear ribosomal internal transcribed spacer of DNA extracts of the cultured fungi as well as total DNA extracts of the mites themselves, also containing fungal DNA, were amplified and sequenced. Based on phylogenetic analysis, a total of 31 fungal species from major fungal groups were found to be associated with oribatid mites, indicating that mites do not selectively disperse specific species or species groups. The detected taxa were mainly saprobiontic, cosmopolitan (e.g., Alternaria tenuissima), but also parasitic fungi (Beauveria bassiana) for whose dispersal oribatid mites might play an important role. In contrast, no mycorrhizal fungi were detected in association with oribatid mites, indicating that their propagules are dispersed in a different way. In addition, fungi that are known to be a preferred food for oribatid mites such as the Dematiacea were not detected in high numbers. Results of this study point to the potential of oribatid mites to disperse fungal taxa in soil and indicate that co-evolutionary patterns between oribatid mites and their associated fungi might be rare or even missing in most cases, since we only detected ubiquitous taxa attached to the mites.     

BEHAVIORAL CONVERGENCE AND ADAPTIVE RADIATION: EFFECTS OF HABITAT USE ON TERRITORIAL BEHAVIOR IN ANOLIS LIZARDS

Most studies of adaptive radiations focus on morphological aspects of differentiation, yet behavior is also an important component of evolutionary diversification, often mediating the relationship between animal ecology and morphology. In species within radiations that are convergent in ecology and morphology, we then also expect convergence in behavior. Here, we examined 13 Anolis lizard species to determine whether territorial strategies have evolved convergently with morphology and habitat use. We evaluated two aspects of territoriality: behavioral defense of space via territorial displays, and territory overlap within and between sexes. Controlling for the phylogenetic relationships of the taxa in our study, we found that species similar in perch height and diameter convergently evolved patterns of territory overlap, whereas species similar in habitat visibility (the proportion of space that can be seen from a perch) convergently evolved display behavior. We also found that species with greater display time have more extensive male–male territory overlap. This study provides strong evidence for the role of habitat in the evolution of territoriality and suggests that the social structure of a species ultimately evolves in concert with habitat use and morphology.     

Non‐trophic effects of oribatid mites on cord‐forming basidiomycetes in soil microcosms

1. Saprotrophic cord‐forming basidiomycetes are the primary agents of decomposition in forest ecosystems. Collembola and oribatid mites affect fungal growth and foraging, and therefore decomposition, through direct mycelial grazing. 2. Grazing on the fungal species Hypholoma fasciculare, Resinicium bicolor and Phanerochaete velutina by the collembola Folsomia candida, and the oribatid mites Steganacarus magnus, Euzetes globulus and Hermannia gibba was investigated in soil microcosms. 3. Folsomia candida grazed on all fungal species: radial extent of R. bicolor, hyphal coverage of all fungal species, and fractal dimension of R. bicolor and P. velutina were all reduced. Oribatid mites did not graze the fungi but did affect mycelial morphology. Steganacarus magnus caused a reduction in the radial extent of H. fasciculare, and the hyphal coverage and fractal dimension in both H. fasciculare and R. bicolor. Euzetes globulus and H. gibba reduced the hyphal coverage of P. velutina. 4. Oribatid mites are associated with a cornucopia of chemical secretions with possible anti‐fungal properties. Chemical analysis of H. gibba opisthonotal secretions revealed four main compounds, all of which are new to the known spectrum of opisthonotal components. The most abundant was (E)‐β‐farnesene. 5. Treatment effects were species‐specific in terms of both fungal and invertebrate species. This study provides the first evidence of non‐grazing effects of oribatid mites on fungal growth and morphology. This could potentially influence the spatial organisation of mycelium in forest soils and therefore the ability of fungi to locate, colonise and decompose dead organic matter.     

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.     

Inducible Defenses with a "Twist": Daphnia barbata Abandons Bilateral Symmetry in Response to an Ancient Predator

Predation is one of the most important drivers of natural selection. In consequence a huge variety of anti-predator defenses have evolved in prey species. Under unpredictable and temporally variable predation pressure, the evolution of phenotypically plastic defensive traits is favored. These “inducible defenses”, range from changes in behavior, life history, physiology to morphology and can be found in almost all taxa from bacteria to vertebrates. An important group of model organisms in ecological, evolutionary and environmental research, water fleas of the genus Daphnia (Crustacea: Cladocera), are well known for their ability to respond to predators with an enormous variety of inducible morphological defenses. Here we report on the “twist”, a body torsion, as a so far unrecognized inducible morphological defense in Daphnia, expressed by Daphnia barbata exposed to the predatory tadpole shrimp Triops cancriformis. This defense is realized by a twisted carapace with the helmet and the tail spine deviating from the body axis into opposing directions, resulting in a complete abolishment of bilateral symmetry. The twisted morphotype should considerably interfere with the feeding apparatus of the predator, contributing to the effectiveness of the array of defensive traits in D. barbata. As such this study does not only describe a completely novel inducible defense in the genus Daphnia but also presents the first report of a free living Bilateria to flexibly respond to predation risk by abandoning bilateral symmetry.     

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。     

小麦-玉米轮作农田土壤螨多样性空间分布格局

以农田土壤动物长期监测样地为平台, 阐明土壤动物物种和功能多样性空间分布格局, 是揭示农田土壤动物多样性维持机制、提高农田土壤质量的重要基础。本试验于2020年10月, 对河南商丘农田土壤动物大型固定样地(9 ha)的210个采样点进行土壤样品野外采集和室内分离, 将土壤螨样品鉴定到种并测量其体长体宽数据, 以说明小麦-玉米轮作农田土壤螨多样性及其体长体宽的空间分布格局。结果表明: (1)共捕获成螨个体17,256头, 其中甲螨亚目为优势类群, 其个体数占总捕获量的94.67%; MGP分析表明样地甲螨群落属于P型, 说明受人为因素影响强烈; 生态位宽度和重叠度分析表明, 进化程度越高甲螨的生态位宽度越宽, 进化程度越相近甲螨之间的竞争越激烈。(2) Moran’s I分析显示, 在20-100 m的空间尺度上, 土壤螨群落、优势种的个体数和体长体宽多为显著正相关; 在220-300 m的空间尺度上, 部分为显著负的空间自相关。半方差函数结果表明, 甲螨群落物种数、个体数和体长体宽的空间变异主要受确定性过程影响, 中气门螨群落的空间变异由确定性和随机性过程共同影响。(3)土壤螨个体数与体长体宽存在显著弱的负相关关系, 这种关系普遍存在于土壤螨各群落与优势种中。本研究建议同时开展物种多样性和以体长体宽为代表的功能多样性空间格局研究, 对揭示土壤螨群落维持机制、保护土壤螨多样性具有重要意义。     

Awesome or ordinary? Global diversity patterns of oribatid mites

Diversity of most above-ground organisms increases with decreasing latitude, but the biogeographical and macroecological diversity patterns of below-ground animals have been poorly studied. We investigated the latitudinal diversity gradient in a primarily below-ground living soil taxon, oribatid mites. Furthermore, oribatid mite species richness from islands and mainlainds was tested for correlation with the size of the respective area (island or mainland) to evaluate if their species–area relationships are similar to those of above-ground taxa. The results suggest that for oribatid mites 1) diversity increases from the boreal to the warm temperate region but not further to the tropics, and 2) species–area relationships for islands and mainlands are similar to those of above-ground taxa, but this is mainly caused by very small islands, such as Cocos islands, and very large islands, such as Madagascar. When these islands are excluded the species–area relationship strongly differs from those of typical islands. The results support the view that below-ground animal taxa are generalists that inhabit wide niches. Most small islands have relatively rich oribatid mite faunas, supporting the observation that a large number of species can coexist in a small area (high α-diversity).     

Sequestration of nematocysts by divergent cnidarian predators: mechanism,function, and evolution

Animals have evolved diverse mechanisms to protect themselves from predators. Although such defenses are typically generated endogenously, some species have evolved the ability to acquire defenses by sequestering defensive chemicals or structures from other species. Chemical sequestration is widespread among animals, but the ability to sequester entire structures, such as organelles, appears to be rare. Here, we review information on the sequestration of functional nematocysts, the stinging organelles produced by Cnidaria, by divergent predators. Nematocyst sequestration has evolved multiple times, having been documented in Ctenophora, Acoelomorpha, Platyhelminthes, and Mollusca. For each of these phyla, we review the phylogenetic distribution, mechanisms, and possible functions of nematocyst sequestration. We estimate that nematocyst sequestration has evolved 9–17 times across these four phyla. Although data on the mechanism of sequestration remain limited, similarities across several groups are evident. For example, in multiple groups, nematocysts are transported within cells from the gut to peripheral tissues, and certain types of nematocysts are selectively sequestered over others, suggesting convergent evolution in some aspects of the sequestration process across phyla. Similarly, although the function of nematocyst sequestration has not been well documented, several studies do suggest that the nematocysts sequestered by these groups are effective for defense. We highlight several traits that are common to Ctenophora, Acoelomorpha, Platyhelminthes, and Mollusca and suggest hypotheses for how these traits could have played a role in the evolution of nematocyst sequestration. Finally, we propose a generalized working model for the steps that may lead to the evolution of nematocyst sequestration and discuss important areas for future research.     

“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.”     

Living in the danger zone: Exposure to predators and the evolution of spines and body armor in mammals

Mammals have independently evolved a wide variety of morphological adaptations for use in avoiding death by predation, including spines, quills, dermal plates, and noxious sprays. Although these traits appear to protect their bearer from predatory attack, it is less obvious why some species evolved them and others have not. We investigated the ecological correlates favoring the evolution of specialized defenses in mammals, focusing on conspicuousness to predators due to body size and openness of habitat. We scored species for the degree to which they are protected by spines, quills, dermal plating, and sprays and used phylogenetic comparative analyses to study the morphological and ecological factors that may favor their evolution. We show that medium‐sized insectivorous mammals (～800 g to 9 kg) that live in open habitats are more likely to possess one of these defensive traits to reduce predation. Smaller species (<200 g) and those in closed habitats can typically rely on crypsis to avoid predators, and larger species (>10 kg) are less susceptible to predation by most small‐ to medium‐sized predators. We discuss how diet, metabolic rate, and defensive strategy evolve in concert to allow species to exploit this ecomorphological “danger zone” niche.     

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.     

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.     

Global diversity of oribatids (Oribatida: Acari: Arachnida)

Oribatid mites are primarily terrestrial. Only about 90 species (less than 1% of all known oribatid species) from 10 genera are truly aquatic, with reproduction and all stages of their life cycle living in freshwater. Adaptation to aquatic conditions evolved independently in different taxa. However, many terrestrial species can also be found in aquatic habitats, either as chance stragglers from the surrounding habitats, or from periodic or unpredictable floodings, where they can survive for long periods. In spite of their low species richness aquatic oribatids can be very abundant in different freshwater habitats as in lentic (pools, lakes, water-filled microhabitats) or flowing waters (springs, rivers, streams), mainly on submerged plants. The heavily sclerotized exoskeletons of several species enables subfossil or fossil preservation in lakes or bog sediments. Guest editors: E. V. Balian, C. Lévêque, H. Segers & K. Martens Freshwater Animal Diversity Assessment     

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.