Hunger for sex: Abundant, heterogeneous resources select for sexual reproduction in the field.

Major hypotheses on sex evolution predict that resource abundance and heterogeneity should either select for or against sexual reproduction. However, seldom have these predictions been explicitly tested in the field. Here, we investigated this question using soil oribatid mites, a diverse and abundant group of soil arthropods whose local communities can be dominated by either sexual or asexual species. First, we refined theoretical predictions by addressing how the effects of resource abundance, heterogeneity and abiotic conditions could modify each other. Then, we estimated the strength of selection for sexual species in local communities while controlling for phylogeny and neutral processes (ecological drift and dispersal), and tested its relation to resource and abiotic gradients. We show that sexual species tended to be favoured with increasing litter amount, a measure of basal resource abundance. Further, there was some evidence that this response occurred mainly under higher tree species richness, a measure of basal resource heterogeneity. This response to resources is unlikely to reflect niche partitioning between reproductive modes, as sexual and asexual species overlapped in trophic niche according to a comparative analysis using literature data on stable isotope ratios. Rather, these findings are consistent with the hypothesis that sex facilitates adaptation by breaking unfavourable genetic associations, an advantage that should increase with effective population size when many loci are under selection and, thus, with resource abundance.

Modelling selection, drift, dispersal and their interactions in the community assembly of Amazonian soil mites.

Three processes can explain contemporary community assembly: natural selection, ecological drift and dispersal. However, quantifying their effects has been complicated by confusion between different processes and neglect of expected interactions among them. One possible solution is to simultaneously model the expected effects of each process within species, across communities and across species, thus providing more integrative tests of ecological theory. Here, we used generalized linear mixed models to assess the effects of selection, drift and dispersal on the occurrence probability of 135 soil oribatid mite species across 55 sites over an Amazonian rainforest landscape (64 km2). We tested for interactions between process-related factors and partitioned the explained variation among them. We found that occurrence probability (1) responded to soil P content and litter mass depending on body size and reproductive mode (sexual or parthenogenetic), respectively (selection); (2) increased with community size (drift); and (3) decreased with distance to the nearest source population, and more so in rare species (dispersal limitation). Processes did not interact significantly, and our best model explained 67% of the overall variation in species occurrence probability. However, most of the variation was attributable to dispersal limitation (55%). Our results challenge the seldom-tested theoretical prediction that ecological processes should interact. Rather, they suggest that dispersal limitation overrides the signatures of drift and selection at the landscape level, thus rendering soil microarthropod species ecologically equivalent and possibly contributing to the maintenance of metacommunity diversity.

How land-use intensity affects sexual and parthenogenetic oribatid mites in temperate forests and grasslands in Germany.

Intensive land use has been shown to alter the composition and functioning of soil communities. Due to their low dispersal ability, oribatid mites are particularly vulnerable to land-use intensification and species which are not adjusted to management-related disturbances become less abundant. We investigated how different land-use parameters in forests and grasslands affect oribatid mite diversity and abundance, with a focus on: (1) species-level impacts, by classifying species as increasing ('winners') or decreasing ('losers') in abundance with higher land-use intensity, and (2) reproductive impact, by investigating whether sexual and parthenogenetic species react differently. We collected 32,542 adult oribatid mites in 60 forests and grasslands of known land-use intensity in two regions of Germany. Diversity and total abundance as well as the proportion of sexual species were higher in forests than in grasslands. Diversity declined with higher land-use intensity in forests, but increased with higher mowing and fertilization in grasslands. Depending on land-use parameter and region, abundance either declined or remained unaffected by increasing intensity. Gravidity was higher in sexual than in parthenogenetic species and sexuals had 1.6× more eggs per gravid female. Proportions of sexual species and gravid females decreased with land-use intensity in forests, but increased with mowing in grasslands. At the species level, 75% of sexuals and 87.5% of parthenogens were 'losers' of higher percentages of dead wood originating from management-related disturbances. Across land-use parameters and habitats, a similar proportion of sexual and parthenogenetic oribatid mite species were 'losers' of high land-use intensity. However, 'winner' species were more common among sexuals.

Storage and release of hydrogen cyanide in a chelicerate (Oribatula tibialis).

Cyanogenesis denotes a chemical defensive strategy where hydrogen cyanide (HCN, hydrocyanic or prussic acid) is produced, stored, and released toward an attacking enemy. The high toxicity and volatility of HCN requires both chemical stabilization for storage and prevention of accidental self-poisoning. The few known cyanogenic animals are exclusively mandibulate arthropods (certain myriapods and insects) that store HCN as cyanogenic glycosides, lipids, or cyanohydrins. Here, we show that cyanogenesis has also evolved in the speciose Chelicerata. The oribatid mite Oribatula tibialis uses the cyanogenic aromatic ester mandelonitrile hexanoate (MNH) for HCN storage, which degrades via two different pathways, both of which release HCN. MNH is emitted from exocrine opisthonotal oil glands, which are potent organs for chemical defense in most oribatid mites.

A tropical arthropod unravels local and global environmental dependence of seasonal temperature-size response.

In most ectotherms, adult body size decreases with warming, the so-called 'temperature-size rule' (TSR). However, the extent to which the strength of the TSR varies naturally within species is little known, and the significance of this phenomenon for tropical biota has been largely neglected. Here, we show that the adult body mass of the soil mite Rostrozetes ovulum declined as maximum temperature increased over seasons in a central Amazonian rainforest. Further, per cent decline per °C was fourfold higher in riparian than in upland forests, possibly reflecting differences in oxygen and/or resource supply. Adding our results to a global dataset revealed that, across terrestrial arthropods, the seasonal TSR is generally stronger in hotter environments. Our study suggests that size thermal dependence varies predictably with the environment both locally and globally.

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.

Catalogue of juvenile instars of oribatid mites (Acari: Oribatida)the next decade (20142023).

In its traditional, paraphyletic context (sans Astigmata), oribatid mites comprise more than 11,000 known species (Subas 2022). They are largely fungivores and decomposers in organic horizons of soil and have a plesiotypic ontogeny that includes four active, free-living juvenile instars. In a taxonomically organized 2014 catalogue, we summarized literature resources concerning the 805 species for which ontogenetic data (mainly morphological) were available. Herein, we supplement that catalogue with all data known to us that were published during the intervening decade. These relate to 267 species, of which 165 were described prior to 2014. As in the 2014 catalogue, representation is strongest among: the middle-derivative hyporder Nothrina; brachypyline superfamilies that are affiliated with aquatic, semiaquatic or intertidal environments (Limnozetoidea, Ameronothroidea); some eupheredermous groups (Plateremaeoidea, Damaeoidea); and Ceratozetoidea. Also as in 2014, groups that are underreprented, based on their high known diversity, are the ptyctimous Mixonomata (Euphthiracaroidea, Phthiracaroidea) and the brachypyline superfamilies Oppioidea and Oripodoidea.

Psammochthoniidae n. fam., a paedomorphic family of oribatid mites (Oribatida: Enarthronota) from sandy soil in Thailand, Brazil and the U.S.A.

A new genus and species of enarthronote oribatid mite, Psammochthonius kethleyi n.g., n. sp., is described and illustrated based on adult and immature specimens collected from coastal sandy soil in Thailand (Phangnga), Brazil (SAo Paulo) and the U.S.A. (Mississippi). Analysis shows that it is a member of Hypochthonioidea, but not of any named family, so the monobasic family Psammochthoniidae n. fain. is proposed. Traits that are unique among hypochthonioid mites include an adult body length under 250 microm, a functionally trichoid body form (postpedal flexing), strong lateral displacement of setae in row e, which insert on a unique form of transverse scissure, subcapitular stenarthry, leg IV vestiges and a possible precocious genital swelling in the larva, apparent absence of an anal segment, and a highly regressed leg setation. The latter two, and some other traits that were previously unknown in Hypochthonioidea, suggest that Psammochthoniidae represents the first clearly paedomorphic lineage in this diverse superfamily. Like all other known hypochthonioid mites, P. kethleyi appears to be thelytokous.

Convergent evolution of defense mechanisms in oribatid mites (Acari, Oribatida) shows no "ghosts of predation past".

Oribatid mites are diverse and abundant terrestrial soil arthropods that are involved in decomposition of organic matter and nutrient cycling. As indicated by fossils starting from the Devonian, they evolved varied mechanisms and structures for defense from predators. We investigated four of these defensive structures (ptychoid body, hologastry, mineralization and opisthonotal glands) and used ancestral character state reconstruction to determine whether they evolved convergently and how many times this may have happened. Phylogenetic trees based on 18S rDNA were constructed for 42 oribatid mite species and two outgroup taxa using likelihood and Bayesian algorithms. The results suggest that at least three of the four defensive structures evolved convergently several times; for opisthonotal glands convergent evolution remains equivocal. This high level of convergence indicates that predation has been an important factor throughout the evolution of oribatid mites, contributing to morphological diversity and potentially also to species richness, as there are indications that some taxa radiated after the evolution of defense structures. Despite the ancientness of oribatid mites, defense structures seems to have been rarely lost, suggesting that they still are functional and necessary to reduce predation, rather than being 'ghosts of predation past'.

Tasty but protected--first evidence of chemical defense in oribatid mites.

Oribatid mites (Acari, Oribatida) represent one of the most abundant and speciose groups of microarthropods in the decomposer food webs of soils, but little is known of their top-down regulation by predators. Oribatids are relatively long-lived and have numerous morphological defensive adaptations, and so have been proposed to live in 'enemy-free space'. Most also possess a pair of large exocrine oil glands that produce species-specific mixtures of hydrocarbons, terpenes, aromatics, and alkaloids with presumably allomonal functions, although their adaptive value has never been tested empirically. We developed a protocol that discharges the oil glands of the model oribatid species, Archegozetes longisetosus. and offered 'disarmed' individuals as prey to polyphagous Stenus beetles (Staphylinidae), using untreated mites as controls. Stenus juno fed on disarmed mites with behavioral sequences and success rates similar to those observed when they prey on springtails, a common prey. In contrast, mites from the control group with full glands were almost completely rejected; contact with the gland region elicited a strong reaction and cleaning behavior in the beetle. This is the first evidence of an adaptive value of oribatid mite oil gland secretions for chemical defense. The protocol of discharging oil glands should facilitate future studies on top-down control of oribatid mites that aim to differentiate between morphological and chemical aspects of defensive strategies.

Alkaloids in the mite Scheloribates laevigatus: further alkaloids common to oribatid mites and poison frogs.

Poison frogs are chemically defended from predators by diverse alkaloids, almost all of which are sequestered unchanged from alkaloid-containing arthropods in the frog diet. Oribatid mites recently have been proposed as a major dietary source of poison frog alkaloids. Here, we report on alkaloids common to an oribatid mite and poison frogs. Gas chromatographic-mass spectrometric analysis of methanol extracts of adult Scheloribates laevigatus (Oribatida: Scheloribatidae) revealed nine alkaloids. Five of these have been detected previously in the skin glands of poison frogs: two isomers of the pumiliotoxin 291G, two isomers of the 5,6,8-trisubstituted indolizidine 209C, and the 5,6,8-trisubstituted indolizidine 195G. The other four alkaloids, a pumiliotoxin, a tricyclic (coccinelline-like), and two isomers of an izidine, were not previously known, but are similar in structure to alkaloids found in poison frogs. Alkaloids were not detected in immature S. laevigatus, suggesting that they are adult-specific and possibly the result of mite biosynthesis. Although most of the alkaloids detected in S. laevigatus are common to poison frogs, the geographic distributions of these organisms are not sympatric. The findings of this study indicate that oribatid mites, and in particular, members of the genus Scheloribates, represent a relatively unexplored arthropod repository for alkaloids and a significant dietary source of alkaloids in poison frogs.

Redescriptions of North American Epidamaeus (Acari, Oribatida, Damaeidae) species proposed by N. Banks, H.E. Ewing, A.P. Jacot, and J.W. Wilson.

Early American acarologists proposed several species that have been recombined to the oribatid mite genus Epidamaeus (Damaeidae), but none has been redescribed to modern standards. These include E. puritanicus (Banks, 1906), E. michaeli (Ewing, 1909), E. globifer (Ewing, 1913), E. florida (Wilson, 1936), E. olitor (Jacot, 1937), and E. craigheadi (Jacot, 1939). We redescribe and illustrate these species, based on available type specimens and other material in various collections, and give an indication of their overall distributions. Juvenile instars are at least partly described for all except E. florida and E. olitor. Since these six collectively include the most encountered Epidamaeus species in the eastern USA, a diagnostic key is presented that also includes E. arcticola (Hammer, 1952), which is reliably recorded from eastern North America (New Hampshire) for the first time. A neotype is selected for E. michaeli, which is considered a senior subjective synonym of E. canadensis (Banks, 1909) [new syn.]. Synonymy of E. puritanicus with E. grandjeani (Bulanova-Zachvatkina, 1957) is rejected.

Arthropod colonization of land--linking molecules and fossils in oribatid mites (Acari, Oribatida).

Terrestrial fossils that document the early colonization of land are scarce for >100 my after the Cambrian explosion. This raises the question whether life on land did not exist or just did not fossilize. With a molecular dating technique, we analyzed the origin of terrestrial chelicerate microarthropods (Acari, Oribatida) which have a fossil record since the Middle Devonian that is exceptional among soil animals. Our results suggest that oribatid mites originated in the Precambrian (571+/-37 mya) and that the radiation of basal groups coincides with the gap in the terrestrial fossil record between the Cambrian explosion and the earliest fossilized records of continental ecosystems. Further, they suggest that the colonization of land started via the interstitial, approximately 150 my earlier than the oldest fossils of terrestrial ecosystems. Overall, the results imply that omnivorous and detritivorous arthropods formed a major component in early terrestrial food webs, thereby facilitating the invasion of terrestrial habitats by later colonizers of higher trophic levels.

Density and community structure of soil- and bark-dwelling microarthropods along an altitudinal gradient in a tropical montane rainforest.

Microarthropod communities in the soil and on the bark of trees were investigated along an elevation gradient (1,850, 2,000, 2,150, 2,300 m) in a tropical montane rain forest in southern Ecuador. We hypothesised that the density of microarthropods declines with depth in soil and increases with increasing altitude mainly due to the availability of resources, i.e. organic matter. In addition, we expected bark and soil communities to differ strongly, since the bark of trees is more exposed to harsher factors. In contrast to our hypothesis, the density of major microarthropod groups (Collembola, Oribatida, Gamasina, Uropodina) was generally low and decreased with altitude. However, as we predicted the density of each of the groups decreased with soil depth. Density of microarthropods on tree bark was lower than in soil. Overall, 43 species of oribatid mites were found, with the most abundant higher taxa being Poronota, pycnonotic Apheredermata, Mixonomata and Eupheredermata. The oribatid mite community on bark did not differ significantly from that in soil. The number of oribatid mite species declined with altitude (24, 23, 17 and 13 species at 1,850, 2,000, 2,150 and 2,300 m, respectively). Rarefaction curves indicate that overall about 50 oribatid mite species are to be expected along the studied altitudinal gradient. Results of this study indicate (1) that microarthropods may be limited by the quality of resources at high altitudes and by the amount of resources at deeper soil layers, and (2) that the bark of trees and the soil are habitats of similar quality for 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.

Anderemaeus (Acari, Oribatida)-overview, three new species from South America and reassessment of Anderemaeidae supported by ontogeny.

Anderemaeus is a genus of Gondwanan soil-dwelling oribatid mites with seven of the eight previously known species being South American. We propose two new species from Chile- A. sidorchukae sp. nov. and A. dentatus sp. nov.-and a third from Ecuador, A. mataderoensis sp. nov. Juveniles of the former two species are described, comprising the first such data for Anderemaeidae: nymphs notably lack both exuvial scalps and centrodorsal gastronotic setae, and the opisthonotal gland opens on a distinct stalk. The generic description is revised and expanded and a key to known species of Anderemaeus is presented, including A. tridactylus comb. nov. We reject the inclusion of Anderemaeus in a broad concept of Caleremaeidae and the implied subsumption of Anderemaeidae, as there are no synapomorphies linking the taxa. Anderemaeus species possess derived traits-e.g. adult with circumpedal carina and nymphs with smooth cuticle and no scalp retention-that are absent from Caleremaeus but are shared with more derived brachypyline taxa. The higher classification of Anderemaeus is reviewed: an analysis of known traits is inconclusive regarding both the generic composition of Anderemaeidae and its superfamilial relationships. However, on the strength of juvenile morphology, we propose the transfer of Anderemaeidae to Gustavioidea.

The Neotropical mite genus Neopilizetes (Oribatida: Galumnidae): redescription of N. neotropicus, rediagnosis of the genus, and descriptions of four new species.

Neopilizetes is a genus of the oribatid mite family Galumnidae that was first characterized by having well-developed setae, a dorsosejugal suture, and raised striae on the prodorsum and notogaster. It was proposed to accommodate Pilizetes neotropicus Balogh Mahunka 1978 from Brazil-the only supposed Pilizetes species known outside the Ethiopian Region-and it has remained monotypic. We found adults of N. neotropicus and four related species in soil-litter interface samples of a single cacao plantation in Ilhéus, Bahia, Brazil. Herein we: redescribe N. neotropicus; describe Neopilizetes thoracicus n. sp., Neopilizetes tigris n. sp., Neopilizetes triumnasus n. sp. and Neopilizetes unumnasus n. sp., and use this information to develop a new concept and expanded diagnosis of Neopilizetes. A key to the five species is provided, and we offer reasons for rejecting the classification of Neopilizetes as a subgenus of Pilizetes.

Identity of the oribatid mite Oribata curva and transfer to Trichogalumna (Acari, Oribatida, Galumnidae), with discussion of nomenclatural and biogeographical issues in the 'curva' species-group.

Based on the study of type material, other historical specimens, and new collections, the adult of the thelytokous oribatid mite Oribata curva Ewing, 1907 (Galumnidae) is redescribed and the name is recombined to Trichogalumna curva (Ewing, 1907) comb. nov. A confusing history of synonymies and misidentifications is traced in detail, and their effect on published statements about biogeography is assessed. Reliable records of T. curva are only those from North America. The tropical mite Pergalumna ventralis (Willmann, 1932) is not a subspecies of T. curva. The widely-reported Trichogalumna nipponica (Aoki, 1966) and other similar species form a complex with T. curva that needs further morphological and molecular assessment.

A checklist of the oribatid mite species (Acari: Oribatida) of Brazil.

A checklist of the oribatid mite species reported in Brazil is presented, including all published records up to 2015. A total of 576 described species in 206 genera and 83 families is presented. Information includes the names by which each species was reported in the Brazilian literature, its general known distribution and by Brazilian States, references, and remarks, when needed. As with most countries, there was a slow early accumulation of knowledge but in recent decades the pace of description has been relatively high. A graphical overview of the number of described oribatid mite species from Brazil in different decades is given. The proportion contributed by each of the major oribatid groups is generally similar to that of the overall world fauna, with a composition that reflects the South American fauna and all of the Neotropics in general. There is a relatively low percentage of primitive mites (Palaeosomata, Enarthronota) other than Lohmanniidae and Mesoplophoridae, which are quite diverse. The Brachypylina comprises about 68% of the oribatid mite fauna. In the checklist, 41% of the species are known only from Brazil, 37% from the Neotropical region, 13.5% have a wider distribution in the global tropical and subtropical regions, and 8.5% are considered cosmopolitan or semicosmopolitan species. The number of descriptions of new species since 2000 from Brazil (73 spp.) and South America (230) is high, but the oribatid mite fauna of these countries remains poorly known. Only continued studies can determine if the high number of species known only from Brazil is an indication of high endemism.