Radiation in Sexual and Parthenogenetic Oribatid Mites (Oribatida, Acari) As Indicated by Genetic Divergence of Closely Related Species

Experimental & Applied Acarology. 2003  |  Pubmed ID: 14635813

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.

Molecular Phylogeny of Oribatid Mites (Oribatida, Acari): Evidence for Multiple Radiations of Parthenogenetic Lineages

Experimental & Applied Acarology. 2004  |  Pubmed ID: 15347023

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

Imaging Applications of Synchrotron X-ray Phase-contrast Microtomography in Biological Morphology and Biomaterials Science. I. General Aspects of the Technique and Its Advantages in the Analysis of Millimetre-sized Arthropod Structure

Journal of Microscopy. Jul, 2007  |  Pubmed ID: 17635659

Synchrotron-generated X-rays provide scientists with a multitude of investigative techniques well suited for the analysis of the composition and structure of all types of materials and specimens. Here, we describe the properties of synchrotron-generated X-rays and the advantages that they provide for qualitative morphological research of millimetre-sized biological organisms and biomaterials. Case studies of the anatomy of insect heads, of whole microarthropods and of the three-dimensional reconstruction of the cuticular tendons of jumping beetles, all performed at the beamline ID19 of the European Synchrotron Radiation Facility (ESRF), are presented to illustrate the techniques of phase-contrast tomography available for anatomical and structural investigations. Various sample preparation techniques are described and compared and experimental settings that we have found to be particularly successful are given. On comparing the strengths and weaknesses of the technique with traditional histological thin sectioning, we conclude that synchrotron radiation microtomography has a great potential in biological microanatomy.

Small but Powerful: the Oribatid Mite Archegozetes Longisetosus Aoki (Acari, Oribatida) Produces Disproportionately High Forces

The Journal of Experimental Biology. Sep, 2007  |  Pubmed ID: 17704078

We investigated the holding and pulling forces generated by claws of the microarthropod Archegozetes longisetosus (Chelicerata, Acari, Oribatida) on three substrates with different roughness (R(a)=0.05 microm, 1 microm, 30 microm). Holding forces were measured perpendicular to the substrate using a strain gage force transducer; pulling forces were measured parallel to the substrate using an analytical scale. We found a significant positive correlation of surface roughness and the forces generated. Mites produced holding forces on horizontal rough surfaces (R(a)=30 microm) of up to 1180 times their weight; on vertical rough surfaces (R(a)=30 microm) they can pull with 530 times their weight, effectively involving only two pairs of legs. The relative forces are five times higher than theoretically expected for organisms of this size (<1 mm, 100 microg) and higher than any relative forces reported for insect claws. Muscles involved in claw action produced stresses up to 1170 kN m(-2), a value that is only excelled by decapod crustacean claw closer muscles. Ours is the first study of performance by chelicerate apoteles and claws and also the first to measure forces generated by any microarthropod.

Role of Musculature During Defecation in a Particle-feeding Arachnid, Archegozetes Longisetosus (Acari, Oribatida)

Journal of Morphology. Jan, 2009  |  Pubmed ID: 18726923

Most extant Chelicerata are characterized by external digestion and the ingestion of fluid food. Exceptions include the marine taxa, most Opiliones, and the mite groups Opilioacarida (Parasitiformes) and Sarcoptiformes (Acariformes), which ingest particulate food. This leads to different physiological and morphological adaptations for food processing, including the production and extrusion of solid fecal pellets, which are rather large in sarcoptiform mites. Few studies have investigated the defecation of such large fecal pellets, and available information is contradictory. We use a combination of non invasive microscopical techniques and in vivo examination to investigate the complex functional morphology of the anal region of the oribatid mite Archegozetes longisetosus Aoki. The opening of the anus is at least initiated by indirect muscular action via an increase of hemolymph pressure, through the action of dorsoventral muscles (dvm). Extrusion of the fecal pellet is accomplished by the prerectal muscle collar, with full opening of the anus and rotation and bowing of the plates probably resulting from pressure of the pellet. The sequential nature of these actions was demonstrated by many observations in which the anus opened only partially; these were concomitant with dmv contraction but pellets were not being extruded. All muscles directly connected to the anal and adanal regions assist in keeping the anus closed; they are antagonists to hydrostatic forces that are necessary for normal activity. Based on the literature, no obvious similarities were noted with defecation musculature in other particle-feeding chelicerates, but most muscles can be homologized with those of more specialized oribatid mites. The function of the outer anal muscles has been modified in both Euphthiracaridae and Brachypylina to assist in providing general hemolymph pressure.

The Ptychoid Defensive Mechanism in Euphthiracaroidea (Acari: Oribatida): a Comparison of Muscular Elements with Functional Considerations

Arthropod Structure & Development. Nov, 2009  |  Pubmed ID: 19595788

Ptychoidy is a mechanical predator defence in some groups of Oribatida (Acari), where the animals can retract their legs into the idiosoma and encapsulate. This mechanism is enabled by a number of morphological adaptations. We used the non-invasive technique of synchrotron X-ray microtomography to compare muscular elements involved in ptychoidy of two species from the Euphthiracaroidea (Oribotritia banksi and Rhysotritia ardua) which differ in a number of cuticular elements involved in ptychoidy. We hypothesize that a strong functional correlation exists between these cuticular structures and their corresponding musculature. We found a number of distinct differences concerning quantitative and qualitative muscle morphology. For testing the functional impact of different muscle configurations we simulated two conditions in silico (encapsulated / opened) and analysed the spatial relative force vectors of the prodorsum lateral adjustor muscles (pla) which are responsible for retraction and adjustment of the prodorsum during encapsulation. We show that the functional morphology of these muscles strongly differs between the two species and that this can be explained by the structure of corresponding cuticular elements. Furthermore, the dynamics of pla, as measured by the extent of contraction during encapsulation, is more than two times higher in R. ardua than in O. banksi.

First Cleavages, Preblastula and Blastula in the Parthenogenetic Mite Archegozetes Longisetosus (Acari, Oribatida) Indicate Holoblastic Rather Than Superficial Cleavage

Arthropod Structure & Development. Jul, 2010  |  Pubmed ID: 20153841

The mode of cleavage in the Acari is generalized as superficial or intralecithal, with a preceding phase of total (holoblastic) cleavage, but the knowledge is fragmentary and conclusions have been inconsistent, even when relating to the same species. Since no data about early embryology is available for the speciose group Oribatida, we studied Archegozetes longisetosus using transmission electron microscopy. We focused on early cleavages and the formation of the blastula, as these are the important and controversial points in early embryology of the Acari. We expected, as postulated for other acarine eggs, the early cleavages to be holoblastic and followed by a superficial preblastoderm stage. The early cleavages of A. longisetosus are holoblastic and blastomeres give rise to yolk-free micromeres and macromeres containing all the yolk. In contrast to expectations, the micromeres do not form a superficial preblastoderm layer. They are scattered along the embryonic surface and form an external, monocellular layer that covers the whole surface of the embryo. Since each of the existing TEM studies of mites shows this same pattern, and since this specialized form of total cleavage seems to be unique in Chelicerata, it may be the general mode of cleavage in Acari. However, the question will require much more investigation, especially since most data relate to the Actinotrichida and very few are currently available for species in the other major group, the Anactinotrichida.

Integrative Taxonomy: Combining Morphological, Molecular and Chemical Data for Species Delineation in the Parthenogenetic Trhypochthonius Tectorum Complex (Acari, Oribatida, Trhypochthoniidae)

Frontiers in Zoology. 2011  |  Pubmed ID: 21303503

There is a long-standing controversial about how parthenogenetic species can be defined in absence of a generally accepted species concept for this reproductive mode. An integrative approach was suggested, combining molecular and morphological data to identify distinct monophyletic entities. Using this approach, speciation of parthenogenetic lineages was recently demonstrated for groups of bdelloid rotifers and oribatid mites. Trhypochthonius tectorum, an oribatid mite from the entirely parthenogenetic desmonomatan family Trhypochthoniidae, is traditionally treated as a single species in Central Europe. However, two new morphological lineages were recently proposed for some Austrian populations of T. tectorum, and were described as novel subspecies (T. silvestris europaeus) or form (T. japonicus forma occidentalis). We used the morphological and morphometrical data which led to this separation, and added mitochondrial and nuclear DNA sequences and the chemical composition of complex exocrine oil gland secretions to test this taxonomical hypothesis. This is the first attempt to combine these three types of data for integrative taxonomical investigations of oribatid mites.

Oribatid Mites and Skin Alkaloids in Poison Frogs

Biology Letters. Aug, 2011  |  Pubmed ID: 21345855

Fine Structure of the Gnathosoma of Archegozetes Longisetus Aoki (acari: Oribatida, Trhypochthoniidae)

Journal of Morphology. Sep, 2011  |  Pubmed ID: 21630320

Oribatida are one of the main groups of Acari comprising mostly important decomposers in soils. Most species are particle feeders, an exceptional mode of nutrition in Arachnida. Hence, their feeding organs, the gnathosoma, are of special functional interest. We studied nearly all components using scanning and transmission electron microscopies as well as reconstructions based on synchrotron X-ray microtomography from the model oribatid Archegozetes longisetosus. Besides cuticular structures, we describe the full set of muscles and confirm the presence of a trochanter remnant at the base of the chelicera. Setae on the prodorsum and the anterior and posterior infracapitular setae are mechanoreceptors innervated by two dendrites ending with tubular bodies. Dendrites of adoral setae, anterior setae of the chelicerae, and the supracoxal setae show neither obvious tubular bodies nor wall or terminal pores. Thus their function remains obscure. For the first time, a muscular proprioreceptor has been found in Arachnida. It likely monitors the actions of muscles moving the movable digit of the chelicera. Glandular structures within and associated with the gnathosoma are described. Dermal glands represented by secretory porose areas are found within the infracapitulum. More complex associated glands comprise the podocephalic glands and the infracapitular glands, the ducts of which were traced completely for the first time. The components described are mostly fundamental for the gnathosoma of Actinotrichida (Acariformes), one of the two lineages of Acari, to which Oribatida belong. The gnathosoma is generally considered the most relevant putative synapomorphy of Acari. Since the monophyly of Acari has become more and more questionable during the last decades, a thorough reinvestigation of this body part is necessary for a comprehensive understanding of acarine and even arachnid phylogeny and evolution. This article provides a starting point of such a re-evaluation of the gnathosoma.

Tasty but Protected--first Evidence of Chemical Defense in Oribatid Mites

Journal of Chemical Ecology. Sep, 2011  |  Pubmed ID: 21898169

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.

Expanding the 'enemy-free Space' for Oribatid Mites: Evidence for Chemical Defense of Juvenile Archegozetes Longisetosus Against the Rove Beetle Stenus Juno

Experimental & Applied Acarology. Feb, 2012  |  Pubmed ID: 22048786

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.

Triggering Chemical Defense in an Oribatid Mite Using Artificial Stimuli

Experimental & Applied Acarology. Jan, 2012  |  Pubmed ID: 22286143

Most oribatid mites are well known for their exocrine oil gland secretions, from which more than a hundred different chemical components (hydrocarbons, terpenes, aromatics and alkaloids) have been described. The biological functions of these secretions have remained enigmatic for most species, but alarm-pheromonal and allomonal functions have been hypothesized, and demonstrated in some cases. Here, we tested different experimental stimuli to induce the release of defensive secretions in the model oribatid mite Archegozetes longisetosus Aoki. Whereas various mechanical stimuli did not result in a reproducible and complete expulsion of oil gland secretions, repeated treatments with hexane led to complete discharge. Life history parameters such as survival, development and reproduction were not influenced by the hexane treatment. Repeated hexane treatments also resulted in a complete depletion of oil glands in Euphthiracarus cribrarius Berlese.