Elements of classification and phylogeny of the Oribatida

Introduction

The Oribatida and the other mites form a monophyletic group among the Arachnida characterized notably by the following characters (Grandjean, 1969a, pp. 801-825; Shultz, 1989, pp. 38-52):

• segments of the body bearing the legs (or podosoma) almost completely suppressed dorsally.
  
  
• movable and terminal mouthparts consisting of the cheliceral and pedipalpal segments (i.e. acquisition of a gnathosoma).
  
  
• a pair of manducatory blade-like scrapers (or rutella) present in various genera.
  
  
• upper buccal lip (or labrum) denticulate in some genera.
  
  
• prelarva with three pairs of vestigial legs in several genera.
  
  
• larvae usually with three pairs of legs (see e.g. Fig. 2 in Grandjean, 1966a).
  
  
• femora of the legs divided into basifemur and telofemur during postembryonic development in some genera.
  
  
• patello-tibial joints in the legs made up of a dorsal transverse hinge (note: in the legs in the Oribatida, similar dorsal hinge joints are found between the femora and the patellae, and between the tibiae and the tarsi; by contrast, the femur (or the basifemur when the segment is bipartite) joins the trochanter at a roughly vertical bicondylar joint while the ambulacrum joins the tarsus at a roughly transverse joint; see e.g. Figs 2E, 6E and 10E in Shultz, 1989) (see Lindquist, 1984, for a discussion about these and other characters; see also Shultz, 1990, for other traits).

 

Habitats

The Oribatida are typical soil-dwelling animals (for instance, the iteroparous parthenogenetic, fungivorous circumdehiscent oppiid, Oppiella nova, is probably the most common arthropod on earth; Norton & Palmer, 1991; see also Siepel, 1994, for reproduction tactics). Their highest diversity and abundance are found in forest litter where densities have been estimated at 55,000-425,000 per square meter (Anderson, 1978; Wallwork, 1983).

In addition, a lot of species belonging notably to the families Euphthiracaridae, Camisiidae, Damaeidae, Cepheidae, Liacaridae, Carabodidae, Ceratozetidae, Mycobatidae, Scheloribatidae, and Oribatulidae are known to spend all of their life cycle on trees (cf. e.g. Travé, 1963, pp. 116-141; André, 1984; Ito, 1986; Winchester et al., 1999; Walter & Behan-Pelletier, 1999; Behan-Pelletier et al., 2001). Several other species inhabit lichens and mosses on rocks of terrestrial environments as well as of the intertidal zone (Travé, 1963, pp. 116-141; Talker et al., 1981; Colloff, 1983). In contrast, fewer oribatid mite lineages have made the transition to caves (Hippa et al., 1988; Bruckner, 1995), to arid deserts (Wallwork, 1972; Wallwork et al., 1986), to freshwater (Grandjean, 1948a; Fernández & Athias-Binche, 1986; Norton et al., 1988) and to salt-marshes (Luxton, 1967; Woodring, 1973; Polderman, 1974).

 

Characters of Actinotrichida and Oribatida

The Oribatida constitute a suborder among the Acarida. Together with the Prostigmata and Astigmata, they belong to the order Actinotrichida, the monophyly of which is supported by the following characters:

• setae (i.e. hair or bristle-like phaneres) with a birefringent axis of chitin (i.e. actinopilin).
  
  
• originally six pairs of setae borne by the prodorsum (i.e. dorsal region of the prosoma).
  
  
• pedipalps without claw.
  
  
• coxae of the legs absent.
  
  
• one lyrifissure (i.e. slit sense organs or strain detectors; cf. Barth, 1981) in the tarsi of the legs (see Grandjean, 1969a, pp. 807-815, and van der Hammen, 1972, p. 276, for other characters).
  
  

Traditionally, in comparison with the Endeostigmata (primitive Prostigmata) which are assumed to be a legitimate outgroup of the Oribatida (Grandjean, 1937a, p. 397; 1939a, pp. 15-50; 1942a, pp. 94-135), the oribatid mites were characterized by the following features:

• dorsal region of the opisthosoma (or notogaster) partly or completely sclerified, rarely completely soft in adults (see e.g. Figs 1 and 3 in Strenzke, 1963).
  
  
• solenidia (i.e. hollow, monorefringent seta-like organs) never borne by the femora but by the patellae, tibiae and tarsi in the appendages.
  
  
• prodorsum with a pair of trichobothria (i.e. compound organs made up of a cavity, or bothridium, and a seta, or sensillus) (click 12), rarely without trichobothria (see e.g. Fig. 4A in Grandjean, 1962a).
  
  
• claws of the legs (or apoteles or ambulacra) with sclerotized, sickle-like ungues, never bearing tenent hairs (see e.g. Fig. 6A in Grandjean, 1937b) and rarely with a pulvillus (see e.g. Figs 2C, 2D, 2E, 5B and 5C in Grandjean, 1951a).

 

Clades and classification of Oribatida

The oribatid mites are divided into six series (Grandjean, 1953a, p. 426; Grandjean, 1969b, pp. 141-144), namely the Palaeosomata, Enarthronota, Parhyposomata, Mixonomata, Desmonomata and Circumdehiscentiae (see Travé et al., 1996, pp. 45-51, for morphological traits used for the diagnosis of the series; see also Evans, 1992, pp. 464, 465, for a key).

A recent phylogenetic study of the series (Norton, 1998) suggested that the Astigmata are derived from some Desmonomata. In fact, a phylogenetic reconstruction obtained from 14 morphological traits regarded as synapomorphies of both Oribatida and Astigmata (Fig. 2A in Norton, 1998) revealed that ‘glandulate’ oribatids (i.e. the Parhyposomata, Mixonomata, Desmonomata and Circumdehiscetiae) form a clade. Conversely, the Palaeosomata and Enarthronota constituted a second clade when the analysis was performed on both previous characters and 12 other traits regarded as exceptional plesiomorphies and apomorphies in the Oribatida (Grandjean, 1969b, p. 147, 148).

The series Circumdehiscentiae comprises a large number of families which are divided into six subdivisions (Grandjean, 1953a, pp. 433-440; 1958a, p. 139; 1965a):

• ‘Opsiopheredermal Pycnonoticae’: adults bearing the dorsal part of the exuviae (or scalp) of their tritonymphs, without paired areae porosae on the opisthosoma (i.e. usually small, more or less rounded elements, traversed by pore canals, with a secretory function; cf. Norton & Alberti, 1997), and nymphs not covered with the exuviae of preceding instars. This subdivision consists of the only family Hermanniellidae. It should be noted that in view of the existence of paired large, notogasteral apophyses on which the exocrine, latero-abdominal glands (or ‘lateral opisthosomal glands’ or ‘oil glands’) open, and of a peculiar, apodemato-acetabular tracheal system, the Hermanniellidae and Plasmobatidae are assigned to the superfamily Hermannielloidea by Grandjean (1962b, p. 271), although it is clear that they have no direct ancestor.
  
  
• ‘Eupheredermal Pycnonoticae’: nymphs bearing the scalp of preceding instars (i.e. protonymphs with one scalp, deutonymphs with two scalps and tritonymphs with three scalps), with at least paired centrodorsal setae (i.e. setae labelled d1, d2, e1) and paired dorsal setae f1 suppressed on the opisthosoma, and adults usually not covered with the exuviae of preceding instars and not equipped with paired areae porosae on the opisthosoma.
  
  Eupherederma are divided into two groups:
  
  • nymphs with tracheal organs associated with the podosoma (see Fig. 8B in Grandjean, 1934a) as e.g. in the Liodidae, Plasmobatidae, Eremaeoidea and Niphocepheidae.
  • nymphs without podosomatic tracheal organs as e.g. in the Polypterozetidae, Gymnodamaeoidea, Damaeidae, Cepheidae, Microzetidae, Gustaviidae, Tenuialidae, Caleremaeidae and Eremuloidea (cf. Grandjean, 1965a).
    
    
• ‘Apheredermal Pycnonoticae with dorsodeficient nymphs’ (not illustrated): nymphs not covered with the exuviae of preceding instars, with paired centrodorsal setae and one or two other dorsal pairs of setae suppressed on the opisthosoma, and adults without paired areae porosae on the opisthosoma.
  This subdivision consists of the families Ceratoppiidae and Liacaridae.
  
  
• ‘Normal Apheredermal Pycnonoticae’: nymphs not covered with the exuviae of preceding instars, with the dorsal cuticle of the opisthosoma completely smooth or partially plicate, with one or two pairs of dorsal setae suppressed on the opisthosoma, and adults without paired areae porosae on the opisthosoma.
  
  Normal Apherederma are divided into two groups:
  • nymphs with the dorsal opisthosomatic cuticle smooth as e.g. in the Carabodidae, Oppiidae, Suctobelbidae, Autognetidae, Thyrisomidae, Hydrozetidae and Limnozetidae. It should be noted that in nymphs in some Autognetidae and Hydrozetidae, the dorsal cuticle of the opisthosoma is completely plicate (see e.g. Fig. 5 in Grandjean, 1963a, and Fig. 1C in Fernández & Travé, 1984, respectively).
  • nymphs with the dorsal opisthosomatic cuticle partially plicate (see Fig. 2A in Grandjean, 1966b) as e.g. in the Selenoribatidae, Fortuyniidae and Micreremidae.
    
    
• ‘Circumdehiscentiae with wrinkled nymphs’: nymphs not covered with the exuviae of preceding instars, with the dorsal opisthosomatic cuticle completely plicate, usually with one pair of dorsal setae suppressed on the opisthosoma, and adult without areae porosae or with a maximum of eight paired areae porosae or derived organs on the opisthosoma (‘octotaxic system’). Altough the oribatid mites belonging to this subdivision were assigned in Grandjean’s (1953) classification to two distinct groups, their inclusion in one taxon, in consideration of their distinctive traits, seems as appropriate as it is convenient (see Grandjean, 1958a, p. 139, for a discussion).
  
  This subdivision comprises two groups:
  • adults without pteromorphs (i.e. a pair of lateral, anterior, movable or immovable cuticular folds on the opisthosoma) as e.g. in the Ameronothridae, Charassobatidae, Licneremaeidae, Tectocepheidae, Podacaridae, Cymbaeremaeidae, Scutovertexidae and Passalozetidae.
  • adults with pteromorphs as e.g. in the Achipteriidae, Tegoribatidae and Pelopsidae.
    
    
• ‘Poronoticae’: nymphs not covered with the exuviae of preceding instars (except in the Oribatellidae), with the dorsal opisthosomatic cuticle not plicate but equipped with shields or microsclerites, with one pair, more rarely two pairs of dorsal setae suppressed on the opisthosoma, and adult usually with eight paired areae porosae or sacculi. It should be noted that the alignments depicted by three pairs of areae porosae or sacculi, namely the anterior pair Aa and the two posterior pairs A2 and A3, correspond to the circumdorsal alignments demonstrated by the insertions of dorsoventral muscles. These muscles are arranged like a curtain. They are homologous to those found in many Arthropoda (cf. Grandjean, 1959a, pp. 250-252).
  
  Poronoticae consist of four groups:
  • ‘Apopheredermal Poronoticae’: scalps borne by nymphs similar to a canopy carried by all or some centrodorsal setae. This group consists of only the family Oribatellidae (see Fig. 2A in Grandjean, 1953c).
  • nymphs with the opisthosoma naked and equipped dorsally and laterally with large, clearly delimited sclerites as e.g. in the Ceratozetidae, Euzetidae, Mycobatidae, Chamobatidae and Galumnidae (see e.g. Figs 1 and 2 in Grandjean, 1951b, and Fig. 4 in Travé, 1970).
  • nymphs with the opisthosoma naked and equipped with both large sclerites and small, porous, roughly rounded or oval microsclerites, on the border of which a seta is inserted (i.e. microsclerites ‘thrown off centre’). This group consists of only the family Humerobatidae.
  • nymphs with the opisthosoma naked and equipped only with microsclerites thrown off centre (see e.g. Figs 3A, 3B and 3C in Grandjean, 1950a) as e.g. in the Mochlozetidae, Sellnickiidae, Stelechoribatidae, Parakalumnidae and Scheloribatoidea.

 

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