Zootaxa, Comparative morphology and species ...

Zootaxa 2213: 1–46 (2009) www.mapress.com / zootaxa/

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ZOOTAXA ISSN 1175-5334 (online edition)

Comparative morphology and species-groups of the oribatid mite genus Scapheremaeus (Acari: Oribatida: Cymbaeremaeidae), with new species from South Australia MATTHEW J. COLLOFF CSIRO Entomology, GPO Box 1700, Canberra, ACT 2601, Australia. E-mail: [email protected]

Table of contents Abstract ............................................................................................................................................................................... 1 Introduction ......................................................................................................................................................................... 2 Morphology of the Adults ................................................................................................................................................... 4 Species groups ................................................................................................................................................................... 10 New diagnosis of Scapheremaeus ..................................................................................................................................... 12 Scapheremaeus Berlese, 1910 ........................................................................................................................................... 12 Materials and methods ...................................................................................................................................................... 13 Descriptions of new species .............................................................................................................................................. 13 Scapheremaeus cheloniella sp. nov. .......................................................................................................................... 13 Scapheremaeus pulleni sp. nov. ................................................................................................................................. 16 Scapheremaeus angusi sp. nov. ................................................................................................................................. 19 Scapheremaeus lambieae sp. nov. ............................................................................................................................. 22 Scapheremaeus ewani sp. nov.................................................................................................................................... 24 Discussion ......................................................................................................................................................................... 26 Acknowledgements ........................................................................................................................................................... 27 References ......................................................................................................................................................................... 27

Abstract The morphology of the genus Scapheremaeus Berlese, 1910 is reviewed and characters of taxonomic utility delineated. Based on the morphological review, some 13 species-groups are outlined based on major morphotypes. There are two main categories: i) species that have a complete circumdorsal scissure with plicate microsculpture on the circumnotogastral plate and strongly contrasting microsculpture (foveolae, ridges or tubercles) on the centrodorsal plate (plicate species-groups), and ii) species with the circumdorsal scissure complete, incomplete or absent but with little or no contrast in microsculpture between the central and lateral regions: typically both regions foveolate or reticulate (nonplicate species-groups). A catalogue of world species of Scapheremaeus is provided. Scapheremaeus petrophagus (Banks, 1906) is not a Scapheremaeus but belongs to an undetermined genus in the Ameronothroidea. Cymbaeremaeus cyclops Oudemans, 1915 is recombined to Scapheremaeus. Five new species are described (S. angusi sp. nov., S. cheloniella sp. nov., S. ewani sp. nov., S. lambieae sp. nov., and S. pulleni sp. nov.) from soil and litter habitats in semi-arid Mallee eucalypt vegetation at Bookmark Biosphere Reserve, South Australia. These are the first members of the genus Scapheremaeus to be described from Australia, though undescribed species have been recorded previously. All the new species are morphologically closely-related and belong to a single species-group: Carinatus. Key words: mite, taxonomy, morphology, Oribatida, systematics

Accepted by H. Schatz: 27 Jul. 2009; published: 28 Aug. 2009

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Introduction The genus Scapheremaeus Berlese, 1910 contains 97 species recognised herein (including new species described below, and two fossils; Appendix 1). Subías (2004) listed 81 species, of which 59 have been described since 1960. The genus has a cosmopolitan distribution, with 37 spp. previously described from localities in the Neotropics, 13 from the Palaearctic, 21 from the Oriental and 10 from the Afrotropical regions, six from the Nearctic and five from the Australasian regions. This paper adds five new species from this latter region. Scapheremaeus is placed in the Cymbaeremaeidae, as defined by Behan-Pelletier (1989b), together with Ametroproctus Higgins & Woolley, 1968 (7 spp.; Behan-Pelletier, 1987), including the subgenus Coropoculia Aoki & Fujikawa (1972); Bulleremaeus Hammer, 1966 (2 spp.); Cymbaeremaeus Berlese, 1896 (1 sp.); Glanderemaeus Balogh & Csiszár, 1963 (1 sp.); Seteremaeus Hammer, 1971 (1 sp.) and Scapuleremaeus Behan-Pelletier, 1989a (1 sp.). Scapheremaeus represents something of a challenge to oribatid taxonomists. Many species are less than 500 μ m long, heavily sclerotised and with intricate patterns of microsculpture on the prodorsum and notogaster, which may obscure surface structures such as the often minute setae, lyrifissurae and glands. The addition of cerotegument and its adherent debris adds to the problem. Thus some species descriptions, especially older ones, may illustrate an incomplete chaetome, or contain inaccurate or stylistic depictions of cuticular microsculpture (tubercles drawn as foveolae, or vice versa). For example, Norton and Kethley (1989) found the type of S. pulchellus (Berlese, 1910) has irregular ridges on the centrodorsal region, not uniform reticulations as depicted by Berlese (1910). In order to fully describe the gross morphology, as well as specific details, is necessary to illustrate specimens in lateral view. Several characters of taxonomic value can only be seen in this orientation, including the humeral processes, exobothridial setae (where present), the pedotecta and the relative positions of the posterior notogastral setae. This requires micro-dissection of the legs and careful positioning of the specimen as a temporary mount in a cavity slide. Some structures are hard to see even under a high power oil immersion lens. The combination of temporary mount preparations and immersion oil can be messy and frustrating. Members of the genus Scapheremaeus display a lack of strongly correlated characters, and many have polytypic states. In other words, character states occur in various matrix-type combinations. For example, elongated lamellar apophyses occur in groups of species that have a circumdorsal scissure that is either complete (6 spp.) or incomplete (5 spp.). The notogastral chaetome may be 10–14 pairs in the former group or 14 pairs in the latter. Both groups contain species with either foveolae or reticulate centrodorsal microsculpture, with 1 or 3 claws, and humeral processes and interlamellar setae either present or absent. As stated by Travé & Fernandez (1986) of Scapheremaeus, ‘…nous avons affaire à un ensemble héterogène.’ Hence it is no great surprise that the genus has a reputation as difficult, and no revision has been attempted. Berlese (1886, 1908, 1910a, 1910b) described six species and, in the 1910a paper, created Scapheremaeus as a subgenus of Cymbaeremaeus. The type species S. patella (Berlese, 1886), S. guerini (Berlese, 1908) and S. reticulatus (Berlese, 1910) have been redescribed by Mahunka & Mahunka-Papp (1995). Of the latter species, they consider the redescription by Balogh (1943) probably represents a different species. Scapheremaeus patella was previously redescribed by Mahunka (1977) and Pérez-Iñigo (1995). S. patella (Berlese) sensu Hammer, 1966 differs from the type species in several respects (Hammer, 1966), and was regarded as a separate species and named S. hammerae by Balogh & Balogh (2002). Scapheremaeus corniger (Berlese, 1908) was redescribed by Pérez-Iñigo & Subías (1974), though not based on type material. Scapheremaeus marmoratus (Berlese, 1910) and S. pulchellus remain unrevised. New species were added by other authors during the late 19th and early 20th Centuries (Michael, 1890; Banks, 1896; 1906; 1909; Hall, 1911; Oudemans, 1915; 1917; Willmann, 1936; 1939), including two fossil species from Baltic amber (Sellnick, 1918; 1931), but the genus remained relatively neglected until the work by Marie Hammer who, between 1958 and 1982, described 15 species from Argentina and Bolivia (1958), Peru (1961), New Zealand (1966), Fiji (1971), Java (1979) and Bali (1982). The majority of other new species described around this time were in publications by the Hungarian acarologists, J. Balogh (1958, 1970; 3 spp.),

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Mahunka (1978; 1983a; 1983b; 1984a; 1984b; 1985a; 1985b; 1986; 1987a; 1987b; 1988; 14 spp.) and Balogh & Mahunka (1967; 1978; 3 spp.). The work of Hammer and the Hungarians focussed heavily on the description of diverse oribatid taxa in collections from particular countries, and represents a strongly faunistic approach, rather than being primarily taxon-orientated. As a result, there was a tendency for species to accumulate within the genus Scapheremaeus with little comparative or diagnostic analysis of their character states. The first review of the genus (Aoki, 1964) consisted of an examination of seven characters for some 16 species, mostly described by Berlese and by Hammer. Aoki & Fujikawa (1972) provided further details of character states of Scapheremaeus in comparison with the genera Coropoculia, Charassobates, Topalia, Cymbaeremaeus, Glanderemaeus and Micreremus. In a subsequent review, Pérez-Iñigo & Subías (1974) provided the first detailed morphological account of a Scapheremaeus species with their redescription of S. corniger (Berlese, 1908), including the lateral aspect, precise details of notogastral, ventral and leg chaetotaxy, as well as immatures and a comparison with other species with well-developed humeral spine-like processes. Engelbrecht (1975) described two species from South Africa and included a morphological comparison with other ameronothroid genera, while Travé & Fernandez (1986) provided a comparably detailed morphology of S. argentinensis and comparison with S. patella, including clarification of several important characters such as the arrangement of scissures and plates of the notogaster. These publications represent benchmarks for subsequent detailed descriptions and redescriptions (Fernandez & Cleva, 1997; Ramani & Haq, 1998; Fujikawa, 2002; 2005) which form the basis of our understanding of the comparative morphology of Scapheremaeus. Assemblages of Scapheremaeus species have been described by Ríos & Palacios-Vargas (1998) from sites in Mexico, several in rainforest, (15 spp.) and Wang (1998) from China, mostly from Fujian Province (5 spp.). Very little is known of the ecology or natural history of members of the genus. They appear to be catholic in habitat, having been found to be surface-active in arid conditions such as those encountered in Eucalyptus and Banksia litter, creosote bushes in semi-desert and in dry moss and maritime lichens. But they are also found in moist or aquatic habitats such as stems and leaves of rainforest trees, corticolous lichens and mosses, in samples obtained by insecticide fogging of the forest canopy (Ríos & Palacios-Vargas, 1998), in forest litter and in temporary pools. Several species are associated with epiphytic plants, especially bromeliads, from wet tropical and sub-tropical forest, (S. flamiferus Palacios-Vargas & Ríos, 1998; S. hungarorum Mahunka, 1986, S. ornatus Balogh & Mahunka, 1978, S. simplex Palacios-Vargas & Ríos, 1998; S. tillandsiae Fernandez & Cleva, 1997; and S. tillandsiophilus Palacios-Vargas & Ríos, 1998). No species of Scapheremaeus have been described from Australia hitherto. Two un-named species were recorded by Lee (1986), three each by Walter et al. (1994), and Walter (1995), one by Walter & BehanPelletier (1993) and a fossil species is known from leaf domatia of Sloanea australis parvifolia (Elaeocarpaceae) from Eocene clay lenses in Victoria (O'Dowd et al., 1991). Additionally, I have collected some 16 other undescribed species from Queensland, Victoria, Tasmania, New South Wales, South Australia and Western Australia. It would appear, even from these preliminary observations, that the Scapheremaeus fauna of Australia is quite diverse. Despite this diversity, I have never found these mites to be particularly abundant in their natural habitat. Scapheremaeus is now a large genus in need of revision, but many of the species require redescription, especially those described during the late 19th and early 20th Centuries. In the interim, a consideration of species groups and morphotypes represents a first step in the process. The purpose of this paper is to provide a review of the genus Scapheremaeus by presenting a morphological survey to highlight characters that are of taxonomic value, determining species groups within Scapheremaeus based on published descriptions, providing a redefinition of the genus, and describing five new species from soil and litter in Mallee eucalypt vegetation in South Australia. Mallee eucalypt woodland is a vegetation type unique to semi-arid regions of Southern Australia in rainfall zones of ca. 200–300 mm. Mallee refers both to the vegetation type and to the multi-stemmed growth form of the constituent Eucalyptus species, which include E. gracilis, E. dumosa, E. incrassata, E. oleosa and SCAPHEREMAEUS (ACARI: ORIBATIDA)

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E. socialis as common species (Noble et al., 1990; Costermans, 1994; Yen et al., 2006). The trees are often relatively short (no more than 10 m, usually less than 4 m), with stems emerging from a prominent lignotuber. For a vegetation community in the semi-arid zone, the diversity of the invertebrate fauna of the South Australian mallee is remarkably high, as revealed by an intensive survey of invertebrate biodiversity at the Bookmark Biosphere Reserve conducted by CSIRO Entomology staff between 1995 and 1996 (Pullen, 1997). The material described herein was collected during that survey.

Morphology of the Adults Immatures have been described only for S. glaber by Hammer (1958), S. corniger by Perez-Iñigo & Subías (1974), S. argentinensis by Travé & Fernandez (1986) and S. tillandsiae by Fernandez & Cleva (1997). Consequently, the morphological review is based on adults. Prodorsum. The absence of interlamellar setae in adults of Scapheremaeus is the norm. They are present in only about a quarter of species: S. cheloniella sp. nov.; S. corniger; S. cornutus Balogh, 1958; S. demeteri Mahunka, 1983; S. digitatus Wang, 1998; S. humeratus Balogh & Mahunka, 1967; S. latus Mahunka, 1985; S. longicuspis Mahunka, 1984; S. longilamellatus Mahunka, 1985; S. madeirensis Willmann, 1939; S. palustris (Sellnick, 1924) (redescribed by Mahunka, 1987b); S. polysetosus Sitnikova, 1975; S. pseudoreticulatus Mahunka, 1984; S. quadrireticulatus Wang, 1998; S. retestriatus Wang, 1998; S. subcorniger Perez-Iñigo & Peña, 1996; S. subglaber Balogh & Mahunka, 1978 and S. tosaensis Fujikawa, 2005. All species with a unideficient notogastral chaetome (i.e. 15 pairs) have interlamellar setae. Behan-Pelletier (1989b) considers their presence to be plesiomorphic for Cymbaeremaeidae, with numerous subsequent losses within genera. True lamellae are absent in Scapheremaeus. Instead, they have lamellar costulae, i.e. cuticular ridges, not bladelike or having an apical cusp, but terminating with the apophyses of the lamellar setae. Aoki (1972, his Fig. 16) illustrated some of the different arrangements of costulae and other prodorsal ridges of Scapheremaeus, though not the condition whereby the lamellar apophyses are greatly extended. A tutorium, i.e. a raised blade-like ridge with an apical cusp lateral to each lamella is found in Ametroproctus but not Scapheremaeus (Behan-Pelletier, 1989b), though many species of Scapheremaeus have well-developed carina lateral to the costula. Exobothridial setae are typically absent in the adult, though present in the tritonymphs (Fernandez & Cleva, 1997). There are a few exceptions (e.g. S. hammerae, S. tosaensis). Notogastral plates and scissures. The typical arrangement of notogastral plates is an oval centrodorsal plate encompassed by a pair of lateral circumnotogastral plates and separated from them by the circumdorsal furrow (Travé & Fernandez, 1986) or scissure. The two lateral plates, one ventral and one dorsal, have a circumferential scissure, and the ventral circumnotogastral plate is joined to the ventral plate at the circumgastric scissure (Fig. 4; Travé & Fernandez, 1986; their Fig. 10c). Variations on this arrangement include diagonal extensions of the circumdorsal scissure into the humeral region (Figs. 4a, 8a); an incomplete circumdorsal scissure (Figs. 2f, 3e, 3f, 3g; S. bituberculatus Wang, 1998, S. cornutus, S. striatomarginatus Hammer, 1979), sometimes accompanied by a transverse ridge posterior of the lenticulus ( as in S. alvarezius Ríos & Palacios-Vargas, 1998 and S. humeratus); or modifications of the centrodorsal shield in the caudal region including a V- or U-shaped medial ridge (S. clavifer Hammer, 1958, S. magdalenae Ríos & PalaciosVargas, 1998, S. volcanicus Ríos & Palacios-Vargas, 1998) or a raised sub-circular structure (S. baloghius Ríos & Palacios-Vargas, 1998, S. corniger, S. fimbriatus (Michael, 1890), S. sinuosus Aoki, 1964). In species depicted with an incomplete circumdorsal scissure, without descriptive supporting text, it is sometimes difficult to determine whether the scissure is truly incomplete or it simply has been drawn as such. Regarding other genera of the Cymberemaeidae, the circumdorsal scissure is absent in Bulleremaeus, Glanderemaeus, and Cymberemaeus. In Scapuleremaeus kobauensis there is no scissure, but the centrodorsal region is clearly differentiated from the marginal region by different microsculpture (Behan-Pelletier, 1989a), as it is in Ametroproctus oresbios and A. tuberculosus (Behan-Pelletier, 1987; her Figs. 3 and 16 respectively),

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but not in other Ametroproctus spp. In Seteremaeus the circumdorsal scissure is complete and well-developed (Hammer, 1971).

FIGURE 1. Character states of the notogastral chaetotaxy of Scapheremaeus spp., indicating setal deficiencies (dotted circles) and examples of species with the particular character state illustrated.

Notogastral chaetotaxy. Notogastral chaetotaxy is one of the more troublesome characters because of the small size of the setae, combined with a sclerotised, heavily-ornamented notogaster. Indeed, several authors have remarked upon the difficulty in providing a definitive description of notogastral chaetotaxy, and it is as uncertain in the descriptions of S. arcuatus Hammer, 1971; S. clavifer; S. convexus Hammer, 1979; S. fisheri Aoki, 1966; S. guerini; S. striatomarginatus and S. trirugis Hammer, 1958. Older descriptions rarely include SCAPHEREMAEUS (ACARI: ORIBATIDA)


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mention of the notogastral chaetotaxy and it is not known for S. fimbriatus; S. hieroglyphicus (Hall, 1911); S. marginalis (Banks, 1896); S. marmoratus; S. parvula (Banks, 1909) and S. pulchellus (Berlese, 1910). The plesiomorphic condition in Scapheremaeus is probably the unideficient state of fifteen pairs, as for many brachypyline oribatid mites (Fig. 1a). A small group of species from China and eastern Russia have this character state, comprising a full complement of setae in the c series, including seta c3 positioned on, or adjacent to, the humeral process (S. digitatus; S. polysetosus; S. quadrireticulatus; and S. retestriatus). In the bi-deficient state, i.e. fourteen pairs, this pair of setae is lost (Fig. 1b). Notogastral chaetotaxy may have been misinterpreted in some descriptions. Common examples of this are where seven or eleven pairs are cited but where the p series, which may be reduced in size compared with the h series, are positioned on the ventral circumnotogastral plate and have been overlooked. These setae are not visible from above and sometimes difficult to see even with the specimen in ventral view (contrast Figs. 11a and 12a). The positioning of setae of the p series either on the ventral or dorsal circumnotogastral plate is potentially a useful taxonomic character. Including the ‘missing’ p series brings the total to ten (Fig. 1e) or fourteen pairs (Fig. 1b), the most common character states for the genus. The h series are highly conserved, although one or more of the p series may be significantly reduced or absent, but accompanied by the loss of adanal setae. For example, one undescribed species from New South Wales lacks p3, together with ad3 . Another, from Queensland, lacks the entire p series, plus ad2 and ad3. However S. palaciosi Ríos, 1996 has eight pairs of notogastral setae (p2 and p3 are missing) but possesses a full complement of adanal setae. There are no examples of species which posess seven pairs of notochaetae and in which all the p series are definitively present (Fig. 1f). Species in which ten pairs of setae are probably present, but seven pairs have been cited (and which have the full complement of adanal setae), are S. arboreus Corpuz-Raros, 1979; S. bicornutus Hammer, 1971; S. crassus Mahunka, 1988; S. cuspidatus Pérez-Iñigo, 1981; S. ornatus Balogh & Mahunka, 1968; S. sinuosus Aoki, 1964; and S. subglaber Balogh & Mahunka, 1978. Species which have been described with 7 pairs of notogastral setae but where adanal setal reduction is unclear include S. emarginatus Hammer, 1966 and S. stratus Hammer, 1958. In addition, S. carinatus Willmann, 1936 and S. madeirensis Willmann, 1939 are described as having nine pairs and S. latirostris Willmann, 1936 as 8 pairs, but all three probably have ten pairs. Scapheremaeus longilamellatus Mahunka, 1985 appears to have seven pairs of notogastral setae, missing p2, p3 and lm. Species where fourteen pairs of setae are probably present, but eleven pairs have been cited include S. glaber Hammer, 1958 and S. semiornatus Hammer, 1979. Additional species with a complement of fourteen pairs are S. longicuspis Mahunka, 1984, described with 10 pairs, and S. bituberculatus Wang, 1998, described with 13 pairs. Only two species, S. bituberculatus and S. clavisetus Mahunka, 1978, appears to have a chaetome of 13 pairs (Fig. 1c), and only one, S. humeratus Balogh & Mahunka, 1967 appears to have a chaetome of twelve pairs (Fig. 1d; though described with eleven pairs). No species could be confirmed with eleven pairs of notogastral setae. Notogastral microsculpture. The microsculpture of the centrodorsal plate usually consists either of foveolae or ridges with foveolae. More rarely, the centrodorsal plate is tuberculate (S. alvarezius; S. argentinensis; S. chaac Ríos & Palacios-Vargas, 1998; S. cheloniella sp. nov., S. clavifer, S. hectorperezius Ríos & Palacios-Vargas, 1998; S. johnsi Balogh, 1970; S. lambieae sp. nov.; S. pisacensis Hammer, 1961; S. schatzi Ríos & Palacios-Vargas, 1998). These character states represent some simple variations on a theme. The basic state is foveolae separated by ridges and it is probable that some of the other states are derived from this. As the ridges narrow and foveolae broaden, it gives the appearance of an entirely foveolate centrodorsal plate. As ridges become fragmented they form ovoid or round tubercles. Only two species are smooth: S. glaber Hammer, 1958 and S. subglaber. The microsculpture of the circumnotogastral plates is either plicate or similar to that on the centrodorsal region, typically consisting of foveolae. Lyrifissurae. The full complement of lyrifissurae is often unclear because it is hard to locate them amonst similarly-shaped elements of the notogastral microsculpture. Lyrifissurae ip, ips, im and ih are usually visible,

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whereas ia is often very hard to locate and is positioned either dorsally in the humeral region amongst the plicate folds or on the ventral circumnotogastral plate and not readily visible in dorsal view. Cerotegument and setae. The cerotegument of Scapheremaeus is dark brown and usually only weaklydeveloped. An example of dense cerotegument is found in S. tonatiuh Palacios-Vargas, Ríos & Vázquez, 1998. Notogastral and lamellar setae that appear club-shaped and black are covered with a thick layer of cerotegument apically. Notogastral setae are typically spiniform or bacilliform. There are no examples where the setae themselves are club-shaped. On a single specimen, certain setae have a club-shaped cerotegumental covering and others do not. The amount of cerotegument may vary between setae of the same pair. Considering that cerotegument is a cuticular secretion, the amount and degree of development is contingent in part on the age of the adult. Black, club-shaped setae have been regarded as a definitive character of taxonomic value, but should be regarded as more variable. In some specimens, notogastral setae may have several layers of cerotegument (darkest innermost, lightest outermost), indicating multiple deposition/ secretion events.

FIGURE 2. Plicate species groups of Scapheremaeus: a) Carinatus group; b) Petrosus group; c) Fimbriatus group; d) Patella group; e) Polysetosus group; f) Longilamellatus group. SCAPHEREMAEUS (ACARI: ORIBATIDA)


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FIGURE 3. Non-plicate groups of Scapheremaeus: a) Alveolatus group; b) Simplex group; c) Palustris group; d) Argentinensis group; e) Cuspidatus group; f) Humeratus group; g) Emarginatus group.

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Humeral processes. Humeral processes are ventrally-directed extensions of the cuticle on the anteriolateral margins of the circumnotogastral plate. They can only be seen fully in lateral view. In dorsal view, these structures can be confused with the acetabulum of leg II but they can be visible when they are long, spine-like and curved (as in S. arboreus Corpuz-Raros, 1979; S. baloghius; S. bituberculatus; S. carinatus Willmann, 1936; S. convexus Hammer, 1979; S. corniger; S. cornutus; S. hungarorum, S. palaciosi Ríos, 1996; S. pundamiliaensis Engelbrecht, 1975; S. subcorniger; S. taurus J. Balogh, 1970; S. yamashitai Aoki, 1970). In other species they are short and broadly triangular (S. tillandsiophilus). Humeral processes are absent in some species (S. alvarezius; S. argentinensis; S. chaac; S. eugenius Ríos & Palacios-Vargas, 1998; S. flamiferus; S. fungisetosus Ríos & Palacios-Vargas, 1998; S. grahamius Ríos & Palacios-Vargas, 1998; S. mahunkaius Ríos & Palacios-Vargas, 1998; S. morulisensillatus Ríos & Palacios-Vargas, 1998; S. nogueraius Ríos & Palacios-Vargas, 1998; S. schatzi; S. simplex Ríos & Palacios-Vargas, 1998; S. volcanicus). Ventral plate. Epimeral setae: The most common epimeral setal formula is 3-1-2-2 (47 spp.) then rarely 3-1-3-3 (S. baloghius; S. cuspidatus; S. nashiroi Nakatamari, 1989; S. schatzi and S. subcorniger); 3-1-2-3 (S. hungarorum); 3-1-1-2 (S. pundamiliaensis); 2-1-2-2 (S. bisculpturatus Mahunka, 1984a) or 2-1-2-1 (S. pseudoreticulatus). Genital setae: there are six pairs of genital setae (though an undescribed species from Queensland has five) and they are deployed either in a linear fashion or with g1 lateral to the rest (cf. Aoki, 1972, his Fig. 19). Anal setae: The positioning of the anal setae along the medial edges of the anal plates is regarded by Behan-Pelletier (1989) as a synapomorphy for the genera Ametroproctus and Cymbaeremaeus, but this character state occurs in Scapheremaeus also. Pre-anal organ: A triangular pre-anal organ is present in many species (evident in illustrations of the ventral surface of S. argentinensis, S. cuspidatus, S. johnsi, S. longicuspis, S. nashiroi, S. palaciosi, S. retestriatus, S. subglaber, S. taurus and S. tosaensis. This structure is also present in other cymbaeremaeid taxa including Cymbaeremaeus (Walzl, 2000; Fujikawa, 2002), Ametroproctus, Coropoculia (Aoki, 1972; Behan-Pelletier, 1987), Seteremaeus Hammer, 1971 and Bulleremaeus Hammer, 1966. Subcapitulum. The subcapitulum is diarthric, with one pair each of setae h, m and a (Fig. 9). The plesiomorphic condition for the palp setal formula in the Brachypylina, and found in the cymbaeremaeid genera Ametroproctus, Cymbaeremaeus and Scapheremaeus is 0-2-1-3-9 (Behan-Pelletier, 1989b). Descriptions of palp setal formulae of Scapheremaeus spp. are few. Travé & Fernandez (1986) give the plesiomorphic condition for S. patella. Scapheremaeus nuciferosa Ramani & Haq, 1998 has 0-2-1-3-10. Travé & Fernandez, 1986 give the formula 0-2-1-2-9 (missing tibial seta v) for S. argentinensis. Fernandez & Cleva (1997) give the same formula for Scapheremaeus tillandsiae. Legs. The legs of Scapheremaeus are characterised by a squat, attenuated tarsus I and II, typically less than twice as long as broad. The anteriodorsal margin is strongly sloped or convex. Tibia I has a long tubular apophysis from which emerges solenidion φ1. Solenidion φ2 may be paired with φ1 (S. argentinensis, S. nuciferosa, S. tonatiuh), or positioned posterior of it (S. corniger, S. tillandsiae, S. palaciosi, S. tosaensis). The solenidial apophysis overhangs the tarsus for most of its length (Fig. 5a). Some 34 species are monodactylous and 27 are heterotridactylous. Partial or complete leg chaetotaxy is only known for 12 species of Scapheremaeus including those new species described herein. The most common setal formula for leg I is 14-2-4-14 (solenidiotaxy 1-2-2). Leg tracheal organs in the form of sacculi or porose areas are found on certain leg segments of Cymbaeremaeidae (Behan-Pelletier, 1989b). In Scapheremaeus the pattern of their arrangement is variable. They are present on trochantera and femora I-IV in S. argentinensis and S. pulleni sp. nov., on these and tibia IV in S. cheloniella sp. nov., but only on trochantera and femora III and IV in S. tillandsiae and I and II in S. angusi sp. nov.

Species-groups Species-groups, whilst having no formal taxonomic validity, represent a useful category between genus and species. The species-group is particularly valuable as a precursor to generic revisions, to better define SCAPHEREMAEUS (ACARI: ORIBATIDA)


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characters, and in formulating working hypotheses on phylogenetic and zoogeographical relationships of large, heterogeneous, cosmopolitan genera (Colloff, 2009). Species-groups allow for a flexible classification without the formality of establishing subspecies names. Despite Scapheremaeus having a reputation as a difficult genus, some species-groups are discernible based on morphotypes. The genus can be divided into two main categories: i) species that have a complete circumdorsal scissure with plicate microsculpture on the circumnotogastral plate and strongly contrasting microsculpture (foveolae, ridges or tubercles) on the centrodorsal plate (plicate species-groups), and ii) species with the circumdorsal scissure complete, incomplete or absent but with little or no contrast in microsculpture between the central and lateral regions: typically both regions foveolate or reticulate (nonplicate species-groups). Plicate species-groups 1. Carinatus group (Fig. 2a): with a complete circumdorsal scissure; prodorsum usually with well-developed lateral carinae. Typically there are 10 pairs of notogastral setae. Often with a centrodorsal ridge on the notogaster. Generally there is a trans-costular ridge and the lamellar setae and their apophyses are either short, not extending as far as the rostrum (S. arboreus, S. angusi sp. nov., S. bisculpturatus, S. lambieae sp. nov., S. carinatus, S. cheloniella sp. nov., S. clavifer, S. convexus, S. ewani sp. nov., S. latirostris Willmann, 1936, S. latus, S. nogueraius, S. nuciferosa Ramani and Haq, 1998, S. ornatus, S. pulleni sp. nov., S. pundamiliaensis, S. schatzi and S. tillandsiae), or alternatively, the lamellar apophyses and setae are long, well-developed, often extending as far as the rostrum (S. balazsi, S. bicornutus, S. chaac, S. fisheri, S. insularis, S. semiconvexus and S. uncinatus). 2. Petrosus group (Fig. 2b): with a complete circumdorsal scissure; prodorsum without lateral carinae. Costulae, lamellar setae and their apophyses are short (S. guerinii, S. johnsi, S. palaciosi, S. petrosus, S. taurus and S. yamashitai). Typically there are 10 pairs of notogastral setae, with those of p series positioned on the ventral circumnotogastral plate. 3. Fimbriatus group (Fig. 2c): with a complete circumdorsal scissure; prodorsum without lateral carinae. Costulae, lamellar setae and their apophyses short. With long, dagger-like humeral processes. Typically with 14 pairs of notogastral setae.With circular mound-like structure medially on the circumnotogastral plate in the caudal region (S. corniger, S. fimbriatus, S. madeirensis and S. subcorniger). 4. Patella group (Fig. 2d): with a complete circumdorsal scissure; prodorsum without lateral carinae. Costulae, lamellar setae and their apophyses short. Humeral processes sub-triangular; typically with 14 pairs of notogastral setae. Setae of p series on the dorsal circumnotogastral plate (S. clavisetus, S. demeteri, S. patella and S. rustenburgensis). 5. Polysetosus group (Fig. 2e): with a complete circumdorsal scissure and 15 pairs of notogastral setae. Circumnotogastral plate plicate, centrodorsal plate with transverse ridges and foveae. Humeral process very strongly developed, visible from above and with seta c3 positioned on or adjacent to humeral process. Lamellar apophyses elongated (S. digitatus, S. polysetosus, S. quadrireticulatus and S. retestriatus). 6. Longilamellatus group (Fig. 2f): with a poorly-developed or incomplete circumdorsal scissure but with a plicate margin and alveolate centrodorsal plate; with costulae and lamellar apophyses long and prominent, extending well beyond rostrum. Notogastral and lamellar setae with black, club-shaped cerotegument. Angular humeral margins, but without well-developed humeral processes. Caudal region scalloped. With 10 or 14 pairs of notogastral setae. (S. cellulatifer, S. longicuspis and S. longilamellatus).

Non-plicate species-groups 7.

Alveolatus group (Fig. 3a): circumdorsal scissure complete; with circumnotogastral and centrodorsal plates typically ornamented with foveolae, but centrodorsal plate with one, two, three or four longitudinal

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ridges. Typically with 10 pairs of notogastral setae.. With caudal indentation. Prodorsum with welldeveloped lateral carinae, linked to costulae by posterior trans costular ridges in the form of a diagonal cross, or simple, transverse (S. alveolatus, S. arcuatus, S. eugenius, S. flamiferus, S. foveolatus, S. grahamius, S. nashiroi, S. obliteratus, S. pisacensis, S. quadrilineatus, S. sinuosus, S. tillandsiophilus, S. tosaensis, S. tricarinatus, S. trirugis and S. undosus). 8. Simplex group (Fig. 3b): circumdorsal scissure complete; circumnotogastral and centrodorsal plates typically ornamented with foveolae; centrodorsal longitudinal ridges absent. Broad centrodorsal ridge extended posteriorly. Humeral processes present or absent. With 10 or 14 pairs of notogastral setae. Without caudal indentation. Prodorsum with lamellar costulae anterior transcostula and well-developed lateral carinae with apical projections in the shape of an inverted V; sensillus large, elongated (S. baloghius, S. hectorperezius, S. magdalenae and S. simplex). 9. Palustris group (Fig. 3c): circumdorsal scissure complete; circumnotogastral and centrodorsal plates typically ornamented with foveolae; No longitudinal dorsal ridges, though broad centrodorsal ridge may be extended posteriorly. Humeral processes absent or small, sub-triangular. With caudal indentation. With 10 pairs of notogastral setae. Prodorsum with lamellar costulae, anterior transcostula and often posterior transcostula. Without well-developed lateral carinae. (S. fungisetosus, S. mahunkaius, S. morulisensillatus, S. palustris S. pseudoreticulatus, S. reticulatus, S. tonatiuh and S. volcanicus). 10. Argentinensis group (Fig. 3d): circumdorsal scissure often faint and incomplete; circumnotogastral and centrodorsal plates typically ornamented with foveolae or tubercles; centrodorsal longitudinal ridges absent. Humeral processes absent and humeral margin rounded, continuous. With 10 or 14 pairs of notogastral setae. With caudal indentation. No longitudinal dorsal ridges, though broad centrodorsal ridge may be extended posteriorly. Prodorsum with lamellar costulae and anterior transcostula. No lateral carinae (S. argentinensis, S. glaber, S subglaber and S. stratus). 11. Cuspidatus group (Fig. 3e): with poorly-developed or incomplete circumnotogastral scissure (except S. crassus Mahunka, 1988); with costulae and lamellar apophyses long and prominent, extending as far as the rostrum, or beyond. With anterior and posterior transcostula. Notogastral and lamellar setae often with black, club-shaped cerotegument. and well-developed humeral processes - often long and curved. Typically with 10 pairs of notogastral setae. (S. cornutus, S. crassus, S. cuspidatus, S. semiornatusand S. tonatiuh). 12. Humeratus group (Fig. 3f): anterior portion of circumnotogastral scissure is present as a long lateral scissure, extending into the humeral regions, sometimes with lateral vestiges present. With highly modified, reduced circumnotogastral scissure and no contrast in microsculpture between centrodorsal and marginal regions. With 10, 12 or 14 pairs of notogastral setae. Sensillus markedly elongated. With welldeveloped ridge-like lateral carinae. Lamellar setae and their apophyses short (S. alvarezius, S. bituberculatus, S. humeratus and S. hungarorum). 13. Emarginatus group (Fig. 3g): with no circumnotogastric scissure, and centrodorsal and marginal notogastral regions indistinguishable; notogaster sub-rectangular or U-shaped; with foveolate or polygonate notogastral microsculpture; with diffuse, complex series of prodorsal ridges, including lateral carinae. With 10 pairs of notogastral setae. Lamellar apophyses short. Humeral processes absent or reduced (S. emarginatus, S. morenoi and S. striatomarginatus). Scapheremaeus acuminatus, S. marginalis and the species recombined with Scapheremaeus by Marshall et al. (1987) (S. hieroglyphicus, S. marmoratus, S. parvula and S. pulchellus) could not be placed within speciesgroups. Norton & Poinar (1993) referred the fossil species from miocene amber, Scapheremaeus brevitarsus Woolley 1971, to Liodes, then Heethoff et al. (2009) recombined it with Neoliodes. Cymbaeremaeus cyclops Oudemans 1915 is hereby recombined with Scapheremaeus on the basis of it lacking interlamellar and lamellar setae but possessing a lenticulus; short, black, club-shaped sensilla; costulae; circumnotogastral plates; a complete dorsosejugal suture with humeral projections; prodorsal and

SCAPHEREMAEUS (ACARI: ORIBATIDA)


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centrodorsal foveolae; six pairs of genital setae with g2 lateral of g1; and two pairs of anal setae positioned on central region of plates. It cannot be assigned to a species group without a redescription. Scapheremaeus petrophagus (Banks 1906) was redescribed and recombined from Scutovertex by Jacot (1934), who mentioned that the species was "…not typical of the genus Scapheremaeus [but]…sufficiently close to warrant its inclusion." The species is lenticulate but lacks a circumdorsal scisssure. Scapheremaeus petrophagus differs from other members of the genus as follows: i) the dorsosejugal suture is much more acute, projecting forward as far as the anterior margins of the bothridia, and abutting them. In other Scapheremaeus spp. the dorsosejugal suture is transverse, slightly bowed anteriorly, and the bothridia are usually positioned some distance anterior of the dorsosejugal furrow; ii) Scapheremaeus petrophagus has 15 pairs of notogastral setae: the most common condition for Ameronothridae. Although 15 pairs are present in Scapheremaeus as outlined above, 14 pairs are more common; iii) the centrodorsal microsculpture is ridged and reticulate and that of the marginal region is tuberculate. The only other Scapheremaeus with clearly tuberculate microsculpture is S. johnsi, although the centrodorsal region of S. clavifer Hammer was described with ‘irregular indistinct knobs, perhaps due to the secretion’ (i.e. cerotegument); iv) the epimeral setal formula is 2-1-2-2, whereas in other Scapheremaeus spp. it is 3-1-2-2, or more rarely 3-1-2-3, except S. pseudoreticulatus where it is 2-1-2-1. In Jacot's habitus drawing (1934, Fig 4), the species looks much more like an ameronothrid based on the ratio of the length of the legs to the body and the unideficient chaetome (only found in the Polysetosus group). Scapheremaeus petrophagus has complete prodorsal-notogastral separation (i.e. an entire dorsosejugal suture), as do several ameronothrid genera including Chudalupia and Aquanothrus, but it does not fit existing ameronthrid genera and is a member of a new genus (R.A. Norton, personal communication, 2009).

New diagnosis of Scapheremaeus Scapheremaeus Berlese, 1910 Scapheremaeus Berlese, 1910 Type-species: Cymbaeremaeus (Scapheremeus) patella Berlese, 1910, p. 226. Mulvius Sellnick, 1918: Sellnick, 1931, p. 164; Norton, 2006, p. 112.

Diagnosis. Adults of this genus have the following unique combination of character states within the Cymbaeremaeidae: ornamented notogaster with a lenticulus and a complete or incomplete circumdorsal scissure, occasionally reduced. Sensillus typically darkly pigmented, globose, club-shaped apically. Genital and anal apertures not closely adjacent. Description (Adult). Brachypyline oribatid mites. Notched tutorium absent, but a lateral carina may be present in some species. Lamellar setae setiform, bacilliform and/or darkly pigmented and club-shaped due to cerotegument; lamellar seta associated with lamellar ridge or costula, emerging from well-developed lamellar apophysis. Interlamellar setae setiform or peg-like, represented only by alveoli or absent. Bothridium welldeveloped, not obscured by notogaster in dorsal view; sensillus rounded in cross section; sensillus clavate, smooth or ornamented with spicules, longitudinal pleats or tubercles; often darkly pigmented. Exobothridial setae present or absent. Dorsosejugal suture continuous, convex; notogaster with complete or incomplete circumdorsal scissure, separating centrodorsal plate from dorsal circumnotogastral plate; occasionally scissure reduced so as to be visible only by difference in lateral and centrodorsal microsculpture; rarely absent. Usually 3 pairs of lyrifissurae concentrated posteriorly on circumnotogastral rim (where present); central part of notogaster with microsculpture of tubercles and/or ridges or alveoli, partly cerotegumental in origin. Lenticulus present on notogaster anteriomedially. With typically 10, 14 or 15 pairs of setiform notogastral setae, more rarely 7, 12 or 13 pairs. With or without humeral spines extending ventrally. Epimeral

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setal formula 3-1-2-2 or rarely 3-1-3-3 (2-1-2-1, 3-1-1-2 are known in individual species). Junction of epimere IV and ventral plate without enantiophyses; caudal margin of venter U-shaped, not V-shaped; genital and anal plates separated by distance equivalent to between that of the width of the genital aperture and half its width; 4, 6 or 7 pairs of genital setae; one pair of aggenital setae; two pairs of anal setae situated on median edge of plates or some distance from median edge; 3 pairs of adanal setae. With triangular pre-anal sclerite. Legs monodactylous or tridactylous.

Materials and methods Generally, the morphological terms used herein are those of Grandjean (cf. Travé and Vachon, 1975 for references), Hunt et al. (1998) and Norton and Behan-Pelletier (2009). Body length (in micrometres) was measured in ventral view from the tip of the rostrum to the posterior margin of the dorsal notogastral plate. Breadths were measured in dorsal view at the point of the widest part of the notogaster. Length and breadth are given for all types. Other body measurements are for the holotype only. Except in the case of Scapheremaeus ewani sp. nov. and S. lambieae sp. nov., the legs were dissected and mounted in Heinze PVA in order to examine the chaetotaxy. Lengths of leg segments, including portions inserted into the next proximal segment, are given in sequence from femur to tarsus. Lengths of trochantera are excluded. Setal formulae of the legs include the number of solenidia in parentheses. The famulus is included in the tarsal setal formula. All descriptions refer to adult specimens. All type material is deposited in the Australian National Insect Collection (ANIC), CSIRO Entomology, Canberra.

Descriptions of New Species Scapheremaeus cheloniella sp. nov. (Figs. 4, 5) Dimensions: Length: holotype: 529; paratype: 533. Breadth: holotype: 280; paratype: 293. Prodorsum. Tuberculate laterally, with rugose pattern of poorly-defined ridges medially between costulae. Rostrum pointed, rostral setae (ro) smooth, curved, ca. 19 long; lamellar setae (le; ca. 16 long) on small tubercles (ca. 5 long) slightly longer than broad, connected by translamella. Costulae well developed, moreor-less parallel, bowed laterally, not extending posteriorly as far as bothridia. Alveoli of interlamellar setae (in) present. Sensillus club-shaped, darkly-pigmented; head ca. 20 broad with tuberculate microsculpture, apex extending beyond prodorsal margin. Notogaster. Length 430 μm. Circumdorsal scissure entire, with diagonal extensions into humeral region. Lenticulus present (ca. 38 long, 30 broad). Cerotegument consisting of dark, coarse nodules. Centrodorsal region 398 long, 205 broad; with microsculpture of linearly-arranged tubercles of cerotegument overlaying colliculate plaques. Microsculpture of region lateral of circumdorsal scissure consisting of parallel ridges and troughs posteriorly and scattered tubercles anteriorly. Ten pairs of smooth, spiniform notogastral setae borne on short tubercles; lm and lp positioned on centrodorsal region. Caudal region indented slightly. Lyrifissurae ih, ips and ip arranged radially on posterior half of laterodorsal region. Ventral region. Epimeral setal formula, numbers of aggenital, anal and adanal setae typical of genus; 6 pairs of genital setae; g2 offset laterally from g1. Ventral microsculpture similar to that of centrodorsal region. Lyrifissurae iad in para-anal position.



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FIGURE 4. Scapheremaeus cheloniella sp. nov. a) dorsal aspect; b) ventral aspect; c) lateral aspect. Cd = centrodorsal plate; Cf = circumferential scissure; Cgs = circumgastric scissure; Cnd = dorsal circumnotogastral plate; Cnv = ventral circumnotogastral plate; Cs = circumdorsal scissure; Hu = humeral extension; Vp = ventral plate.

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FIGURE 5. Scapheremaeus cheloniella sp. nov. a): right leg I, antiaxial view; b): right leg II, antiaxial view; c): right leg III, paraxial view; d): right leg IV, paraxial view.

Lateral aspect. Prodorsal microsculpture composed of round, oval and elongate tubercles arranged in linear patterns. Costulae projecting markedly above surface of prodorsum; tubercle of lamellar seta free of prodorsal surface. Carina an angular ridge ventral of lamella; cusp or free projection absent. Pedotectum I SCAPHEREMAEUS (ACARI: ORIBATIDA)


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auriculate, covering acetabulum I; pedoctectum II midway between acetabula II and III, apex directly ventral of short humeral spine. Gnathosoma. Typical of family. Tectum of mentum slight, not extending anteriorly as far as bases of setae a. Legs. Setal formulae: legs I: 1-4-2(1)-4(2)-14(2); legs II: 1-3-3(1)-3(1)-12(1); legs III: 1-3-2-3(1)-12; legs IV: 1-2-2-3(1)-12. Lengths of leg segments (femur to tarsus): legs I: 100, 33, 67, 32; legs II: 91, 33, 58, 30; legs III: 70, 32, 57, 34; legs IV: 77, 32, 68, 36. Femora I-IV with tuberculate microsculpture on antiaxial surface and series of parallel ridges on paraxial surface; circular porose areas present on paraxial surfaces. Elongate porose area present on paraxial surface of tibia IV. Claws heterotridactylous. Material examined: Holotype and paratype: pitfall trap, Mallee eucalypt vegetation on dune, 14 km WNW Renmark, 34°07’S 140°37’E, South Australia; coll. A. Lambie, 4-24 July, 1995. Types in ANIC. Etymology: The specific name, cheloniella, is Latinised Greek for ‘little tortoise’. Remarks. Scapheremaeus cheloniella sp. nov. can be distinguished from other members of the genus by the following combination of characters: 1) microtuberculate, tesellated centrodorsal microsculpture; 2) a complete circumdorsal scissure with humeral extensions; 3) 10 pairs of setiform notogastral setae, those of the p series on dorsal circumnotogastral plate; 4) alveolus of interlamellar seta present; 5) caudal margin indented; 6) prodorsal microsculpture tuberculate; 7) humeral process short, sub-triangular. This species is most similar morphologically to S. lambieae in the pattern of the centrodorsal microsculpture, but the latter species has no caudal indentation, a posterior centrodorsal rige with lateral concavities and tuberculate protuberances on either side of the lenticulus.

Scapheremaeus pulleni sp. nov. (Figs. 6, 7) Dimensions: Length: holotype: 430; paratype: 455. Breadth: holotype: 242; paratype: 268. Integument. Cerotegument consisting of fine dark nodules. All surfaces of the body with ornate microsculpture Prodorsum. Tuberculate laterally with pattern of diagonal ridges medially between costulae. Rostrum pointed, rostral setae (ro) smooth, curved, ca. 13 long; lamellar setae (le; ca. 5 long) as long as the tubercles from which they emerge; translamella present. Costulae well developed, more-or-less parallel, extending posteriorly as far as bothridia. Interlamellar setae absent. Sensillus club-shaped, darkly-pigmented, head ca. 12 broad, with tuberculate microsculpture, apex barely extending beyond prodorsal margin. Notogaster. Length 323 . Circumdorsal scissure entire, with diagonal extensions into humeral region. Lenticulus present (ca. 35 long, 26 broad), flanked by tuberculate protuberances. Centrodorsal region 253 long, 177 broad; with microsculpture consisting of linearly-arranged elongate plaques. Microsculpture of region lateral of circumdorsal scissure consisting of parallel ridges and troughs posteriorly and scattered tubercles anteriorly. Ten pairs of smooth, minute (ca. 5–7 long) spiniform notogastral setae; those of h and ps series borne on short tubercles; lm and lp positioned on centrodorsal region. Caudal region convex. Lyrifissurae ih, ips and ip present on posterior half of laterodorsal region; ih and ips parallel, oblique. Lateral aspect. Microsculpture composed of round, oval and elongate tubercules arranged in linear patterns. Costulae projecting above surface of prodorsum, but tubercle of lamellar setae not free. Tutorium consisting of angular ridge ventral of lamella; cusp or free projection absent. Pedotectum I auriculate, covering acetabulum I; pedoctectum II bladelike, positioned slightly posterior of acetabula II. Humeral spine short, broad, with two blunted apices. Ventral region. Epimeral setal formula, numbers of aggenital, anal and adanal setae typical of genus; 6 pairs of genital setae; g2 offset laterally from g1. Ventral microsculpture similar to that of lateral region. Lyrifissurae iad in para-anal position.

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FIGURE 6. Scapheremaeus pulleni sp. nov. a): dorsal aspect; b) ventral aspect; c) lateral aspect.



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FIGURE 7. Scapheremaeus pulleni sp. nov.; right legs, paraxial view: a) leg I; b) leg II; c) leg III; d) leg IV.

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Gnathosoma. Typical of family. Tectum of mentum slight, not extending anteriorly as far as bases of setae a. Legs. Setal formulae: legs I: 1-3-2(1)-4(2)-14(2); legs II: 1-3-2(1)-2(1)-13(1); legs III: 1-2-1(1)-3(1)-12; legs IV: 1-2-1(1)-3(1)-12. Lengths of leg segments (femur to tarsus): legs I: 86, 28, 58, 26; legs II: 83, 22, 51, 22; legs III: 70, 19, 55, 24; legs IV: 68, 23, 51, 31. Femora I-IV with tuberculate microsculpture on antiaxial surface and series of parallel ridges on paraxial surface; circular porose areas present on paraxial surfaces. Claws heterotridactylous. Material examined: Holotype and paratype: pitfall trap, Mallee eucalypt vegetation on dune-swale system, 14 km WNW Renmark, 34°07’S 140°37’E, South Australia; coll. K. Pullen, 2 May–7 June, 1995. Types in ANIC. Etymology. This species is named after its collector, Kimberi R. Pullen (CSIRO Entomology) in recognition of his contribution to Australian entomology. Remarks. Scapheremaeus pulleni sp. nov. can be distinguished from other members of the genus by the following combination of characters: 1) the linear arrangement of elongated plaques on the centrodorsal region; 2) a complete circumdorsal scissure with humeral extensions; 3) 10 pairs of setiform notogastral setae, those of the p series on dorsal circumnotogastral plate; 4) a diagonal cross-shaped posterior transcostular ridge; 5) prodorsal microsculpture tuberculate; 6) humeral process short, sub-triangular, notched apically. This species is most similar morphologically to S. cheloniella but differs in the pattern of the centrodorsal microsculpture, and the shape of the posterior transcostular ridge and humeral process.

Scapheremaeus angusi sp. nov. (Figs. 8–10) Dimensions: Length: holotype: 531; paratypes (mean and range; n = 7): 623 (545–678). Breadth: holotype: 343; paratypes (mean and range; n = 7): 368 (308–418). Integument. Dark, granular cerotegument present. All surfaces of body with ornate microsculpture Prodorsum. Tuberculate laterally, with rugose pattern of poorly defined ridges medially between costulae. Rostrum pointed, rostral setae (ro) smooth, pointed, ca. 11 long; lamellar setae (le; ca. 13 long) with clubshaped tips of cerotegument, on small tubercles (ca. 8 long) slightly longer than broad, connected by translamela. Costulae well developed, more-or-less parallel, extending posteriorly as far as bothridia. Alveoli of interlamellar setae (in) present. Sensillus club-shaped, darkly-pigmented, tuberculate, head ca. 19 broad, apex extending well beyond prodorsal margin; stalk ca 25 long. Notogaster. Length 450. Circumdorsal scissure entire, with diagonal extensions into humeral region. Lenticulus present (ca. 41 long, 38 broad) flanked by tuberculate protuberances. Centrodorsal region 348 long, 233 broad; with microsculpture consisting of randomly-arranged minute foveolae. Anterior two thirds of centrodorsal region strongly ridged. Microsculpture of region lateral of circumdorsal scissure consisting of parallel ridges and troughs. Ten pairs of smooth, short (8–14 long) spiniform notogastral setae borne on short tubercles; lm and lp positioned on centrodorsal region. Caudal region indented slightly. Lyrifissurae ih, ips and ip arranged radially on posterior half of laterodorsal region. Lateral aspect. Prodorsal microsculpture composed of round, oval and elongate tubercles arranged in linear patterns. Costulae projecting markedly above surface of prodorsum; tubercle of lamellar seta free of prodorsal surface. Carina an angular ridge ventral of lamella; cusp or free projection absent. Pedotectum I auriculate, covering acetabulum I; pedoctectum II narrow, bladelike, midway between acetabula II and III, apex directly ventral of short humeral spine. Ventral region. Epimeral setal formula, numbers of aggenital, anal and adanal setae typical of genus; 6 pairs of genital setae; g2 offset laterally from g1. Ventral microsculpture consisting of series of ridges and plaques. Lyrifissurae iad in para-anal position.



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Subcapitulum. Mentum with ridged microsculpture; tectum of mentum slight, not extending anteriorly as far as bases of setae a. Setal formula of palp: 1-1-3-9(1); tarsal formula including 4 eupathidia. Tarsus with prominent tubercle bearing eupathidium acm; eupathidium not attached to solenidion.

FIGURE 8. Scapheremaeus angusi sp. nov. a) dorsal aspect; b) ventral aspect; c) lateral aspect.

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FIGURE 9. Scapheremaeus angusi sp. nov., subcapitulum.

Legs. Setal formulae: legs I: 1-4-2(1)-4(2)-14(2); legs II: 1-3-2(1)-3(1)-13(1); legs III: 1-2-1(1)-3(1)-12; legs IV: 1-2-1(1)-3(1)-11. Lengths of leg segments: legs I: 110, 38, 70, 32; legs II: 116, 44, 90, 40; legs III: 86, 30, 76, 38; legs IV: 80, 36, 82, 46. Femora I-IV with tuberculate microsculpture on antiaxial surface and series of parallel ridges on paraxial surface; circular porose areas present on paraxial surfaces. Claws heterotridactylous. Material examined: Holotype: pitfall trap, Casuarina woodland, 79 km NNW Renmark, 33°31’S 140°24’E; coll. A. Lambie, 3 May - 6 June, 1995. Paratypes: 1 adult, as above, coll. K. Pullen, 29 March–3 May, 1995. 1 adult, gutter of flight intercept trap, Mallee eucalypt vegetation on dune-swale system, 14 km WNW Renmark, 34°.07’S 140°37’E; coll. K. Pullen, 13 Dec, 1995–25 Jan, 1996. 1 adult, gutter of flight intercept trap, Mallee eucalypt vegetation on dune-swale system, 14 km WNW Renmark, 34°.07’S 140°37’; coll. K. Pullen, 13 Dec, 1995–25 Jan, 1996. 3 adults, data as above, 28 February –28 March, 1995. 1 adult, gutter of flight intercept trap, Mallee eucalypt vegetation on dune-swale system, 31 km NW Renmark, 33°59’S 140°30’; coll. K. Pullen, 30 March - 2nd May, 1995. 1 adult, malaise trap, Mallee eucalypt vegetation on dune-swale system, 32 km NW Renmark, 33°59’S 140°30’; coll. K. Pullen, 11 October–9 November, 1995. Types in ANIC. Etymology. This species is named after my youngest son, Angus John Colloff. Remarks. Scapheremaeus angusi sp. nov. can be distinguished from other members of the genus by the following combination of characters: 1) the minutely foveolate centrodorsal region; 2) a complete circumdorsal scissure with humeral extensions; 3) the massive centrodorsal ridge and posterior concave region; 4) the rugose inter-costular microsculpture; 5) short, sub-triangular humeral processes; 6) 10 pairs of setiform notogastral setae, those of the p series on dorsal circumnotogastral plate; 6) with palp setal formula 01-1-3-9; missing femoral seta l". This species is most similar morphologically to S. cheloniella but differs in the pattern of the centrodorsal microsculpture, and the shape of the posterior transcostular ridge and humeral process. SCAPHEREMAEUS (ACARI: ORIBATIDA)


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Scapheremaeus lambieae sp. nov. (Fig. 11) Dimensions: Holotype: length: 440; breadth: 255. Integument. Cerotegument consisting of dark, coarse nodules. All surfaces of body with ornate microsculpture.

FIGURE 10. Scapheremaeus angusi sp. nov., legs (left, antiaxial aspect). a) Leg I; b) Leg II; c) Leg III; d) Leg IV.

Prodorsum. Striate laterally and between costulae. Rostrum rounded, rostral setae (ro) smooth, straight, spinose, ca. 16 long; lamellar setae (le; ca. 13 long) on small tubercles, shorter than broad, connected by translamella. Costulae convergent anteriorly, consisting of thin ridges, extending posteriorly as far as

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bothridia. Interlamellar setae and their alveoli absent. Sensillus club-shaped, darkly-pigmented, head ca. 20 broad, with tuberculate microsculpture, apex not extending beyond prodorsal margin.

FIGURE 11. Scapheremaeus lambieae sp. nov. a) dorsal aspect; b) ventral aspect; c) lateral aspect.

Notogaster. Length 350. Circumdorsal scissure entire, with diagonal extensions into humeral region. Lenticulus present (ca. 30 long, 19 broad), flanked by tuberculate protuberances. Centrodorsal region 260 long, 167 broad; with microsculpture consisting of randomly-arranged tubercles. Microsculpture of region lateral of circumdorsal scissure consisting of parallel ridges and troughs and scattered tubercles. Ten pairs of SCAPHEREMAEUS (ACARI: ORIBATIDA)


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smooth, spiniform notogastral setae borne on short tubercles, covered in cerotegument giving them clubshaped appearance; lm and lp positioned on centrodorsal region. Caudal region indented slightly. Lyrifissurae ih, ips and ip present on posterior half of laterodorsal region; ih and ips parallel, oblique. Lateral aspect. Costulae barely projecting above surface of prodorsum; tubercle of lamellar seta on prodorsal surface, not free. Carina an angular ridge ventral of lamella; cusp or free projection absent. Pedotectum I auriculate, covering acetabulum I; pedoctectum II bladelike, midway between acetabula II and III, apex directly ventral of short, cusp-like humeral spine. Ventral region. Epimeral setal formula, numbers of aggenital, anal and adanal setae typical of genus; 6 pairs of genital setae; g2 offset laterally from g1. Ventral microsculpture consisting of series of ridges arranged in radial manner. Lyrifissurae iad in para-anal position. Gnathosoma. Typical of family. Tectum of mentum slight, not extending anteriorly as far as bases of setae a. Legs. Claws heterotridactylous. Chaetotaxy not examined in detail. Only lengths of leg segments I and II measurable without dissection: leg I 79, 31, 56, 29; leg II 74, 26, 49, 24. Material examined: Holotype: pitfall trap, Casuarina woodland, 79 km NNW Renmark, 33°31’S 140°24’E; coll. A. Lambie, 3 May–6 June, 1995. Holotype in ANIC. Etymology. This species is named after Anne Lambie (ex. Bookmark Biosphere Reserve, SA), who collected it. Remarks. Scapheremaeus lambieae sp. nov. can be distinguished from other members of the genus by the following combination of characters: 1) the tuberculate centrodorsal region; 2) a complete circumdorsal scissure with humeral extensions; 3) the posterior centrodorsal ridge with lateral concavities; 4) the striate prodorsum; 5) short, sub-triangular humeral processes; 6) 10 pairs of setiform notogastral setae with clubshaped cerotegument, those of the p series on dorsal circumnotogastral plate. This species is most similar morphologically to S. cheloniella (see remarks section for this species above).

Scapheremaeus ewani sp. nov. (Fig. 12) Dimensions: Length: holotype: 390; paratype: 394. Breadth: holotype: 220; paratype: 212. Integument. Cerotegument consisting of dark, coarse nodules. All surfaces of the body with ornate microsculpture Prodorsum. Tuberculate laterally, with rugose pattern of poorly defined ridges medially between costulae. Rostrum pointed, rostral setae (ro) smooth, curved, ca. 17 long; lamellar setae (le; ca. 8 long) bacilliform, on squat, poorly-defined tubercles. Costulae with translamellate structure in posterior half; converging anteriorly; ending posteriorly as far as bothridia. Interlamellar setae and their alveoli absent. Sensillus club-shaped, darkly-pigmented, head ca. 18 broad, tuberculate; apex not extending beyond prodorsal margin. Notogaster. Length 354. Circumdorsal scissure entire, with diagonal extensions into humeral region. Lenticulus present (ca. 31 long, 18 broad). Centrodorsal region 275 long, 157 broad with microsculpture consisting of transversely-directed ridges interspersed with tubercles. Microsculpture of region lateral of circumdorsal scissure consisting of parallel irregular ridges and troughs. Ten pairs of spiniform notogastral setae covered in dark cerotegument, giving them a club-shaped appearance; only seven pairs visible in dorsal view; lm and lp positioned on centrodorsal region. Caudal region convex. Lyrifissurae ih, ips and ip arranged radially on posterior half of laterodorsal region. Lateral aspect. Prodorsal microsculpture composed of round, oval and elongate tubercles arranged in linear patterns. Costulae projecting markedly above surface of prodorsum; tubercle of lamellar seta free of prodorsal surface. Carina an angular ridge ventral of lamella; cusp or free projection absent. Pedotectum I auriculate, covering acetabulum I; pedoctectum II midway between acetabula II and III, apex directly ventral of short humeral spine.

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Ventral region. Epimeral setal formula, numbers of aggenital, and anal setae typical of genus; 3 pairs of spiniform, darkened adanal setae; 6 pairs of genital setae; g2 in line with g1 along medial edge of genital plate. Caudal region trilobed. Ventral microsculpture consisting of series of ridges and tubercles. similar to that of centrodorsal region. Lyrifissure iad in para-anal position.

FIGURE 12. Scapheremaeus ewani sp. nov. (E) a) dorsal aspect; b) ventral aspect; c) lateral aspect.



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Gnathosoma. Typical of family. Tectum of mentum slight, not extending anteriorly as far as bases of setae a. Legs. Claws heterotridactylous. Material examined: Holotype and paratype: gutter of flight intercept trap, Casuarina woodland, 79 km NNW Renmark, 33°31’S 140°24’E; coll. K. Pullen, 29 March –3 May, 1995. Types in ANIC. Etymology. This species is named after my eldest son, Ewan MacLeod Colloff. Remarks. Scapheremaeus ewani sp. nov. can be distinguished from other members of the genus by the following combination of characters: 1) long curved rostral setae and bacilliform lamellar setae, 2) the incurved shape of the costulae with both a medial and posterior transcostular ridge; 3) a complete circumdorsal scissure with humeral extensions; 4) striae lateral of the costulae; 5) short, sub-triangular humeral processes; 6) 10 pairs of setiform notogastral setae with black, club-shaped cerotegument, those of the p series on ventral circumnotogastral plate; 7) a prominent centrodorsal ridge; 8) the centrodorsal microsculpture consisting of well-developed lateral ridges. This species is most similar morphologically to S. lambieae (see remarks section for this species above), but differs in the morphology of the costulae and the centrodorsal microsculpture.

Discussion The purpose of this paper is to provide a review, as the basis for a revision, of a large, complex, cosmopolitan genus containg several classical species. The question then rises why not simply undertake the full revision? The main reasons are because this task would take a long time, be logistically complex, and because new species are being described at a relatively rapid rate, and many others await description. This review may serve to provide some interim basic structure and comparative context for those researchers undertaking descriptions of new species. The species-groups presented herein thus provide a working hypothesis of the taxonomic relationships within the genus. It is anticipated that with revisionary work, especially of classical species, a clearer picture of the relatedness of species will emerge in time that can be linked more cogently to phylogeny and biogeography. In this review, emphasis is placed on the patterns of notogastral microsculpture as a basis for recognition of species-groups. The two major divisions are those species-groups with plicate microsculpture on the dorsal circumnotogastral plate and strongly contrasting microsculpture (foveolae, ridges or tubercles) on the centrodorsal plate and those species-groups with the circumdorsal scissure complete, incomplete or absent but with little or no contrast in microsculpture between the central and lateral regions: typically both regions foveolate or reticulate. Centrodorsal microsculpture appears to be a reliable character, showing relatively little variation in basic structure and form, and those new species described herein show consistent patterns of microtubercles (S. cheloniella sp. nov.), tubercles (S. lambieae sp. nov.), elongate linear plaques (S. pulleni sp. nov.), foveolae (S. angusi sp. nov.), and transverse irregular ridges and troughs (S. ewani sp. nov.) amongst all individuals of each species examined. The species-groups outlined herein show varying degrees of biogeographic consistency amongst their component species. The most consistent are Simplex (all Neotropical), Fimbriatus (all Palaearctic) and Polysetosus (all Oriental). Less consistent are Argentinensis (Neotropical, but with one Afrotropical species), Emarginatus (New Zealand and Java), Palustris group (Palaearctic and Neotropical), Longilamellatus (Oriental and Neotropical), Alveolatus group (predominantly Oriental and Neotropical), Cuspidatus and Humeratus (both circumtropical). Other groups show cosmopolitan distribution (Carinatus) or no clear biogeographic grouping (Caudalis and Petrosus). As a result of the present review of the adult characters of Scapheremaeus, the diagnosis of the family Cymbaeremaeidae by Behan-Pelletier (1987) requires only minor modification: notogastral setation usually 10, 13, 14 or 15 pairs (rarely 7 or 12 pairs); c1, c3 and d series present or absent. Cymbaeremaeidae are closely related to the Ameronothridae, Passalozetidae, Podacaridae and Scutoverticidae, all of which have immatures

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with plicate cuticle (Grandjean, 1954). Differentiation of Cymbaeremaeidae from these other families based on adult characters rests on the eupathidium acm being free of the solenidion and borne on a large tubercle on the palp tarsus, and the presence of a tectum on the mentum (Behan-Pelletier, 1989b). Given the extent of these differences, the elevation of Cymbaeremaeidae to superfamily status by Subías (2004) seems hard to justify.

Acknowledgements The specimens described herein were collected during the course of a survey by the CSIRO Entomology funded by the then Australian Nature Conservation Agency (now the Australian Government Department of Environment, Water, Heritage and the Arts). I thank Kim Pullen (CSIRO Entomology) and Anne Lambie (formerly of Australian Landscape Trust, Calperum Station, South Australia) for the collection and donation of specimens. I am very grateful to Dr Heinrich Schatz (Institut für Zoologie, Leopold-Franzens-Universität Innsbruck) for editing the manuscript and for bringing records of Scapheremaeus spp. to my attention. Finally, I thank Dr Valerie Behan-Pelletier (Agriculture and Agrifood Canada, Ottawa) and Professor Roy Norton (College of Environmental Science and Forestry, State University of New York, Syracuse) for their unflagging assistance, support and encouragement.

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Eos, 57, 201–212. Pérez-Iñigo, C. (1995) Oribátidos (Acari, Oribatei) hallados sobre plantas en la region de Los Monegros (Aragon, España). Miscellania Zoologica, 18, 41–46. Pérez-Iñigo, C. & Peña, M.A. (1995) Oribátidos (Acari, Oribatei) de Fuerteventura (Islas Canarias). Graellsia, 51, 143– 164. Perez-Iñigo, C. & Peña, M.A. (1996) Oribátidos edáficos (Acari, Oribatei) de Gran Canaria (II). Boletin de la Asociacion española de Entomologia, 20(1–2), 201–219. Pérez-Iñigo, C. & Subías, L.S. (1974) Redescription de Scapheremaeus corniger (Berlese, 1908) (Acari, Oribatei). Acarologia, 16(4), 739–745. Pullen, K.R. (1997) A Survey of the Invertebrates of Calperum Station, Bookmark Biosphere Reserve, South Australia. Report prepared on behalf of Environment Australia. CSIRO Entomology, Canberra. 31 pp. Ramani, N. & Haq, M.A. (1998) Oribatid mites from coconut palm-5. A new species of Scapheremaeus Berlese, 1910 (Acari: Cymbaeremaeidae) from Kerala, India. Entomon, 23(1), 55–60. Ríos, G. (1996) A peculiar new Scapheremaeus species from Mexico (Acari: Oribatida). Folia Entomologica Hungarica, 62, 237–241. Ríos, G. & Palacios-Vargas, J.G. (1998) Especies nuevas de Scapheremaeus (Acari: Oribatei: Cymbaeremaeidae) de Mexico. Anales Instituto de Biologia Universidad Nacional Autonomia de Mexico, Serie Zoologia, 69(2), 181-215. Schatz, H. (1983) U.-Ordn.: Oribatei, Hornmilben. Catalogus Faunae Austriae Teil IXi. Verlag der Österreichischen Akademie der Wissenschaften, Wien. 118 pp. Sellnick M. (1918) Die Oribatiden der Bernsteinsammlung der Universität Königsberg i. Pr. Schriften der physikalischökonomischen Gesellschaft zu Königsberg, 59, 21–42. Sellnick, M. (1924) Oribatiden. In: Dampf, A. Zur Kenntnis der estländischen Hochmoorfauna. Sitzungsberichte der Naturforscher Gesellschaft zu Dorpat, 31(1–2), 65–71. Sitnikova, L.G. (1975) Superfamily Cymbaeremaeoidea, pp. 236–242 In: Ghilyarov, M.S. (Ed.). A key to the Soilinhabiting Mites Sarcoptiformes. Nauka, Moscow, 491 pp. [in Russian]. Subías, L.S. (2004) Listado sistemático sinonímico y biogeográfico de los ácaros oribátidos (Acariformes, Oribatida) del mundo (1758–2002). Graellsia, 60 (suppl.), 3–305. Subías, L.S. & Gil-Martin, J. (1997) Systematic and biogeographic checklist of oribatids from Western Mediterranean (Acari, Oribatida) Annali del Museo Civico di Storia Naturale “G. Doria”, 91, 459–498. Travé, J. & Fernandez, N. (1986) Contribution à la connaissance du genre Scapheremaeus: S. argentinensis, n. sp. (Oribate). Acarologia, 27(4), 349–359. Travé, J. & Vachon, M. (1975) François Grandjean 1882–1975 (Notice biographique et bibliographique). Acarologia, 17, 1–19. Walter, D.E. (1995) Dancing on the head of a pin: mites in the rainforest canopy. Records of the Western Australian Museum, Supplement Number 52, 49–53. Walter, D.E. & Behan-Pelletier, V.M. (1993) Systematics and ecology of Adhaesozetes polyphyllos sp. nov. (Acari: Oribatida: Licneremaeioidea), a leaf-inhabiting mite from Australian rainforests. Canadian Journal of Zoology, 71, 1024–1040. Walter, D.E., O'Dowd, D. & Barnes, V. (1994) The forgotten arthropods: foliar mites in the forest canopy. Memoirs of the Queensland Museum, 36(1), 221–226. Wang, H-F. (1998) Studies on the genus Scapheremaeus in China (Acari: Oribatida: Cymbaeremaeidae) Acta Zootaxonomica Sinica, 23(3), 252–263. [in Chinese] Willmann, C. (1936) Zoologische Ergebnisse einer Reise nach Bonaire, Curaçao und Aruba im Jahre 1930. Zoologische Jahrbücher Abteilungen für Systematik, 67(5–6), 429–442. Willmann, C. (1939) Die Arthropodenfauna von Madeira nach den Ergebnissen der Reise von Prof. Dr. O. Lundblad. Juli-August 1935. Arkiv für Zoologie, series A., 31(10), 1–42. Yen, A.L., Ewart, D. & Walker, K. (2006) Mallee eucalypts, hummock grasses and social insects - key elements of the Victorian mallee. Proceedings of the Royal Society of Victoria, 118, 281–293.

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Appendix 1. Catalogue of Scapheremaeus species of the world described prior to the present work. †= fossil species. †Scapheremaeus acuminatus Sellnick, 1931 Scapheremaeus acuminatus Sellnick, 1931, p. 164, fig. 20. Type depository: Holotype was in the Geologische-paläontologisches Institut, Universität Konigsberg, but is either lost or in the Geowissenschaftliches Zentrum der Universität Göttingen (Norton, 2006, p. 119). Habitat: in Baltic amber.

Scapheremaeus alvarezius Rios & Palacios-Vargas, 1998 Scapheremaeus alvarezius Rios & Palacios-Vargas, 1998, 211, fig. 16. Type depository: Department of Biology, Universidad Nacional Autonomia de Mexico, Mexico City. Type locality: Los Tuxtlas, Veracruz, MEXICO. Habitat: from leaf litter of Nectandra ambigens and Ficus yoponensis in the canopy of Astrocarium mexicanum.

Scapheremaeus alveolatus Hammer, 1961 Scapheremaeus alveolatus Hammer, 1961, 31, fig. 23. Type depository: Zoologisk Museum, Copenhagen. Type locality: Machu Picchu, PERU. Habitat: moist Polytrichum on wall.

Scapheremaeus arboreus Corpuz-Raros, 1979 Scapheremaeus arboreus Corpuz-Raros, 1979, 50, fig. 22A–D. Type depository: Department of Entomology, University of the Philippines at Los Baños. Type locality: National Botanic Gardens, Siniloan, Laguna, PHILIPPINES. Habitat: from leaves of Gonocaryum calleryanum (holotype); paratypes all collected from foliage also.

Scapheremaeus arcuatus Hammer, 1971 Scapheremaeus arcuatus Hammer, 1971, 31, fig. 33. Type depository: Zoologisk Museum, Copenhagen. Type locality: Viti Levu, FIJI. Habitat: in wet Polytrichum on a rock, about 1–2 cm. above the water level in a brook in rain forest.

Scapheremaeus argentinensis Travé & Fernandez, 1986 Scapheremaeus argentinensis Travé & Fernandez, 1986, 349, figs. 1–4. Type depository: Laboratoire Arago, Baynuls-sur-Mer. Type locality: Cordon del Plata, Vallecitos, Mendoza Province, ARGENTINA. Habitat: amongst saxicolous lichens.

Scapheremaeus balazsi Mahunka, 1983 Scapheremaeus balazsi Mahunka, 1983, 217, figs. 29–32. SCAPHEREMAEUS (ACARI: ORIBATIDA)


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Type depository: Hungarian Natural History Museum, Budapest. Type locality: Albina, SURINAM. Habitat: soil.

Scapheremaeus baloghius Rios & Palacios-Vargas, 1998 Scapheremaeus baloghius Rios & Palacios-Vargas, 1998, 96, figs. 8, 9. Type depository: Department of Biology, Universidad Nacional Autonomia de Mexico, Mexico City. Type locality: “Chamela” Biological Station, Jalisco, MEXICO. Habitat: from canopy (fogging), tropical dry forest.

Scapheremaeus bicornutus Hammer, 1971 Scapheremaeus bicornutus Hammer, 1971, 30, fig. 32. Scapheremaeus bicornutus: Hammer, 1972, 34. Type depository: Zoologisk Museum, Copenhagen. Type locality: Viti Levu, FIJI. Habitat: on Hibiscus sp. [imported from Viti Levu] found at the quarantine station, Washington, D.C. (Hammer, 1971); rotten leaves on a lawn (Hammer, 1972) Distribution: Fiji (Hammer, 1971); Tahiti (Hammer, 1972)

Scapheremaeus bisculpturatus Mahunka, 1984 Scapheremaeus bisculpturatus Mahunka, 1984, 130, figs. 47–50. Type depository: Museum d'Histoire Naturelle, Geneva and Hungarian Natural History Museum, Budapest. Type locality: Estancia Viancho Postillon, 5 km E. Puerto Max, PARAGUAY. Habitat: sievings from forest gallery.

Scapheremaeus bituberculatus Wang, 1998 Scapheremaeus bituberculatus Wang, 1998, 255, fig. 4. Type depository: Institute of Zoology, Chinese Academy of Sciences, Beijing. Type locality: Mount Meihua, Liancheng, Fujian Province, CHINA. Habitat: plant litter.

Scapheremaeus carinatus Willmann, 1936 Scapheremaeus carinatus Willmann, 1936, 431, pl. 14, fig. 1. Type depository: [?] Zoologische Staatssamlung, Munich. Type locality: Bonaire, LESSER ANTILLES. Habitat: in dead coral in a salt lake.

Scapheremaeus cellulatifer Mahunka, 1987 Scapheremaeus cellulatifer Mahunka, 1987, 262, figs. 17–21. Type depository: Museum d'Histoire Naturelle, Geneva and Hungarian Natural History Museum, Budapest. Type locality: Ha San Binh, Tan Lac (20 km. from Hoa Bin), VIETNAM.

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Habitat: material netted from a stream valley.

Scapheremaeus chaac Rios & Palacios-Vargas, 1998 Scapheremaeus chaac Rios & Palacios-Vargas, 1998, 184, fig. 2. Type depository: Department of Biology, Universidad Nacional Autonomia de Mexico, Mexico City. Type locality: Uxmal, Yucatan, MEXICO. Habitat: in rocky habitat on an archaeological site.

Scapheremaeus clavifer Hammer, 1958 Scapheremaeus clavifer Hammer, 1958, 36, fig. 36. Type depository: Zoologisk Museum, Copenhagen. Type locality: Rio Caldera Valley nr. Salta, ARGENTINA. Habitat: in thin layer of slightly moist moss on chalk cliff, sheltered by trees.

Scapheremaeus clavisetus Mahunka, 1978 Scapheremaeus clavisetus Mahunka, 1978, 322, figs. 26, 27. Type depository: Museum d'Histoire Naturelle, Geneva and Hungarian Natural History Museum, Budapest. Type locality: Île Ronde, MAURITIUS. Habitat: under litter of Pandanus leaves in dry stream bed.

Scapheremaeus convexus Hammer, 1979 Scapheremaeus convexus Hammer, 1979, 48, fig. 86. Type depository: Zoologisk Museum, Copenhagen. Type locality: Botanical Gardens, Bogor, JAVA. Habitat: amongst dead leaves, mosses, lichens, liverworts, Selaginella and dead leaves and debris under very tall, dark trees, dense under-vegetation and numerous bushes and flowers.

Scapheremaeus corniger (Berlese, 1908) Cymbaeremaeus corniger Berlese, 1908, 11. Cymbaeremaeus (Scapheremaeus) corniger Berlese, 1910, 227, pl. 19, fig. 62. Scapheremaeus corniger: Perez-Iñigo & Subías, 1974, 739, figs. 1–6. Type depository: Berlese Acaroteca, Firenze. Type locality: Portici, ITALY. Habitat: no data given (Berlese, 1908); xerophytic shrubs amongst littoral sand dunes (Perez-Iñigo & Subías, 1974). Distribution: Italy (Berlese, 1908); Spain (Perez-Iñigo & Subías, 1974); Canary Islands (Perez-Iñigo & Subías, 1974); Spain (Kahwash et al., 1991); Portugal (Gil & Subías, 1990).

Scapheremaeus cornutus J. Balogh, 1958 Scapheremaeus cornutus J. Balogh, 1958, 8. Scapheremaeus cornutus: Mahunka, 1985, 246, figs. 40–45. Type depository: Holotype was in Musée Royal de l'Afrique Centrale, Tervuren, but is lost according to Mahunka



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(1985); paratype in Hungarian Natural History Museum. Type locality: no data given (Balogh,1958); Riv. Cuile, Alto Chicapa, ANGOLA (Mahunka, 1985) Habitat: soil (Balogh, 1958); ‘galerie forestiere des sources’ (Mahunka, 1985).

Scapheremaeus crassus Mahunka, 1988 Scapheremaeus crassus Mahunka, 1988, 232, figs. 42–46. Type depository: Hungarian Natural History Museum, Budapest. Type locality: Ha Son Bin, Hoa Binh, VIETNAM. Habitat: decaying debris, roots and litter from cracks of a steep slope of a rocky wall.

Scapheremaeus cuspidatus Perez-Iñigo, 1982 Scapheremaeus cuspidatus Perez-Iñigo, 1982, 209, figs. 15–18. Type depository: Instituto Español de Entomologia, Madrid. Type locality: Annobón, EQUATORIAL GUINEA Habitat: no data given.

Scapheremaeus cyclops (Oudemans, 1915) comb. nov. Cymberemaeus cyclops Oudemans, 1915, 193. Cymberemaeus cyclops: Oudemans, 1917, 18, figs. 39–44. Type depository: Rijksmuseet voor Naturhistorie, Leiden. Type locality: SRI LANKA. Habitat: moss and soil.

Scapheremaeus demeteri Mahunka, 1983 Scapheremaeus demeteri Mahunka, 1983, 178, figs. 88–90. Type depository: Museum d'Histoire Naturelle, Geneva and Hungarian Natural History Museum, Budapest. Type locality: bank of Awash River, 1400 m., Sodere, ETHIOPIA. Habitat: wet soil.

Scapheremaeus digitatus Wang, 1998 Scapheremaeus digitatus Wang, 1998, 256, figs. 5a,b Type depository: Institute of Zoology, Chinese Academy of Sciences, Beijing. Type locality: Mount Jianfeng, Hainan Province, CHINA. Habitat: soil.

Scapheremaeus emarginatus Hammer, 1966 ?Scapheremaeus emarginatus Hammer, 1966, 44, fig. 54. Type depository: Zoologisk Museum, Copenhagen. Type locality: Mirror Lake, Rotorua; Millford Sound, NEW ZEALAND. Habitat: moist moss under ferns and tall trees, Rotorua; thick, moist moss and dead branches in Nothofagus forest, Millford Sound.

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Scapheremaeus eugenius Rios & Palacios-Vargas, 1998 Scapheremaeus eugenius Rios & Palacios-Vargas, 1998, 209, fig. 14. Type depository: Department of Biology, Universidad Nacional Autonomia de Mexico, Mexico City. Type locality: Puerto “Los Mazos”, Sierra de Manantlan, Jalisco, MEXICO. Habitat: from mosses, 1450 masl.

Scapheremaeus fimbriatus (Michael, 1890) Eremaeus fimbriatus Michael, 1890, 422, pl. 37, fig. 6. Scapheremaeus fimbriatus: Grandjean, 1934, 114, Fig. 5e. Type depository: Natural History Museum, London. Type locality: Algiers, ALGERIA. Habitat: no data given.

Scapheremaeus fisheri Aoki, 1966 Scapheremaeus fisheri Aoki, 1966, 772, fig. 8. Scapheremaeus fisheri: Chakrabarti & Bhaduri, 1972, 500. Type depository: Bishop Museum, Honolulu. Type locality: Midway Island, HAWAIIAN ARCHIPELAGO. Habitat: from nest of Diomedea immutabilis (Aoki, 1966); from humus and decaying leaves of an ornamental tree (Polyanthia longifolia) (Chakrabarti & Bhaduri, 1972). Distribution: Midway Island (Aoki, 1966); India (Chakrabarti & Bhaduri, 1972).

Scapheremaeus flamiferus Palacios-Vargas & Rios, 1998 Scapheremaeus flamiferus Palacios-Vargas & Rios, 30, figs. 1–4. Type depository: Museo Entomologico, Leon, Nicaragua. Type locality: “Las Delicias”, Leon, NICARAGUA. Habitat: ex. Tillandsia sp. on Crescentia alata, tropical forest. Distribution: Nicaragua, Mexico.

Scapheremaeus foveolatus Mahunka, 1987 Scapheremaeus foveolatus Mahunka, 1987, 262: figs. 22–25. Type depository: Museum d'Histoire Naturelle, Geneva; Hungarian Natural History Museum, Budapest; Collection of the Scientific Centre of Vietnam. Type locality: Thanh Hoa, Tho Xuan, VIETNAM. Habitat: moss growing on living trees.

Scapheremaeus fungisetosus Rios & Palacios-Vargas, 1998 Scapheremaeus mahunkaius Rios & Palacios-Vargas, 1998, 205, fig. 13. Type depository: Department of Biology, Universidad Nacional Autonomia de Mexico, Mexico City. Type locality: “Chamela” Biological Station, Jalisco, MEXICO. Habitat: from canopy (fogging), tropical dry forest.



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Scapheremaeus glaber Hammer, 1958 Scapheremaeus glaber Hammer, 1958, 38, fig. 38. Type depository: Zoologisk Museum, Copenhagen. Type locality: Cumbre, BOLIVIA. Habitat: in thin layer of wet Polytrichum on almost bare soil among cliffs.

Scapheremaeus grahamius Rios & Palacios-Vargas, 1998 Scapheremaeus grahamius Rios & Palacios-Vargas, 1998, 188, fig. 4. Type depository: Department of Biology, Universidad Nacional Autonomia de Mexico, Mexico City. Type locality: Pedregal de San Angel Ecological Reserve, Distrito Federal, MEXICO. Habitat: canopy knockdown.

Scapheremaeus guerini (Berlese, 1908) Cymberemaeus Guerinii Berlese, 1908, 11. Cymbaeremaeus (Scapheremaeus) guerini Berlese, 1910, 227, fig. 65. Cymbaeremaeus (Scapheremaeus) guerinii: Castagnoli & Pegazzano, 1985, 173. Scapheremaeus guerini: Mahunka & Mahunka-Papp, 1995, pp. 62, 160, fig. 74. [redescription] Type depository: Berlese Acaroteca, Firenze. Type locality: none given, FRANCE. Habitat: no data given (Berlese, 1908); ‘galle di Doon. Macrophylla’(of type; Castagnoli & Pegazzano, 1985).

Scapheremaeus hammerae Balogh & Balogh, 2002 Scapheremaeus ? patella Hammer, 1966, figs. 52, 52a. Scapheremaeus hammerae Balogh & Balogh, 2002 Type depository: Zoologisk Museum, Copenhagen. Type locality: Rotorua, NEW ZEALAND. Habitat: dry moss on ground under Manuka shrub in thermal [springs] area. Note: S. patella Hammer, 1966 is a junior homonym of S. patella (Berlese, 1886) and was renamed by Balogh and Balogh (2002)

Scapheremaeus hectorperezius Rios & Palacios-Vargas, 1998 Scapheremaeus hectorperezius Rios & Palacios-Vargas, 1998, 203, fig. 12. Type depository: Department of Biology, Universidad Nacional Autonomia de Mexico, Mexico City. Type locality: Los Tuxtlas, Veracruz, MEXICO. Habitat: from lichens.

Scapheremaeus hieroglyphicus (Hall, 1911) Hermannia hieroglyphicus Hall, 1911, 646, fig. 216. Scapheremaeus hieroglyphicus: Marshall, Reeves & Norton, 1987, 236. [recombination] Type depository: type unknown. Type locality: Claremont, California, USA. Habitat: under black scale (Saissetia olei).

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Scapheremaeus humeratus Balogh & Mahunka, 1967 Scapheremaeus humeratus Balogh & Mahunka, 1967, 35, figs. 1–2. Type depository: Hungarian Natural History Museum, Budapest. Type locality: nr. Zanzi, CONGO. Habitat: decaying leaves in a clump of bamboo.

Scapheremaeus hungarorum Mahunka, 1986 Scapheremaeus hungarorum Mahunka, 1986, 306, figs. 18–21. Type depository: Hungarian Natural History Museum, Budapest Type locality: Usambara Mountains, TANZANIA. Habitat: fallen epiphytes, submontane rain forest.

Scapheremaeus insularis Hammer, 1966 Scapheremaeus insularis Hammer, 1966, 43, fig. 53. Type depository: Zoologisk Museum, Copenhagen. Type locality: Kerikeri, NEW ZEALAND. Habitat: moist-wet luxuriant moss on ground (Hammer, 1966).

Scapheremaeus johnsi Balogh, 1970 Scapheremaeus johnsi Balogh, 1970, 306, figs. 40, 41. Type depository: Zoosystematical Institute, Eotvos Lorand University, Budapest. Type locality: Mt. Wilhelm, PAPUA NEW GUINEA. Habitat: litter of Coprosoma divergens mixed with moss, from tussock grassland with subalpine shrubs.

Scapheremaeus latirostris Willmann, 1936 Scapheremaeus latirostris Willmann, 1936, 432, pl. 14, fig. 2. Type depository: [?] Zoologische Staatssamlung, Munich. Type locality: Bonaire, LESSER ANTILLES. Habitat: in dead coral in a salt lake.

Scapheremaeus latus Mahunka, 1985 Scapheremaeus latus Mahunka, 1985, 137, figs. 44, 45. Type depository: Museum d'Histoire Naturelle, Geneva and Hungarian Natural History Museum, Budapest. Type locality: Grand-Terre, GUADELOUPE. Habitat: moist soil.

Scapheremaeus longicuspis Mahunka, 1984 Scapheremaeus longicuspis Mahunka, 1984, 133, figs. 51, 52. Type depository: Museum d'Histoire Naturelle, Geneva and Hungarian Natural History Museum, Budapest.



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Type locality: Estancia Garay Cué, Concepción Province, PARAGUAY. Habitat: sievings from rotten stumps in dry forest.

Scapheremaeus longilamellatus Mahunka, 1985 Scapheremaeus longilamellatus Mahunka, 1985, 162, figs. 99–102. Type depository: Museum d'Histoire Naturelle, Geneva and Hungarian Natural History Museum, Budapest. Type locality: Castries, Piton Flore, ST. LUCIA. Habitat: various types of rotten material, banana.

Scapheremaeus madeirensis Willmann, 1939 Scapheremaeus madeirensis Willmann, 1939, 19, figs. 13a, b. Type depository: [?] Zoologische Staatssamlung, Munich. Type locality: Rabacal, MADEIRA. Habitat: sweep-netted from grass.

Scapheremaeus magdalenae Rios & Palacios-Vargas, 1998 Scapheremaeus magdalenae Rios & Palacios-Vargas, 1998, 192, fig. 6. Type depository: Department of Biology, Universidad Nacional Autonomia de Mexico, Mexico City. Type locality: Isla Cedro, Baja California, MEXICO. Habitat: mosses on branches in Cupressus forest.

Scapheremaeus mahunkaius Rios & Palacios-Vargas, 1998 Scapheremaeus mahunkaius Rios & Palacios-Vargas, 1998, 186, fig. 3. Type depository: Department of Biology, Universidad Nacional Autonomia de Mexico, Mexico City. Type locality: “Chamela” Biological Station, Jalisco, MEXICO. Habitat: from canopy (fogging), tropical dry forest.

Scapheremaeus marginalis (Banks, 1896) Eremaeus marginalis Banks, 1896, 76. Cymbaeremaeus marginalis: Banks, 1904, 72, fig. 138. Cepheus marginalis: Banks, 1915, 99. Scapheremaeus marginalis: Johnston, 1965, 59. Scapheremaeus marginalis: Marshall, Reeves & Norton, 1987, 237. Type depository: Museum of Comparative Zoology, Harvard. Type locality: Sea Cliff, New York, USA. Habitat: under lichens on bark of apple trees (Banks, 1896).

Scapheremaeus marmoratus (Berlese, 1910) Cymbaeremaeus (Scapheremaeus) marmoratus Berlese, 1910, 227, fig. 64. Cymbaeremaeus (Scapheremaeus) marmoratus: Castagnoli & Pegazzano, 1985, 242. Scapheremaeus marmoratus: Marshall, Reeves & Norton, 1987, 237.

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Scapheremaeus marmoratus: Norton & Kethley, 1989, 430. Scapheremaeus marmoratus: Mahunka & Mahunka-Papp, 1995, 160. Type depository: Berlese Acaroteca, Firenze; topotype in Field Museum of Natural History, Chicago. Type locality: Lake City, Florida, USA. Habitat: in mosses.

Scapheremaeus morenoi (Balogh & Mahunka, 1974) Scutovertex morenoi Balogh & Mahunka, 1974, 19, fig 13. Scapheremaeus morenoi: Subías, 2004, 158. [recombination] Type depository: Zoological Institute of the Academy of Sciences, Havana, Cuba; Hungarian Natural History Museum, Budapest. Type locality: Pico Suecica (1730 masl), Sierra Maestra, Oriente Province, CUBA. Habitat: from moss forest.

Scapheremaeus morulisensillatus Rios & Palacios-Vargas, 1998 Scapheremaeus morulisensillatus Rios & Palacios-Vargas, 1998, 190, fig. 5. Type depository: Department of Biology, Universidad Nacional Autonomia de Mexico, Mexico City. Type locality: “Los Dinamos” National Park, Distrito Federal, MEXICO. Habitat: mosses on rocks.

Scapheremaeus nashiroi Nakatamari, 1989 Scapheremaeus nashiroi Nakatamari, 1989, 1. Scapheremaeus nashiroi: Wang, 1998, 256, figs. 6a,b. Type depository: not known. Type locality: Okinawa. Habitat: not known. Distribution: Japan (Nakatamari, 1989); China (Wang,1998).

Scapheremaeus nogueraius Rios & Palacios-Vargas, 1998 Scapheremaeus nogueraius Rios & Palacios-Vargas, 1998, 201, fig. 11. Type depository: Department of Biology, Universidad Nacional Autonomia de Mexico, Mexico City. Type locality: “Chamela” Biological Station, Jalisco, MEXICO. Habitat: from canopy (fogging), tropical dry forest. Scapheremaeus nuciferosa Ramani & Haq, 1998 Scapheremaesus nuciferosa Ramani & Haq, 1998, 55, fig. 30 Type depository: Department of Zoology, University of Calicut, Kerala. Type locality: Calicut, Kerala, INDIA Habitat: from fresh foliage of coconut palm.

Scapheremaeus obliteratus Hammer, 1961 Scapheremaeus obliteratus Hammer, 1961, 32, fig. 24.



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Type depository: Zoologisk Museum, Copenhagen. Type locality: pass between Cusco and Pisac, PERU. Habitat: in thin lichen and moss on clay between broken-off cliffs, on the ground.

Scapheremaeus ornatus Balogh & Mahunka, 1968 Scapheremaeus ornatus Balogh & Mahunka, 1968, 330, figs 23, 24. Type depository: Hungarian Natural History Museum, Budapest. Type locality: Sierra de Córdoba, ARGENTINA. Habitat: epiphytes on shrubs.

Scapheremaeus palaciosi Ríos, 1996 Scapheremaeus sp.: Norton & Palacios-Vargas, 1996, 87. Scapheremaeus palaciosi Ríos, 1996, 237, figs. 1–8. Type depository: Microarthropod collection, Faculty of Sciences, Universidad Nacional Autonómia de México. Type locality: “Los Dínamos”, Mexico City; Popocatépetl Volcano; Nexapa, Amecameca, MEXICO. Habitat: mosses (“Los Dínbamos”; Ríos, 1996), epiphytic mosses and lichens (Popocatépetl; Norton & Palacios-Vargas, 1996), soil under Pinus hartureguii (Nexapa; Ríos, 1996). Remarks: Norton & Palacios-Vargas (1987) consider this species to be specific to corticolous epiphytes (as opposed to underlying soil and leaf litter) at Popocatépetl.

Scapheremaeus palustris (Sellnick, 1924) Cymbaeremaeus (Scapheremaeus) palustris Sellnick, 1924, 68, figs 6–9. Scapheremaeus palustris: Marshall, Reeves & Norton, 1987, 237. Scapheremaeus palustris: Mahunka, 1987, 375, figs. 36–39. Type depository: type considered destroyed. Type locality: ESTONIA. Habitat: rotting moss, leaf-litter and wood in wet forests of Fraxino pannonicae-ulmetum association (Mahunka, 1987). Distribution: Fennoscandia (Niemi et al., 1997); Eastern Baltic, Western Russia (Karppinen & Krivolutsky, 1982); Caucusus (Karppinen et al., 1986), Ukraine (Karppinen et al., 1992); USA and Canada (Marshall, Reeves & Norton, 1987).

Scapheremaeus parvula (Banks, 1909) Cymbaeremaeus parvula Banks, 1909, 141, pl. 13, fig. 26. Scapheremaeus parvula: Marshall, Reeves & Norton, 1987, 237. [recombination] Type depository: Museum of Comparative Zoology, Harvard. Type locality: Guelph, Ontario, CANADA. Habitat: under bark of ironwood.

Scapheremaeus patella (Berlese, 1886) Eremaeus patella Berlese, 1886, fasc. 33, no. 10. Cymbaeremaeus (Scapheremaeus) patella: Berlese, 1910, 226, fig. 63. Scapheremaeus patella: Mahunka, 1977, 914, figs. 14, 15. Cymbaeremaeus (Scapheremaeus) patella: Castagnoli & Pegazzano, 1985, 309. Scapheremaeus patella: Mahunka & Mahunka-Papp, 1995, pp. 62, 160, fig. 75. Scapheremaeus patella: Perez-Iñigo, 1996, 43. [redescription]

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Type depository: Berlese Acaroteca, Firenze. Type locality: Firenze, ITALY. Habitat: thin layer of dry moss on ground under Manuka shrub in thermal area (Hammer, 1966); soil at base of rocks near entrance to cave (Mahunka, 1977) Distribution: Italy (Berlese, 1910); Greece (Mahunka, 1977); Austria (Schatz, 1983); Canary Islands (Pérez-Iñigo & Peña, 1996). Note: S. patella Hammer, 1966 is a junior homonym of S. patella (Berlese, 1886) and was renamed S. hammerae (cf. Above) by Balogh and Balogh (2002).

Scapheremaeus petrosus Sitnikova, 1975 Scapheremaeus petrosus Sitnikova, 1975, 240, fig. 558. Type depository: [?] A.A. Severtzov Institute of Evolutionary Morphology and Ecology of Animals, Moscow. Type locality: Primorye Territory, RUSSIA. Habitat: no data given.

Scapheremaeus pisacensis Hammer, 1961 Scapheremaeus pisacensis Hammer, 1961, 33, fig. 25. Type depository: Zoologisk Museum, Copenhagen. Type locality: pass between Cusco and Pisac, PERU. Habitat: thin cover of lichen and moss on clay on the ground.

Scapheremaeus polysetosus Sitnikova, 1975 Scapheremaeus polysetosus Sitnikova, 1975, 237, figs. 557a, b. Type depository: [?] A.A. Severtzov Institute of Evolutionary Morphology and Ecology of Animals, Moscow. Type locality: Primorye Territory, RUSSIA. Habitat: no data given. Distribution: Siberia, Russian Far East (Golosova et al., 1983).

Scapheremaeus pseudoreticulatus Mahunka, 1984 Scapheremaeus pseudoreticulatus Mahunka, 1984, 673, figs. 10–14. Type depository: Musée Royal de l'Afrique Centrale, Tervuren. Type locality: Île Mahé, SEYCHELLES Habitat: no data given. Scapheremaeus pulchellus (Berlese, 1910) Cymbaeremaeus (Scapheremaeus) pulchellus Berlese, 1910, 227, fig. 66. Scapheremaeus pulchellus: Perez-Iñigo & Subías, 1974, 745. Cymbaeremaeus (Scapheremaeus) pulchellus: Castagnoli & Pegazzano, 1985, 339. Scapheremaeus pulchellus: Marshall, Reeves & Norton, 1987, 238. Scapheremaeus pulchellus: Norton & Kethley, 1989, 474. Scapheremaeus pulchellus: Mahunka & Mahunka-Papp, 1995, 160. Type depository: Berlese Acaroteca, Firenze. Type locality: Lake City, Florida, USA. Habitat: in mosses.



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Remarks: Perez-Iñigo & Subías (1974) consider this species to be a junior synonym of S. marginalis but Norton & Kethley (1989) regard it as a valid species.

Scapheremaeus pundamiliaensis Engelbrecht, 1975 Scapheremaeus pundamiliaensis Engelbrecht, 1975, 79, figs. 48, 49. Type depository: Nasionale Museum, Bloemfontein. Type locality: Pafuri, Kruger National Park, REPUBLIC OF SOUTH AFRICA. Habitat: underneath fever tree (holotype); apple orchard, Tzaneen (paratypes); under sicklebush, Punda Milia (paratype).

Scapheremaeus quadrilineatus Mahunka, 1978 Scapheremaeus quadrilineatus Mahunka, 1978, 324, figs. 28, 29. Type depository: Museum d'Histoire Naturelle, Geneva and Hungarian Natural History Museum, Budapest. Type locality: Île Ronde, MAURITIUS. Habitat: no data given.

Scapheremaeus quadrireticulatus Wang, 1998 Scapheremaeus quadrireticulatus Wang, 1998, 255, fig. 3. Type depository: Institute of Zoology, Chinese Academy of Sciences, Beijing. Type locality: Beijing, CHINA. Habitat: plant litter.

Scapheremaeus retestriatus Wang, 1998 Scapheremaeus retestriatus Wang, 1998, 252, figs. 1a,b. Type depository: Institute of Zoology, Chinese Academy of Sciences, Beijing. Type locality: Liancheng, Fujian Province, CHINA. Habitat: no data given.

Scapheremaeus reticulatus (Berlese, 1910) Cymbaeremaeus reticulatus Berlese, 1910, 381. Scapheremaeus reticulatus: Balogh, 1943, 32, pl. 6. fig. 4. Cymbaeremaeus reticulatus: Castagnoli & Pegazzano, 1985, 309. Scapheremaeus reticulatus: Mahunka & Mahunka-Papp, 1995, pp. 63, 163. Type depository: Berlese Acaroteca, Firenze. Type locality: Ceresole d'Alba, ITALY. Habitat: in soil (Berlese, 1910); no data given (Balogh, 1943). Distribution: Italy (Berlese, 1910); Hungary (Balogh, 1943); Austria (Franz, 1954); Bulgaria (Kunst, 1961). Remarks: Mahunka & Mahunka-Papp (1995, p. 63) state that it is uncertain whether Scapheremaeus reticulatus sensu Balogh, 1943 is the same as the species described by Berlese.

Scapheremaeus rustenburgensis Engelbrecht, 1975 Scapheremaeus rustenburgensis Engelbrecht, 1975, 73, figs. 41–47.

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Type depository: Nasionale Museum, Bloemfontein. Type locality: Groenkloof, Rustenberg, REPUBLIC OF SOUTH AFRICA. Habitat: no data given for holotype; other material from underneath poplars.

Scapheremaeus schatzi Rios & Palacios-Vargas, 1998 Scapheremaeus schatzi Rios & Palacios-Vargas, 1998, 194, fig. 7. Type depository: Department of Biology, Universidad Nacional Autonomia de Mexico, Mexico City. Type locality: “Chamela” Biological Station, Jalisco, MEXICO. Habitat: from canopy (fogging), tropical dry forest.

Scapheremaeus semiconvexus Hammer, 1982 Scapheremaeus semiconvexus Hammer, 1982, 460, fig. 17. Type depository: Zoologisk Museum, Copenhagen. Type locality: Uluwatu, BALI. Habitat: dead leaves and debris.

Scapheremaeus semiornatus Hammer, 1979 Scapheremaeus semiornatus Hammer, 1979, 47, fig. 84. Type depository: Zoologisk Museum, Copenhagen. Type locality: Tangkuban Prahu, ca. 28 km N. Bandung, JAVA. Habitat: amongst dead leaves in wet, deciduous tropical forest with luxurious undervegetation of bamboo, shrubs, ferns, mosses (Hammer, 1979).

Scapheremaeus simplex Rios & Palacios-Vargas, 1998 Scapheremaeus simplex Rios & Palacios-Vargas, 1998, 209, fig. 15. Type depository: Department of Biology, Universidad Nacional Autonomia de Mexico, Mexico City. Type locality: Chichinautzin, Morelos, MEXICO. Habitat: ex. Tillandsia sp. on lava flow.

Scapheremaeus sinuosus Aoki, 1964 Scapheremaeus sinuosus Aoki, 1964, 649, figs. 1–5. Type depository: Bishop Museum, Honolulu. Type locality: Laysan Island, HAWAIIAN ARCHIPELAGO. Habitat: from roots of bunch grass, and resting place of wedge-tailed shearwater.

Scapheremaeus stratus Hammer, 1958 Scapheremaeus stratus Hammer, 1958, 39, fig. 39. Type depository: Zoologisk Museum, Copenhagen. Type locality: Cumbre, BOLIVIA. Habitat: in thin layer of wet Polytrichum on almost bare soil among cliffs.



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Scapheremaeus striatomarginatus Hammer, 1979 Scapheremaeus striatomarginatus Hammer, 1979, 48, fig. 85. Type depository: Zoologisk Museum, Copenhagen. Type locality: Tangkuban Prahu, ca. 28 km N. Bandung, JAVA. Habitat: amongst dead leaves in wet, deciduous tropical forest with luxurious undervegetation of bamboo, shrubs, ferns, mosses.

Scapheremaeus subcorniger Perez-Iñigo & Peña, 1996 Scapheremaeus subcorniger Perez-Iñigo & Peña, 1996, 155, figs. 26–29. Type depository: Museo Nacional de Ciencias Naturales, Madrid. Type locality: El Fraile, Fuerteventura, Canary Islands. Habitat: Lichens (Rochella and Ramalina) on basalt.

Scapheremaeus subglaber J. Balogh & Mahunka, 1978 Scapheremaeus subglaber Balogh & Mahunka, 1978, 285, figs. 13A, B. Type depository: Hungarian Natural History Museum, Budapest Type locality: Botanical Gardens, Asunción, PARAGUAY. Habitat: litter at foot of tree.

Scapheremaeus taurus Balogh, 1970 Scapheremaeus taurus Balogh, 1970, 305, figs. 38, 39. Type depository: Zoosystematical Institute, Eotvos Lorand University, Budapest. Type locality: Mt. Kaindi, Wau, PAPUA NEW GUINEA. Habitat: moss from tree trunk in moss forest.

Scapheremaeus tillandsiae Fernandez & Cleva, 1997 Scapheremaeus tillandsiae Fernandez & Cleva, 1997, 289, figs. 1–4. Type depository: Centro de Investigaciones Científicas y de Transferencia de Technología a la Producción, Diamante, Entre Rios, Argentina. Type locality: Divisadero Largo, Mendoza Province, ARGENTINA Habitat: ex. Tillandsia sp.

Scapheremaeus tillandsiophilus Rios & Palacios-Vargas, 1998 Scapheremaeus tillandsiophilus Rios & Palacios-Vargas, 1998, 199, fig. 10. Type depository: Department of Biology, Universidad Nacional Autonomia de Mexico, Mexico City. Type locality: Chichinautzin, Morelos, MEXICO. Habitat: ex. Tillandsia sp. on lava flow.

Scapheremaeus tonatiuh Palacios-Vargas, Rios & Vazquez, 1998 Scapheremaeus tonatiuh Palacios-Vargas, Rios & Vazquez, 1998, 384, figs. 1–11.

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Type depository: Department of Biology, Universidad Nacional Autonomia de Mexico, Mexico City. Type locality: “Chamela” Biological Station, Jalisco, MEXICO. Habitat: from canopy (fogging), tropical dry forest.

Scapheremaeus tosaensis Fujikawa, 2005 Scapheremaeus tosaensis Fujikawa, 2005, 1, figs. 1 & 2. Type depository: National Science Museum, Tokyo. Type locality: Tanemaji Temple, Shikoku Is., JAPAN. Habitat: soil from garden, graveyard and forest.

Scapheremaeus tricarinatus Sitnikova, 1975 Scapheremaeus tricarinatus Sitnikova, 1975, 240, fig. 559. Type depository: [?] A.A. Severtzov Institute of Evolutionary Morphology and Ecology of Animals, Moscow. Type locality: North Ossetian ASSR, RUSSIA. Habitat: no data given.

Scapheremaeus trirugis Hammer, 1958 Scapheremaeus trirugis Hammer, 1958, 37, fig. 37. Type depository: Zoologisk Museum, Copenhagen. Type locality: Rio Caldera Valley, nr. Salta, ARGENTINA. Habitat: on mosses on a cliff with oozing water.

†Scapheremaeus undosus (Sellnick, 1918) Mulvius undosus Sellnick, 1918, p. 36, fig. 17. Scapheremaeus undosus (Sellnick, 1918): Sellnick, 1931, p. 164; Norton, 2006, p. 112. Type depository: Holotype was in the Geologische-paläontologisches Institut, Universität Konigsberg, but is either lost or in the Geowissenschaftliches Zentrum der Universität Göttingen (Norton, 2006, p. 119). Habitat: in Baltic amber.

Scapheremaeus uncinatus Wang, 1998 Scapheremaeus uncinatus Wang, 1998, 254, fig. 2. Type depository: Institute of Zoology, Chinese Academy of Sciences, Beijing. Type locality: Mount Wuyi, Fujian Province, CHINA. Habitat: under bark; from moss on bark.

Scapheremaeus volcanicus Rios & Palacios-Vargas, 1998 Scapheremaeus volcanicus Rios & Palacios-Vargas, 1998, 182, fig. 1. Type depository: Department of Biology, Universidad Nacional Autonomia de Mexico, Mexico City. Type locality: Popocatepetl Volcano, MEXICO. Habitat: in mosses and epiphytic lichens, 3,000 masl.



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Scapheremaeus yamashitai Aoki, 1970 Scapheremaeus yamashitai Aoki, 1970, 595, figs. 28–31. Scapheremaeus yamashitai: Fujikawa, 2002, 69: 20. [redescription] Type depository: National Science Museum, Tokyo. Type locality: Mt. Ishizuchi, Ehime Prefecture, JAPAN. Habitat: insecticide knock-down from Fagus crenata.

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