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Steganacarus (Tropacarus) Ewing, 1917 S. (T.) brevipilus (Berlese, 1923), S. (T.) carinatus (C.L.Koch, 1841), Published as part of Niedba��a, Wojciech, 2015, Supplement to the knowledge of ptyctimous mites (Acari, Oribatida) from Palaearctic Region, pp. 301-339 in Zootaxa 4057 (3) on page 303, DOI: 10.11646/zootaxa.4057.3.1, http://zenodo.org/record/238226

Steganacarus (Steganacarus) Ewing, 1917 S. (S.) applicatus (Sellnick, 1920), S. (S.) balearicus P��rez-I��igo, 1969, S. (S.) crassisetosus (Willmann, 1939), S. (S.) guanarteme P��rez-I��igo & Pe��a, 1996, S. (S.) magnus (Nicolet, 1855), S. (S.) patruelis Niedbała, 1983, S. (S.) similis (Willmann, 1939), S. (S.) spinosus (Sellnick, 1920), Published as part of Niedba��a, Wojciech, 2015, Supplement to the knowledge of ptyctimous mites (Acari, Oribatida) from Palaearctic Region, pp. 301-339 in Zootaxa 4057 (3) on page 303, DOI: 10.11646/zootaxa.4057.3.1, http://zenodo.org/record/238226

Steganacarus (Tropacarus) brevipilus (Berlese, 1923) Iran localities. Nowshahr (Akrami et al. 2006). Rasht (Mortazavi et al. 2010). Shabestar, Shendabad (Mirzaie et al. 2011 b). Arasbaran (Gheblealivand & Haddad Irani-Nejad 2014). Distribution. Western Palaearctic (except North) and Iran., Published as part of Akrami, Mohammad Ali, 2015, An annotated checklist of oribatid mites (Acari: Oribatida) of Iran, pp. 451-501 in Zootaxa 3963 (4) on page 463, DOI: 10.11646/zootaxa.3963.4.1, http://zenodo.org/record/239832, {"references":["Mirzaie, M., Haddad Irani-Nejad, K. & Akrami, M. A. (2011 b) New records of primitive oribatid mites (Acari: Oribatida) from the Shendabad region (East Azerbaijan province), Iran. Zoology in the Middle East, 53, 137. http: // dx. doi. org / 10.1080 / 09397140.2011.10648874","Gheblealivand, S. S. & Haddad Irani-Nejad, K. (2014) Introducing some of Arasbaran region's oribatid mites (Acari: Oribatida), with new records for Iran's and East Azerbaijan province fauna. Persian Journal of Acarology, 3 (3), 241 - 247."]}

Steganacarus (Steganacarus) inaestimabilis Niedbała, 1984 Iran localities. Chalus (Niedbała 1984). Chalus road (185 km. after Karaj) (Akrami et al. 2006). Distribution. Iran., Published as part of Akrami, Mohammad Ali, 2015, An annotated checklist of oribatid mites (Acari: Oribatida) of Iran, pp. 451-501 in Zootaxa 3963 (4) on page 464, DOI: 10.11646/zootaxa.3963.4.1, http://zenodo.org/record/239832, {"references":["Niedbala, W. (1984) Phthiracaridae (Acari, Oribatida) nouveaux d'Asie Occidentale. Annales Zoologici, 38 (10), 225 - 241."]}

Steganacarus (Steganacarus) balearicus P��rez-I��igo, 1969 (Figs 5 A���F) Measurements of one of the larger specimens from Slovakia: prodorsum: length 293, width 197, height 126, setae: ss 71, in 48, le 20, ro 30, ex 23; notogaster: length 581, width 794, height 328, setae: c 1 53, c 1 / c 1 - h 1 = 0.4, h 1 and ps 1 56. All characters are similar to holotype, only the interlamellar setae are sligtly longer and setae h of mentum are as long as the distance between them., Published as part of Niedba��a, Wojciech, 2015, Supplement to the knowledge of ptyctimous mites (Acari, Oribatida) from Palaearctic Region, pp. 301-339 in Zootaxa 4057 (3) on page 312, DOI: 10.11646/zootaxa.4057.3.1, http://zenodo.org/record/238226

Earlier published and newly determined Swiss moss mites belonging to the "ptychoid" groups are listed, discussed or described. From the 20 named species in the literature 10 are confirmed after critical discussion. New identifications of a total of 42 taxa are given, 3 of which (Phthiracarus besuchetianus sp. n., Steganacarus (S.) antennatus sp. n. and Steganacarus (S.) schweizeri sp. n.) are new to science and 30 new for the Fauna of Switzerland. Some new or additional morphological data as well as some notes on the taxonomy and distribution of this group are given.

Steganacarus (Rhacaplacarus) quaternarius sp. nov. (Fig. 7 A���H) Measurements of holotype. Prodorsum: length 278, height 111, width 207; prodorsal setae: sensillus 96, interlamellar (in) 139, lamellar (le) 43, rostral (ro) 56, exobothridial (ex) 15; notogaster: length 555, height 404, width 353; notogastral setae: c 1 126, h 1 144, ps 1 109; genitoaggenital plate 121 �� 109, anoadanal plate 215 �� 111. Description. Species with neotrichy of adanal setae. Colour light brown. Integument finely pitted. Prodorsum with sigillar fields distinct, median longer than laterals. Lateral carinae with some traces visible of distal part. Sensilli rather long, with narrow stalk and slightly fusiform head covered with very small spines. Interlamellar setae long, erect, rigid, covered sparsely with small spines in distal half, similar to notogastral setae; lamellar and rostral setae short, spiniform, and rough. Notogaster with notogastral setae of medium length (c 1 / c 1 - d 1 =1.0), rather pointed distally, covered with small spines on their distal half. Notogastral setae c 1 and c 3 slightly remote from anterior notogastral margin, setae c 2 considerably further from margin. Vestigial setae f 1 situated anteriorly of setae h 1. Two pairs of lyrifissures, ia and im, present. Ventral region. Setae h of mentum longer than distance between their insertion points. Genitoaggenital plates with nine pairs of setae on right plate or 10 pairs on left plate with formula: 6 (4 + 2): 3 or 7 (5 + 2): 3. Anoadanal plates each with seven setae; three anal setae at the paraxial row and four adanal setae remote from paraxial border, adanal setae ad 1 and ad 2 the longest. Legs. Formulae of leg setae and solenidia of complete type. Setae d on femora I positioned at distal end of article. Material examined. Holotype is deposited at DATE from the locus typicus: South���West Province, Mt. Koup Ecological Reserve, Mt. Koup at Nyasoso, 19���21. V. 2006, about 6 km N of Loum, latitude 04�� 49 '09.18" N, longitude 09�� 42 ' 27.18 " E, altitude 1550 m, submontane foggy forest, litter and soil sifting sample, leg V. Grebennikov. Etymology. The specific name quaternarius comes from Latin for ���consisting of four, arranged in four��� alluding to the four pairs of adanal setae. Comparison. The new species is well characterisable by the presence of four pairs of adanal setae remote from paraxial border of plates. Other Afrotropical Rhacaplacarus species have two pairs of these setae. Some Neotropical species of subgenus Rhacaplacarus with presence also of four pairs of adanal setae remote from paraxial border of plates are morphologically completely different. The most similar Afrotropical Steganacarus (Rhacaplacarus) spinatus Niedbała, 2001 by the length of interlamellar, lamellar and notogastral setae is distinguishable by the presence of flagellate sensilli (versus slightly fusiform sensilli, obtuse distally) and notogastral setae obtuse distally (versus rather pointed distally)., Published as part of Niedba��a, Wojciech & Star��, Josef, 2014, New and little known species of ptyctimous mites (Acari, Oribatida) from Cameroon, pp. 31-57 in Zootaxa 3889 (1) on page 44, DOI: 10.11646/zootaxa.3889.1.2, http://zenodo.org/record/228762, {"references":["Niedbala, W. (2001) Study on the diversity of Ptyctimous mites (Acari, Oribatida) and quest for centres of its origin: the fauna of the Ethiopian Region. Monographs of the Upper Silesian Museum, 3, 1 - 245."]}

Steganacarus (Rhacaplacarus) quaternarius sp. nov. (Fig. 7 A–H) Measurements of holotype. Prodorsum: length 278, height 111, width 207; prodorsal setae: sensillus 96, interlamellar (in) 139, lamellar (le) 43, rostral (ro) 56, exobothridial (ex) 15; notogaster: length 555, height 404, width 353; notogastral setae: c 1 126, h 1 144, ps 1 109; genitoaggenital plate 121 × 109, anoadanal plate 215 × 111. Description. Species with neotrichy of adanal setae. Colour light brown. Integument finely pitted. Prodorsum with sigillar fields distinct, median longer than laterals. Lateral carinae with some traces visible of distal part. Sensilli rather long, with narrow stalk and slightly fusiform head covered with very small spines. Interlamellar setae long, erect, rigid, covered sparsely with small spines in distal half, similar to notogastral setae; lamellar and rostral setae short, spiniform, and rough. Notogaster with notogastral setae of medium length (c 1 / c 1 - d 1 =1.0), rather pointed distally, covered with small spines on their distal half. Notogastral setae c 1 and c 3 slightly remote from anterior notogastral margin, setae c 2 considerably further from margin. Vestigial setae f 1 situated anteriorly of setae h 1. Two pairs of lyrifissures, ia and im, present. Ventral region. Setae h of mentum longer than distance between their insertion points. Genitoaggenital plates with nine pairs of setae on right plate or 10 pairs on left plate with formula: 6 (4 + 2): 3 or 7 (5 + 2): 3. Anoadanal plates each with seven setae; three anal setae at the paraxial row and four adanal setae remote from paraxial border, adanal setae ad 1 and ad 2 the longest. Legs. Formulae of leg setae and solenidia of complete type. Setae d on femora I positioned at distal end of article. Material examined. Holotype is deposited at DATE from the locus typicus: South–West Province, Mt. Koup Ecological Reserve, Mt. Koup at Nyasoso, 19–21. V. 2006, about 6 km N of Loum, latitude 04° 49 '09.18" N, longitude 09° 42 ' 27.18 " E, altitude 1550 m, submontane foggy forest, litter and soil sifting sample, leg V. Grebennikov. Etymology. The specific name quaternarius comes from Latin for “consisting of four, arranged in four” alluding to the four pairs of adanal setae. Comparison. The new species is well characterisable by the presence of four pairs of adanal setae remote from paraxial border of plates. Other Afrotropical Rhacaplacarus species have two pairs of these setae. Some Neotropical species of subgenus Rhacaplacarus with presence also of four pairs of adanal setae remote from paraxial border of plates are morphologically completely different. The most similar Afrotropical Steganacarus (Rhacaplacarus) spinatus Niedbała, 2001 by the length of interlamellar, lamellar and notogastral setae is distinguishable by the presence of flagellate sensilli (versus slightly fusiform sensilli, obtuse distally) and notogastral setae obtuse distally (versus rather pointed distally).

Steganacarus (Rhacaplacarus) spinus Niedbała sp. nov. (Fig. 5 A���I) Description. Measurements of holotype. Prodorsum: length 220, width 152, height 88. Prodorsal setae: sensillus 28, in, le 20, ro 15. Notogaster: length 384, width 257, height 227. Notogastral setae: c 1, h 1, p 1 20. Plates: genitoaggenital 88 �� 88, anoadanal 131 �� 86. Measurements of paratype. Prodorsum: length 263, height 111. Notogaster: length 515, height 288. Integument. Body strongly sculptured. Integument distinctly foveolate, foveoles in anterior part of prodorsum bigger than in posterior part. Prodorsum with long, narrow sigillar fields, weakly visible in lateral aspect. Lateral carinae absent. Posterior furrows well developed. Sensilli short, with narrow stalk and rounded head, rough. Setae (except vestigial exobothridial) short, spinose, rough. Notogaster with very short (c 1 / c 1 ��� d 1 = 0.27), thick, spinose, rough setae. Setae c 1 and c 3 some distance from anterior margin, setae c 2 far from margin. Vestigial setae f 1 invisible. Two pairs of lyrifissures ia and im present. Ventral region. Arrangement of genital setae: 7 (4 + 3): 2. Anoadanal plates with short, spinose setae, three at paraxial border and two adanal setae only slightly remote from border, setae ad 2 situated at level between anal setae. Legs. Formulae of setae and solenidia of ���complete type ���. Setae d on femora I a little remote from distal end of article. Material examined. Holotype: locality 3; one paratype: locality 4. Comparison. Only one oriental species of this subgenus, Steganacarus (Rhacaplacarus) spiniger (Aoki, 1980 (a)) from Japan, has similar spinose setae, but distinguish from the new species by the presence of lateral carinae of prodorsum and longer sensilli with broadened head covered with spines in one side. Another one species with similar notogastral setae is know from Tanzania��� S. (R.) spiniferus (Mahunka, 1993), but sensilli are sicle shaped on setae of anoadanal plates are longer (ad 1 and anal setae are longer than distance between them). Etymology. The specific epithet is derived from Latin spine meaning ���spike, thorn, barb��� referring to the shape of body setae. FIGURE 5 A���I. Steganacarus (Rhacaplacarus) spinus Niedbała sp. nov.: A, prodorsum, dorsal view; B, prodorsum, lateral view; C, opisthosoma, lateral view; D, interlamellar seta; E, seta c 1; F, left genitoaggenital plate: G, left anoadanal plate; H, trochanter and femur of leg I; I, tibia of leg IV., Published as part of Niedba��a, Wojciech & Ermilov, Sergey G., 2014, Ptyctimous mites (Acari, Oribatida) from the Joint Russian-Vietnamese Biological Expedition (October 2013 ��� April 2014), pp. 156-168 in Zootaxa 3884 (2) on pages 164-165, DOI: 10.11646/zootaxa.3884.2.4, http://zenodo.org/record/227242, {"references":["Aoki, J. (1980 a) A revision of the Oribatid mites of Japan. I. The families Phthiracaridae and Oribotritiidae. Bulletin Institute of Environmental Science and Technology, Yokohama, 6 (2), 1 - 88.","Mahunka, S. (1993) A new series of publication on new or little known Oribatid taxa from Africa (Acari), I. Acta Zoologica Hungarica, 39 (1 - 4), 91 - 119."]}

Steganacarus (Steganacarus) similis Willmann, 1939 (Fig. 3 A ��� G) Steganacarus similis: Bernini and Magari 1993 Measurements of specimen from Raba��al, Madeira. Prodorsum: length 257, height 111, width 202; setae of prodorsum: sensillum 96, interlamellar 139, lamellar 101, rostral 45, exobothridial 20; notogaster: length 596, height 353, width 364, setae of notogaster: c 1 88, c 1 / c 1 ��� d 1 = 0.7, h 1 and p 1 101; genitoaggenital plates: 156 �� 106, anaoadanal plates: 207 �� 116. Diagnosis. Surface covered with feeble concavities. Colour brown. Prodorsum: median field narrow and longer than the laterals; lateral carinae absent; sensilla long, narrow, sickle-shaped covered with spines; setae, other than the exobothridial covered with distinct spines in distal half, rostral setae directed inward. Notogaster with rigid, short (c 1 1 ��� d 1) setae covered with spines in distal half; setae c 1 and c 3 near the anterior margin, setae c 2 far away; vestigial setae f 1 slightly posteriorly of setae h 1; two pairs of lyrifissures ia and im. Ventral region: h > h ��� h; formula of genital setae: 6 (4 + 2): 3, anoadanal plate with fairly long setae, ad 1 and ad 2 are longest, an / not = 0.3. Chaetome of legs reduced; setae s on tarsi I are missing; v ���/ v��� = 4.4. Comparison. This species is very similar to the Paneuropean species S. (S.) spinosus (Fig. 3 H ��� N). The comparison was made with the specimens from the population collected in Poland (Sudety, Wielka Sowa). Measurements of specimen of S. (S.) spinosus (Sellnick, 1920) from Wielka Sowa, Poland. Prodorsum: length 303, height 126, width 212; setae of prodorsum: sensillum 94, interlamellar 152, lamellar 63, rostral 43, exobothridial 40; notogaster: length 606, height 404, width 434, setae of notogaster: c 1 121, c 1 / c 1 ��� d 1 = 0.66, h 1 and p 1 101; genitoaggenital plates: 167 �� 126, anaoadanal plates: 202 �� 131. Steganacarus (S.) similis differs from S. (S.) spinosus also by two main morphological features. S. (S.) similis from Madeira has longer lamellar setae (le / in > 0.7), S. (S.) spinosus has much shorter lamellar setae (le / in S. (S.) similis are diriged inward but rostral setae of S. (S.) spinosus are diriged forward. Willmann (1939) differentiated S. (S.) similis from S. (S.) spinosus ��� see also Willmann (1931), but in the comparison he has put too much emphasis on the body size and he was incorrect in claiming that the setae of S. (S.) spinosus are smooth and pointed distally. Bernini and Magari (1993) have determined the neotype of S. (S.) similis. They have pointed to some small morphological differences between S. (S.) similis with S. (S.) spinosus: in the shape of sensilla, shape of anoadanal setae, lack of setae a on the first tarsi, the length of anoadanal setae decreases gradually from ad 1 to an 2 in S. (S.) similis but in S. (S.) spinosus setae ad 1 and ad 2 are longer than setae an 1 and an 2. In the material for comparison performed in this study, i.e. from the S. (S.) similis population from Madeira and S. (S.) spinosus from Poland, these differences were not confirmed; the shapes of sensilla and anoadanal setae are similar in both species and also setae ad 1 and ad 2 are the longest. In view of a considerable morphological similarity of these two species, an attempt was made to establish their genetic relationship. Molecular analysis of the species status of S. (S.) crassisetosus, S. (S.) similis, S. (S.) spinosus, and S. (S.) applicatus The COI alignment for the distance calculations comprised 620 bp of unambiguous sequence data for 22 specimens of four Steganacarus species: S. (S.) applicatus (5), S. (S.) crassisetosus (6), S. (S.) similis (5), and S. (S.) spinosus (6). The number of variable sites was 207 and the estimated base frequencies were as follows: T = 0.4419, C = 0.1750, A = 0.2196, G = 0.1635. The average transition to transversion ratio (R) = 1.29 for all variable sites. NJ analysis of the COI sequences revealed four well-supported clades corresponding to the tested species (not shown). Estimates of average evolutionary divergence (K 2 P) were about one percent within clades grouping: S. (S.) applicatus (1.02 %, SE 0.28), S. (S.) spinosus (1.21 %, SE 0.27), and S. (S.) similis (1.26 %, SE 0.34). Nucleotide substitutions observed in the individual clades were synonymous and did not change the amino acid sequence. The highest divergence was found in S. (S.) crassisetosus clade (3.78 %, SE 0.55) but even in this case all nucleotide substitutions were synonymous. The average genetic distance among reconstructed clades was 23.03 % (SE 2.15) and ranged from 20.14 % (SE 1.94) between S. (S.) crassisetosus and S. (S.) applicatus to 27.17 % (SE 2.47) between S. (S.) spinosus and S. (S.) similis. This level of genetic distance outnumbered intraspecific divergence and strongly support the species status of the analysed taxa. The COI nucleotide sequences were published in GenBank as DNA barcodes for S. (S.) similis (Acc. JX 891540 - 43), S. (S.) applicatus (Acc. JX 891544 - 48), S. (S.) spinosus (Acc. JX 891549 - 54), and S. (S.) crassisetosus (Acc. JX 891555 - 60). The analysis of the nuclear D 2 marker corroborated the COI results. More than five specimens from each species have been sequenced in D 2 region of 28 S rDNA and no intraspecific polymorphism was found among sequences from individuals belonging to the same species. The final D 2 28 S rDNA alignment for the distance calculations comprised 373 nucleotide sites in which 34 were variable. The mean base frequencies were: A = 0.2347, C = 0.2043, G = 0.2737, T = 0.2872, and transition to transversion ratio (R) was 2.35. Estimates of average genetic distance over sequence pairs representing each species showed that S. (S.) spinosus is the most diverged from the three other species (8.37 %, SE 1.50 %), while the average divergence among S. (S.) crassisetosus, S. (S.) similis, and S. (S.) applicatus was 2.03 % (SE 0.73). The genetic distances between S. (S.) crassisetosus and S. (S.) applicatus as well as between S. (S.) spinosus and S. (S.) similis did not prove unambiguously their relationship. To find out the evolutionary relations among them, a phylogenetic analysis was made on the basis of greater representation of the species belonging to Steganacaridae. Phylogenetic relationships among S. (S.) crassisetosus, S. (S.) similis, S. (S.) spinosus, S. (S.) applicatus, and other Steganacaridae The alignment for phylogenetic tree reconstruction using COI sequence data comprised of 129 amino acid positions for 51 sequences. Twenty two sequences from this study were compared with the other known COI sequences for Steganacaridae: several species inhabiting Canary Islands and Sardinia (Salomone et al. 2002), one representative of European species S. (S.) magnus (JF 264105), and Phthiracarus longulus used as the outgroup (for details see Table 1). Maximum likelihood (ML) analysis based on COI amino acid data confidently recovered all analysed species (Fig. 4). Although most of the recovered evolutionary relationships among analysed Steganacaridae remain unresolved due to a low or no nodal support, the ML analysis of the COI sequences showed that S. (S.) spinosus is the most distant species from the others and that there is no support for a close relationship between S. (S.) applicatus and S. (S.) crassisetosus. The ML COI tree revealed S. (S.) spinosus basally to wellsupported (LR-ELW edge support = 93, BS= 83) clade containing the remainder of the taxa. In this group, European Steganacaridae were placed basally to island clades, however this placement was not supported by the tests. Canary Island species formed well-supported monophyletic clades corresponding to each island, with the exception of the position of Steganacarus sp. from La Gomera (GenBank Acc. no AJ 414186) which grouped with two species from Madeira and one from Sardinia, but there was lack of support for this relationship. The COI amino acid sequences provided weak support (57 /-) for close relationship of S. (S.) crassisetosus and S. (S.) similis. Both species were found in a clade with the S. (S.) carusoi, but the latter relationship remains unresolved due to the lack of support for S. (S.) carusoi + S. (S.) similis clade. The alignment for phylogenetic analysis using the sequence data from the D 2 region of 28 S rDNA comprised of 375 nucleotide positions for 6 species: S. (S.) spinosus, S. (S.) crassisetosus, S. (S.) similis, S. (S.) applicatus, S. (S.) magnus, and Phthiracarus longulus used as the outgroup. The number of variable sites was 62. Phylogenetic analysis of the D 2 marker performed using both the ML and NJ methods confirmed a distant relationship of S. (S.) spinosus to the other analysed Steganacaride species. In general, the two methods generated trees with the same topology (Fig. 5) where S. (S.) spinosus was placed basally to well-supported (BS = NJ 91 %, ML 56 %) clade (S. (S.) magnus, (S. (S.) similis, (S. (S.) crassisetosus, S. (S.) applicatus)). Moreover, both analyses recovered with moderate support (NJ 67 %, ML 65 %) a closer relationship between S. (S.) applicatus and S. (S.) crassisetosus., Published as part of Niedba��a, Wojciech & Dabert, Miroslawa, 2013, Madeira's ptyctimous mites (Acari, Oribatida), pp. 571-585 in Zootaxa 3664 (4) on pages 576-581, DOI: 10.11646/zootaxa.3664.4.9, http://zenodo.org/record/215799

Steganacarus (Steganacarus) crassisetosus (Willmann, 1939) (Figs. 2 A���G) Steganacarus applicatum (Sell.) var. crassisetosus n. var. Willmann, 1939 Measurements of specimen from Balcoes, Madeira. Prodorsum: length 379, height 192, width 288; setae of prodorsum: sensillum 101, interlamellar 177, lamellar 126, rostral 76, exobothridial 30; notogaster: length 818, height 566, width 555, setae of notogaster: c 1 185, c 1 / c 1 ��� d 1 =1.0, h 1 106, p 1 91; genitoaggenital plates: 227 �� 192, anaoadanal plates: 268 �� 172. Diagnosis. Large species. Colour brown. Integumental body sculpture slightly areolate. Prodorsum with distinct median crista; median and lateral fields very long; lateral carinae absent; sensilla long, thick covered with spines, sickle-shaped; interlamellar and lamellar setae resemble gastronotic setae covered with small spines, rostral setae long, rough, curved sagittally, in > le > ro > ex. Notogaster with flattened setae covered with spines, setae h 1 and p 1 located perpendicularly to the surface of notogaster, the remaining setae procumbent; setae h 1 and p 1 considerably thicker and shorter, about the half of length of procumbent setae; setae c 1 = c 1 ��� d 1; setae c 3 near the anterior margin, c 1 further away and c 2 in the most distant position; vestigial setae f 1 posteriorly of setae h 1; two pairs of lyrifissures ia and im. Ventral region: h > h ��� h; genitoaggenital plates with setal formula: 6 (4 + 2): 3; anoadanal plate with four setae at proximal margin which diminish towards the anterior margin, setae ad 3 very short, spiniform, an / not = 0.3. Chaetome of legs complete, setad d on femora I slightly remote from anterior margin. Comparison. Willmann (1939) has particularly emphasised one character differentiating crassisetosus from the main form applicatus (Fig. 2 H ��� M), that is short, thick and perpendicular setae h 1 and p 1 much shorter than the others; in applicatus the length of these setae is similar to that of the other ones. Moreover, Willmann claims that the size of crassisetosus is larger, however, in view of great variation in the size of phthiracaroid mites individuals, this feature should be treated with caution. There are two main morphological features differing S. (S.) applicatus and S. (S.) crassisetosus; the perpendicular setae h 1 and p 1 in S. (S.) crassisetosus are thick and shorter (h 1: e 1 = 0.55) than procumbent setae, while in S. (S.) applicatus, the setae h 1 and p 1 are of the same thickness and similar length (h 1, e 1 = 0.85) as procumbent setae. Rostral setae in S. (S.) crassisetosus are long, rough, curved sagittally but rostral setae in S. (S.) applicatus are shorter, spiniform and diriged forward. Steganacarus (S.) crassisetosus is also similar to S. (S. ) carusoi (Bernini & Avanzati, 1989) (Western Palaearctic species, reported from the Maghrebian and Atlanto-Mediterranean region). These species have perpendicular setae h 1 and p 1 and rostral setae of similar length directed inward but S. (S. ) carusoi has shorter lamellar setae, perpendicular setae h 1 and p 1 of the same length as the other procumbent setae, setae c 1 of notogaster situated slightly nearer to anterior border than c 3 setae and setae ad 3 in distal half are rugose or covered with small spines (versus shorter perpendicular setae than procumbent setae, greater distance of c 1 setae from anterior border and small, spiniform ad 3 setae in S. (S.) crassisetosus)., Published as part of Niedba��a, Wojciech & Dabert, Miroslawa, 2013, Madeira's ptyctimous mites (Acari, Oribatida), pp. 571-585 in Zootaxa 3664 (4) on page 576, DOI: 10.11646/zootaxa.3664.4.9, http://zenodo.org/record/215799

Steganacarus (Steganacarus) sol Balogh, 1958 (Figs 4. A, B) Measurements of specimen from locality 30���33. Prodorsum: length 429, height 127, sensillus 121, setae: interlamellar 338, lamellar and rostral 50; notogaster: length 757, height 505, c 1 293, d 1 252, d 2 141, e 1 353, e 2 252, h 1 404, h 2 318, p 1 429, p 2 192, p 3 101, p 4 91. Remarks. Interlamellar and notogastral setae thick, not flagellate, covered with small cilia along its length. Notogastral setae of different length. Setae of anoadanal plates rough. Four pairs of lrifissures: ia, im, ip and ips present. This species differs from S. (S.) werneri Mahunka, 1993 by the shorter lamellar and rostral setae., Published as part of Niedba��a, Wojciech & Ermilov, Sergey G., 2011, New and little known species of ptyctimous mites (Acari, Oribatida) from Ethiopia, pp. 60-68 in Zootaxa 2739 on page 65, DOI: 10.5281/zenodo.202962, {"references":["Balogh, J. (1958) Oribates nouvelles de l'Afrique tropicale. Revue de zoologie et de botanique africaines, 58, 1 - 34.","Mahunka, S. (1993) Some data to the Oribatid Fauna of Rwanda (Acari: Oribatida). Revue suisse de Zoologie, 100, 75 - 89."]}

The classical species, Steganacarus (Steganacarus) similis Willmann, 1939, is redescribed on the basis of material from the type locality, the island of Madeira. The diagnostic character-states indicate that this species belongs to the magnus group of steganacarid taxa. Another species, S. (Rhacaplacarus) ortizi Perez-Inigo, 1969, is reported for the first time on Madeira. The origin of these species on the island is discussed.

Steganacarus Ewing, 1917 Das System der Familie Steganacaridae Niedbala, 1986 basiert teilweise auf wenig hochwertigen Merkmalen im Sinne einer phylogenetischen Systematik (vgl. Niedbala 1992) und ist m. E. nicht hinreichend gesichert. Deshalb wird hier ein konservativer Standpunkt vertreten, indem die im Untersuchungsgebiet vorkommenden Arten alle der Gattung Steganacarus zugeordnet werden. Die weitere Einteilung in Untergattungen folgt weitgehend der morphologisch und enzymatologisch begründeten Verwandschaftsanalyse von Bernini & Avanzati (1991), die auch von Niedbala (1994a) übernommen wurde. Danach gelten die Untergattungen Steganacarus Ewing, 1917 und Tropacarus Ewing, 1917 als anerkannt. Atropacarus Ewing, 1917 wird mit zwei Untergattungen von Niedbala (1994a) in eine eigene Unterfamilie gestellt, womit die trennenden Merkmale m. E. überbewertet werden. Hier wird Atropacarus jedoch nur als weitere Untergattung zu Steganacarus gestellt. Für zwei Steganacarus-Arten konnten Bernini und Mitarbeiter ein sehr breites Variabilitätsspektrum belegen, das jeweils zwei bisher beschriebene Arten umfaßt: Steganacarus (Tropacarus) carinatus hat in der typischen Morphe einen wenig vorgewölbten vorderen Notogasteroberrand (forma carinata). Bei der forma pulcherrima, die bisher als eigene Art Tropacarus pulcherrimus (Berlese, 1887) galt, ragt der Notogasterrand weit über das Prodorsum. Morphologisch vergleichbar gibt es die typische Form, forma magna, von Steganacarus (Steganacarus) magnus mit einem normalen Notogasterrand und die forma anomala mit vorgewölbtem Rand, die als Steganacarus anomalus (Berlese, 1883) geführt wurde. Bei beiden Arten sind die Morphen durch kontinuierliche Zwischenformen verbunden. Im Bearbeitungsgebiet gibt es drei Untergattungen: Steganacarus (Steganacarus) Ewing, 1917 Typ: Hoploderma anomala Berlese, 1883 Diagnose: Körper mit Kutikulargruben, 15 ng, G am Rand mit 4 Borsten in einer Reihe, davor eine Borste (ad3) vom Rand entfernt. Solenidium auf TiIV ohne begleitende Borste d, NG ohne dorsomedianen Längskiel. Steganacarus (Tropacarus) Ewing, 1917 Typ: Hoplophora carinata C.L. Koch, 1841 Diagnose: wie S. (S.), aber mit dorsomedianem Längskiel auf gesamtem NG. Steganacarus (Atropacarus) Ewing, 1917 Typ: Hoplophora stricula C.L. Koch, 1836 Diagnose: wie S. (S.), aber Solenidium auf TiIV mit kleiner begleitender Borste d [64e]. - Einige Arten mit mehr als 16 ng. Schlüssel für alle Steganacarus-Arten: 1. Ano-Adanaldeckel mit (3-)4 kurzen, randständigen Borsten, 1 (-2) kurze Borsten etwas entfernt..................................................................................................................2 - Ano-Adanaldeckel mit (3-)4 langen, randständigen Borsten, deutlich länger als die 1 (- 2) entfernteren Borsten........................................................................................... 5 2. (1) Mit 19-20 Paar kurzen Notogasterborsten, Lamellar- und Interlamellarborste kurz und beborstelt. (+) Sensillus lang gebogen, außen beborstelt; schwacher ProdorsumKiel; Notogaster 390-500 µm, mit großen Gruben, ohne Kiel; Prodorsum 190-250 um; grau. [64g]..................................... S. (Atropacarus) clavigerus (Berlese, 1904) - Mit 15-16 Notogasterborsten und alle Prodorsumborsten kurz, spitz und glatt..................................3 3. (2) Notogaster mit deutlichem Längskiel über die ganze Länge..............................4 - Notogaster ohne oder mit nur schwach entwickeltem Längskiel im hinteren Bereich. (+) Sensillus stabförmig, am Ende mit kaum verbreiterter Membran (stumpi bis spitz). Prodorsum mit Längskiel; Lamellar- und Interlamellarborste kurz (um 15 bzw. um 30 um); 15 Paar Notogasterborsten kurz (30-50 µm); Analborsten und Adanalborsten kurz; Notogaster 520-660 µm, Prodorsum 250-350 µm (mitteleurop. Tiere); hellbraun. [66a,b]................................................... S. (Tropacarus) brevipilus (Berlese, 1923) 4. (3) Notogaster vorn dorsal mit Vorwölbung ("Kapuze"), die über Prodorsum ragt, darauf c. [65f].................................. forma pulcherrima (Berlese, 1887) von S. (Tropacarus) carinatus - Notogaster ohne Vorwölbung ("Kapuze"). (+) Sensillus borstenförmig, lang gebogen. Prodorsum mit Längskiel, im Profil fast rechtwinklig aufgewölbt. Körper rauh skulpturiert; 15 Paar Notogasterborsten + kurz, gebogen und spitz; Sensillus mit schlanker Endspindel, kaum beborstelt; Lamellar- und Interlamellarborste kurz; Notogaster 420- 725 µm, Prodorsum 210-350 µm; dunkelbraun. [65d,e]......................................................................forma carinata von S. (Tropacarus) carinatus (C. L. Koch, 1841) 5. (1) 16 Paar Notogasterborsten breit blattförmig. (+) Analborsten und Adanalborsten dünn; Prodorsum mit hohem Kiel; Sensillus lang gebogen, beborstelt; Prodorsum-Borsten± kurz, glatt; Notogaster grob skulpturiert mit Netzmuster; Notogaster 310- 475 µm, Prodorsum 200-250 µm; gelb (Österreich). [64f]................................................................................................ S. (Atropacarus) phyllophorus (Berlese, 1904) - Notogasterborsten nicht breit blattförmig...............................................................6 6. (5) Notogaster dorsal mit Vorwölbung, die über Prodorsum ragt. (+) Dorsomediane Notogasterborsten aufrecht abstehend, seitliche Notogasterborsten anliegend (vgl. auch Punkt 11). [65c]................................. forma anomala (Berlese, 1883) von S. magnus (Nicolet, 1855) - Notogaster dorsal ohne Vorwölbung.......................................................................7 7. (6) 15 Paar Notogasterborsten, h1 und ps1 besonders lang, die anderen kurz; Notogaster und Prodorsum vorn grob granuliert. (+) Prodorsum mit Längskiel; Interlamellarborste lang nach vorn anliegend; Sensillus dick borstenförmig, lang, gebogen; Körper braunschwarz; Notogaster 700-825 µm, Prodorsum 380-450 µm (Österreich). [66e-g].................................................................... Steganacarus herculeanus Willmann, 1953 - 15-16 Paar Notogasterborsten ± gleich lang; Notogaster nicht grob granuliert, teils mit grubiger Struktur....................................................................................................8 8. (7) alle Notogasterborsten ± abstehend..................................................................9 - einige Notogasterborsten anliegend...................................................................... 11 9. (8) 15 Paar Notogasterborsten stabförmig, rauh beborstelt, um 90 µm lang; Interlamellarborste lang (um 140 µm), länger als halber Abstand in-ro, dick stabförmig beborstelt; Tibia IV ohne kurze Borste d am Solenidium. (+) Prodorsum mit Längskiel; Prodorsum und Notogaster mit Gruben und Netzstruktur; Sensillus dünn borstenförmig, lang gebogen; Notogaster 450-600 µm, Prodorsum 300-380 µm; hell gelbbraun. [64a-c]........................................................ Steganacarus spinosus (Sellnick, 1920) - 16 Paar Notogasterborsten stabförmig oder borstenförmig (zusätzliche Borste x [64d]); Tibia IV mit winziger Borste d, die am Solenidium inseriert [64e]......................... 10 10. (9) Notogasterborsten stabförmig, rauhbeborstelt, um 90 µm lang. (+) Interlamellarborste in um 30-40 µm lang, kürzer als halber Abstand in-ro; Prodorsum mit Längskiel; Prodorsum und Notogaster mit Gruben und Netzstruktur; Sensillus lang borstenförmig, gebogen, beborstelt; Notogaster 340-590 µm, Prodorsum 170-290 µm lang; hell gelbbraun bis rosa. [64d,e]................. S. (Atropacarus) striculus (C.L. Koch, 1835) - Notogasterborsten borstenförmig, spitz endend und fein beborstelt. (+) Sensillus lang borstenförmig, gebogen und beborstelt; Prodorsum und Notogaster grubig skulpturiert; Sensillus lang borstenförmig, gebogen, beborstelt; Notogaster um 380 µm, Prodorsum um 207 µm lang; gelblich (bisher in Polen, Dänemark, England, Frankreich, Spanien)................................................. S. (Atropacarus) ivandae Niedbala, 1981 11. (8) 15 Paar Notogasterborsten, h1 und ps1 steif abstehend, beborstelt, um 170 µm; andere Notogasterborsten lang, anliegend, glatt, spitz und teils median verdickt. (+) Prodorsum vorn grubig; Interlamellarborste länger als Lamellarborste; Sensillus lang gebogen, distal beborstelt; Notogaster 740-760 µm, Prodorsum 390-400 µm; braun. [66c,d]..................................................... Steganacarus applicatus (Sellnick, 1920) - Seitliche der 15 Paar Notogasterborsten anliegend, dorso-mediane aufrecht, um 300 µm. (+) Sensillus stabförmig, geschwungen, mäßig lang; Prodorsum mit Längskiel; Körper grubig skulpturiert; Interlamellarborste und Lamellarborste lang, anliegend; Notogaster 700-1200 µm, Prodorsum 310-550 µm; dunkelbraun. [65a,b]........................................................ forma magna von Steganacarus magnus (Nicolet, 1855), Published as part of Weigmann, G. & Miko, L., 2006, Hornmilben (Oribatida) [pages 102 to 148], pp. 102-148 in Hornmilben (Oribatida) [Dahl, Tierwelt Deutschlands, Teil 76], Keltern :Goecke & Evers on pages 123-127

Steganacarus (Steganacarus) spinosus (Sellnick, 1920) [64a-c] Syn., Tax.: Phthiracarus spinosus Sellnick, 1920. Hoploderma spinosum: Willmann 1931. Steganacarus sp.: Balogh & Mahunka 1983; Niedbala 1992 (B). - Steganacarus punctulatus Sergenko, 1985. Ökologie: In Wäldern und feuchten moorigen Böden. Verbreitung: Europa., Published as part of Weigmann, G. & Miko, L., 2006, Hornmilben (Oribatida) [pages 102 to 148], pp. 102-148 in Hornmilben (Oribatida) [Dahl, Tierwelt Deutschlands, Teil 76], Keltern :Goecke & Evers on page 128

Steganacarus (Tropacarus) carinatus (C.L. Koch, 1841) [65d,e; 65f] Die Art tritt in 2 Morphen auf, die kontinuierlich miteinander verbunden sind; die geographischen Verbreitungen überschneiden sich partiell: S. carinatus forma carinata und forma pulcherrima (fehlt im nördlichen Europa). Syn., Tax.: Hoplophora carinata C.L. Koch, 1841 (CMA 32.9). Hoploderma c.: Michael 1898. Tropacarus c.: Willmann 1931. Steganacarus c.: Balogh & Mahunka 1983 (B); Niedbala 1992 (B). Steganacarus (Tropacarus) c.: Bernini & Avanzati 1989a (B). - Hoplophora carinata var. pulcherrima Berlese., 1887. Hoploderma pulcherrimum: Michael 1898. Tropacarus p.: Sellnick 1960. Steganacarus p.: Balogh & Mahunka 1983 (B); Niedbala 1992 (B). Steganacarus (Tropacarus) carinatus forma pulcherrima: Bernini & Avanzati 1988b (B): Niedbala 1994b. Ökologie: In Waldböden. Verbreitung: Paläarktis., Published as part of Weigmann, G. & Miko, L., 2006, Hornmilben (Oribatida) [pages 102 to 148], pp. 102-148 in Hornmilben (Oribatida) [Dahl, Tierwelt Deutschlands, Teil 76], Keltern :Goecke & Evers on pages 128-129

Steganacarus (Tropacarus) brevipilus (Berlese, 1923) [66a,b] Syn., Tax.: Phthiracarus (Trachychoplophora) b. Berlese, 1923. Tropacarus b.: Van der Hammen 1959. Steganacarus b.: Bernini 1971 (B); Mahunka 1991c (B); Niedbala 1992 (B). St. (Tropacarus) b.: Bernini & Avanzati 1989b (B); Niedbala 1994b (B). - Tropacarus curtipilus: Strenzke 1952. Ökologie: In moorigen Böden (selten). Verbreitung: Paläarktis, Asien., Published as part of Weigmann, G. & Miko, L., 2006, Hornmilben (Oribatida) [pages 102 to 148], pp. 102-148 in Hornmilben (Oribatida) [Dahl, Tierwelt Deutschlands, Teil 76], Keltern :Goecke & Evers on page 128

Steganacarus (Steganacarus) herculeanus Willmann, 1953 [66e-g] Syn., Tax.: Steganacarus herculeanus Willmann, 1953. Balogh & Mahunka 1983 (B); Niedbala 1992 (B). Ökologie: Waldböden; auch alpin Verbreitung: Frankreich, Österreich., Published as part of Weigmann, G. & Miko, L., 2006, Hornmilben (Oribatida) [pages 102 to 148], pp. 102-148 in Hornmilben (Oribatida) [Dahl, Tierwelt Deutschlands, Teil 76], Keltern :Goecke & Evers on page 128

Steganacarus (Atropacarus) clavigerus (Berlese, 1904) [64g] Syn., Tax.: Hoploderma clavigerum Berlese, 1904. Steganacarus clavigerus: Balogh & Mahunka 1983 (B). Atropacarus c.: Kamill & Baker 1980. Atropacarus (Atropacarus) c.: Niedbala 1992 (B), 1994b (B). Ökologie: Überwiegend in Wäldern. Verbreitung: Östliche Paläarktis, mediterran. In Mitteleuropa selten., Published as part of Weigmann, G. & Miko, L., 2006, Hornmilben (Oribatida) [pages 102 to 148], pp. 102-148 in Hornmilben (Oribatida) [Dahl, Tierwelt Deutschlands, Teil 76], Keltern :Goecke & Evers on page 129

Steganacarus (Atropacarus) phyllophorus (Berlese, 1904) [64f] Syn., Tax.: Hoploderma phyllophorum Berlese, 1904. Steganacarus phyllophorus: Schuster 1957 (B); Balogh & Mahunka 1983 (B). Atropacarus p.: Kamill & Baker 1980. Atropacarus (Atropacarus) p.: Niedbala 1992 (B), 1994b. - Steganacarus ropalus Feider & Suciu, 1957. Ökologie: Wald- und Heideböden. Verbreitung: Österreich, Südeuropa, mediterran., Published as part of Weigmann, G. & Miko, L., 2006, Hornmilben (Oribatida) [pages 102 to 148], pp. 102-148 in Hornmilben (Oribatida) [Dahl, Tierwelt Deutschlands, Teil 76], Keltern :Goecke & Evers on page 129

Steganacarus (Steganacarus) magnus (Nicolet, 1855) [65a,b; 65c] Die Art tritt in 2 Morphen auf, die kontinuierlich miteinander verbunden sind; die geographischen Verbreitungen überschneiden sich: St. magnus forma magna und forma anomala. Syn., Tax.: Hoplophora magna Nicolet, 1855. Hoploderma m.: Willmann 1931. Steganacarus m.: Van der Hammen 1952; Balogh & Mahunka 1983 (B); Bernini & Avanzati 1988a (B); Niedbala 1992 (B), 1994b. - Hoplophora anomala Berlese, 1883. Steganacarus a.: Balogh & Mahunka 1983 (B); Niedbala 1992 (B). Steganacarus magnus forma anomala: Bernini & Avanzati 1988a (B). Ökologie: In Wäldern. Verbreitung: Europa., Published as part of Weigmann, G. & Miko, L., 2006, Hornmilben (Oribatida) [pages 102 to 148], pp. 102-148 in Hornmilben (Oribatida) [Dahl, Tierwelt Deutschlands, Teil 76], Keltern :Goecke & Evers on page 128

Steganacarus (Atropacarus) striculus (C.L. Koch, 1835) [64d,e] Syn., Tax.: Hoplophora stricula C.L. Koch, 1835 (CMA 2.10). Steganacarus s.: Balogh & Mahunka 1983 (B). Atropacarus s.: Aoki 1980a (B); Kamill & Baker 1980; Niedbala 1992 (B), dort weitere Synonyme. Ökologie: Frische bis nasse Wald-, Wiesen- und Moorböden. Verbreitung: Holarktis., Published as part of Weigmann, G. & Miko, L., 2006, Hornmilben (Oribatida) [pages 102 to 148], pp. 102-148 in Hornmilben (Oribatida) [Dahl, Tierwelt Deutschlands, Teil 76], Keltern :Goecke & Evers on page 129

Steganacarus (Steganacarus) applicatus (Sellnick, 1920) [66c,d] Syn., Tax.: Phthiracarus a. Sellnick, 1920. Hoploderma applicatum: Willmann 1931. Steganacarus a.: Sellnick 1960; Balogh. & Mahunka 1983 (B); Bernini & Avanzati 1989c (B); Niedbala 1992 (B). Ökologie: In Waldböden. Verbreitung: Europa, Algerien., Published as part of Weigmann, G. & Miko, L., 2006, Hornmilben (Oribatida) [pages 102 to 148], pp. 102-148 in Hornmilben (Oribatida) [Dahl, Tierwelt Deutschlands, Teil 76], Keltern :Goecke & Evers on page 127

Steganacarus (Atropacarus) wandae Niedbala, 1981 Syn., Tax.: Steganacarus wandae Niedbala, 1981. Atropacarus (Atropacarus) w.: Niedbala 1992 (B). Ökologie: unklar Verbreitung: Europa (von Spanien bis Polen und Dänemark). Bisher noch nicht im Bearbeitungsgebiet gefunden., Published as part of Weigmann, G. & Miko, L., 2006, Hornmilben (Oribatida) [pages 102 to 148], pp. 102-148 in Hornmilben (Oribatida) [Dahl, Tierwelt Deutschlands, Teil 76], Keltern :Goecke & Evers on page 129

Three new species of the family Phthiracaridae, Austrophthiracarus longisetosus sp. nov., Phthiracarus allocotos sp. nov., Protophthiracarus amboroensis sp. nov. from Bolivia are described and figured. A comparison of morphological similarities with the most closely related species is presented. Additional descriptions and taxonomical notes for three ptyctimous species: Acrotritia peruensis (Hammer, 1961), Acrotritia vestita (Berlese, 1913), and Steganacarus (Rhacaplacarus) sedecimus Niedbala, 2004 are added. A list of twenty six ptyctimous species from Bolivia is presented, ten of these species are new records for the fauna of Bolivia. A key to all species of ptyctimous mites of Bolivia is presented.

An annotated checklist of ptyctimous mite taxa, collected from the Joint Russian-Vietnamese Biological expedition (October 2013–April 2014) in four forest zones (Dong Nai Biosphere Reserve, Dong Nai Culture and Nature Reserve, Bu Gia Map National Park, Bi Dup—Nui Ba National Park) is provided. It includes 28 species, 16 genera and six families. One new subgenus, Euphthiracarus ( Parapocsia ) Niedbala subgen. nov., and five new species, Acrotritia paragranulata Niedbala sp. nov., Hoplophthiracarus clavatus Niedbala sp. nov., Steganacarus ( Rhacaplacarus ) spinus Niedbala sp. nov., Euphthiracarus ( Euphthiracarus ) quasitakahashii Niedbala sp. nov. and Euphthiracarus ( Parapocsia ) medius Niedbala sp. nov., are described. Supplementary descriptions of Apoplophora minuscula Niedbala, 2013 (see Niedbala & Ermilov 2013), Arphthicarus parasentus Niedbala, 2000 and Atropacarus ( Hoplophorella ) stilifer (Hammer, 1961) are presented.

Three new species of oribatid mites of the genus Plonaphacarus , Plonaphacarus sidorchukae sp. nov., Plonaphacarus rybalovi sp. nov. and Plonaphacarus hamulus sp. nov., are described from Bale Mountains National Park (Afrotropical region, Ethiopia). Morphological and taxonomical remarks about Steganacarus ( Steganacarus ) sol and Steganacarus ( Steganacarus ) vestitus are presented.

Steganacarus magnus (NICOLET, 1855), in Bodenstreu und anderen Mikrohabitaten sehr selten, Published as part of Beck, L. & Woas, S., 1991, Die Oribatiden-Arten (Acari) eines suedwestdeutschen Buchenwaldes I., pp. 37-82 in carolinea 49 on page 43

Steganacarus herculeanus WILLMANN, 1953, h��ufigste Phthiracaride in der Bodenstreu, Published as part of Beck, L. & Woas, S., 1991, Die Oribatiden-Arten (Acari) eines suedwestdeutschen Buchenwaldes I., pp. 37-82 in carolinea 49 on page 43

The classical taxon, Steganacarus (Steganacarus) magnus (Nicolet, 1855) is redescribed on the basis of topotypic material. The study and the comparison of many European and North African populations reveals a variation in the size of the anterior notogastral tectum and consequently enables another classic entity, S. (Steganacarus) anomalus (Berlese, 1883), to be put into synonymy. These two different ecophenotypes, in the authors' opinion, seem to be linked to the presence of xeric low-altitude southern environments (S. (S.) magnus f. anomala) or cool higher-altitude mid-northern environments (S. (S.) magnus f. magna).

The immatures of this oribatid fed specifically upon the decaying schlerenchyma and endoderm of conifer needles and the nonvascular portion of leaf petioles from deciduous species. The adults usually fed upon the same foodstuff but frequently wandered and fed upon decaying filter paper when not engaged in reproductive activity. Immatures were removed from the plant tissues during most of the months of the year in their laboratory environment. As many as five immatures were found in sections of conifer needles of 8–10 mm. length. At the completion of growth from egg to adult requiring approximately 5 months, a perforated cuticle and vascular bundles remained while most of the schlerenchyma and endoderm were reduced to particles of 1 µ to 20 µ size.

The classical taxon, Steganacarus (Steganacarus) applicatus (Sellnick, 1920), is redescribed on the basis of the type material found in Berlese's Collection. Analysis of many European and North African populations reveals the existence of at least another closely related new species with a geographic distribution restricted to the west-Mediterranean countries and testifying ancient land connections.

Morphological, ecological, biogeographic and genetic evidence is presented to suggest the synonymy of Steganacarus (Tropacants) pulcherrimus (Berlese) and the earlier S. (T.) carinatus (C.L.Koch).

Two new species closely related to Steganacarus (Steganacarus) magnus (Nicolet) are described. S. (S.) michaeli can also be genetically differentiated from known congeneric species such as S. (S.) hirsutus Perez-Inigo by biochemical methods. Some biogeographic observations are made about the collection sites of the two new entities.

Steganacarus clavigerus (Berlese, 1904) Hoploderma clavigerum Berlese, 1904a, p. 27.S; 1913, p. 104, pl. 8 fig. 104. Phthiracarus clavigerus, Lombardini, 1936, p. 47. In the Berlese Collection the following slides are present: 141/5, 8 from the "Gardino di Boboli", Florcnce; and 141/9 from Pisa. The last-mentioned specimen is designated as type, although Berlese (1904a) mentioned "Boboli" as type-locality; I collected indeed topotypes in the "Gardino". I do not know if Sellnick's (1928, p. 40) description of Hoploderma clavigera indeed represents the species; the same applies to Phthiracarus lutulentus Sellnick (1920, p. 37)., Published as part of van der Hammen, L., 1959, Berlese's Primitive Oribatid Mites, pp. 1-93 in Zoologische Verhandelingen 40 on page 44

Steganacarus applicatus (Sellnick, 1920) Phthiracarus applicatus Sellnick, 1920, p. 36. Hoplophora stricula, Berlese, 1896, fasc. 78 (3); 1896b, p. 19. Phthiracarus berlesei Oudemans, 1915, p. 214; Lombardini, 1936, p. 47. Hoploderma applicata, Sellnick, 1928, p. 40. Hoploderma applictum, Willmann, 1931, p. 191, fig. 349. The species described by Berlese (1896) as Hoplophora stricula is certainly not identical with Hoplophora stricula C. L. Koch (1836). For this reason Oudemans (1915) gave the new name berlesei to stricula Berlese (non Koch); he overlooked, however, that Michael (1898) already used the name Phthiracarus berlesei for Tritia decumana sensu Berlese (1883), so that P. berlesei Oudemans is a primary homonym. In the Collection I did not see the specimens from Vallombrosa on which Berlese's 1896 description was based, but slide no. 31/23 from Tiarno bears the label " Phtiracarus Berlesei Oudemans "; it contains Steganacarus applicatus. Two cotypes of S. applicatus Sellnick from L��tzen are also present in the Collection; on the label Berlese added the name " Berlesei Michael " (certainly a lapsus for " Berlesei Oudemans "!) as a synonym. For these reasons I place Berlese's 1896 record of Hoplophora stricula in the synonymy of S. applicatus; the figure is, however, certainly inaccurate because the two posterior pairs of dorsal hairs are not drawn as strikingly erect., Published as part of van der Hammen, L., 1959, Berlese's Primitive Oribatid Mites, pp. 1-93 in Zoologische Verhandelingen 40 on page 43

Steganacarus vitrinum (Berlese, 1913) Hoploderma vitrinum 1) Berlese, 1913, p. 103, pl. 8 figs. 100, 100a, 100b 2). The present species was not mentioned by Lombardini; I do not know if the type-material is indeed no more present because the species was omitted from my private catalogue when I visited Florence. The figure shows a Phthiracarid mite with convex anal covers, which points to Steganacarus; the structure of the tegument (although vague), the pale colour, and the shape of the notogastral hairs point to the striculus-group. 1) I do not know if vitrinum (from vitrum) is an adjective. 2) In the explanation of pl. 8 erroneous reference is made to figs. 99a, 99b., Published as part of van der Hammen, L., 1959, Berlese's Primitive Oribatid Mites, pp. 1-93 in Zoologische Verhandelingen 40 on page 44

Steganacarus somalicus (Berlese, 1923) Phthiracarus (Trachyhoplophora) somalicus Berlese, 1923, p. 258. Berlese described the present species as closely related to S. striculus, possibly even a variety. In the Berlese Collection the original material (nos. 208/ 9-14, no type-selection) from "Somalia italiana, Foci del Giuba" is labelled as " Phthiracarus echidninus Berl. var. somalicus "; apparently echidninus is an unpublished synonym of striculus. As mentioned by Berlese, the species differs from striculus by the rather short sensillus that is thickened towards the end. The notogastral hairs appear to be slightly longer than in striculus; they are stif f and coarse at the end., Published as part of van der Hammen, L., 1959, Berlese's Primitive Oribatid Mites, pp. 1-93 in Zoologische Verhandelingen 40 on page 45

Steganacarus magnus (Nicolet, 1855) Hoplophora magna Nicolet, 1855, p. 472, pl. 10 fig. 4; Berlese, 1892a, fasc. 67 (9); 1896b, p. 19. Hoploderma magnum, Sellnick, 1928, p. 140; Willmann, 1931, p. 191, figs. 347, 348. This common European species was recorded by Berlese from Vallombrosa. I have seen no specimen in the Berlese Collection., Published as part of van der Hammen, L., 1959, Berlese's Primitive Oribatid Mites, pp. 1-93 in Zoologische Verhandelingen 40 on pages 42-43

Steganacarus Ewing, 1917 Hoplophora, Berlese, 1892d, fasc. 67 (5). Steganacarus Ewing, 1917, pp. 125, 130. Atropacarus Ewing, 1917, pp. 125, 130. Trachyhoplophora Berlese, 1923, p. 257. Species of the present genus have been listed with the genera Hoplophora C. L. Koch, Hoploderma Michael, Steganacarus Ewing, Atropacarus Ewing, and Trachyhoplophora Berlese. C. L. Koch used the name Hoplophora in 1836 when he described two species: Hoplophora decumana (now Oribotritia) and Hoplophora stricula (now Steganacarus). In 1842 C. L. Koch published a diagnosis of the genus:, and designated Hoplophora laevigata C. L. Koch (1841) as type. Because the name Hoplophora appeared to be preocupied (Hoplophora Perty, Orthoptera), Michael (1898) gave the new name Hoploderma to the genus. In the present case it is of no importance that Michael regarded laevigata as a possible synonym of " Phthiracarus " nudus, so that we can follow the general use that laevigata is also the type of Hoploderma. Opinion 204 of the "Opinions and declarations rendered by the International Commission on Zoological Nomenclature" deals with Koch's designation of types. In the present case (type-species described in 1841, two other species in 1836) it will be necessary to submit an application to the Commission for the use of the Plenary Powers, because adoption of the rules (i.e. to place Oribotritia in the synonymy of Hoploderma) would lead to instability and confusion; the possibility of this use is mentioned in the opinion in question. Because Hoploderma Michael (1898) has laevigata as type, it must be placed in the synonymy of Phthiracarus Perty (1839) of which, according to Jacot (1930, p. 211), the type (P. contractilis Perty, 1841) is a synonym of laevigata. Further notes on the genus Phthiracarus are given below. Although the name Hoploderma has been in use for some time for the present genus, most authors followed Jacot (1930) in accepting the correct name Steganacarus Ewing, 1917 (type: Hoplophora anomala Berlese). Ewing (1917) also described the genus Atropacarus (type: Hoplophora stricula C. L. Koch); this genus is not yet sufficiently characterized. In 1923 Berlese created Trachyhoplophora, a subgenus of Phthiracarus (!), with Hoplophora magna Nicolet as type; because magna is related to anomala, Trachyhoplophora becomes a synonym of Steganacarus. Berlese's species can be arranged in some groups that are based on the general habitus: anomalus, magnus, applicatus (Steganacarus s.str.); phyllophorus (related to Hoplophorella, but dif ferent in the arrangement of the anal hairs); clavigerus, vitrinum, remigerus, somalicus (? Atropacarus); pardinus and rapax (which have a superficial resemblance to anomalus and magnus, but are different in several characters)., Published as part of van der Hammen, L., 1959, Berlese's Primitive Oribatid Mites, pp. 1-93 in Zoologische Verhandelingen 40 on pages 41-42

Steganacarus pardinus (Berlese, 1916) Hoploderma pardinum Berlese, 1916b, p. 337. The present species was described after a specimen from East Africa; the type (no. 183/47) is still present in the Collection. Berlese compares the species with magnus and anomalus and in general shape it reminds indeed of these species. The notogastral hairs are, however, very small, hardly visible, slightly pointing backwards, and irregularly curved. The type-specimen is closed (as Berlese mentioned already); the colour is dark-brown., Published as part of van der Hammen, L., 1959, Berlese's Primitive Oribatid Mites, pp. 1-93 in Zoologische Verhandelingen 40 on page 44

Steganacarus remigerus (Berlese, 1923) Phthiracarus (Trachyhoplophora) remigerus Berlese, 1923, p. 258. The present species is related to Steganacarus striculum. The notogaster shows similar pits. The notogastral hairs are thickened towards the end, although the top is rounded or somewhat pointed. The sensillus is curved, slightly thickened towards the end. The original slides are still present in the Berlese Collection, viz. the nos. 208/15 (type), 208/16-18; the specimens originate from "Somalia italiana, Foci del Giuba''., Published as part of van der Hammen, L., 1959, Berlese's Primitive Oribatid Mites, pp. 1-93 in Zoologische Verhandelingen 40 on page 45