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May 5, 2017 - Lohmanniidae (Acari, Oribatida) from South Africa ... This work is part of our continuing study of oribatid mites (Acari, Oribatida) of South Africa ( ...
Systematic & Applied Acarology 22(5): 666–682 (2017) http://doi.org/10.11158/saa.22.5.6 Article

ISSN 1362-1971 (print) ISSN 2056-6069 (online)

http://zoobank.org/urn:lsid:zoobank.org:pub:102D38BB-F1B7-4DE4-9374-EFACE87A9700

Contribution to the knowledge of oribatid mites of the family Lohmanniidae (Acari, Oribatida) from South Africa SERGEY G. ERMILOV1*, ELIZABETH A. HUGO-COETZEE2,3 & ALEXANDER A. KHAUSTOV1 1

Tyumen State University, Tyumen, Russia. E-mail: [email protected], [email protected] National Museum, Bloemfontein, South Africa. E-mail: [email protected] 3 Department of Zoology and Entomology, University of the Free State, Bloemfontein, South Africa * Corresponding author 2

Abstract The genus Lohmannia (Oribatida, Lohmanniidae) is recorded for the first time in South Africa. A new species of Lohmannia is described from the nest of termites of the Franklin Game Reserve on Naval Hill, Bloemfontein, based on the adult and tritonymphal instar. Lohmannia (Lohmannia) lerallana sp. nov. differs from L. (Lohmannia) turcmenica Bulanova-Zachvatkina, 1960 by the presence of distinct macrofoveolate ornamentation on the body, setiform subcapitular setae m1 and long lateral and posterior notogastral setae. A supplementary description of Papillacarus angulatus Wallwork, 1962 is presented based on the South African specimens. Information on distribution and ecology of known lohmanniids in South Africa is provided. Key words: oribatid mites, Lohmannia, Papillacarus, new species, supplementary description, Africa

Introduction This work is part of our continuing study of oribatid mites (Acari, Oribatida) of South Africa (e.g. Ermilov et al. 2011, 2017; Ermilov & Hugo-Coetzee 2012a, b; Hugo-Coetzee 2013, 2014, 2016) and includes data on the family Lohmanniidae. At present, Lohmaniidae of South Africa are poorly known, and only six species, of which two are fossils, have been recorded. These species are the extant species, Annectacarus eksteeni Coetzee, 2001, Paulianacarus groblerae Coetzee, 2001, P. barlowi Coetzee, 2001, and Cryptacarus promecus Grandjean, 1950, and the fossil species from Holocene deposits, Papillacarus brinki Coetzee 2001 and Torpacarus cf. omittens (Coetzee 2001a, b; Hugo-Coetzee & Avenant 2011). During taxonomic identification of material collected from the nest of termites of the Franklin Game Reserve on Naval Hill (Bloemfontein, South Africa), we found two species, one of which is new for science, belonging to the genus Lohmannia Michael, 1898, and the other species Papillacarus angulatus Wallwork, 1962, which is distributed in the Ethiopian and southern Palaearctic regions and Brazil (Subías 2004, online version 2017) of which extant species was previously unknown in South Africa. The genus Lohmannia is recorded for the first time in South Africa. The primary goal of the paper is to describe and illustrate a new species under the name Lohmannia (Lohmannia) lerallana sp. nov., based on the adult and tritonymphal instar. Lohmannia was proposed by Michael (1898) with Michaelia paradoxa Haller, 1884 as type species. At present, this genus comprises two subgenera and 29 species, which are distributed in the tropical and subtropical regions (Subías 2004, online version 2017; Ermilov 2016, 2017). The main generic traits were summarized by Balogh (1961) and Grandjean (1950). Data on the morphology of juvenile instars of Lohmannia are known for 10 species (Norton & Ermilov 2014; Ermilov 2017), and a 666

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comparative morphological analysis for the majority of these species is given in Ermilov et al. (2014). The secondary goal of the paper is to present a supplementary description of Papillacarus angulatus based on the South African specimens. The original description (Wallwork 1962) is incomplete, partially lacking measurements and morphological detail of some structures. In addition, the data on distribution and ecology of known Lohmanniidae species in South Africa are provided.

Material and methods Material. The collection locality and habitat for each studied lohmanniid species are given in the respective "Material examined" section. Methods. Specimens were mounted in lactic acid on temporary cavity slides for measurement and illustration. Body length was measured in lateral view, from the tip of the rostrum to the posterior edge of the ventral plate. Notogastral width refers to the maximum width in dorsal aspect. Lengths of body setae were measured in lateral aspect. All body measurements are presented in micrometers (μm). Formulas for leg setation are given in parentheses according to the sequence trochanter– femur–genu–tibia–tarsus (famulus included). Formulas for leg solenidia are given in square brackets according to the sequence genu–tibia–tarsus. Morphological terminology used in this paper follows that of F. Grandjean (e.g. 1950): see Travé & Vachon (1975) for general references, Norton (1977) for leg setal nomenclature, and Norton & Behan-Pelletier (2009) for overview. Drawings were made with a camera lucida using a Carl Zeiss transmission light microscope “Axioskop-2 Plus”.

Systematics Lohmannia (Lohmannia) lerallana sp. nov. (Figs 1–13) Diagnosis. Adult and tritonymphal instar. Body size: adult 898–946 × 365–415, tritonymph 763– 792 × 315–332. Body surface micro- and macrofoveolate. Lateral parts of prodorsum undulate, with four to six small teeth basally. Rostral setae widely phylliform, lamellar, interlamellar and anterior exobothridial setae narrowly phylliform, posterior exobothridial setae disk-like. Bothridial setae pectinate, with 11 to 12 branches on one side and several barbs on opposite sides. Notogastral setae narrowly phylliform, lateral and posterior setae with long, thin tips, c1, d1, e1, h1 shortest, c3, d3, f2, h2, h3, p1–p3 longest. Eight transverse bands present on notogaster, S7, S8 and S9 hardly visible, other bands distinct, S2 complete in adult and interrupted medially in tritonymph. Subcapitular setae a and m1 setiform, thickened, m2 and h phylliform. Epimeral setae short, narrowly phylliform, 2a, 3a and 4a shortest. Genital setae slightly phylliform. Anal setae setiform. Adanal setae slightly phylliform, with long, thin tips. Adult (Figs 1–11) Description. Measurements. Large species. Body length: 898 (holotype), 898–946 (five paratypes); notogaster width: 365 (holotype), 365–415 (five paratypes). 2017

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Integument (Figs 1–5, 8, 9, 11). Body color yellow-brownish. Body surface (including subcapitular mentum, genae, palps, genital, anal and adanal plates) and legs with dense microfoveolae forming mostly micropolygonal ornamentation. Also, dorsal and ventral sides (including subcapitular mentum, genital and adanal plates) and leg femora macrofoveolate (diameter of macrofoveolae up to 10). Body sculpture and reticulate pattern absent.

FIGURE 1. Lohmannia (Lohmannia) lerallana sp. nov., adult: dorsal view (legs not shown). Scale bar 100 μm.

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FIGURE 2. Lohmannia (Lohmannia) lerallana sp. nov., adult: ventral view (gnathosoma and legs except basal parts not shown). Scale bar 100 μm.

Prodorsum (Figs 1–3). Roughly triangular in dorsal view, occupying about 2/5 of dorsal length, distinctly undulate laterally, with four to six small teeth (t) basally. Rostrum rounded or slightly truncate. Rostral setae (ro, 106–110) widely phylliform, barbed. Lamellar (le, 106–110), interlamellar (in, 123–131) and anterior exobothridial (exa, 82) setae narrowly phylliform, barbed, exa directed anteromediad. Posterior exobothridial setae (exp, 36–45) disk-like, barbed. Bothridial 2017

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setae (bs, 118–127) pectinate, with 11 to 12 branches on one side and several barbs on opposite sides each. Postbothridial transverse band (Sb) present, interrupted behind bothridia.

FIGURES 3–7. Lohmannia (Lohmannia) lerallana sp. nov., adult: 3—medioposterior part of prodorsum and anterior part of notogaster, lateral view; 4—medioposterior part of hysterosoma, lateral view; 5—subcapitulum, ventral view; 6—chelicera (basal part not shown), right, antiaxial view; 7—palp, right, antiaxial view. Scale bars 100 μm (3, 4), 50 μm (5, 6; 7).

Notogaster (Figs 1–4). Anterior notogastral margin straight. Sixteen pairs of notogastral setae narrowly phylliform, barbed, lateral and posterior setae with long, thin, smooth tips, c1, d1, e1, h1 (65– 73) shorter than f1 (82–90), c2, d2 (90–98), e2 (110–131) and c3, d3, f2, h2, h3, p1–p3 (155–168). Lyrifissures ia, im, ih and ip distinct, ips not visible. Eight transverse bands present dorsally, S7, S8 and S9 hardly visible, other bands well-developed, S2 complete, others interrupted medially. Two additional pairs of bands (Sva, Svp) located laterally to anogenital region. Gnathosoma (Figs 5–7). Subcapitulum longer than wide (217–233 × 180 –205), with one pair of triangular lateral tubercles. Subcapitular setae a (61–65) and m1 (61–65) setiform, thickened, a smooth, m1 barbed, m2 (57–61) and h (57–61) phylliform, barbed. Adoral setae or1 (36–41) lobeshaped, slightly roughened distally, or2 (45–49) thick, blunt-ended, with one tooth in distal parts, or3 (36–41) thickened, pointed. Palps (102–110) with setation 0-1-0-3-10(+ω). Three distal setae 670

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connected basally. Solenidia similar to palptarsi in length. Postpalpal setae (12–14) spiniform, slightly barbed. Chelicerae (217–233) with numerous small teeth (t) antero-dorsally, setae chb (49– 61) setiform, barbed, cha (8) spiniform, thin, smooth. Trägårdh’s organs (Tg) triangular, rounded distally.

FIGURES 8–11. Lohmannia (Lohmannia) lerallana sp. nov., adult: 8—leg I, without trochanter, right, antiaxial view; 9—femur and genu of leg II, right, antiaxial view; 10—genu and tibia of leg III, left, antiaxial view; 11—leg IV, without trochanter, left, antiaxial view. Scale bars 50 μm.

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Epimeral and lateral podosomal regions (Fig. 2). Epimeral setal formula 3-1-3-4. Setae 2a, 3a, 4a (20–28) shorter than other setae (41–53), all narrowly phylliform, barbed. Anogenital region (Figs 2, 4). Genital setae (41–49) slightly phylliform, barbed. Transverse genital furrows distinct. Two pairs of anal setae (an1, an2, 77–90) setiform, barbed. Four pairs of adanal setae (ad1–ad4, 106–114) slightly phylliform, barbed, with long, thin, smooth tips. Lyrifissures ian and iad not visible. Legs (Figs 2, 8–11). Claw of each leg smooth. Formulas of leg setation and solenidia: leg I (05-3-5-17) [2-1-2], leg II (0-6-3-5-13) [1-1-1], leg III (2-3-2-3-12) [1-1-0], leg IV (2-3-2-2-12) [1-00]; homology of setae and solenidia indicated in Table 1. Solenidia ω1 on tarsi I, ω on tarsi II and φ on tibiae III thickened, blunt-ended; other solenidia thin, setiform. Famuli (ɛ) tubercle-like, inserted posterolateral to ft”. Solenidia ω2 on tarsi I free, not coupled with setae. TABLE 1. Leg setation and solenidia of adult and tritonymph Lohmannia (Lohmannia) lerallana sp. nov. Leg

Tr

Fe

Ge

Ti

Ta

I

-

d, (l), bv”, v”

d, (l), σ”, σ’

d, xt1, xt2, l’, v’, φ

(ft), it’, (tc), (p), (u), (a), s, m, n, (pv), ɛ, ω1, ω2

II

-

d, l’, la”, lp”, bv”, v”

d, (l), σ

d, xt1, xt2, l’, v’, φ

(ft), (tc), (p), (u), (a), s, (pv), ω

III

l', v'

d, l’, ev’

d, l’, σ

d, l’, v’, φ

(ft), (tc), (p), (u), a’, s, (pv)

IV

l', v'

d, l’, ev’

d, l’, σ

d, l’

(ft), (tc), (p), (u), a’, s, (pv)

Note: Roman letters refer to normal setae, Greek letters to solenidia (except ɛ = famulus). Single prime (') marks setae on anterior and double prime (") setae on posterior side of the given leg segment. Parentheses refer to a pair of setae. Tr—trochanter, Fe—femur, Ge—genu, Ti—Tibia, Ta—tarsus.

Tritonymph (Figs 12, 13) Description. Measurements. Body length: 763, 792 (two tritonymphs); notogaster width: 315, 332 (two tritonymphs). Integument (Figs 12, 13). Body color light grey. Generally body surface similar to adult, but anterior part of notogaster densely striate. Prodorsum (Fig. 12). Similar to adult, but setae shorter: ro 82–94, le 110, in 106–118, exa 69–73, exp 28, bs 106–114. Notogaster (Figs 12, 13). Generally similar to adult. Anterior notogastral margin slightly convex medially. Setae c1, d1, e1, h1 (49–53) shorter than f1 (73), c2, d2 (77–90), e2 (90–110) and c3, d3, f2, h2, h3, p1–p3 (135–143). Transverse bands S2 interrupted medially. Gnathosoma (Fig. 13). Similar to adult, but sizes of subcapitulum, palps and chelicerae, their structures and setae smaller. Subcapitulum size 151–165 × 180–205, a, m1 49–53, m2, h 36–45, or1, or3 30–32, or2 36–41, palps 82, postpalpal setae 10, chelicerae 151–165, chb 41–53, cha 6. Epimeral and lateral podosomal regions (Fig. 13). Similar to adult, but setae shorter, 2a, 3a, 4a 16, other setae 32–36. Anogenital region (Fig. 13). Generally similar to adult, but setae shorter, genital plates with eight pairs of setae and lyrifissures iad visible. Genital setae 28–36, an1, an2 53–61, ad1–ad4 82–90. Legs (Fig. 13). Similar to adult. Material examined. Holotype, five paratypes and two tritonymphs: South Africa, Bloemfontein, Franklin Game Reserve on Naval Hill, 29°05’54.9’’S, 26°14’18.9’’E, 1477 m a.s.l., in the nest of termites Trinervitermes trinervoides (Sjöstedt), sample #6, 9.XI.2016 (A.A. Khaustov, S.G. Ermilov & E.A. Hugo-Coetzee).

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FIGURE 12. Lohmannia (Lohmannia) lerallana sp. nov., tritonymph: dorsal view (legs not shown). Scale bar 100 μm.

Type deposition. The holotype (alcohol) and one paratype (alcohol) are deposited in the collection of the National Museum, Bloemfontein, South Africa; one paratype (alcohol) is deposited in the collection of the Senckenberg Institute, Görlitz, Germany; three paratypes (alcohol) and two tritonymphs (alcohol) are deposited in the collection of the Tyumen State University Museum of Zoology, Tyumen, Russia. 2017

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FIGURE 13. Lohmannia (Lohmannia) lerallana sp. nov., tritonymph: ventral view (legs except basal parts not shown). Scale bar 100 μm.

Etymology. The species name ‘lerallana’ is derived from the Sotho (local language) word for hill, ‘leralla’, referring to the type locality, Naval Hill. Remarks. The adult and tritonymphal instar of Lohmannia (Lohmannia) lerallana sp. nov. is morphologically most similar to L. (Lohmannia) turcmenica Bulanova-Zachvatkina, 1960 (see Ermilov et al. 2014; Ermilov 2017) from the Tropical and Subtropical regions in having disk-like posterior exobothridial setae, phylliform prodorsal and notogastral setae, and the absence of sculpture and reticulate ornamentation on the notogaster, however it differs from the latter by the presence of distinct macrofoveolate ornamentation on the body (vs. macrofoveolae absent), setiform 674

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subcapitular setae m1 (vs. phylliform) and long lateral and posterior notogastral setae (vs. medium size). Papillacarus angulatus Wallwork, 1962 (Figs 14–21) Adult Supplementary description (based on specimens from South Africa). Measurements. Medium size. Body length: 547 (four specimens); notogaster width: 232–249 (four specimens). Integument (Figs 14–19, 21). Body color yellow-brownish. Body surface (including subcapitular mentum, genae, palps, genital, anal and adanal plates) and legs with dense microfoveolae forming mostly micropolygonal ornamentation. Also, dorsal and ventral sides (including subcapitular mentum, genae, genital and adanal plates) and leg femora with dense conical papillae (their length up to 8) except basal part of prodorsum. Body sculpture and reticulate pattern absent. Prodorsum (Figs 14, 16). Roughly triangular in dorsal view, occupying about 2/5 of dorsal length. Rostrum and its lateral sides distinctly undulate. Rostral, lamellar, interlamellar and anterior exobothridial (all 53–57) and posterior exobothridial (65–73) setae setiform, barbed. Bothridial setae (82–86) pectinate, with 15 to 18 branches on one sides and several barbs on opposite sides. Postbothridial transverse band present, complete. Notogaster (Figs 14–17). Anterior notogastral margin almost straight. Sixteen pairs of primary notogastral setae and 11 to 12 pairs of additional neotrichal setae present, all setiform, barbed or shortly ciliate. Setae c1, d1, e1, f1, h1 (28–32) shorter than c2, d2 (41–49), e2 (69–73) and c3, d3, f2, h2, h3, p1–p3 (73–82). Neotrichal setae of two types: three pairs long (m, 41–49; n, 69–73; r, 73–82) and eight to nine pairs (1–8[or 9]) short (20–28). Lyrifissures ia, im, ih and ip distinct, ips not visible. Four transverse bands present dorsally, S2 complete, others interrupted medially. One additional pair of bands (Svp) located laterally to anogenital region. Gnathosoma (Fig. 15). Subcapitulum longer than wide (139–143 × 106 –123), with one pair of lateral tubercles. Subcapitular setae setiform, a (32–41) smooth, m1 (32–41) barbed, m2, m3, m4, h (24–28) setiform, shortly ciliate. Adoral setae smooth, or1 (20) lobe-shaped, or2 (24–28) thick, bluntended, or3 (20) thickened, pointed. Palps (61) with setation 0-1-0-3-10(+ω). Distal three setae connected basally. Solenidia similar to palptarsi in length. Postpalpal setae (8) spiniform, smooth. Chelicerae (143–151) antero-dorsally with some small teeth, setae chb (32) setiform, smooth, cha (8) spiniform, thin, smooth. Trägårdh’s organs triangular, pointed distally. Epimeral and lateral podosomal regions (Figs 15, 17). Epimeral setal formula 9-4-3-4. Medial setae 1a, 2a, 3a, 4a and one pair of lateral setae of epimere I minute (12), smooth. Other setae setiform, shortly ciliate, anterior pair of epimere I (32–36) longer than three pairs of lateral setae on epimere II (28) and other setae (20–24). Anogenital region (Figs 15, 17). Genital setae (24–28) setiform, shortly ciliate. Transverse genital furrows distinct. Two pairs of anal (36–41) and four pairs of adanal (49–57) setae setiform, barbed. Lyrifissures ian not visible, iad distinct. Legs (Figs 15, 18–21). Claw of each leg smooth with one small tubercle ventrobasally. Formulas of leg setation and solenidia: leg I (0-5-3-4-17) [2-1-2], leg II (0-6-3-4-13) [1-1-2], leg III (2-4-2-312) [1-1-0], leg IV (2-3-2-3-11) [1-0-0]; homology of setae and solenidia indicated in Table 2. Solenidia ω1 on tarsi I, ω1 and ω2 on tarsi II and φ on tibiae III thickened, blunt-ended; other solenidia thin, setiform. Famuli tubercle-like, inserted laterally to solenidia ω1. Solenidia ω2 coupled with setae ft”. 2017

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FIGURE 14. Papillacarus angulatus Wallwork, 1962, adult: dorsal view (legs not shown). Scale bar 100 μm.

Material examined. Four specimens: South Africa, Bloemfontein, Franklin Game Reserve on Naval Hill, 29°05’54.9’’S, 26°14’18.9’’E, 1477 m a.s.l., in the nest of termites Trinervitermes trinervoides (Sjöstedt), sample #6, 9.XI.2016 (A.A. Khaustov, S.G. Ermilov & E.A. Hugo-Coetzee). Material deposition. Four studied specimens (alcohol) are deposited in the collection of the Tyumen State University Museum of Zoology, Tyumen, Russia.

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Remarks. The specimens of Papillacarus angulatus from South Africa (data above) corresponds morphologically with those from Ghana (see Wallwork 1962), but differs by the length of centrodorsal notogastral setae (similar in morphology and slightly longer than short neotrichal setae vs. thicker and distinctly longer in the Ghanaian specimens from the original description), longer subcapitular setae m1 and the presence of eight to nine pairs of short neotrichal setae (vs. 10 pairs). In our opinion, the above listed differences are intraspecific and may be explained as population variation. Hence, this possibly geographic variability should be considered in any future identification of P. angulatus.

FIGURE 15. Papillacarus angulatus Wallwork, 1962, adult: ventral view (legs except basal parts not shown). Scale bar 100 μm. 2017

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FIGURES 16–17. Papillacarus angulatus Wallwork, 1962, adult: 16—medioposterior part of prodorsum and anterior part of notogaster, lateral view; 17—medioposterior part of hysterosoma, lateral view. Scale bar 100 μm. TABLE 2. Leg setation and solenidia of adult Papillacarus angulatus Wallwork, 1962. Leg

Tr

Fe

Ge

Ti

Ta

I

-

d, (l), bv”, v”

d, (l), σ”, σ’

xt1, xt2, l’, v’, φ

(ft), it’, (tc), (p), (u), (a), s, m, n, (pv), ɛ, ω1, ω2

II

-

d, l’, la”, lp”, bv”, v”

d, (l), σ

xt1, xt2, l’, v’, φ

(ft), (tc), (p), (u), (a), s, (pv), ω1, ω2

III

l', v'

d, la’, lp’, ev’

d, l’, σ

d, l’, v’, φ

(ft), (tc), (p), (u), a’, s, (pv)

IV

l', v'

d, l’, ev’

d, l’, σ

d, l’, v’

(ft), (tc), p’, (u), a’, s, (pv)

Distribution and ecology of Lohmanniidae in South Africa The family Lohmanniidae is not well represented in South Africa. Previously only four living species have been recorded, each with a mostly localized distribution (see map, Fig. 22). Annectacarus eksteeni was recorded from coastal dune forests and natural beach vegetation of KwaZulu-Natal on the north-eastern coast of South Africa, Paulianacarus groblerae from coastal forests in KwaZulu678

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Natal, P. barlowi from coastal cultivated sugar cane fields in KwaZulu-Natal and Cryptacarus promecus from natural grasslands in central South Africa (Coetzee 2001a, b; Hugo-Coetzee & Avenant 2011). The natural grasslands of central South Africa are a much drier habitat than the forest vegetation of the north-eastern coast. Cryptacarus promecus has previously been recorded in dry environments (e.g. dry soil under olive trees (Pérez-Íñigo 1967)), humus under cactus pear (Bernini 1984)) and moist habitats (e.g. soils of moss cushions (Bayoumi & Al-Khalifa 1985), banana plantations (Mahunka 2000), along the shores of the Red Sea (Aoki 1971)), and therefore it seems that this species is adapted to survive in a wide variety of habitats.

FIGURES 18–21. Papillacarus angulatus Wallwork, 1962, adult: 18—leg I, without trochanter, right, antiaxial view; 19—femur and genu of leg II, right, antiaxial view; 20—femur, genu and tibia of leg III, left, antiaxial view; 21—leg IV, without trochanter, left, antiaxial view. Scale bars 50 μm. 2017

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Papillacarus angulatus described here was sampled from humid termitaries in central South Africa. The other record of P. angulatus in the collection of the National Museum, Bloemfontein is from soils in vineyards close to Upington, in the Northern Cape, which is situated next to the Orange River. It seems likely that P. angulatus prefers moist habitats since it was sampled previously only from moist habitats (soil litter in an Atlantic forest (Ermilov & Tolstikov 2015), from alfalfa fields in Iran (Lotfollahi & Haddad Irani-Nejad 2010), and vegetation of the tropical island of Annobon (Pérez-Íñigo 1969)). Like P. angulatus, Lohmannia lerallana sp. nov. was sampled from humid termite mounds in central South Africa.

FIGURES 22. Distribution of known Lohmanniidae species in South Africa (excluding fossils).

Acknowledgements We cordially thank three anonymous reviewers for the valuable comments.

References Aoki, J. (1971) A new species of mite, Thamnacarus moribei from the West coast of the Red Sea, with records of two other species of the family Lohmanniidae. The Japanese Journal of Zoology, 16(3), 127–129. Balogh, J. (1961) An outline of the family Lohmanniidae Berlese, 1916 (Acari: Oribatei). Acta Zoologica Academiae Scientiarum Hungaricae, 7(1–2), 19–44. Bayoumi, B.M. & Al-Khalifa, M.S. (1985) Oribatid Mites (Acari) of Saudi Arabia. Fauna of Saudi Arabia, 7, 66–92. Bernini, F. (1984) Notulae Oribatologicae XXXIII. New records of Cryptacarus promecus Grandjean and Cryptogalumna

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