Acari, Oribatida - BioOne

3 downloads 67 Views 527KB Size Report
(Acari, Oribatida) from Brazil. SERGEY G. ERMILOV & ANDREI V. TOLSTIKOV. Tyumen State University, Tyumen, Russia. E-mail: [email protected].
Systematic & Applied Acarology 20(6): 629–640 (2015) http://dx.doi.org/10.11158/saa.20.6.5 Article

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

http://zoobank.org/urn:lsid:zoobank.org:pub:A45D3EBB-D335-48FA-B90C-222BB4147102

New species and records of mites of the superfamily Oripodoidea (Acari, Oribatida) from Brazil SERGEY G. ERMILOV & ANDREI V. TOLSTIKOV Tyumen State University, Tyumen, Russia. E-mail: [email protected]

Abstract An annotated checklist of the Brazilian Oripodoidea is provided. It includes 14 species from eight genera and three families. Four species, Protoribates paracapucinus, Peloribates perreti, Scheloribates (Bischeloribates) elegantulus and Haplozetes minutus, are recorded for the first time in Brazil; of these, the last species is recorded for the first time in the Neotropical region. Two new species of the family Haplozetidae are described and illustrated on the basis of adult specimens. Protoribates lemensis sp. nov. differs from all representatives of the triaunguis-group by the presence of prolamellar lines and notogastral setae of medium size. Haplozetes biheterodactylus sp. nov. is most similar to H. minimicoma Beck, 1964, however, it differs from the latter by the long interlamellar setae and bidactylous legs. Key words: oribatid mites, Oripodoidea, new species, new record, Brazil

Introduction This work is a part of our study of Brazilian oribatid mite fauna (Acari, Oribatida) and includes data on the superfamily Oripodoidea. During taxonomic identification, we found two new oripodoid species of the family Haplozetidae; one belonging to the genus Protoribates Berlese, 1908, the other to the genus Haplozetes Willmann, 1935. This paper presents an annotated checklist Oripodoidea, new records of the recorded taxa, and descriptions of two new species of Haplozetidae. The genus Protoribates was proposed by Berlese (1908) with Oribata dentata Berlese, 1883 as type species. Currently it comprises 75 species, which have a cosmopolitan distribution. The main generic characters for the genus were summarized by Weigmann et al. (1993), Weigmann (2006), Bayartogtokh (2010) and Walter & Latonas (2013). Identification keys for some species of Protoribates were presented by Balogh & Balogh (2002), Weigmann (2006), Bayartogtokh (2010), Walter & Latonas (2013), and Corpuz-Raros (2014). The genus Haplozetes was proposed by Willmann (1935) with Peloribates vindobonensis Willmann, 1935 as type species. Generic classification is inconsistent (see, for example, Subías (2004, updated 2014) and Weigmann (2006)). We agree with Bayartogtokh’s (2000, 2010) and Weigmann’s (2006) diagnoses of this genus; hence, Haplozetes s. str. comprises about 60 species, which have a cosmopolitan distribution. Identification keys for some species of the genus were presented by Balogh & Balogh (2002), Weigmann (2006), and Bayartogtokh (2010).

Material and methods Our results are based on collections from three sites in Brazil (unknown date and collector; collection at the Tyumen State University, Tyumen, Russia): © Systematic & Applied Acarology Society

629

— Braz-2014–1: Brazil, 22°57'S, 43°12'W, Rio de Janeiro, Corcovado Mountain, Parque Nacional da Tijuca, 683 m.a.s.l., Atlantic forest, soil litter. — Braz-2014–2: Brazil, 23°33'S, 46°39'W, São Paulo, Parque Trianon, 852 m.a.s.l., Atlantic forest, soil litter. — Braz-2014–3: Brazil, 22°57'S, 43°09'W, Rio de Janeiro, Morro do Leme, Forte Duque de Caxias, 91 m a.s.l., Atlantic forest, soil litter. Specimens were mounted in lactic acid on temporary cavity slides for measurement and illustration. The 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. 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. General terminology used in this paper follows that of F. Grandjean (summarized by Norton & Behan-Pelletier 2009). Drawings were prepared with the aid of a drawing tube using a Carl Zeiss compound microscope “Axioskop-2 Plus”.

Checklist of oripodoid mites Specific localities for Oripodoidea, new records and overall known distribution1 are listed. Fourteen species (three not identified and two new to science), eight genera and three families were found. Four species are recorded for the first time in Brazil and of these one species is recorded for the first time in the Neotropical region (see checklist below). Haplozetidae — Protoribates bipilis (Pérez-Íñigo & Baggio, 1980). Locality: Braz-2014–1, Braz-2014–2, Braz2014–3. Distribution: Neotropical region and Iran. — Protoribates lemensis sp. nov. Locality: Braz-2014–3. Distribution: Brazil. — Protoribates paracapucinus (Mahunka, 1988). Locality: Braz-2014–2, Braz-2014–3. Distribution: Palaearctic, Oriental, Ethiopian and Neotropical regions. First record for Brazil. — Peloribates perreti Mahunka, 1984. Locality: Braz-2014–3. Distribution: Neotropical region. First record for Brazil. — Haplozetes biheterodactylus sp. nov. Locality: Braz-2014–3. Distribution: Brazil. — Haplozetes minutus (Tseng, 1984). Locality: Braz-2014–3. Distribution: Taiwan and Brazil. First record for the Neotropical region (it was noted in Shtanchaeva et al. 2014). — Trachyoribates (Rostrozetes) ovulum Berlese, 1908 sensu Beck (see 1965) as Rostrozetes foveolatus Sellnick, 1925. Locality: Braz-2014–3. Distribution: Cosmopolitan. Scheloribatidae — Muliercula femoroserrata (Pérez-Íñigo & Baggio, 1980). Locality: Braz-2014–1, Braz-2014–2, Braz-2014–3. Distribution: Brazil. — Muliercula sp. Locality: Braz-2014–1. Distribution: Brazil. — Scheloribates (Scheloribates) praeincisus acuticlava (Pérez-Íñigo & Baggio, 1986). Locality: Braz-2014–1, Braz-2014–2, Braz-2014–3. Distribution: Neotropical region.

1.

630

See mostly Subías (2004, updated 2014). SYSTEMATIC & APPLIED ACAROLOGY

VOL. 20

— Scheloribates (Bischeloribates) elegantulus (Hammer, 1961). Locality: Braz-2014–2. Distribution: Neotropical region. First record for Brazil. — Scheloribates (Hemileius) laticlava (Pérez-Íñigo & Baggio, 1991). Locality: Braz-2014–1. Distribution: Neotropical region. — Perscheloribates sp. Locality: Braz-2014–2. Distribution: Brazil. Oribatulidae — Eporibatula sp. Locality: Braz-2014–1. Distribution: Palaearctic region.

Descriptions of new species Protoribates lemensis sp. nov. (Figs 1–9) Diagnosis. Large species. Body size: 879–1128 × 614–747. Body surface microtuberculate. Prolamellar lines present. Rostral and lamellar setae setiform, barbed, shorter than interlamellar setae with short attenuate tip, slightly barbed. Bothridial setae spindle form to setiform, ciliate. Notogastral setae of medium size, setiform, with short attenuate tip, smooth. Four pairs of porose areas oval. Epimeral and ano-adanal setae setiform, barbed. Leg tarsi with three claws. Ventral side of femora II with clear ledge. Description. Measurements. Body length: 946 (holotype: female), 879–1128 (four paratypes: two females, two males); notogaster width: 680 (holotype), 614–747 (four paratypes). Integument. Body color brown to black brown. Body surface microtuberculate; tubercles small (up to 2), well visible especially on epimere I and posterior margin of ventral plate. Lateral parts of prodorsum between tutoria and acetabula I with reticulate pattern. Prodorsum. Rostrum rounded. Lamellae (lam) located dorso-laterally, slightly longer than half of prodorsum length (measured in lateral view). Prolamellar lines (plam) well developed. Sublamellar lines (slam) about half the length of lamellae, strong. Sublamellar porose areas (Al) oval (24–36 × 16–24). Tutoria (tu) distinct. Rostral (ro, 57–69) and lamellar (le, 90–110) setae setiform, barbed. Interlamellar setae (in, 213–229) setiform, with short attenuate tip, slightly barbed. Bothridial setae (ss, 184–196) spindle form to setiform, ciliate; stalk long, head short with distinct, thin, apex or absolutely absent. Exobothridial setae (ex, 57–65) thin, slightly barbed. Porose areas Ad not found. Notogaster. Anterior notogastral margin slightly convex anteriorly. Dorsophragmata (D) short, oval. Ten pairs of notogastral setae of medium size (102–123), setiform, with short attenuate tip, smooth. Four pairs of porose areas oval (Aa, 24–36 × 20–28; A1, A2, A3, 16–20 × 14–16). Setae lp inserted near and medially to A1. Lyrifissures (ia, im, ip, ih, ips) and opisthonotal gland openings (gla) distinct. Gnathosoma. Morphology of subcapitulum, palps and chelicerae generally typical for Protoribates (for example, Weigmann et al. 1993; Ermilov & Anichkin 2011; Ermilov et al. 2013). Subcapitulum longer than wide (209–233 × 147–155). Subcapitular setae (h, m, a) setiform, barbed, similar in length (41–49). Two pairs of adoral setae (or, 24–32) thickened, densely bilaterally barbed. Palps (length 123–135) with setation 0–2–1–3–9(+ω). Solenidion attached to eupathidium, both located on dorsal tubercle. Chelicerae (length 209–233) with two barbed setae; cha (61–69) longer than chb (36–41). Trägårdh’s organ long, tapered. Epimeral and lateral podosomal regions. Apodemes 1, 2, 3 and sejugal apodemes distinct. Epimeral setal formula: 3–1–3–2. All setae thin, slightly barbed; 1b, 3c (61–77) longer than 1c (28– 36) and 1a, 2a, 3a, 3b, 4a, 4b (41–49). Pedotecta I (Pd I) large, concave (in dorsal view) and scale2015

ERMILOV & TOLSTIKOV: NEW ORIPODOIDEA (ACARI, ORIBATIDA) FROM BRAZIL

631

like (in lateral view). Pedotecta II (Pd II) smaller, trapezoid, rounded anteriorly (in ventral view) and scale-like (in lateral view). Discidia (dis) triangular, pointed. Circumpedal carinae (cp) distinct, long. Anogenital region. Five pairs of genital setae (g1–g5, 36–45) setiform, barbed. One pair of aggenital (ag, 65–73), two pairs of anal (an1, an2, 69–77) and three pairs of adanal (ad1, ad2, 98– 102; ad3, 73–77) setae setiform, with short attenuate tip, barbed. Lyrifissures iad located close to and parallel anal plates. Legs. Tridactylous, lateral claw thinner than median claw, all with several minute barbs on dorsal side. Ventral side of femora II with well visible ledge, having wavy margin. Formulas of leg setation and solenidia: I (1–5–3–4–20) [1–2–2], II (1–5–3–4–16) [1–1–2], III (2–3–1–3–15) [1–1– 0], IV (1–2–2–3–12) [0–1–0]; homology of setae and solenidia indicated in Table 1. Famuli (ɛ) short, thin, straight, indistinctly dilated distally, inserted posteriorly to solenidion ω2. Solenidia ω1 on tarsus I, ω1 and ω2 on tarsus II and σ on genua III thickened, blunt-ended, other solenidia thinner, setiform.

FIGURE 1. Protoribates lemensis sp. nov., adult: dorsal view. Scale bar 200 μm.

632

SYSTEMATIC & APPLIED ACAROLOGY

VOL. 20

FIGURE 2. Protoribates lemensis sp. nov., adult: ventral view (gnathosoma and legs except trochanters IV not illustrated). Scale bar 200 μm. TABLE 1. Leg setation and solenidia of adult Protoribates lemensis sp. nov. Leg

Trochanter

Femur

Genu

Tibia

Tarsus

I

v'

d, (l), bv'', v''

(l), v', σ

(l), (v), φ1, φ2

(ft), (tc), (it), (p), (u), (a), s, (pv), v', (pl), l'', ɛ, ω1, ω2

II

v'

d, (l), bv'', v''

(l), v', σ

(l), (v), φ

III

l', v'

d, l', ev'

l', σ

l', (v), φ

(ft), (tc), (it), (p), (u), (a), s, (pv)

IV

v'

d, ev'

d, l'

l', (v), φ

ft'', (tc), (p), (u), (a), s, (pv)

(ft), (tc), (it), (p), (u), (a), s, (pv), l'', ω1,

ω2

Roman letters refer to normal setae (ɛ to famulus), Greek letters to solenidia. 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.

2015

ERMILOV & TOLSTIKOV: NEW ORIPODOIDEA (ACARI, ORIBATIDA) FROM BRAZIL

633

FIGURES 3–4. Protoribates lemensis sp. nov., adult: 3 — lateral view of prodorsum and anterior part of notogaster (gnathosoma and legs not illustrated); 4 — posterior view. Scale bar 200 μm.

Material examined. Holotype (female) and four paratypes (two females, two males): Braz2014–3. Type deposition. The holotype is deposited in the collection of the Senckenberg Institution Frankfurt, Germany; four paratypes are deposited in the collection of the Tyumen State University Museum of Zoology, Tyumen, Russia. 634

SYSTEMATIC & APPLIED ACAROLOGY

VOL. 20

FIGURES 5–9. Protoribates lemensis sp. nov., adult: 5 — subcapitulum, right half, ventral view; 6 — leg I, without trochanter, left, antiaxial view; 7 — tibia, genu and femur of leg II, left, antiaxial view; 8 — genu and femur of leg III, left, antiaxial view; 9 — leg IV, left, antiaxial view. Scale bar 100 μm.

Etymology. The species is named after Leme, a neighborhood in the South Zone of the City of Rio de Janeiro, Brazil, which, in turn, takes its name from a nearby rock formation whose shape resembles the rudder of a ship ("leme" in Portuguese language). Remarks. Protoribates lemensis sp. nov. can be included to triaunguis-group (species with two and three leg claws). It clearly differs from all representatives of this group by the presence of strong prolamellar lines (versus absent) and notogastral setae of medium size (versus minute or absent). The presence of prolamellar lines and well developed notogastral setae are unusual traits for Protoribates. Hence, these characters should be considered in any future diagnosis of this genus. 2015

ERMILOV & TOLSTIKOV: NEW ORIPODOIDEA (ACARI, ORIBATIDA) FROM BRAZIL

635

In having prolamellar lines, Protoribates lemensis sp. nov. is similar to the haplozetid genus Octodurozetes Mahunka, 1983; it is monotypic and was proposed with Octodurozetes berndhauseri Mahunka, 1983 from Switzerland as type species. However, the new species clearly differs from the latter by the notogastral setae of medium size (versus short), lamellar setae inserted on lamellae (versus on prodorsum) and rounded posterior part of notogaster (versus truncated).

Haplozetes biheterodactylus sp. nov. (Figs 10–20) Diagnosis. Small species. Body size: 274–282 × 172–176. Body surface smooth. Rostrum rounded. Prodorsal setae setiform, barbed; rostral and lamellar setae shorter than interlamellar setae. Bothridial setae spindle form, ciliate. Notogastral setae short, smooth. One seta (c) inserted near to pteromorphal hinge. Sacculi Sa with longitudinally elongated channel. Four pairs of genital setae present. Epimeral and ano-adanal setae short, smooth. Leg tarsi with two claws.

FIGURE 10. Haplozetes biheterodactylus sp. nov., adult: dorsal view view (left bothridial seta not illustrated). Scale bar 50 μm.

636

SYSTEMATIC & APPLIED ACAROLOGY

VOL. 20

Description. Measurements. Body length: 282 (holotype: male), 274 (one paratype: male); notogaster width: 172 (holotype), 176 (one paratype). Integument. Body color light brown. Body surface smooth. Trochanters and femora of legs slightly microfoveolate. Prodorsum. Rostrum rounded. Lamellae located dorso-laterally, slightly longer than half the length the prodorsum length (measured in lateral view). Prolamellar lines absent. Sublamellar lines short. Sublamellar porose areas rounded, very small (2). Tutoria strong, with distal tooth. Rostral (30–32), lamellar (45) and interlamellar (61–69) setae setiform, barbed. Bothridial setae (77–86) spindle form, ciliate, with long stalk, shorter head and distinct, thin, apex. Exobothridial setae (6) thin, smooth. Porose areas Ad not found. Notogaster. Anterior notogastral margin slightly convex. Dorsophragmata of medium size, slightly branched apically. Ten pairs of notogastral setae short (4–6), thin, smooth. One seta (c) inserted near to pteromorphal hinge. Four pairs of sacculi present: Sa with longitudinally elongated channel, with slightly dilated base, S1 and S3 with slightly elongated channel, S2 with well dilated base. All lyrifissures and opisthonotal gland openings distinct.

FIGURE 11. Haplozetes biheterodactylus sp. nov., adult: ventral view (legs except trochanters IV not illustrated). Scale bar 50 μm.

2015

ERMILOV & TOLSTIKOV: NEW ORIPODOIDEA (ACARI, ORIBATIDA) FROM BRAZIL

637

Gnathosoma. Morphology of subcapitulum, palps and chelicerae generally typical for Haplozetes (for example, Beck 1964; Bayartogtokh 2000; Shtanchaeva et al. 2014). Subcapitulum longer than wide (61–69 × 45–49). Subcapitular setae setiform, slightly barbed, h and m (12) longer than a (8). Two pairs of adoral setae short (6), barbed. Palps (length 41–45) with setation 0–2–1–3– 9(+ω). Solenidion attached to eupathidium, both located on dorsal tubercle. Chelicerae (length 65– 69) with two barbed setae; cha (20) longer than chb (12). Trägårdh’s organ long, tapered. Epimeral and lateral podosomal regions. Apodemes 1, 2, 3 and sejugal apodemes distinct. Epimeral setal formula: 3–1–3–3. All setae thin, smooth, similar in length (8–12). Pedotecta I concave (in dorsal view) and scale-like (in lateral view). Pedotecta II smaller, trapezoid, rounded anteriorly, with ventral tubercle (in ventral view) and scale-like (in lateral view). Discidia roundly triangular. Circumpedal carinae distinct, long.

FIGURES 14–20. Haplozetes biheterodactylus sp. nov., adult: 14 — bothridial seta; 15 — part of podosomal and epimeral regions; 16 — genital plate, right, ventral view; 17 — tarsus and tibia of leg I, left, antiaxial view; 18 — genu and femur of leg II, right, antiaxial view; 19 — genu, femur and trochanter of leg III, left, antiaxial view; 20 — leg IV, left, antiaxial view. Scale bar 20 μm.

638

SYSTEMATIC & APPLIED ACAROLOGY

VOL. 20

Anogenital region. Four pairs of genital setae (g1, 6; g2–g4, 4), one pair of aggenital (4–6), two pairs of anal (4–6) and three pairs of adanal (4–6) setae thin, smooth. Lyrifissures iad located close to and parallel anal plates. Postanal porose area (Ap) transverse, very thin furrow (28–41 × 4). Legs. Bidactylous, paraxial claw (on tarsi I, II) and antiaxial claw (in tarsi III, IV) thinner than the other, all claws smooth. Formulas of leg setation and solenidia: I (1–5–3–4–19) [1–2–2], II (1– 5–3–4–15) [1–1–2], III (2–3–1–3–15) [1–1–0], IV (1–2–2–3–12) [0–1–0]; homology of setae and solenidia indicated in Table 2. Famuli short, thin, straight, indistinctly dilated distally, inserted posteriorly to solenidion ω2. Solenidia ω1 on tarsus I, ω1 and ω2 on tarsus II and σ on genua III thickened, blunt-ended, other solenidia thinner, setiform.

TABLE 2. Leg setation and solenidia of adult Haplozetes biheterodactylus sp. nov. Leg

Trochanter

Femur

Genu

Tibia

Tarsus

I

v'

d, (l), bv'', v''

(l), v', σ

(l), (v), φ1, φ2

(ft), (tc), (it), (p), (u), (a), s, (pv), v', (pl), ɛ, ω1, ω2

II

v'

d, (l), bv'', v''

(l), v', σ

(l), (v), φ

III

l', v'

d, l', ev'

l', σ

l', (v), φ

(ft), (tc), (it), (p), (u), (a), s, (pv)

IV

v'

d, ev'

d, l'

l', (v), φ

ft'', (tc), (p), (u), (a), s, (pv)

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

ω2

See Table 1 for explanations.

Material examined. Holotype (male) and one paratype (male): Braz-2014–3. Type deposition. The holotype is deposited in the collection of the Senckenberg Institution Frankfurt, Germany; one paratype is deposited in the collection of the Tyumen State University Museum of Zoology, Tyumen, Russia. Etymology. The specific name biheterodactylus refers to the biheterodactylous legs. Remarks. In having small body size, rounded rostrum, spindle form, ciliate bothridial setae, insertion of only one seta (c) near to pteromorphal hinge and four pairs of genital setae, Haplozetes biheterodactylus sp. nov. is most similar to Haplozetes minimicoma Beck, 1964 from the Neotropical region and India. However, it clearly differs from the latter by the long interlamellar setae (versus very short) and bidactylous legs (versus monodactylous).

Acknowledgements The authors are thankful to two anonymous reviewers for valuable comments on the manuscript.

References Balogh, J. & Balogh, P. (2002) Identification keys to the oribatid mites of the Extra-Holarctic regions. Vol. 1. Miskolc, Well-Press Publishing Limited, 453 p. Bayartogtokh, B. (2000) Three species of Haplozetes (Acari: Oribatida: Haplozetidae) from Mongolia. International Journal of Acarology, 26 (1), 11–24. http://dx.doi.org/10.1080/01647950008683631 Bayartogtokh, B. (2010) Oribatid mites of Mongolia (Acari: Oribatida). Moscow, KMK, 372 p. Beck, L. (1964) Beiträge zur Kenntnis der neotropischen Oribatidenfauna 4. Haplozetes und Peloribates 2015

ERMILOV & TOLSTIKOV: NEW ORIPODOIDEA (ACARI, ORIBATIDA) FROM BRAZIL

639

(Arach., Acari). Senckenbergiana Biologica, 45 (2), 161–183. Beck, L. (1965) Über variabilität und wertigkeit morphologischer merkmale bei adulten Oribatiden (Arachnida, Acari) am beispiel der gattung Rostrozetes Sellnick, 1925. Abhandlungen der Senckenbergischen Naturforschenden Gesellschaft, 508, 1–64. Berlese, A. (1883) Acari, Myriapoda et Scorpiones hucusque in Italia reperta. Portici, Padova, 3–9. Berlese, A. (1908) Elenco di generi e specie nuovi di Acari. Redia, 5, 1–15. Corpuz-Raros, L.A. (2014) Additional species of soil mites of the subgenus Protoribates (Triaunguis) Kulijev (Acari: Oribatida: Protoribatidae), with key to Philippine species. Asia Life Sciences, 23 (1), 1–16. Ermilov, S.G. & Anichkin, A.E. (2011) A new species of Protoribates (Acari: Oribatida: Haplozetidae) from southern Vietnam. International Journal of Acarology, 37 (Supplement 1), 244–251. http://dx.doi.org/10.1080/01647954.2011.558852 Ermilov, S.G., Bayartogtokh, B., Sandmann, D., Marian, F. & Maraun, M. (2013) New and little known species of oribatid mites of the family Haplozetidae (Acari, Oribatida) from Ecuador. ZooKeys, 346, 43–57. http://dx.doi.org/10.3897/zookeys.346.6436 Mahunka, S. (1993) Beitrag zur Kenntnis der Höhlen-Oribatiden der Schweiz. Revue suisse de Zoologie, 100 (2), 225–233. Norton, R.A., Behan-Pelletier, V.M. (2009) Oribatida. Chapter 15. In: Krantz, G.W. & Walter, D.E. (Editors.), A Manual of Acarology. Texas Tech University Press, Lubbock, pp. 430–564. Shtanchaeva, U.Ya., Ermilov, S.G., Tolstikov, A.V. & Subías, L.S. (2014) Supplementary description of Indoribates (Haplozetes) minutus (Tseng, 1984) and Muliercula femoroserrata (Pérez-Iñigo et Baggio, 1980) (Acari, Oribatida, Oripodoidea). Acarina, 22 (2), 76–84. Subías, L.S. (2004) Listado sistemático, sinonímico y biogeográfico de los ácaros oribátidos (Acariformes: Oribatida) del mundo (excepto fósiles). Graellsia, 60 (número extraordinario), 3–305. Online version accessed in February 2014, 577 pp.; http://www.ucm.es/info/zoo/Artropodos/Catalogo.pdf Walter, D. E. & Latonas, S. (2013) A review of the ecology and distribution of Protoribates (Oribatida, Oripodoidea, Haplozetidae) of Alberta, Canada, with the description of a new species. Zootaxa, 3620 (3), 483– 499. http://dx.doi.org/10.11646/zootaxa.3620.3.9 Weigmann, G. (2006) Hornmilben (Oribatida). Die Tierwelt Deutschlands. Teil 76. Goecke & Evers, Keltern, 520 p. Weigmann, G., Miko, L. & Nannelli, R. (1993) Redescription of Protoribates dentatus (Berlese, 1883) with remarks about the genus Protoribates (Acarina, Oribatida). Redia, 76 (1), 39–55. Willmann, C. (1935) IV. Die Milbenfauna – Oribatei. In: Jaus I.: Faunistisch-ökologische Studien im Anningergebiet, mit besonderer Berücksichtigung der xerothermen Formen. Zoologische Jahrbücher (Systematik), 66 (5), 331–344. Submitted: 30 Nov. 2014; accepted by Q.-H. Fan: 13 Feb. 2015; published: 30 Sept. 2015

640

SYSTEMATIC & APPLIED ACAROLOGY

VOL. 20