Ermilov, Sergey G. ¹

¹✉ Tyumen State University, Institute of Environmental and Agricultural Biology (X-BIO), Tyumen, Russia.

2024 - Volume: 64 Issue: 3 pages: 976-988

ZooBank LSID: 10DF1694-46A4-4C91-ADFE-386F8FB9263D

Original research

Keywords

Abstract

Introduction

Among the oribatid mite (Acari, Oribatida) materials collected from the Merida State, Western Venezuela, I found two new species of the family Mochlozetidae, one species belonging to the genus Dynatozetes Grandjean, 1960, the other to Uracrobates Balogh and Mahunka, 1967. The main goal of the paper is to describe the two new species.

Dynatozetes was proposed by Grandjean (1960), with Dynatozetes amplus Grandjean, 1960 as type species. The genus comprises four species distributed in the Nearctic and Neotropical regions (Subías 2022, online version 2024). The generic diagnosis and an identification key to the known species were provided by Ermilov and Friedrich (2017a).

Uracrobates was proposed by Balogh and Mahunka (1967), with Uracrobates magniporosus Balogh and Mahunka, 1967 as type species. The genus comprises two subgenera (nominate and Parauracrobates Ermilov and Martens, 2015) and eight species distributed in tropics (Subías 2022, online version 2024). The generic and subgeneric diagnoses as well as an identification key to the known species were provided by Ermilov and Martens (2015) and revised by Ermilov and Mary (2019).

Additional goal of the paper is to update identification keys and to summarize data on distribution and habitat to representatives of Dynatozetes and Uracrobates.

Prior to this study, two identified species of Dynatozetes have been registered in the Venezuelan fauna, D. amplus Grandjean, 1960 (see Călugăr 1990; Behan-Pelletier et al. 1993) and D. obesus Grandjean, 1960 (see Grandjean 1960; Ermilov 2016). Uracrobates has not yet been registered in Venezuela.

Methods

Observation and documentation — For measurement and illustration, specimens were mounted in lactic acid on temporary cavity slides. All body measurements are presented in micrometers (µm); body length was measured in lateral view, from the tip of the rostrum to the posterior edge of the notogaster; body width refers to the maximum width at pteromorph level in dorsal aspect and to the maximum width at ventral plate level in ventral view; the lengths of body setae were measured in lateral aspect. 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. Drawings were made with a camera lucida using a Leica DM 2500 light microscope.

Terminology — Morphological terminology used in this paper mostly follows that of papers on Mochlozetidae (e.g., Grandjean 1959, 1960; Norton 1983, 1984; Ermilov and Yurtaev 2023); Norton (1977) for leg setal nomenclature and Norton and Behan-Pelletier (2009) for overview.

Abbreviations — Prodorsum: lam = lamella; tlam = translamella; plam = prolamella; slam = sublamella; Al = sublamellar porose area; tu = tutorium; ro, le, in, bs, ex = rostral, lamellar, interlamellar, bothridial, and exobothridial setae, respectively; Ad = dorsosejugal porose area; D = dorsophragma; P = pleurophragma. Notogaster: c, la, lm, lp, h, p = notogastral setae; Aa, A1, A2, A3, A₁–A₄ = porose areas; im, ip, ih, ips = lyrifissures; gla = opisthonotal gland opening. Gnathosoma: a, m, h = subcapitular setae; or = adoral seta; d, l, cm, acm, ul, su, lt, vt, inf, sup = palp setae; ω = palp solenidion; ep = postpalpal seta; cha, chb = cheliceral setae; Tg = Trägårdh's organ. Epimeral and lateral podosomal regions: 1a, 1b, 1c, 2a, 3a, 3b, 3c, 4a, 4b, 4c = epimeral setae; Am, Ah = humeral porose areas; PdI, PdII = pedotecta I and II, respectively; cus = custodium; dis = discidium; cir = circumpedal carina; z = aperture of supracoxal gland. Anogenital region: g, ag, an, ad = genital, aggenital, anal, and adanal setae, respectively; iad = adanal lyrifissure; Amar = marginal porose area; po = preanal organ. Legs: Tr, Fe, Ge, Ti, Ta = trochanter, femur, genu, tibia, and tarsus, respectively; trp = trochanter process; pa = porose area; ω, σ, φ = solenidia; ɛ = famulus; d, l, v, ev, bv, ft, tc, it, p, u, a, s, pv, pl = setae.

Taxonomy

Family Mochlozetidae Grandjean, 1960

Genus Dynatozetes Grandjean, 1960

Type species: Dynatozetes amplus Grandjean, 1960

Dynatozetes asetosus n. sp.

ZOOBANK: 14A566E5-E4C6-4447-A2C0-22B6200C39E9

(Figures 1, 2)

Material examined — Holotype (female) and two paratypes (two females): Western Venezuela, Merida State, 2300 m a.s.l., 10 km E Tabay, sweeping in cloud forest, 28.IV.1981 (L. Masner and Marsh).

Type deposition — The holotype is deposited in the collection of the Canadian National Collection, Ottawa, Canada; two paratypes are in the collection of the Tyumen State University Museum of Zoology, Tyumen, Russia. All specimens are preserved in 70% solution of ethanol with a drop of glycerol.

Diagnosis — Body length: 855–930. Rostrum with small, semi-oval protruding. Prolamella present. Rostral, lamellar and bothridial setae long, setiform, barbed; ro and le longer than bs; interlamellar seta represented by alveolus. Notogaster and anogenital region foveolate. Six pairs of notogastral porose areas, of these, four pairs in posteroperipheral position; Aa oval, others rounded. Epimeral, genital and aggenital setae setiform, thin, slightly barbed; anal and adanal setae short, setiform, thin, roughened. Leg trochanter IV with anterodorsal process. Dorsal porose area on tarsi I, II not observable.

Description of adult — Measurements – Body length: 930 (holotype), 855, 915 (female paratypes); notogaster width (level of pteromorphs): 840 (holotype), 825 (female paratypes); ventral plate width: 750 (holotype), 705 (female paratypes).

Integument – Body color brown. Surface with dense microgranules (visible under high magnification, 10 × 100); notogaster and anogenital region with small poorly observable foveolae; lateral side of body partially with dense microgranulate cerotegument.

Prodorsum (Figs 1a, 1c, 2a) – Rostrum with small, semi-oval protruding (visible in anterior view). Lamella (including cusp) about 2/3 length of prodorsum, with short cusp having strong lateral tooth; translamella absent; prolamella lineate, located medial to insertions of rostral and lamellar setae; tutorium ridge-like, with small tooth distally. Sublamellar porose area oval (26–30 × 19–22). Rostral (112–131), lamellar (112–131) and bothridial (101–106) setae setiform, barbed; interlamellar seta represented by alveolus; exobothridial seta (45–49) setiform, thin, slightly barbed. Dorsosejugal porose area diffuse, located posterolateral to interlamellar seta.

Notogaster (Figs 1a, 1c, 1d) – Dorsosejugal suture interrupted medially. Pteromorph broadly rounded laterally. Six pairs of porose areas, with four pairs in posteroperipheral position; Aa oval (41–52 × 19–26), others (15–26) rounded; A1 anteromedial to A₁. All notogastral setae represented by alveoli. Opisthonotal gland opening and all lyrifissures (except ia) distinct.

Gnathosoma (Figs 2b–2d) – Subcapitulum size: 195–202 × 172–184; subcapitular seta a (22–26) setiform, brushed distally; subcapitular setae m (75–86) and h (52–60) and both adoral setae (26) setiform, barbed. Palp length: 157–165; setation: 0–2–1–3–9(+ω); postpalpal seta (7–9) spiniform, smooth. Chelicera length: 206–225; setae (cha: 71–75; chb: 56–60) setiform, barbed.

Epimeral and lateral podosomal regions (Figs 1b, 1c) – Epimeral setal formula: 3–1–3–3; all setae (1a, 1c, 2a, 3a: 45–49; others: 71–90) setiform, thin, slightly barbed. Humeral porose areas Am and Ah elongate oval, poorly observable. Custodium long, thorn-like. Discidium triangular. Circumpedal carina long, distally fused to custodium.

Anogenital region (Figs 1b–1d) – Anogenital setal formula: 6–1–2–3; genital and aggenital setae (45–49) setiform, thin, slightly barbed; anal and adanal setae (19) setiform, thin, roughened. Adanal lyrifissure located close and slightly oblique to anterior half of anal plate. Marginal porose area represented by numerous oval and elongate oval parts.

Legs (Figs 2e–2h) – Median claw distinctly thicker than lateral ones, all slightly barbed dorsally; each lateral claw with small tooth ventrodistally. Trochanter IV with strong anterodorsal process. Dorsoparaxial porose area on femora I–IV, trochanters III, IV, ventrodistal porose area on tibia I–IV and proximoventral porose area on tarsi I–IV well visible; dorsal porose area on tarsi I, II not observable. Formulas of leg setation and solenidia: I (1–5–3–4–20) [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 1.

Remarks — The new species differs from all species of the genus in the presence of the prolamella (versus prolamella absent) and the setiform bothridial seta (versus bothridial seta with well-developed head). Distinctive characters between species within of the genus Dynatozetes can be found in the identification key below.

Etymology — The species name asetosus refers to the absence of interlamellar setae.

Key to the known species of Dynatozetes

1. Bothridial seta setiform; body length: 855–930
...... D. asetosus n. sp.

— Bothridial seta with well-developed head
...... 2

2. Interlamellar seta well-developed; notogastral porose area Aa distinctly elongate oval; body length: 763–863
...... D. hexaporosus Ermilov, 2017 (in Ermilov and Friedrich 2017a)

— Interlamellar seta represented by alveolus; notogastral porose area Aa rounded/oval
...... 3

3. Dorsal porose area on leg tarsi I and II absent; lamellar seta as long as distance le–le; body length: 934–964
...... D. magnus (Banks, 1895) (see also Norton 1984)

— Dorsal porose area on leg tarsi I and II present; lamellar seta shorter than distance le–le
...... 4

4. Five or six pairs of posteroperipheral notogastral porose areas present; leg trochanter III with one seta (v′ absent); body length: 770–870
...... D. amplus Grandjean, 1960

— Four pairs of posteroperipheral notogastral porose areas present; leg trochanter III with two setae (v′ present); body length: 800–930
...... D. obesus Grandjean, 1960

Genus Uracrobates Balogh and Mahunka, 1967

Subgenus Uracrobates (Uracrobates) Balogh and Mahunka, 1967

Type species: Uracrobates magniporosus Balogh and Mahunka, 1967

Uracrobates (Uracrobates) paramasneri n. sp.

ZOOBANK: 24D88AE8-C0F5-4B16-A5DC-210E345D57F8

(Figures 3, 4)

Material examined — Holotype (female) and four paratypes (two females): Western Venezuela, Merida State, 2300 m a.s.l., 10 km E Tabay, sweeping in cloud forest, 28.IV.1981 (L. Masner and Marsh).

Type deposition — The holotype is deposited in the collection of the Canadian National Collection, Ottawa, Canada; four paratypes are in the collection of the Tyumen State University Museum of Zoology, Tyumen, Russia. All specimens are preserved in 70% solution of ethanol with a drop of glycerol.

Diagnosis — Body length: 930–1020. Body without foveolae. Rostrum pointed. Lamellar cusp with strong lateral tooth. Rostral, lamellar and interlamellar setae long, setiform, barbed; ro shortest, in longest; bothridial seta medium-sized, clavate, barbed. Notogastral porose areas Aa, A2 and A3 strongly elongate oval (nearly band-like), A1 rounded. Epimeral, genital and aggenital setae setiform, thin, slightly barbed; anal and adanal setae short, setiform, thin, roughened. Leg dorsal porose area on tarsi I, II present.

Description of adult — Measurements – Body length: 930 (holotype), 945–1020 (female paratypes); notogaster width (level of pteromorphs): 720 (holotype), 720–765 (female paratypes); ventral plate width: 615 (holotype), 630–705 (female paratypes).

Integument – Body color brown. Surface with dense microgranules (visible under high magnification, 10 × 100); pedotectum I partially striate; lateral side of body partially with dense microgranulate cerotegument.

Prodorsum (Figs 3a, 3c, 4a) – Rostrum pointed. Lamella (including cusp) about 1/2 length of prodorsum, with long cusp having strong lateral tooth; translamella well-developed, indistinctly convex medially; tutorium ridge-like, with small tooth distally. Sublamellar porose area nearly rounded (15–17). Rostral (131–139), lamellar (180–187) and interlamellar (236–240) setae setiform, barbed; bothridial seta (79–82) clavate, barbed, stalk and head similar in length or stalk slightly longer; exobothridial seta (60–67) setiform, thin, slightly barbed. Dorsosejugal porose area diffuse, located posterolateral to interlamellar seta.

Notogaster (Figs 3a, 3c, 3d) – Dorsosejugal suture interrupted medially. Posterior margin distinctly narrowed in dorsal and ventral aspects. Pteromorph broadly rounded laterally. Four pairs of porose areas; A1 rounded (26–37), others strongly elongate oval, nearly band-like (Aa: 161–176 × 7–19; A2, A3: 90–146 × 7–19). All notogastral setae represented by alveoli. Opisthonotal gland opening and all lyrifissures (except ia) distinct.

Gnathosoma (Figs 4b–4d) – Subcapitulum size: 206–229 × 187–206; subcapitular setae (a: 37–41; m: 64–71; h: 56–60) and adoral (19) setae setiform, barbed. Palp length: 169–180; setation: 0–2–1–3–9(+ω); postpalpal seta (7) spiniform, smooth. Chelicera length: 225–236; setae (cha: 71–75; chb: 56–60) setiform, barbed.

Epimeral and lateral podosomal regions (Figs 3b, 3c) – Epimeral setal formula: 3–1–3–3; all setae (1a, 2a, 3a, 4c: 34–37; 1c, 4a, 4b: 45–49; 1b, 3b, 3c: 67–71) setiform, thin, slightly barbed. Humeral porose areas Am and Ah elongate oval, poorly observable. Custodium long, thorn-like. Discidium triangular. Circumpedal carina long, distally fused to custodium.

Anogenital region (Figs 3b–3d) – Anogenital setal formula: 6–1–2–3; genital and aggenital setae (34–37) setiform, thin, slightly barbed; anal and adanal setae (19) setiform, thin, roughened. Adanal lyrifissure located close and slightly oblique to anterior half of anal plate. Marginal porose area complete, band-like.

Legs (Figs 4e–4h) – Median claw distinctly thicker than lateral ones, all slightly barbed dorsally; each lateral claw with small tooth ventrodistally. Dorsoparaxial porose area on femora I–IV, trochanters III, IV, ventrodistal porose area on tibia I–IV, proximoventral porose area on tarsi I–IV, and dorsal porose area on tarsi I, II well visible. Formulas of leg setation and solenidia: I (1–5–3–4–20) [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 1.

Remarks — The new species differs from all species of the genus in the presence of the strongly elongate oval (nearly band-like) notogastral porose area Aa (versus Aa rounded/oval). Distinctive characters between species within of the genus Uracrobates can be found in the identification key below.

Etymology — The species name paramasneri refers to the similarity of the new species and U. (U.) masneri Ermilov, 2019.

Key to the known species of Uracrobates

1. Notogaster truncated posteriorly (trapezoid form); notogastral porose area A3 located in corners of the posterior notogastral margin
...... 2. Subgenus Uracrobates (Parauracrobates) Ermilov and Martens, 2015

— Notogaster narrowed posteriorly (triangular form); notogastral porose area A3 located dorsally on the notogaster
...... 3. Subgenus Uracrobates (Uracrobates) Balogh and Mahunka, 1967

2. Lamellar cusp truncated; notogaster and anogenital region foveolate; anal and adanal setae medium-sized; body length: 664–680
...... U. (P.) newtaipeiensis Ermilov and Liao, 2017

— Lamellar cusp with strong lateral tooth, notogaster and anogenital region not foveolate; anal and adanal setae short; body length: 680–780
...... U. (P.) truncatus Ermilov and Martens, 2015

3. Rostrum pointed; notogastral porose areas A2 and A3 strongly elongate oval
...... 4

— Rostrum rounded or bidentate; notogastral porose areas A2 and A3 rounded
...... 5

4. Notogastral porose area Aa strongly elongate oval; body length: 930–1020
...... U. (U.) paramasneri n. sp.

— Notogastral porose area Aa rounded; body length: 730–879
...... U. (U.) masneri Ermilov, 2019

5. Rostrum rounded; lamellar cusp with minute lateral and medial teeth; body length: 730–796
...... U. (U.) mayottensis Ermilov and Mary, 2019

— Rostrum bidentate; lamellar cusp with strong lateral tooth, medial tooth absent
...... 6

6. Adanal setae ad₁ and ad₂ medium-sized, longer than short ad₃; adanal lyrifissure inverse apoanal; body length: 620
...... U. (U.) pygiseta (Hammer, 1973)

— All adanal setae short, similar in length; adanal lyrifissure paraanal
...... 7

7. Notogaster smooth; body length: 480–500
...... U. (U.) africanus Mahunka, 1988 (see also Mahunka 2001)

— Notogaster foveolate
...... 8

8. Posterior triangular part of notogaster extended; body length: 676–687
...... U. (U.) magniporosus Balogh and Mahunka, 1967 (see also Mahunka 1988)

— Posterior triangular part of notogaster not extended; body length: 548–675
...... U. (U.) indicus Ramani and Haq, 1990

General remarks

Ermilov and Friedrich (2017a) proposed a detailed generic diagnosis of the genus Dynatozetes. Their data were based only on morphology of four species. In light of the description of the new (fifth) species, some generic traits need to be adjusted.

The generic diagnosis indicated, e.g.: integument without ornamentation; prolamella absent; bothridial seta clavate to lanceolate (Ermilov and Friedrich 2017a). However, specimens of the new species have: foveolate notogaster and anogenital region; lineate prolamella; setiform bothridial seta. Hence, the generic diagnosis should include these morphological additions.

Distribution and habitat of Dynatozetes and Uracrobates

Species of Dynatozetes are known from the Nearctic and Neotropical region; species of Uracrobates are known from the Oriental, Afrotropical, Neotropical, and Australasian regions. According to the summarized data (Tables 2, 3), they inhabit mainly soil/litter in the tropical forests but some species are probably arboricolous.

Acknowledgements

I cordially thank Dr. Julia Baumann and two anonymous reviewers for valuable comments, and Dr. W. Knee for facilitating the loan of Venezuelan Acari material from the Canadian National Collection of Insects, Arachnids and Nematodes, Ottawa, Canada.

References

1. Balogh J., Mahunka S. 1967. New oribatids (Acari) from Vietnam. Acta Zool. Acad. Sci. Hung., 13: 39-74.
2. Banks N. 1895. On the Oribatoidea of the United States. American Ent. Soc., 22: 1-16. https://www.jstor.org/stable/25076633
3. Behan-Pelletier V.M., Paoletti M.G., Bissett B., Stinner B.R. 1993. Oribatid mites of forest habitats in northern Venezuela. Trop. Zool., Spec. Issue 1: 39-54.
4. Călugăr M. 1990. Oribátidos nuevos de Venezuela (Acari, Oribatida). Bol. Soc. Venez. Espeleolog., 24: 15-20.
5. Chu Y.I., Aoki J. 1997. Fauna of oribatid mite at Fu-shan forest litter and humus layer. Zhong. Kunch., 17: 172-178.
6. Corpuz-Raros L., Ermilov S.G. 2020. Catalogue of oribatid mites (Acari: Oribatida) from Continental Southeast Asia. Zootaxa, 4893: 1-216. https://doi.org/10.11646/zootaxa.4893.1.1
7. Ermilov S.G. 2016. Additions to the oribatid mite fauna of Venezuela, with description of two new species of the family Achipteriidae (Acari, Oribatida). Syst. Appl. Acarol., 21: 1591-1603. https://doi.org/10.11158/saa.21.12.1
8. Ermilov S.G. 2017. Contribution to the knowledge of oribatid mites of Colombia, with description of two new species of Galumna (Acari, Oribatida, Galumnidae). Syst. Appl. Acarol., 22: 102-114. https://doi.org/10.11158/saa.22.1.11
9. Ermilov S.G. 2019. New findings of oribatid mites of the family Mochlozetidae (Acari: Oribatida) from Ecuador. Biologia, 74: 1313-1324. https://doi.org/10.2478/s11756-019-00248-5
10. Ermilov S.G. 2024. Additions to the oribatid mite (Acari, Oribatida) fauna of Guatemala, with description of a new species of Pergalumna. Persian Journal of Acarology. In Press. https://doi.org/10.1080/01647954.2024.2318366
11. Ermilov S.G., Friedrich S. 2017a. Contribution to knowledge of the oribatid mite genus Dynatozetes (Acari, Oribatida, Mochlozetidae). Spixiana, 40: 23-28. https://doi.org/10.11158/saa.23.4.6
12. Ermilov S.G., Friedrich S. 2017b. New faunistic and taxonomic data on oribatid mites (Acari, Oribatida) of Peru. Acarina, 25: 3-13. https://doi.org/10.21684/0132-8077-2017-25-1-3-13
13. Ermilov S.G., Liao J.-R. 2017. New faunistic and taxonomic data on oribatid mites (Acari, Oribatida) of Taiwan. Syst. Appl. Acarol., 22: 824-840. https://doi.org/10.11158/saa.22.6.8
14. Ermilov S.G., Martens J. 2015. The oribatid mite genus Uracrobates (Acari, Oribatida, Mochlozetidae). Syst. Appl. Acarol., 20: 188-194. https://doi.org/10.11158/saa.20.2.5
15. Ermilov S.G., Mary N. 2019. To the knowledge of oribatid mites (Acari, Oribatida) of the Comoro lslands. Syst. Appl. Acarol., 24: 624-637. https://doi.org/10.11158/saa.24.4.8
16. Ermilov S.G., Smit H. 2017. Additions to the oribatid mite fauna of the Caribbean, with a description of a new species of Epidamaeus (Acari, Oribatida, Damaeidae). Acarologia, 57: 791-804. https://doi.org/10.24349/acarologia/20174194
17. Ermilov S.G., Yurtaev A.A. 2023. Unguizetes (Knorozovia) ershovae (Acari, Oribatida, Mochlozetidae), a new oribatid mite subgenus and species from Mexico. Acarologia, 63: 1187-1196. https://doi.org/10.24349/r0vq-kaq5
18. Grandjean F. 1959. Sur le genre Mochlozetes Grandj. 1930 (Oribate). Acarologia, 1: 452-474.
19. Grandjean F. 1960. Les Mochlozetidae n. fam. (Oribates). Acarologia, 2: 101-148.
20. Hammer M. 1973. Oribatids from Tongatapu and Eua, the Tonga Islands, and from Upolu, Western Samoa. Det Kong. Dansk. Vidensk. Selsk. Biol. Skr., 20: 1-70.
21. Mahunka S. 1983. Neue und interessante Milben aus dem Genfer Museum XLVII. Oribatida Americana 7: Guatemala II. Rev. suisse Zool., 90: 709-724. https://doi.org/10.5962/bhl.part.82009
22. Mahunka S. 1988. The oribatid fauna of Tanzania. Ann. Hist. Nat. Mus. Nat. Hung., 80: 189-213.
23. Mahunka S. 2001. Arboricolous oribatid mites (Acari: Oribatida) from Kenya. Folia Ent. Hung., 62: 11-22.
24. Mahunka S. 2006. Some interesting beetle mites from Pacific Islands collected by Antonius van Harten (Acari: Oribatida). (Acarologica Genavensia CVIII). Rev. suisse Zool., 113: 579-593. https://doi.org/10.5962/bhl.part.80364
25. Norton R.A. 1977. A review of F. Grandjean's system of leg chaetotaxy in the Oribatei (Acari) and its application to the family Damaeidae. In: Dindal D.L. (Ed.). Biology of oribatid mites. Syracuse: SUNY Col. Environ. Sci. Forestry. pp. 33-61.
26. Norton R.A. 1983. Redefinition of Mochloribatula (Acari: Mochlozetidae), with new species, recombinations, and notes on plant associations. Acarologia, 24: 449-464.
27. Norton R.A. 1984. Notes on Nathan Banks' and Henry Ewings' species of Mochlozetidae (Acari: Sarcoptiformes) with the proposal of a new genus. Acarologia, 25: 397-406.
28. Norton R.A., Behan-Pelletier V.M. 2009. Oribatida. In: Krantz G.W., Walter D.E. (Eds). A Manual of Acarology (TX). Lubbock: Texas Univ. Press. Chapter 15. pp. 430-564.
29. Oliveira A.R., Norton R.A., Moraes, G.J. de. 2005. Edaphic and plant inhabiting oribatid mites (Acari: Oribatida) from Cerrado and Mata Atlântica ecosystems in the State of São Paulo, southeast Brazil. Zootaxa, 1049: 49-68. https://doi.org/10.11646/zootaxa.1049.1.4
30. Pérez-Íñigo C., Baggio D. 1994. Oribates édaphiques du Brésil (VIII). Oribates de l′État de São Paulo (cinquieme partie). Acarologia, 35: 181-198.
31. Ramani N., Haq M.A. 1988. Developmental studies of Uracrobates indicus (Acari: Oribatei) inhabiting Mangifera indica. In: Channabasavanna G.P. and Viraktamath C.A. (eds.). Progress in Acarology. Brill, Leiden, vol. 1, p. 483-490.
32. Ramani N., Haq M.A. 1990. Oribatid mites from coconut palm 1. A new species of Uracrobates (Acari: Oribatei) from Kerala (India). Acarologia, 31: 297-302.
33. Saporito R.A., Donnelly M.A., Norton R.A., Garraffo H.M., Spande T.F., Daly J.W. 2007. Oribatid mites as a major dietary source for alkaloids in poison frogs. Proc. Nat. Acad. Sci. USA, 104: 8885-8890. https://doi.org/10.1073/pnas.0702851104
34. Schatz H. 2006. Catalogue of known oribatid mite species (Acari Oribatida) from the Central American landbridge (First part). Trop. Zool., 19: 209-288.
35. Sheela K., Haq M.A. 1991. Oribatid mites as bioindicators of soil moisture. In: Veeresh G.K., Rajagopal D. and Viraktamath C.A. (eds.). Advances in management and conservation of soil fauna. Oxford and IBH Publishing Co. Pvt. Ltd., New Delhi, p. 871-876.
36. Subías L.S. 2022. Listado sistemático, sinonímico y biogeográfico de los ácaros oribátidos (Acariformes: Oribatida) del mundo (excepto fósiles). Monogr. Electr. S.E.A., 12: 1-538.
37. Subías L.S. 2024. Listado sistemático, sinonímico y biogeográfico de los ácaros oribátidos (Acariformes: Oribatida) del mundo (excepto fósiles), 19ª actualización: 1-545. Available from: http://bba.bioucm.es/cont/docs/RO_1.pdf (accessed January 2024).
38. Subías L.S., Zaballos J.P., Banda E., Fontal-Cazalla F., Nieves-Aldrey J.L. 2004. Ácaros oribátidos (Acari, Oribatei) del Parque Nacional de la isla de Coiba, Panamá. Rev. Biol. Trop., 52: 85-95. https://doi.org/10.15517/rbt.v52i1.14755
39. Vázquez M. 1999. Catálogo de los ácaros oribátidos edáficos de Sian Ka′an, Q. ROO, México. 1st Edition. Universidad de Quintana Roo, Chetumal, Quintana Roo, 126 pp.
40. Vázquez M., May D., Alamilla E. 2016. Riqueza específica y biodiversidad de Cozumel, Quintana Roo, México. Teoría y Praxis, 12: 137-171. https://doi.org/10.22403/UQROOMX/TYP19/07

instructions