1✉ University of Tyumen, Institute of Environmental and Agricultural Biology (X-BIO), Tyumen, Russia.
2025 - Volume: 65 Issue: 2 pages: 405-416
https://doi.org/10.24349/8znh-f82iThe oribatid mites (Acari, Oribatida) of the Amhara region in Central Ethiopia have been insufficiently studied (Bayartogtokh et al. 2021; Niedbała and Ermilov 2022; Ermilov and Rybalov 2024a). During taxonomic identification of new materials from this region, two new species of the superfamily Oripodoidea belonging to the genera Zygoribatula Berlese, 1916 (Oribatulidae) and Scheloribates Berlese, 1908 (Scheloribatidae) are found. The main objective of the paper is to describe new species under the names Zygoribatula birhanensis n. sp. and Scheloribates (Scheloribates) debreensis n. sp.
The genus Zygoribatula was proposed by Berlese (1916b) as a subgenus of the genus Oribatula Berlese 1896, with Oribatula connexa Berlese, 1904 as type species. I support the generic independence of this taxon, following Weigmann (2006), Bayartogtokh and Smelyansky (2008), Bayartogtokh (2010). The genus comprises about 100 species, which have a cosmopolitan distribution collectively (Subías 2022, online version 2024). The main generic characters can be found in: Bulanova-Zachvatkina (1975), Lee (1992), Grobler and Kok (1993), Pérez-Íñigo (1993), Weigmann (2006), Subías and Shtanchaeva (2023). The identification keys to selective species of Zygoribatula were given by: Bulanova-Zachvatkina (1975), Lee (1992), Pérez-Íñigo (1993), Balogh and Balogh (2002), Weigmann (2006), Franklin et al. (2008), Bayartogtokh (2010), Ermilov (2023).
The genus Scheloribates was proposed by Berlese (1908), with Zetes latipes Koch, 1844 as type species. The nominate subgenus comprises more than 210 species, which have a cosmopolitan distribution collectively (Subías 2022, online version 2024). The main subgeneric characters can be found in: Pletzen van (1963), Coetzer (1968), Corpuz-Raros (1980), Balogh and Balogh (1992), Weigmann (2006), Bayartogtokh (2010), Ermilov and Anichkin (2014). The identification keys to selective species of Scheloribates (Scheloribates) were given by: Shaldybina (1975), Balogh and Balogh (2002), Weigmann (2006), Bayartogtokh (2010), Ermilov et al. (2011), Ermilov and Starý (2017).
Prior to this study, four species of Zygoribatula and 22 species of S. (Scheloribates) have been recorded in the Ethiopian fauna (Ermilov and Rybalov 2013a, b, 2022, 2024b, c; Ermilov et al. 2024).
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 of the notogaster in dorsal view in Zygoribatula, and to the maximum width at pteromorph level in dorsal aspect and to the maximum width at ventral plate level in ventral view in Scheloribates; 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.
Morphological terminology used in this paper mostly follows that of papers on Zygoribatula and Scheloribates (e.g. Ermilov 2023; Ermilov et al. 2024); also, Norton (1977) for leg setal nomenclature and Norton and Behan-Pelletier (2009) for overview are used.
Prodorsum: lam = lamella; plam = prolamella; tlam = translamella; slam = sublamella; Al = sublamellar porose area; tu = tutorium; kf = keel-shaped ridge; ro, le, in, bs, ex = rostral, lamellar, interlamellar, bothridial, and exobothridial setae, respectively; Ad = dorsosejugal porose area; D = dorsophragma; P = pleurophragma. Notogaster: pcar = pleural carina; c, da, la, dm, lm, dp, lp, h, p = setae; Aa, A1, A2, A3, A4 = porose areas; Sa, S1, S2, S3 = saccules; ia, 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; cha, chb = cheliceral setae; Tg = Trägårdh's organ. Epimeral and lateral podosomal regions: 1a, 1b, 1c, 2a, 3a, 3b, 4a, 4b, 4c = epimeral setae; z = aperture of supracoxal gland; Am = humeral porose area; PdI, PdII = pedotecta I and II, respectively; dis = discidium; cir = circumpedal carina. Anogenital region: g, ag, an, ad = genital, aggenital, anal, and adanal setae, respectively; iad = adanal lyrifissure; Ap = postanal porose area; Amar = marginal porose area; po = preanal organ. Legs: Tr, Fe, Ge, Ti, Ta = trochanter, femur, genu, tibia, and tarsus, respectively; pa = porose area; e = famulus; d, l, v, ev, bv, ft, tc, it, p, u, a, s, pv, pl = setae; ω, σ, φ = solenidia.
ZOOBANK: 3C994735-9F32-40CE-9150-D117BAB1470C
(Figures 1, 2)
Type material — Holotype (female) and eight paratypes (three males and five females): Central Ethiopia, Amhara Region, Semien Shewa Zone, 9°51′32.4″N, 39°43′51.2″E, 10 km NE of Debre Birhan, 2900 m a. s. l., forest with Erica arborea, sifted moss and litter, 24.V.2014 (L. B. Rybalov).
The holotype is deposited in the collection of the Senckenberg Museum of Natural History, Görlitz, Germany; eight paratypes are in the collection of the University of Tyumen, Museum of Zoology, Tyumen, Russia. All specimens are preserved in 70% solution of ethanol with a drop of glycerol.
Diagnosis — Body length: 420–480. Body surface densely microsculpturing tuberculate; anogenital region slightly foveolate. Rostrum with triangular medial mucro. Lamella with small trapezoid cusp; translamella thin, straight; tutorium present. Sublamellar porose area absent. Rostral, lamellar and interlamellar setae slightly differs in length, long, setiform, barbed; bothridial seta with short stalk and globose, roughened head. Fourteen pairs of short, slightly dilated in median part, barbed setae. Five pairs of rounded notogastral porose areas. Epimeral, aggenital, anal, and adanal setae short, setiform, slightly barbed; all genital setae short, setiform, nearly smooth; leg femora I, II with four setae (l″ absent); genua I, II with two setae (v′ absent).
Description — Measurements. Body length: 465 (holotype), 420–480 (paratypes); notogaster width: 270 (holotype), 255–300 (paratypes). No difference between males and females in body size.
Integument — Body light brown to brown. Surface densely microsculpturing tuberculate (well observable under low magnification, × 400); additionally, anogenital region with slight sparse foveolae (diameter up to 4).
Prodorsum (Figs 1(a, c)) — Rostrum with strongly projecting triangular medial mucro. Lamella about 1/3 length of prodorsum, narrow, with small trapezoid cusp; translamella thin, straight; tutorium developed, ridge-like. Sublamellar porose area absent. Rostral (67–71), lamellar (67–75), interlamellar (60–67), and exobothridial (30–34) setae setiform, barbed; ex thinnest; in erect; bothridial seta (26) with short (7), smooth stalk and globose, roughened head (19). Dorsosejugal porose area slightly observable, oval.
Notogaster (Figs 1(a, c), 2(a)) — Anterior margin distinctly convex and rounded medially. Fourteen pairs of setae (15–19) slightly dilated in median part, barbed. Five pairs of rounded porose areas (7–15; sizes vary in specimens, but Aa frequently largest). Opisthonotal gland opening and all lyrifissures well observable.
Gnathosoma (Figs 2(b–d)) — Subcapitulum size: 101–105 × 75–82; subcapitular (22–26) and adoral (13–15) setae setiform, barbed. Palp length: 64–67; setation: 0–2–1–3–9(+ω); postpalpal seta (7) spiniform, nearly smooth. Chelicera length: 116–120; setae (cha: 34; chb: 22) setiform, barbed.
Epimeral and lateral podosomal regions (Figs 1(b, c)) — Epimeral formula: 3–1–3–3; all setae (1a, 2a, 3a: 17–19; others: 22–26) setiform, slightly barbed. Humeral porose area Am slightly observable, oval. Discidium not developed. Circumpedal carina short.
Anogenital region (Figs 1(b, c), 2(a)) — Anogenital formula: 4–1–2–3; all genital setae (15) setiform, nearly smooth; aggenital (19), anal (15) and adanal (19) setae setiform, slightly barbed. Adanal lyrifissure close and anterior to anal plate. Postanal porose area band-like. Ovipositor typical for Oribatulidae (Ermilov 2010); length of blade: 71; length of distal section (beyond middle fold): 105; width of distal section: 52; each of the three blades with four smooth setae: ψ1 ≈ τ1 (37–41) setiform; ψ2 ≈ τa ≈ τb ≈ τc (11–15) narrowly thorn-like; coronal setae absent.
Download as Note: Roman letters refer to normal setae; Greek letters to solenidia; single prime (‘) marks setae on the anterior and double prime (’’) setae on the posterior side of a given leg segment; parentheses refer to a pair of setae.
Leg
Tr
Fe
Ge
Ti
Ta
I
v’
d,* l’, bv’‘, v’’*
(l),* *σ
(l),* (v)*, φ1, φ2
(ft),* (tc), (it), (p), (u), (a), s, (pv), v’, (pv), e*, ω1, ω2
II
v’
d,* l’, bv’‘, v’’*
(l),* *σ
(l), (v), φ
(ft),* (tc), (it), (p), (u), (a), s, (pv)*, ω1, ω2
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)*
Legs (Figs 2(e–h)) — Median claw thicker than lateral claws; all slightly barbed on dorsal side; each lateral claw with tubercle distoventrally. Proximoventral porose area on tarsi I–IV, distoventral porose area on tibiae I–IV and dorsoparaxial porose area on femora I–IV and on trochanters III, IV well observable. Formulas of leg setation and solenidia: I (1–4–2–4–19) [1–2–2], II (1–4–2–4–15) [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; seta s eupathidial on tarsus I.
Remarks — In having medium-sized body (about 450), pointed rostrum, short cusp on the lamella, short and globose bothridial seta, rounded notogastral porose areas, and 14 pairs of short, barbed notogastral setae, Z. birhanensis n. sp. is similar to Z. josefstaryi Ermilov and Rybalov, 2013 from Ethiopia. The new species can be distinguished from the latter by the morphology of the rostrum (with triangular medial mucro versus mucro absent), the length of the interlamellar seta (distinctly shorter than prodorsum, and not longer than rostral and lamellar setae versus as long as prodorsum or longer, and distinctly longer than rostral and lamellar setae), the presence of five pairs (versus four pairs) of notogastral porose areas, the morphology and lengths of notogastral setae (slightly dilated in median part, similar in length versus setiform, c1, c2 , da, la distinctly longer than others), and the leg setation (femora I, II with four setae, l″ absent; genu I with two setae, v′ absent versus femora I, II with five setae, l″ present; genu I with three setae, v′ present).
Etymology — The species name birhanensis refers to vicinities of the place of origin, Debre Birhan.
ZOOBANK: 1B3957F5-249E-42A9-AC65-A69D55CBB280
(Figures 3, 4)
Type material — Holotype (male) and eight paratypes (five males and three females): Central Ethiopia, Amhara Region, Semien Shewa Zone, 9°51′32.4″N, 39°43′51.2″E, 10 km NE of Debre Birhan, 2900 m a. s. l., forest with Erica arborea, sifted moss and litter, 24.V.2014 (L. B. Rybalov).
The holotype is deposited in the collection of the Senckenberg Museum of Natural History, Görlitz, Germany; eight paratypes are in the collection of the University of Tyumen, Museum of Zoology, Tyumen, Russia. All specimens are preserved in 70% solution of ethanol with a drop of glycerol.
Diagnosis — Body length: 480–525. Rostrum rounded. Translamella represented by two short indistinct lines near lamellae; prolamella complete (reaching insertion of rostral seta); lateral keel-shaped ridge slightly arch-like. Rostral, lamellar and interlamellar setae long, setiform, barbed; ro shortest; in longest; bothridial seta long, lanceolate, barbed. Pteromorph large, rounded laterally, its anterior margin slightly oblique. All notogastral setae short, setiform, with flexible tip, smooth. Epimeral, aggenital, anal, and adanal setae short, setiform, slightly barbed; three pairs of genital setae short, setiform, nearly smooth. Leg tarsi I and II with two longitudinal ridges and two teeth distodorsally; femora I and II large.
Description — Measurements. Body length: 525 (holotype), 480–525 (paratypes); body width (level of pteromorph): 285 (holotype), 240–285; width of ventral plate: 240 (holotype), 210–240 (paratypes). No difference between males and females in body size.
Integument — Body color brown. Body surface partially microfoveolate (foveolae sparsely located, slightly observable).
Prodorsum (Figs 3(a, c)) — Rostrum rounded. Lamella about 1/2 length of prodorsum; translamella represented by two short indistinct lines near lamellae; prolamella complete (reaching insertion of rostral seta); lateral keel-shaped ridge slightly arch-like, located above acetabulum I. Sublamellar porose area nearly rounded (9–11). Rostral (37–49), lamellar (52–64), interlamellar (71–79), and exobothridial (30–34) setae setiform, barbed; ex thinnest; in erect; bothridial seta (86–101) lanceolate, barbed, directed posterolaterad; stalk longer than head. Dorsosejugal porose area not observable.
Notogaster (Figs 3(a, c), 4(a)) — Anterior margin distinctly convex and rounded medially. Pteromorph large, rounded laterally, its anterior margin slightly oblique (not perpendicular to longitudinal axis of body in dorsal aspect). Ten pairs of setae (19–22) setiform, with flexible tip, smooth. Four pairs of saccules with small opening and drop-like channel. Opisthonotal gland opening and all lyrifissures well observable.
Gnathosoma (Figs 4(b–d)) — Subcapitulum size: 112–120 × 79–86; subcapitular (a: 22–26; m: 11; h: 26) and adoral (13–15) setae setiform, slightly barbed. Palp length: 67–75; setation: 0–2–1–3–9(+ω); postpalpal seta (7) spiniform, truncated; roughened. Chelicera length: 120–124; setae (cha: 37–41; chb: 26) setiform, barbed.
Epimeral and lateral podosomal regions (Figs 3(b, c)) — Epimeral formula: 3–1–3–3; all setae (1a, 2a, 3a: 15; 4a, 4b: 19; others: 19–22) setiform, slightly barbed. Humeral porose areas Am and Ah not observable. Pedotectum II quadrangular distally in ventral view, with posterolateral tooth. Circumpedal carina short. Discidium slightly developed, broadly rounded.
Anogenital region (Figs 3(b, c), 4(a)) — Anogenital formula: 3–1–2–3; genital (11–13) and aggenital (15) setae setiform, nearly smooth; anal and adanal setae (19–22) setiform, slightly barbed. Adanal lyrifissure close and parallel to anterior half of anal plate. Marginal porose area complete, band-like. Ovipositor typical for Scheloribatidae (Ermilov 2010); length of blade: 75; length of distal section (beyond middle fold): 90; width of distal section: 37; each of the three blades with four smooth setae: ψ1 ≈ τ1 (30–34) setiform; ψ2 ≈ τa ≈ τb ≈ τc (11–13) and six coronal setae (7) narrowly thorn-like.
Download as Note: See Table 1 for explanations.
Leg
Tr
Fe
Ge
Ti
Ta
I
v’
d,* (l), bv’‘, v’’*
(l),* *σ
(l),* (v)*, φ1, φ2
(ft),* (tc), (it), (p), (u), (a), s, (pv), v’, (pv), e*, ω1, ω2
II
v’
d,* (l), bv’‘, v’’*
(l),* *σ
(l), (v), φ
(ft),* (tc), (it), (p), (u), (a), s, (pv)*, ω1, ω2
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)*
Legs (Figs 4(e–h)) — Median claw thick, with small tubercle ventrobasally; both lateral claws thin, with tubercle distoventrally; all claws slightly barbed on dorsal side. Tarsi I and II with two longitudinal ridges and two teeth distodorsally; tarsus III with slight longitudinal ridges, without teeth. Femora I and II large. Proximoventral porose area on tarsi I–IV, distoventral porose area on tibiae I–IV and dorsoparaxial porose area on femora I–IV and on trochanters III, IV well observable. Formulas of leg setation and solenidia: I (1–5–2–4–19) [1–2–2], II (1–5–2–4–15) [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; seta s setiform, barbed on tarsus I.
Remarks — In having medium-sized body (about 500) and strong teeth on the leg tarsi I, II, S. (S.) debreensis n. sp. is similar to S. (S.) diversidactylus (Hammer, 1961) from the Neotropical region and India, S. (S.) potchefstroomensis Ermilov, Hugo-Coetzee, Khaustov and Theron, 2017 from South Africa, S. (S.) sudafricanus Subías, 2018 (=Urubambates calcaratus Mahunka, 1984) from South Africa, and S. (S.) tricarinus Coetzer, 1968 from the Afrotropical region. The new species can be clearly distinguished from S. (S.) diversidactylus, S. (S.) sudafricanus, and S. (S.) tricarinus by the morphology of the bothridial seta (lanceolate versus setiform), and the presence of three pairs of the genital setae (versus four pairs). The new species can be distinguished from S. (S.) potchefstroomensis by the length of the lanceolate bothridial head (long versus short), the direction of the prolamella (directed to insertion of rostral seta versus directed to lateral side of prodorsum), the morphology of the notogastral, anal and adanal setae (with flexible mediodistal part versus setiform), the presence of three pairs of genital setae (versus four pairs), and longer anal and adanal setae.
The representatives of Scheloribates (Scheloribates) and Hammerabates Balogh, 1970 are similar, but differs from each other mainly by the number of genital setae (four pairs versus three pairs). The new species has three pairs of genital setae (as species of Hammerabates), however, it is placed in Scheloribates (Scheloribates) based on its morphological similarities with group of the Scheloribates (Scheloribates) species (see above) having strong teeth on the leg tarsi I, II, as well as on the location of the genital setae (one seta in anterior part of plate, two setae in posterior part of plate in S. (S.) debreensis versus two setae in anterior part of plate, one seta in posterior part of plate in Hammerabates). Thus, this unique case should be taken into account in the future when updating the diagnosis of Scheloribates (Scheloribates).
Etymology — The species name debreensis refers to vicinities of the place of origin, Debre Birhan.
Pérez-Íñigo (1983) proposed the new scheloribatid genus Annobonzetes Pérez-Íñigo, 1983, with Annobonzetes sphaericus Pérez-Íñigo, 1983 as type species. Later, Mahunka (1994) presented a supplementary description of Podoribates latissimus Berlese, 1916 (in 1916a) and included this species in Annobonzetes. The genera Annobonzetes and Scheloribates are generally similar morphologically, and based only on original generic diagnosis (Pérez-Íñigo 1983), there are difficulties in distinguishing them. However, additional data presented by Mahunka (1994) regarding A. latissimus allow to preliminarily support the generic status of Annobonzetes due mainly to the presence of a strong ridge-like translamella (complete or interrupted medially), which is absent or represented by a simple line in representatives of Scheloribates. The other morphological characters of Annobonzetes (e.g. large body size, spherical notogaster in dorsal aspect, presence of setal alveoli instead of setae etc.) appear to be species-level within Scheloribatidae, but may perhaps be considered in conjunction with the main generic character (presence of strong ridge-like translamella).
Subías (2004) included Oribata lata Warburton, 1912 in Annobonzetes. However, the description and the schematic figure in the original description (Warburton 1912) do not allow understanding the true generic and family placement of this species. For example, the presence of a strong translamella and two longitudinal ridges extending from it are a characteristic trait of some representatives of Mochlozetidae (e.g. Unguizetes Sellnick, 1925).
Wan et al. (2022) described Annobonzetes hamatus Wan, Tan, Yan and Xie, 2022. They excluded A. latissimus from Annobonzetes based on the original description only, and added Scheloribates acutirostrum Ermilov, Rybalov and Franke, 2011 and S. helenensis Wallwork, 1977 in Annobonzetes based on the presence of spherical notogaster, the absence of prolamella, and the presence of short notogastral setae or their alveoli. Also, they presented the new generic diagnosis of Annobonzetes. First, the morphological characters (e.g. notogaster form, presence/absence of prolamella and notogastral setae) that the authors used (Wan et al. 2022) for transfer of S. acutirostrum and S. helenensis are species-level characters within Scheloribates (see original descriptions and redescriptions of representatives of the genus). Second, they (Wan et al. 2022) omitted Mahunka's paper (1994) with a supplementary description of A. latissimus, which did not allow them to combine the common morphological characters of Annobonzetes and understand the main differences between Scheloribates and Annobonzetes. Third, the generic diagnosis of Annobonzetes (Wan et al. 2022) includes a set of typical morphological characters that can be applied to several supraspecies taxa within Oripodoidea including Scheloribates. Fourth, A. hamatus has thin translamellar lines near lamellae (instead strong ridge-like translamella as in Annobonzetes species) which are typical for Scheloribates species.
Hence, based on above listed explanations, the following conclusions are proposed: Scheloribates (Scheloribates) hamatus (Wan, Tan, Yan and Xie, 2022) n. comb. (from Annobonzetes); initial generic placement of S. acutirostrum and S. helenensis in Scheloribates (Scheloribates) is supported; further morphological and taxonomic investigations on Oribata lata are needed; the generic diagnosis of Annobonzetes in Wan et al. (2022) is inapplicable.
I thank Dr. Gezahegn Degefe for supporting our field studies and organizing laboratory work, Dr. Leonid B. Rybalov for sampling assistance and two anonymous reviewers for valuable comments. The work was performed within the framework of the Joint Russian-Ethiopian Biological Expedition, financially supported by the Russian Academy of Sciences. The collection of materials was conducted under the Agreement between the Russian Academy of Sciences and the Ministry of Science and Technology (Ministry of Innovation and Technology) of the Federal Democratic Republic of Ethiopia.