Ermilov, Sergey G. ¹ and Makarova, Olga L. ²

¹✉ Institute of Environmental and Agricultural Biology (X-BIO), Tyumen State University, Tyumen, Russia.
²Severtsov Institute of Ecology and Evolution, Russian Academy of Sciences, Moscow, Russia.

2023 - Volume: 63 Issue: 1 pages: 241-252

ZooBank LSID: 1B4DDD34-94AD-48EF-B700-1CE646664E7D

Original research

Keywords

Abstract

Introduction

The oribatid mite family Epimerellidae (Acari, Oribatida, Oppioidea) was proposed by Ayyildiz and Luxton (1989), based on the following morphological traits: medial crista of anterior notogastral margin well developed and produced into prodorsum; epimeres III+IV separated medially, with broad concavity between them; 10–13 pairs of notogastral setae, with shortest seta c; five pairs of genital, one pair of aggenital, two pairs of anal, and three pairs of adanal setae; legs monodactylous. The family includes two genera: Epimerella Kulijev, 1967; and Enisella Ayyildiz and Luxton, 1989. Subias (2009) considered the rather specific Indonesian genus Luxtonia Mahunka, 2001 also the member of Epimerellidae (see Discussion section below).

Epimerella was described by Kulijev (1967), with Oppia smirnovi Kulijev, 1962 as type species; at present, the genus comprises eight species and one subspecies, which are distributed in the southern Palaearctic region (Subías 2022). Enisella was described by Ayyildiz and Luxton (1989), with Enisella turcica Ayyildiz and Luxton, 1989 as type species; at present, the genus comprises one (type) species, which is known only from Turkey (Subías 2022).

During the taxonomic identification of oribatid mites from salty habitats in Kalmykia, Russia (Fig. 1), we found one new species belonging to Epimerella. The main goals of our paper are: to describe and illustrate a new species under the name Epimerella kalmykorum n. sp.; to summarize generic morphological traits; and to present the renewed identification key, distribution, and habitats to known species of the family. Besides, new record of Enisella turcica Ayyildiz and Luxton, 1989 is presented based on samplings in Crimea by N.A. Kuznetsova and M.B. Potapov in 2021.

Presently, two species and one subspecies of Epimerellidae have been registered in the Russian fauna (e.g., Gordeeva and Karppinen 1988; Shtanchaeva and Subías 2010): Epimerella puzanovi Gordeeva and Karppinen, 1988; Ep. smirnovi (Kulijev, 1962); and Ep. smirnovi longisetosa Kulijev, 1967.

Material and methods

Specimens

Mites were collected during the special expedition organized in November 2021 for the testing the assumption about the possible occurence of littoral halophilic species within the Kuma-Manych Depression. This area is the former ancient strait that connected the Black and Caspian Seas in the geological past. Its course is characterized by numerous salt lakes, and the Manych-Gudilo Lake is the largest from them. The climate of this hot and dry during the summer area is considered as moderately continental. The mid-July temperature in this area is 24.4 °C, the mid-January temperature is –6.8 °C, annual precipitation 360 mm (Scientifically-Applied Handbook, 1990), and the salinity of the Manych-Gudilo Lake during last years is more than 40‰ (Bulysheva et al. 2018). In general, 31 samples of turf, litter and soil were taken (about 6 liters) on banks of five lakes and two rivers. And only one sample included two specimens of a new species belonging to Epimerella.

Besides, the enough collection of mites obtained by N.A. Kuznetsova and M.B. Potapov from different saline sites of the Crimea in September 2021 was studied (27 localities, about 40 samples). Among other mites, this yielded one specimen of Enisella turcica.

Samples (each of volume 150–300 ml) were taken from the turf of saline lands, litter of reeds as well as from storm debris on shores of salty lakes and rivers. Mites were extracted from the cores in Moscow using laboratory eclectors by drying the samples for 10 days until their completely dry condition without additional heating or lighting. Microarthropods were fixed with 85–96% alcohol.

Observation and documentation

For measurement and illustration, specimens were mounted in lactic acid on temporary cavity slides. All measurements are in micrometers. Body length was measured in lateral view, from the tip of the rostrum to the posterior edge of the notogaster; other structures were oriented to avoid parallax errors. Notogastral width refers to the maximum in dorsal aspect. Setal lengths were measured perpendicular to their long axis, accounting for curvature. 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 follows that of F. Grandjean: see Travé and Vachon (1975) for references; Norton (1977) for leg setal nomenclature; Norton and Behan-Pelletier (2009) for overview.

Abbreviations

Prodorsum: ro, le, in, bs, ex = rostral, lamellar, interlamellar, bothridial, and exobothridial seta, respectively; exv = vestige of second exobothridial seta; pbt = postbothridial tubercle. Notogaster: ap = apophysis; cr = crista; c, la, lm, lp, h, p = setae; ia, im, ip = lyrifissures; gla = opisthonotal gland opening. Gnathosoma: a, m, h = subcapitular setae; or = adoral seta; sup, inf, d, l, cm, acm, ul, su, vt = palp setae; ω = palp solenidion; cha, chb = cheliceral setae; Tg = Trägårdh's organ. Epimeral and lateral podosomal regions: con = concavity; 1a–1c, 2a, 3a–3c, 4a–4c = epimeral setae; PdI = pedotectum I; dis = discidium. Anogenital region: g, ag, an, ad = genital, aggenital, anal, and adanal seta, respectively; iad = adanal lyrifissure; p.o. = preanal organ. Legs: Tr, Fe, Ge, Ti, Ta = trochanter, femur, genu, tibia, and tarsus, respectively; ω, φ, σ = solenidia; ɛ = famulus; d, l, v, bv, ev, ft, tc, it, p, u, a, s, pv, pl = setae.

Taxonomy

Family diagnosis of Epimerellidae Ayyildiz and Luxton, 1989

The representatives of the family Epimerellidae are characterized within the superfamily Oppioidea by the presence of broad concavity between epimeres III+IV and strong protruding apophysis in anteromedial part of the notogaster.

Generic diagnosis of Epimerella Kulijev, 1967

Adult

With character states of Epimerellidae (as above). Measurements: Small species, length often about 300. Integument. Without heavy ornamentation and sculpturing. Prodorsum. Rostrum rounded, incised or tripartite. Costula, transcostula and lateral ridge absent. Rostral, lamellar, interlamellar, and exobothridial setae well developed, setiform; le inserted closer to in than to ro. Bothridial seta long, with elongate, slightly dilated and ciliate unilaterally head. Interbothridial tubercles absent; postbothridial tubercles present. Notogaster. Anteromedial apophysis bifurcate. Humeral region without tooth but sometimes with indistinct tubercle and short crista. Ten pairs of setiform setae (da, dm and dp absent): c shortest; posterior setae (p₁–p₃) medium-sized; dorsal setae long. Gnathosoma. Subcapitulum diarthric; adoral setae present. Palp solenidion bacilliform, entirely pressed to surface. Chelicera chelate-dentate. Epimeral and lateral podosomal regions. Epimeres I and II medially distinctly or slightly separated, or fused; sometimes epimere I with deep pit-like structure. Epimeral setae setiform. Pedotectum I represented by small lamina. Discidium present. Anogenital region. Five pairs of genital, one pair of aggenital, two pairs of anal, and three pairs of adanal setae; adanal seta ad₁ posterior, ad₂ lateral, ad₃ anterolateral to anal plate, distance ad₃–ad₃ longer than ag–ag and ad₂–ad₂. Adanal lyrifissure located close and parallel to anal aperture. Legs. Tarsus I with 20 setae (l″ and v′ present), tarsus II with 16 setae (l″ present). Tarsus II with two solenidia.

Epimerella kalmykorum n. sp.

ZOOBANK: 95498F39-2317-40DD-B57F-9FADBAC49566

(Figures 2–4)

Diagnosis

Body length: 315–323. Epimeres I, II, mentum, gena of subcapitulum and lateral side of prodorsum tuberculate. Rostrum with median incision. Rostral, lamellar, interlamellar, and exobothridial setae setiform, barbed; ro arch-like, directed anteriorly; bothridial seta with 10 ciliae unilaterally; relative length: bs ˃ in = le ˃ ex ˃ ro. Dorsal notogastral setae long (la, lm and h₃ reaching beyond insertions of h₃, lp and h₂, respectively). Epimere I complete versus epimeres II, III+IV separated medially by large concavity. Epimeral and anogenital setae setiform, barbed; 4c located on top of discidium.

Description

Measurements – Body length: 315 (holotype: female), 323 (one paratype: female); body width: 180 (holotype), 180 (paratype).

Integument – Body color light brown. Body surface mostly smooth but epimeres I, II, mentum and gena of subcapitulum, and lateral side of prodorsum with sparse, distinct, tubercles (diameter up to 2).

Prodorsum – Rostrum with semi-rectangular incision. Rostral (28–30), lamellar (47–49), interlamellar (49), and exobothridial (34–37) setae setiform, barbed; ro arch-like, directed anteriorly; head of bothridial seta (64–67) with 10 ciliae unilaterally. Postbothridial tubercle slightly developed.

Notogaster – Anterior bifurcate notogastral apophysis well-developed. Crista short, slightly visible. All notogastral setae (c: 18–22; p₁–p₃: 60–67; others: 90–96) setiform, barbed; c thinnest, curved backwards; dorsal setae long (la, lm and h₃ continues beyond insertions of h₃, lp and h₂, respectively); in holotype, insertions of lp and h₃ on right side of notogaster located close to each other. Lyrifissures ia, im and ip distinct versus ih and ips not observed.

Gnathosoma – Subcapitulum size: 67 × 47–49; subcapitular and adoral setae setiform, a (11) slightly barbed, m, h (17–19) barbed, or (4) smooth. Palp length: 47–49; setation: 0–2–1–3–8(+ω); solenidion medium-sized (1/2 of tarsus); postpalpal seta (4) spiniform, smooth. Chelicera length: 67; setae (cha: 22; chb: 15) setiform, barbed.

Epimeral and lateral podosomal regions – Epimere I medially fused versus epimeres II, III+IV separated medially by large concavity. Epimeral setae (1a, 2a, 3a: 15–19; 1b, 4b: 28–30; others: 41–49) setiform, barbed; in holotype, left side of epimere IV with extra seta between setae 4a and 4b. Discidium triangular, bearing insertion of seta 4c on the top.

Anogenital region – Genital (g₁: 19–22; others: 15–19), aggenital (19–22), adanal (34–41), and anal (19–22) setae setiform, barbed. Adanal lyrifissure distinct.

Legs – Claw of each leg smooth; claw of leg IV larger than those on legs I–III. Porose area on all segments not observed. Formulas of leg setation and solenidia: I (1–5–2–4–20) [1–2–2, II (1–5–2–4–16) [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. Setae p′ and p″ setiform on tarsus I versus very short, conical on tarsi II–IV; seta s eupathidial on tarsus I; famulus short, erect, slightly swollen distally; solenidion ω₁ on tarsus I, ω₁, ω₂ on tarsus II and σ on genu III slightly bacilliform versus other solenidia setiform.

Type material

Holotype (female) and one paratype (female): European Russia, Republic of Kalmykia, bank of salty Manych-Gudilo Lake, southern shore of the Left Island, floodplain, 45°59′N, 43°32′E, reed debris (Fragmites australis) under Tamarix ramossisima among Salicornia perennans and Bassia prostrata, 13.11.2021 (leg. O.L. Makarova) (Fig. 1).

The holotype and one paratype are deposited in the collection of the Tyumen State University Museum of Zoology, Tyumen, Russia. Both specimens are preserved in 70% solution of ethanol with a drop of glycerol.

Etymology

The species name refers to the place of origin, Republic of Kalmykia.

Remarks

Epimerella kalmykorum n. sp. is morphologically similar to Ep. luxtoni Toluk, Ayyildiz and Baran, 2008 in having semi-rectangular incision of the rostrum and the structure of epimeral region (epimere I complete versus epimeres II–III+IV interrupted medially by large concavity). However, the new species differs from the latter by: anterior notogastral apophysis well developed reaching mid-distance between interlamellar and lamellar setae (versus hardly reaching the transversal level of interlamellar setae); rostral setae arch-like, directed anteriorly (versus straight, divergent); epimeres I, II, mentum and gena of subcapitulum, and lateral side of prodorsum sparsely tuberculate (versus not tuberculate); dorsal notogastral setae long (la, lm and h₃ reaching beyond insertions of h₃, lp, h₂, respectively, versus not reaching these insertions); and notogastral setae c barbed (versus smooth).

Distinctive characters of the new species compared with other members of Epimerellidae can be found in the identification key below.

Generic diagnosis of Enisella Ayyildiz et Luxton, 1989

Adult — With character states of Epimerella (as above) but with 13 pairs of notogastral setae (da, dm, and dp present).

Enisella turcica Ayyildiz and Luxton, 1989

Material — One specimen: Crimea, Leninsky district, Arabatskaya Strelka, bank of Sivash Lake, 45°18′N, 35°27′E, saline land with grasses and Limonium sp., 26.09.2021 (leg. N.A. Kuznetsova, M.B. Potapov). First record of the genus and species from Crimean peninsula.

Discussion

Mahunka (2001) described the genus Luxtonia from Borneo, with Luxtonia hauseri Mahunka, 2001 as type species, and he placed it in the newly described family Luxtoniidae Mahunka, 2001. Subías (2004) did not support the independence of the new family and included Luxtonia in Epimerellidae. Schatz et al. (2011) espoused the Mahunka's opinion. Luxtonia hauseri does not have the specific structure of the epimeral region (epimeres III+IV separated by broad concavity), what is the main morphological trait of Epimerellidae, and also has a unique structure of posterior part of the notogaster (divided into overlapping lobes and connecting with postanal saccular structure). Therefore, we support the initial placement of Luxtonia within Luxtoniidae.

Key to known species of Epimerellidae

1. Thirteen pairs of notogastral setae (da, dm, dp present); body length: 320–360
...... Enisella turcica Ayyildiz and Luxton, 1989

— Ten pairs of notogastral setae (da, dm, dp absent)
...... 2

2. Epimere I medially with deep pit-like structure
...... 3

— Epimere I medially well separated or fused (pit-like structure absent)
...... 5

3. Rostrum tripartite; insertions of rostral setae located close to each other (on the distance about their alveolus size); body length: 310
...... Epimerella puzanovi Gordeeva and Karppinen, 1988

— Rostrum rounded; insertions of rostral setae widely spaced (on the distance much more than their alveolus size)
...... 4

4. Pit-like structure of epimere I posteriorly triangular; epimere II medially fully coalescent; body length: 250–328
...... Epimerella marasensis Toluk and Ayyildiz, 2013

— Pit-like structure of epimere I posteriorly broadly rounded; epimere II medially narrowly separated; body length: 300–330
...... Epimerella ankaraensis Baran, Ayyildiz and Kence, 2012

5. Epimere I medially broadly separated; rostrum with semi-rectangular incision
...... 6

— Epimere I medially fused; rostrum tripartite or with semi-rectangular incision
...... 7

6. Dorsal notogastral setae lm and h₃ reaching beyond insertions of h₃ and h₁, respectively; body length: 360
...... Epimerella distenta Ayyildiz and Luxton, 1989

— Dorsal notogastral setae lm and h₃ shorter, not reaching beyond insertions of h₃ and h₁, respectively; body length: 284–286
...... Epimerella subiasi Toluk, Ayyildiz and Baran, 2008

7. Rostrum with semi-rectangular incision
...... 8

— Rostrum tripartite
...... 9

8. Dorsal notogastral setae la, lm and h₃ reaching beyond insertions of h₃, lp, and h₂, respectively; rostral setae arch-like, directed anteriorly; epimeres I, II, mentum, and lateral side of prodorsum tuberculate; body length: 315–323
...... Epimerella kalmykorum n. sp.

— Dorsal notogastral setae la, lm and h₃ shorter, not reaching beyond insertions of h₃, lp and h₂, respectively; rostral setae straight, divergent; epimeres I, II, mentum and lateral side of prodorsum not tuberculate; body length: 292–336
...... Epimerella luxtoni Toluk, Ayyildiz and Baran, 2008

9. Dorsal notogastral setae la and lm shorter, not reaching beyond insertions of lp and h₂, respectively; body length: 494
...... Epimerella rubeni Khanbekyan et Gordeeva, 1991

— Dorsal notogastral setae la and lm longer, reaching beyond insertions of lp and h₂, respectively
...... 10

10. Head of bothridial seta with 13 ciliae; notogastral setae p₁–p₃ half as long as most dorsal notogastral setae; body length: 340
...... Epimerella smirnovi longisetosa Kulijev, 1967

— Head of bothridial seta with five ciliae; notogastral setae p₁–p₃ fivefold shorter than most dorsal notogastral setae; body length: 340
...... Epimerella smirnovi smirnovi Kulijev, 1962 (see also Kulijev 1967)

Distribution and habitats of Epimerellidae

At present, all 11 representatives of Epimerellidae have been recorded only in the southern Palaearctic region. Except Epimerella smirnovi smirnovi and Ep. smirnovi longisetosa, the other nine species have a highly confined geographic distribution, i.e. are conventionally endemic to a single country.

Six species are known only from Turkey: Enisella turcica was described from forest soil (plum grove); Epimerella distenta from pasture soil (Ayyildiz and Luxton 1989); all other species (Ep. luxtoni, Ep. marasensis, Ep. subiasi, and Ep. ankaraensis) were found mainly within different pine woods, rarely in other forests (Toluk et al. 2008; Baran et al. 2012; Toluk and Ayyildiz 2013; Ay and Ayyildiz 2019b).

Two species are known only from the south areas of the European Russia: Ep. kalmykorum is described from the reed debris nearly salty lake in Kalmykia (data of this paper); the strict habitat in Crimea (Karadag Mts.) for Ep. puzanovi is unknown (Gordeeva and Karppinen 1988).

One species is recorded only from Armenia: Ep. rubeni from litter under acacia trees (Khanbekyan and Gordeeva 1991).

Epimerella smirnovi smirnovi and Ep. smirnovi longisetosa are broadly distributed in the East Mediterranean: the former is known from Azerbaijan, Georgia, Iran, and south of the European Russia; the latter is known from Turkey, Ukraine and south of the European Russia; both species are found mostly in forest soil-litter and moss, as well as in meadow and urban soils (e.g., Kulijev 1962, 1967; Bocharova 1973; Mortazavi et al. 2011; Murvanidze and Mumladze 2016; Ay and Ayyildiz 2019a; Murvanidze et al. 2020).

According to the summarized data, representatives of Epimerellidae prefer mainly the soil-litter sites in forest ecosystems, however, they also occur in the other habitats, such as meadows and urban soils.

Thus, Ep. kalmykorum is unique among congeners dwelling in extremal conditions on the low bank of the hypersaline Manych-Gudilo Lake very close to the water edge (Fig. 1). The salinity of this waterbody in last years varied in average between 40 to 56 ‰ (Bulysheva et al., 2018) and wave height reaches several meters. So, one can conclude that this species is strongly halolerant. Two specimens of a new species co-habited with numerous specimens of predatory mites, Neoseiuslus tervus (Meshkov, 1994) and N. extricatus (Kolodochka, 1991). These species are halotolerant judging from their records by Kolodochka (1991) and Döker et al. (2021). Some eurybiontic mesostigmatic species (Arctoseius cetratus (Sellnick, 1940), Gaeolaelaps spp., and Proctolaelaps pygmaeus Müller, 1859) were also found in this habitation. Besides, the sample in question contained also numerous specimens of oribatid mite Zetomimus acutirostris (Mihelčič, 1957), which has been recorded on extensive farmed crop fields with the high application of mineral fertilizers and biocides (Kulijev 1962) as well.

Enisella turcica Ayyildiz and Luxton, 1989 described from the plum litter in Turkey more than thirty years ago and was not recorded anywhere else till now. The satisfactory sampling in Crimea gave only one individual of this species from the saline soil. It is the first record from Crimean Peninsula.

Both Crimean and Kalmykian expeditions in 2021 were conducted after the hot summer period in September and November, respectively. Probably in autumn, small mites migrate upward within the soil and just this circumstance enabled us to reveal these rare species.

Acknowledgements

We are thankful to N.A. Kuznetsova and M.B. Potapov kindly supplying us by the comprehensive mite collection from Crimea as well as to the huntsman of the State Nature Biosphere Reserve ''Black Lands (Chernye Zemli)'', Yu.V. Babich, for the all-round help during field works in Kalmykia. Our colleagues, O. Joharchi, V.A. Khaustov and N.Yu. Stepanova amiably consulted our identification of some mites and plants; and two anonymous reviewers for valuable comments. This research was partially supported by the grant from the Russian Foundation for Basic Research (No. 20-54-56054 Iran_t).

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