# A new subgenus and two new species of oribatid mites
(Acari, Oribatida) from Madagascar

Sergey G Ermilov, Andrey V Frolov

► To cite this version:

Sergey G Ermilov, Andrey V Frolov. A new subgenus and two new species of oribatid mites (Acari, Oribatida) from Madagascar. Acarologia, 2022, 62 (3), pp.798-810. 10.24349/1m3y-yi0t. hal-03766901

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# A new subgenus and two new species of oribatid mites (Acari, Oribatida) from Madagascar

Sergey G. Ermilov IDª, Andrey V. Frolov IDᵇ

## Original research

### ABSTRACT

This study is based on oribatid mite materials collected by swiping bushes in central Madagascar. Four species are identified; a new subgenus and two new species, *Afreremella* (*Arboreremella*) **n. subgen.**, with type species *Afreremella* (*Arboreremella*) *madagascarensis* **n. sp.** (Eremellidae), and *Lucoppia ankaratraensis* **n. sp.** (Oribatulidae) are described. An identification key to known representatives of *Lucoppia* is provided. The initial subgeneric status of *Eremella* (*Archeremella*) is supported.

**Keywords** acarofauna; taxonomy; generic diagnosis; new taxa; morphology; *Afreremella*; *Lucoppia*; identification key; Afrotropical region

**Zoobank** http://zoobank.org/7038FCA4-E123-4EA6-9622-0EE84CCC9DFA

## Introduction

The oribatid mite (Acari, Oribatida) fauna of Madagascar is insufficiently studied (e.g. Balogh 1962; Mahunka 1993, 2010; Ermilov and Starý 2020). Our work is based on materials which were collected by swiping bushes during a field trip in central Madagascar in 2022. The collected specimens belong to four species (it is theoretically possible that some species are arboreal) of oribatid mites; of these, two species from the genera *Afreremella* Mahunka, 1973 (Eremellidae) and *Lucoppia* Berlese, 1908 (Oribatulidae) are new to science. The main goal of this paper is to describe these two new species and a new subgenus within *Afreremella*.

The oribatid mite genus *Afreremella* was proposed by Mahunka (1973) with *Afreremella luisiae* Mahunka, 1973 as type species. At present, the genus comprises only one (type) species, which is known from Zimbabwe; hence, *Afreremella* is recorded in Madagascar for the first time. The supraspecies system of Eremellidae and the support for independence of *Afreremella* were discussed by Ermilov and Frolov (2019), however, Subías (2004, online version 2022) synonymized this genus with *Eremella* (*Licnocepheus*) Woolley, 1969 without explanations.

The oribatid mite genus *Lucoppia* was proposed by Berlese (1908) with *Notaspis burrowsi* Michael, 1890 (= *Zetes lucorum* Koch sensu Berlese, 1892) as type species. At present, the genus comprises two species, which are distributed collectively in the Holarctic region, Chile and tropics (except Africa); hence, *Lucoppia* is recorded in the Afrotropical region for the first time. Subías (2004, online version 2022) included also *Lucoppia nicora* Dzhaparidze, 1986 (see Dzhaparidze and Gomelauri 1986) in *Lucoppia*, however, we previously support the placement of this species within *Zygoribatula* Berlese, 1916 according to opinion of Murvanidze et al. (2018).

The additional goals of this paper are: to present the revised generic diagnoses for *Afreremella* and *Lucoppia*; to provide an identification key to known representatives of *Lucoppia*; and to discuss the taxonomic status of the genus *Archeremella* Balogh and Mahunka, 1974.

Received 09 May 2022
Accepted 12 August 2022
Published 29 August 2022

Corresponding author
Sergey G. Ermilov ID:
ermilovacari@yandex.ru

Academic editor
Baumann, Julia

https://doi.org/
10.24349/1m3y-yi0t

ISSN 0044-586X (print)
ISSN 2107-7207 (electronic)

<figure><img src="image_7.png" /></figure>

Ermilov S. G. and Frolov A. V.

Licensed under
Creative Commons CC-BY 4.0

<footnote>ªInstitute of Environmental and Agricultural Biology (X-BIO), Tyumen State University, Tyumen, Russia.</footnote>

<footnote>ᵇZoological Institute of Russian Academy of Sciences, Saint Petersburg, Russia.</footnote>
**Material and methods**

**Material** — All mites were collected in one locality by swiping bushes at 1–2 m height with a beat entomological net and extracted from the net with an entomological aspirator and stored in 96% ethanol.

**Observation and documentation** — Specimens were mounted in lactic acid on temporary cavity slides for measurement and illustration. Body length was measured in lateral view, from the tip of the rostrum to the posterior edge of the notogaster. Notogastral width refers to the maximum width of the notogaster in dorsal view. 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. Drawings were made with a camera lucida using a Leica transmission light microscope “Leica DM 2500”.

**Terminology** — Morphological terminology used in this paper follows that of Grandjean: see Travé and Vachon (1975) for references; Norton (1977) for leg setal nomenclature; and Norton and Behan-Pelletier (2009) for overview.

**Abbreviations** — Prodorsum: *lam* = lamella; *cos* = costula; *slam* = sublamella; *tlam* = translamella; *tcos* = transcostula; *tu* = tutorium; *ro*, *le*, *in*, *ex*, *bs* = rostral, lamellar, interlamellar, exobothridial, and bothridial seta, respectively; *Ad* = dorsosejugal porose area; *D* = dorsophragma; *P* = pleurophragma. Notogaster: *c*, *da*, *la*, *dm*, *lm*, *dp*, *lp*, *h*, *p* = setae; *Aa*, *A1*, *A2*, *A3* = porose areas; *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* = pedotectum I and II, respectively; *dis* = discidium; *cir* = circumpedal carina. Anogenital region: *g*, *ag*, *an*, *ad* = genital, aggenital, anal, and adanal seta, respectively; *iad* = adanal lyrifissure; *Ap* = postanal porose area; *p.o.* = preanal organ. Legs: *Tr*, *Fe*, *Ge*, *Ti*, *Ta* = trochanter, femur, genu, tibia, and tarsus, respectively; *p.a.* = porose area; *ε* = famulus; *d*, *l*, *v*, *ev*, *bv*, *ft*, *tc*, *it*, *p*, *u*, *a*, *s*, *pv*, *pl* = setae; *ω*, *σ*, *φ* = solenidia.

**Taxonomy**

**Family Eremellidae**

**Genus *Afreremella* Mahunka, 1973**

Type species: *Afreremella luisiae* Mahunka, 1973

**Generic traits (adult) of *Afreremella***

With character states of Eremellidae (Balogh 1961; Norton and Behan-Pelletier 2009). *Measurements*: Medium-sized (length about 500). *Integument*: Dorsal and ventral sides areolate. *Prodorsum*: Rostrum rounded. Costulae and transcostula of medium length, simple, forming trapezoid structure. Tutorium well developed, illusory forming triangular projection in dorsal aspect. Rostral, lamellar and interlamellar setae medium-sized to long; *ro* setiform, *le* and *in* fusiform or thick; *ro* inserted dorsolaterally on prodorsum, *le* on transcostula, *in* in interbothridial region. Exobothridial seta present, short. Bothridial seta with short stalk and globular or spoon-shaped head. Bothridium cup-shaped. Dorsophragma and pleurophragma present. *Notogaster*: With 10 or 12–13 (if dorsocentral setae *da*, *dm/dp* developed) pairs of notogastral setae; posterior seta *p*₁ short, broadly phylliform; *p*₂ and *p*₃ shortest, dilated distally; others medium-sized or long, fusiform or thick; humeral region with one seta (*c*₂). *Gnathosoma*. Subcapitulum diarthric. Palp setation: 0–2–1–3–9(+ω); solenidion bacilliform, attached to
eupathidium. Axillary saccule absent. Chelicera chelate-dentate. *Epimeral and lateral podosomal regions*: Epimeral setal formula: 3–1–3–3. Pedotectum I represented by medium-sized lamina, pedotectum II represented by small lamina. Discidium developed. *Anogenital region*: Six pairs of genital, one pair of aggenital, two pairs of anal, and three pairs of adanal setae; *ad*₁ and *ad*₂ dilated distally, others setiform; *an*₁ located close to inner margin of anal plate. *Legs*: All legs homotridactylous. Tibia I with dorsodistal apophysis bearing solenidion φ₁. Solenidia ω₁ of tarsus I and ω of tarsus II inserted on tubercles; tarsus II with one solenidion. Porose area present on tarsi I–IV, femora I–IV, trochanters III, IV.

**Subgeneric traits (adult) of *Afreremella* (Afreremella) Mahunka, 1973**

Notogaster with 10 pairs of notogastral setae; dorsocentral part of notogaster without setae (*da*, *dm*, *dp* absent).

**Subgeneric traits (adult) of *Afreremella* (Arboreremella) n. subgen**

Type species: *Afreremella* (*Arboreremella*) *madagascarensis* n. sp.

Notogaster with 12–13 pairs of notogastral setae; dorsocentral part of notogaster with setae (*da*, *dm* and usually also *dp* present).

**Etymology**

The name *Arboreremella* refers to the arboreal place of origin + ‘*eremella*’, a common suffix for generic names in Eremellidae.

**_Afreremella (Arboreremella) madagascarensis n. sp._**

Zoobank: 28C0F62D-8A0C-4FFD-AA96-45A97703D478

(Figures 1–3)

**Diagnosis** — Body length: 547–564. Rostral seta setiform, barbed; lamellar and interlamellar setae thick, heavily barbed; bothridial seta globular, roughened; *in* longest. Notogastral setae (except posterior setae) long, thick, heavily barbed. Epimeral and anogenital setae (except dilated distally, heavily barbed *ad*₁, *ad*₂) setiform, slightly barbed. Adanal lyrifissure diagonal.

**Description of adult** — Measurements – Body length: 547 (holotype, female), 547–564 (three paratype females); body width: 348 (holotype), 348–365 (three paratype females).

Integument – Body light brown to brown. Surface and setae partially covered by thin layer of gel-like cerotegument. Prodorsum, notogaster, subcapitular mentum, epimeral and anogenital regions, genital and anal plates areolate (diameter of areola up to 12); areolae on genital plate indistinct. Lateral part of body partially microgranulate. Antiaxial side of tarsi I–IV and tibiae I, II partially with transverse rugosities; antiaxial side of leg femora I–IV, tibiae I, II and trochanters III, IV with rounded and elongated tubercles.

Prodorsum – Rostrum broadly rounded. Costula about 1/2 length of prodorsum, little not reaching transcostula. Transcostula long, laterally almost reaching tutorium. Tutorium bent mediodistally. Rostral seta (65–73) setiform, barbed, inserted on tubercle; lamellar (69–82) and interlamellar (102–114) setae thick, heavily barbed, inserted on tubercles; exobothridial seta (12–14) setiform, thin, slightly barbed; bothridial seta (32–36) globular, roughened.

Notogaster – Posterior notogastral margin slightly undulate. Thirteen (in holotype and in two paratypes) or twelve (in one paratype; *dp* absent) pairs of notogastral setae; *p*₁ (24–28) broadly phylliform, heavily barbed; *p*₂ and *p*₃ (14–16) dilated distally, heavily barbed; others (77–86) thick, heavily barbed; all setae inserted on tubercles. Opisthonotal gland opening and all lyrifissures (except *ia* not observed) well visible.

Gnathosoma – Subcapitulum size: 123–131 × 98–106. Subcapitular (*a*, *h*: 20–22; *m*: 28–32) and adoral (10) setae setiform, thin, slightly barbed; *m* slightly thicker than others. Palp (length: 73–82) with typical setation: 0–2–1–3–9(+ω). Postpalpal seta (4) spiniform, smooth. Chelicera (length: 135–139) with two setiform, barbed setae (*cha*: 41–45; *chb*: 24–28).
<figure><img src="image_8.png" /><figcaption>**Figure 1** *Aferemella (Arboreremella) madagascarensis* n. sp., adult: a – dorsal view (legs omitted); b – ventral view (gnathosoma and legs omitted); c – lateral view (gnathosoma and legs omitted). Scale bar 120 µm.</figcaption></figure>
<figure><img src="image_9.png" /><figcaption>**Figure 2** *Afreremella (Arboreremella) madagascarensis* n. sp., adult: a – posterior view; b – subcapitulum, ventral view; c – palp, right, antiaxial view; d – chelicera, left, paraxial view. Scale bars 120 µm (a), 25 µm (b–d).</figcaption></figure>

Epimeral and lateral podosomal regions – Epimeral setae (1b: 24; 3b: 20; others: 12) setiform, thin, slightly barbed. Discidium triangular, tubercle-like distally.

Anogenital region – Genital ($g_1$, $g_2$: 10–12; $g_3$–$g_6$: 12–14), aggenital (12–14) and anal (12–14) setae, and adanal seta $ad_3$ (12–14) setiform, thin, slightly barbed. Adanal setae $ad_1$ (12–14) and $ad_2$ (8–12) dilated distally, heavily barbed, inserted on tubercles. Adanal lyrifissure diagonal.

Legs – Median and lateral claws strong, slightly barbed on dorsal side. Proximoventral porose area on tarsi I–IV slightly visible; distoventral porose area on tibiae I–IV not observed; dorsoparaxial porose area on femora I–IV and on trochanters III, IV distinct. Formulas of leg setation and solenidia: I (1–4–3–4–20) [1–2–2], II (1–4–3–4–15) [1–1–1], III (2–2–1–3–15) [1–1–0], IV (1–2–2–3–12) [0–1–0]; homology of setae and solenidia indicated in Table 1. Solenidia $\phi_1$ of tibia I and $\phi$ of tibia II very long, flagellate; $\omega_1$ of tarsus I long, rod-like, coupled with seta *ft"*; $\omega_2$ of tarsus I, $\phi_2$ of tibia I and $\omega$ of tarsus II medium-sized, setiform; $\phi$ of tibiae III, IV and $\sigma$ of genua I–III short, bacilliform, slightly bent mediodistally. Famulus short, erect, slightly swollen distally, inserted between solenidia $\omega_1$ and $\omega_2$. Some setae thick, heavily barbed.

**Material examined** — Holotype (female) and three paratypes (all females): Madagascar, Ankaratra Mt., 19°20'32"S, 47°18'03"E, 1980 m a.s.l., from bushes (swiping), 24.02.2022 (leg. A.V. Frolov).

**Type deposition** — The holotype is deposited in the collection of the Zoological Institute, Russian Academy of Sciences, Saint-Petersburg, Russia; three paratypes are deposited 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.

**Etymology** — The name *madagascarensis* refers to the country of origin, Madagascar.
<figure><img src="image_10.png" /><figcaption>**Figure 3** *Afreremella (Arboreremella) madagascarensis* n. sp., adult: a – leg I, right, antiaxial view; b – leg II, right, antiaxial view; c – leg III, left, antiaxial view; d – leg IV, left, antiaxial view. Scale bar 60 µm.</figcaption></figure>
**Family Oribatulidae**

**Genus *Lucoppia* Berlese, 1908**

Type species: *Notaspis burrowsi* Michael, 1890

**Generic traits (adult) of *Lucoppia***

With character states of Oribatulidae (Balogh and Balogh 1984; Norton and Behan-Pelletier 2009). *Measurements*: Medium-sized to large (length about 450–750). *Integument*: Notogaster reticulate, foveolate or smooth. *Prodorsum*: Rostrum rounded or pointed. Lamella short, simple, without cusp, convergent; translamella short, thin; lamellae and translamella forming trapezoid structure. Sublamella and dorsosejugal porose area present. Prolamella and sublamellar porose area absent. Tutorium long, ridge-like. Rostral, lamellar and interlamellar setae long, setiform; *ro* inserted dorsally on prodorsum, *le* on the end of lamella, *in* in interbothridial region; *le* nearer to *in* than to *ro*. Exobothridial seta present, medium-sized. Bothridial seta short to medium-sized, globular or clavate to fusiform. Bothridium cup-shaped. Dorsophragma small. *Notogaster*: Anterior margin of notogaster distinct, convex medially. Pteromorph and pleural carina completely absent. Octotaxic system with four pairs of porose areas; *Aa* elongate oval, others rounded to oval. With fourteen pairs of long, stiff, barbed notogastral setae; humeral region with one seta (*c*₂). *Gnathosoma*. Subcapitulum diathric. Palp setation: 0–2–1–3–9(+ω); solenidion bacilliform, attached to eupathidium. Axillary saccule absent. Chelicera chelate-dentate. *Epimeral and lateral podosomal regions*: Epimeral setal formula: 3–1–3–3. Apodemes II, III and sejugal apodeme well separated medially. Humeral porose area *Am* present, *Ah* absent. Pedotectum I represented by medium-sized lamina, pedotectum II represented by small lamina. Discidium and circumpedal carina present. *Anogenital region*: Four or five pairs of genital, one pair of aggenital, two pairs of anal, and three pairs of adanal setae. Adanal lyrifissure located in preanal position. Postanal porose area band-like. *Legs*: All legs heterotridactylous. Porose area present on tarsi I–IV, tibiae I–IV, femora I–IV, trochanters III, IV.

**Lucoppia ankaratraensis n. sp.**

Zoobank: AE6B6891-F377-44BB-BA3B-45861BA3DB81

(Figures 4–6)

**Diagnosis** — Body length: 415–481. Notogaster and anogenital region with small foveolae. Rostrum pointed. Rostral, lamellar and interlamellar setae long, setiform, erect, barbed; *ro* shortest and thinnest, *in* longest and thickest; bothridial seta short, globular, roughened. Notogastral setae long, stiff, erect (*c*, *p*₁–*p*₃ shorter than others). Epimeral, genital and aggenital setae short, setiform, thin, slightly barbed; four pairs of genital setae. Anal and adanal setae short, stiff, thin, slightly barbed. Leg tarsus I with 19 setae (*l*'' absent).

**Description of adult** — Measurements – Body length: 415 (holotype, male), 415 (one male paratype); 448–481 (six paratype females); body width: 249 (holotype), 249 (one male

<table><caption>**Table 1** Leg setation and solenidia of adult *Afreremella* (*Arboreremella*) *madagascarensis* n. sp.</caption><thead><tr><th>Leg</th><th><i>Tr</i></th><th><i>Fe</i></th><th><i>Ge</i></th><th><i>Ti</i></th><th><i>Ta</i></th></tr></thead><tbody><tr><td>I</td><td><i>v'</i></td><td><i>d</i>, (<i>l</i>), <i>bv''</i></td><td>(<i>l</i>), <i>v'</i>, <i>σ</i></td><td>(<i>l</i>), (<i>v</i>), <i>φ</i><sub>1</sub>, <i>φ</i><sub>2</sub></td><td>(<i>ft</i>), (<i>tc</i>), (<i>it</i>), (<i>p</i>), (<i>u</i>), (<i>a</i>), <i>s</i>, (<i>pv</i>), <i>v'</i>, (<i>pl</i>), <i>l''</i>, <i>ε</i>, <i>ω</i><sub>1</sub>, <i>ω</i><sub>2</sub></td></tr><tr><td>II</td><td><i>v'</i></td><td><i>d</i>, (<i>l</i>), <i>bv''</i></td><td>(<i>l</i>), <i>v'</i>, <i>σ</i></td><td>(<i>l</i>), (<i>v</i>), <i>φ</i></td><td>(<i>ft</i>), (<i>tc</i>), (<i>it</i>), (<i>p</i>), (<i>u</i>), (<i>a</i>), <i>s</i>, (<i>pv</i>), <i>ω</i></td></tr><tr><td>III</td><td><i>l'</i>, <i>v'</i></td><td><i>d</i>, <i>ev'</i></td><td><i>l'</i>, <i>σ</i></td><td><i>l'</i>, (<i>v</i>), <i>φ</i></td><td>(<i>ft</i>), (<i>tc</i>), (<i>it</i>), (<i>p</i>), (<i>u</i>), (<i>a</i>), <i>s</i>, (<i>pv</i>)</td></tr><tr><td>IV</td><td><i>v'</i></td><td><i>d</i>, <i>ev'</i></td><td><i>d</i>, <i>l'</i></td><td><i>l'</i>, (<i>v</i>), <i>φ</i></td><td><i>ft''</i>, (<i>tc</i>), (<i>p</i>), (<i>u</i>), (<i>a</i>), <i>s</i>, (<i>pv</i>)</td></tr></tbody></table>

Note: Roman letters refer to normal setae (except ε = famulus); Greek letters refer to solenidia. Single quotation mark (') designates seta on the anterior and double quotation mark (") seta on the posterior side of a given leg segment; parentheses refer to a pair of setae.
<figure><img src="image_11.png" /><figcaption>**Figure 4** *Lucoppia ankaratraensis n. sp.*, adult: a – dorsal view (legs omitted); b – ventral view (gnathosoma and legs omitted); c – lateral view (gnathosoma and legs omitted). Scale bar 120 µm.</figcaption></figure>
<figure><img src="image_12.png" /><figcaption>**Figure 5** *Lucoppia ankaratraensis* n. sp., adult: a – posterior view; b – subcapitulum, ventral view; c – palp, left, paraxial view; d – chelicera, left, paraxial view. Scale bars 120 µm (a), 25 µm (b–d).</figcaption></figure>

paratype); 282–298 (six paratype females).

Integument – Body light brown to brown. Surface covered by layer of dense microgranulate cerotegument. Notogaster and anogenital region sparsely foveolate (diameter of foveola up to 4). Region between costulae with indistinct transverse ridges. Antiaxial side of tarsi I–IV and paraxial side of tarsi I–IV, tibiae I–IV and femora I–IV partially with transverse rugosities.

Prodorsum – Rostrum with triangular projection, pointed. Lamella about 1/4 length of prodorsum, connected by thin, straight or slightly convex medially translamella. Tutorium well developed. Rostral seta (73–82) setiform, barbed, inserted on tubercle; lamellar (98–106) and interlamellar (123–131) setae setiform, erect, barbed, inserted on tubercles; *in* slightly thicker than *le*, *le* slightly thicker than *ro*; exobothridial seta (28–36) setiform, thin, slightly barbed; bothridial seta (28–32) globular, roughened. Dorsosejugal porose area oval (12 × 4), located posterior to *in*.

Notogaster – Anterior margin distinctly convex medially. Fourteen pairs of stiff, erect notogastral setae (c: 41–49; $p_1$: 49–65; $p_2$: 45–49; $p_3$: 41–45; others: 69–77). Porose area *Aa* elongate oval (24–26 × 8–10); *Aa* oval (14–16 × 8–10); *A1* and *A2* rounded (10–16). Opisthonotal gland opening and all lysfissures well visible.

Gnathosoma – Subcapitulum size: 123–131 × 98–102. Subcapitular (*a*: 20; *m*, *h*: 28–32) and adoral (10) setae setiform, thin, slightly barbed. Palp (length: 77–86) with typical setation: 0–2–1–3–9(+ω). Postpalpal seta (8) spiniform, roughened. Chelicera (length: 123–131) with two setiform, barbed setae (*cha*: 36–38; *chb*: 24–26).

Epimeral and lateral podosomal regions – Epimeral setae (*1a*, *2a*, *3a*: 16–20; others: 28–32)
<figure><img src="image_13.png" /><figcaption>**Figure 6** *Lucoppia ankaratraensis* n. sp., adult: a – leg I, right, antiaxial view; b – leg II, right, antiaxial view; c – leg III, left, antiaxial view; d – leg IV, left, antiaxial view. Scale bar 60 µm.</figcaption></figure>
setiform, thin, slightly barbed; *Ic*, *3c* and *4c* slightly thicker than others. Discidium triangular, tubercle-like distally. Circumpedal carina distinct.

Anogenital region – Four pairs of genital setae. Genital (16–20) and aggenital (28–32) setae setiform, thin, slightly barbed; anal (16–20) and adanal (16–24) setae stiff, thin, slightly barbed. Adanal lyrifissure diagonal. Postanal porose area distinct.

Legs – Median claw clearly thicker than lateral claws; all slightly barbed on dorsal side; 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 distinct. Formulas of leg setation and solenidia: I (1–4–3–4–19) [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. Solenidia $\phi_1$ of tibia I and $\phi$ of tibiae II–IV very long, setiform; $\phi$ of tibia IV swollen distally; $\sigma$ of genu III short, bacilliform, slightly bent mediodistally; others medium-sized, rod-like. Famulus short, erect, slightly swollen distally, inserted between solenidion $\omega_2$ and seta ft".

**Material examined** — Holotype (male) and seven paratypes (one male and six females): Madagascar, Ankaratra Mt., 19°20'32"S, 47°18'03"E, 1980 m a.s.l., from bushes (swiping), 24.02.2022 (leg. A.V. Frolov).

**Type deposition** — The holotype is deposited in the collection of the Zoological Institute, Russian Academy of Sciences, Saint-Petersburg, Russia; seven paratypes are deposited 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.

**Etymology** — The name *ankaratraensis* refers to the place of origin, Ankaratra Mt.

**Remarks** — Distinctive characters of the new species from the other *Lucoppia* species can be found in the identification key below.

## Key to known species of *Lucoppia*

1. Rostrum pointed; bothridial seta short, globular; notogaster and anogenital region foveolate; four pairs of genital setae; body length: 415–481
*Lucoppia ankaratraensis n. sp.* Distribution: Madagascar.

— Rostrum rounded; bothridial seta medium-sized, clavate to fusiform; notogaster and anogenital region not foveolate; five pairs of genital setae.
2

2. Notogaster with reticulate pattern; body length: 714
*Lucoppia feideri* Subías, 2010 (=Romanobates reticulatus Feider, Vasiliu and Călugăr, 1970). Distribution: eastern Mediterranean, Chile.

— Notogaster without reticulate pattern; body length: 620–750
*Lucoppia burrowsi* (Michael, 1890) (see also e.g. Weigmann 2006; Seniczak and Seniczak 2012). Distribution: Holarctic region, southeastern China, Hawaii, Mexico.

<table><caption>**Table 2** Leg setation and solenidia of adult *Lucoppia ankaratraensis n. sp.*</caption><thead><tr><th>Leg</th><th><i>Tr</i></th><th><i>Fe</i></th><th><i>Ge</i></th><th><i>Ti</i></th><th><i>Ta</i></th></tr></thead><tbody><tr><td>I</td><td><i>v'</i></td><td><i>d, l', bv'', v''</i></td><td><i>(l), v', σ</i></td><td><i>(l), (v), φ<sub>1</sub>, φ<sub>2</sub></i></td><td><i>(ft), (tc), (it), (p), (u), (a), s, (pv), v', (pl), ε, ω<sub>1</sub>, ω<sub>2</sub></i></td></tr><tr><td>II</td><td><i>v'</i></td><td><i>d, (l), bv'', v''</i></td><td><i>(l), σ</i></td><td><i>(l), (v), φ</i></td><td><i>(ft), (tc), (it), (p), (u), (a), s, (pv), ω<sub>1</sub>, ω<sub>2</sub></i></td></tr><tr><td>III</td><td><i>l', v'</i></td><td><i>d, l', ev'</i></td><td><i>l', σ</i></td><td><i>l', (v), φ</i></td><td><i>(ft), (tc), (it), (p), (u), (a), s, (pv)</i></td></tr><tr><td>IV</td><td><i>v'</i></td><td><i>d, ev'</i></td><td><i>d, l'</i></td><td><i>l', (v), φ</i></td><td><i>ft'', (tc), (p), (u), (a), s, (pv)</i></td></tr></tbody></table>

Note: see Table 1 for explanations.
Discussion

Number and localization of notogastral setae have importance for supporting of the supraspecies status in Eremellidae. For example, Balogh and Mahunka (1974) described the eremellid subgenus *Eremella* (*Archeremella*) Balogh and Mahunka, 1974 which differs from the nominate subgenus by the presence of 14 pairs (versus 10 pairs) of notogastral setae and the localization of setae *da, dm, dp* in the dorsocentral part of the notogaster (versus *da, dm, dp* absent; dorsocentral notogastral part without setae). Later, Pérez-Íñigo (1981) suggested to raise the status of *Archeremella* to the generic level, which was supported further by other authors (e.g., Fujikawa 1991; Subías 2004; Mahunka 2008; Ermilov and Frolov 2019). Thus, following the analogy, *Afreremella* (*Arboreremella*) *madagascarensis* n. sp. could also theoretically belong to the new genus *Arboreremella* which differs from the related genus *Afreremella* by the presence of 12–13 pairs (versus 10 pairs) of notogastral setae including *da, dm* and usually also *dp* in dorsocentral notogastral part (versus *da, dm, dp* absent). However, the type species of *Afreremella* (*A. luisiae*) and *A. madagascarensis* are morphologically very similar in other important characters (e.g., body ornamentation; structure of costular-transcostular complex and tutorium; morphological differentiation of notogastral and adanal setae and their lengths), therefore, we have no reason to allocate a new genus, but, in our opinion, the extra number of notogastral setae and the presence of *da, dm, dp* in the dorsocentral notogastral part are sufficient to support a new subgenus.

Based on above listed explanations, we believe that the generic status of *Archeremella* in the current situation is not justified, therefore, we propose to maintain this taxon as subgenus within *Eremella*, as suggested by Balogh and Mahunka (1974) initially. Hence, we support the initial systematic placement of *Eremella* (*Archeremella*) leowae Balogh and Mahunka, 1974 and propose recombinations of two other known species from *Archeremella* to *Eremella* (*Archeremella*): *E. (A.) africana* (Pérez-Íñigo, 1981) **comb. nov.**; and *E. (A.) bartlae* (Mahunka, 2008) **comb. nov.**

Other records

Family Cymbaeremaeidae

*Scapheremaeus anteriorugosus* Mahunka, 2011: 3 ex. Distribution: Madagascar.

Family Humerobatidae

*Humerobates africanus* (Mahunka, 1984): 29 ex. Distribution: Afrotropical region.

Acknowledgements

The authors thank two anonymous reviewers for valuable comments. The study was performed in the framework of the Russian State research project 122031100272-3. Collecting in Madagascar was conducted under a permit No. 443/21/MEDD/SG/DGGE/DAPRNE/SCBE.Re; samples were exported to Russia under a permit No. 036N-EA02/MG21.

ORCID

Sergey G. Ermilov https://orcid.org/0000-0002-0913-131X

Andrey V. Frolov https://orcid.org/0000-0002-6724-6828
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