# New faunistic data on oribatid mites (Acari, Oribatida) from western, southern and southeastern Ethiopia, with description of two new species of the family Carabodidae

Sergey G Ermilov, Leonid B Rybalov

► To cite this version:

Sergey G Ermilov, Leonid B Rybalov. New faunistic data on oribatid mites (Acari, Oribatida) from western, southern and southeastern Ethiopia, with description of two new species of the family Carabodidae. Acarologia, 2025, 65 (2), pp.559-574. 10.24349/ikjq-9q2d. hal-05088066

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# New faunistic data on oribatid mites
(Acari, Oribatida) from western,
southern and southeastern Ethiopia,
with description of two new species of
the family Carabodidae

Sergey G. Ermilov IDª, Leonid B. Rybalov IDᵇ

## Original research

### ABSTRACT

The present study is based on oribatid mite materials collected from two regions in Ethiopia: Oromia and South Ethiopia Regional State. A list of 70 species belonging to 54 genera and 33 families is presented. Two new species of the family Carabodidae— *Carabodes* (*Klapperiches*) *paradilatatus* Ermilov **n. sp.** and *Congocepheus setiformis* Ermilov **n. sp.**—are described from litter and mineral soil in a mixed forest.

**Keywords** fauna; new record; taxonomy; *Carabodes* (*Klapperiches*); *Congocepheus*; morphology; Afrotropical region

**Zoobank** http://zoobank.org/BD53878B-EA2E-4B80-9806-CCB67A658060

### Introduction

Oribatid mites (Acari, Oribatida) of Ethiopia have been actively studied in recent years (e.g, Ermilov and Rybalov 2024; Ermilov *et al.* 2024). The present study is based on materials collected from the Oromia region (western and southeastern Ethiopia) and South Ethiopia Regional State (southern Ethiopia) during 2015–2018. The main goals of the paper are: to present a list of the identified taxa and to describe two new carabodid species (Carabodidae) belonging to *Carabodes* (*Klapperiches*) Mahunka, 1978 and *Congocepheus* Balogh, 1958.

*Carabodes* (*Klapperiches*) was proposed by Mahunka (1978) as an independent genus, with *Klapperiches nigrosetosus* Mahunka, 1978 as type species. Its main subgeneric traits can be found in Mahunka (1978, 1986), Subías and Shtanchaeva (2023). Presently, the subgenus comprises about 50 species, which collectively have a cosmopolitan distribution (Subías 2022, online version 2024; including personal data of the first author). An identification key to the known species of *Carabodes* (*Klapperiches*) from the Afrotropical region has been presented by Ermilov and Kontschán (2022).

*Congocepheus* was proposed by Balogh (1958), with *Congocepheus heterotrichus* Balogh, 1958 as type species. Its main subgeneric traits can be found in Mahunka (1986), Fernandez *et al.* (2014), Subías and Shtanchaeva (2023). Presently, the genus comprises 13 species, which collectively distributed in the Paleotropical region (Subías 2022, online version 2024). Within the genus there are groups of species with distinctive morphological characters in the presence/absence of the specific notogastral ridges as well as the presence/absence, size and deep of the depressions in anterior and posterior parts of the notogaster, so the genus seems heterogeneous, and additional phylogenetic and molecular investigations are necessary.

Received 09 February 2025

Accepted 23 May 2025

Published 27 May 2025

Corresponding author

Sergey G. Ermilov ID: 
ermilovacari@yandex.ru

Academic editor

Baumann, Julia

https://doi.org/10.24349/ikjq-9q2d

ISSN 0044-586X (print)

ISSN 2107-7207 (electronic)

<figure><img src="image_2.png" /></figure>

Ermilov S. G. and Rybalov L. B.

Licensed under
Creative Commons CC-BY 4.0

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

<footnote>ᵇInstitute of Ecology and Evolution, Russian Academy of Sciences, Laboratory of Soil Zoology and General Entomology, Moscow, Russia.</footnote>
Prior to this study, two species of *Carabodes* (*Klapperiches*) and three species of *Congocepheus* have been registered in Ethiopia (Ermilov *et al.* 2010, 2012a, b; Ermilov and Rybalov 2024a, b): *C. (K.) dilatatus* Ermilov, Sidorchuk and Rybalov, 2010; *C. (K.) pocsi* Mahunka, 1983 (in 1983b); *Co. ornatus* Mahunka, 1983 (in 1983a); *Co. rwandensis* Fernández, Theron and Leiva, 2016; *Co. taurus* Balogh, 1961.

**Materials and methods**

**Sampling** — Substrate samples containing oribatid mites were collected using a stainless-steel frame (50×50 cm) with a sieve (mesh size 2×2 cm) (Fig. 1c) in the following locations (Figs 1a–d):

# 12–13. Western Ethiopia, Oromia region, West Wollega Zone, Dhati Walal National Park, 9°23′54.7″N, 34°40′10.7″E, mountains, 1636 m a.s.l., litter (#12) and green mosses on the tree (#13) in forest with *Ficus* sp., *Schefflera abyssinica* and *Coffea arabica* on mollic Nitisols, 02.XI.2017, end of wet season (L.B. Rybalov).

# 14. Western Ethiopia, Oromia region, West Wollega Zone, near Dhati Walal National Park, 9°25′01.2″N, 34°45′21.3″E, mountains, 1761 m a.s.l., litter in a mixed forest on brown eutrophic soil, 02.XI.2017, end of wet season (L.B. Rybalov).

# 15. Southeastern Ethiopia, Oromia Region, Bale Zone, 6°35′24.0″N, 39°43′29.5″E, 1790 m a.s.l., Bale Mountains National Park, southern macroslope, Harenna Forest, broadleaved forest (first layer of forest: *Syzygium guineense* dominates, singly *Polyscias fulva*; second layer of forest: *Teclea nobilis*, *Galiniera saxifraga*, *Pandanus* sp.; among the shrubs the most common is *Maytenus* sp.), litter, 31.X.2015, beginning of dry season (O.G. Gorbunov).

# 16–17 Southern Ethiopia, South Ethiopia Regional State, Gamo Zone, 6°01′28.7″N, 37°35′49.7″E, 1179 m a.s.l., 5 km East from the town Arba Minch, 20–40 m from the lake Abai, litter (#16) and mineral soil (0–5 cm) (#17) in a mixed deciduous forest on alluvial organic soil, wet place, around water spots, 8.XII.2018, dry season (L.B. Rybalov).

# 18–19. Southern Ethiopia, South Ethiopia Regional State, Gamo Zone, 6°01′36.7″N, 37°35′42.7″E, 1170 m a.s.l., 5 km East from the town Arba Minch, 1 km from the lake Abai, litter (#18) and mineral soil (0–5 cm) (#19) in a mixed forest on the terrace (first layer of forest: *Celtis* sp., *Ficus sycomorus*, *Schefflera* sp., *Acacia* sp.; second layer of forest: *Lemon* sp., *Coffea arabica*) on brown (black-brown) soil with light-medium loam soil texture, 7.XII.2018, dry season (L.B. Rybalov).

Mites were extracted into 75% ethanol using Berlese's funnels with electric lamps in laboratory conditions.

**Observation and documentation** — For measurement and illustration, specimens were mounted in lactic acid on temporary cavity slides. Some specimens preliminarily were placed for one day in a 5% potassium hydroxide solution to lighten the body cuticle, dissolve cerotegument, and release adherent debris. 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; 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 of tarsus I 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 is mostly that of F. Grandjean (references in Travé and Vachon 1975). Fundamentals of leg setation were reviewed by Norton (1977).

**Abbreviations** — *Prodorsum*: *lam* = lamella; *tlam* = translamella; *hs* = hump-like structure; *dep* = depression; *ro*, *le*, *in*, *bs* = rostral, lamellar, interlamellar, and bothridial setae, respectively; *tu* = tutorium; *lpr* = lateral prodorsal ridge. *Notogaster*: *c*, *da*, *la*, *dm*, *lm*, *dp*, *lp*, *h*₁–*h*₃, *p*₁–*p*₃ = setae; *ia*, *im*, *ip*, *ih*, *ips* = lyrifissures; *gla* = opisthonotal gland opening. *Gnathosoma*: *a*, *m*, *h* = subcapitular setae; *d*, *l*, *sup*, *inf*, *acm*, *ul*, *su*, *lt*, *vt* = palp setae; *ω* =
<figure><img src="image_3.png" /><figcaption>**Figure 1** Photos of some sampling locations: a -location 14; b -location 15; c -sifting litter by Dr. Leonid B. Rybalov in location 16-17; d -location 18-19.</figcaption></figure>
solenidion; *cha*, *chb* = cheliceral setae; *Tg* = Trägårdh's organ. *Epimeral and lateral podosomal regions: 1a-1c, 2a, 3a-3c, 4a-4c* = epimeral setae; *PdI, PdII* = pedotecta I, II, respectively; *dis* = discidium. *Anogenital region: g, ag, an, ad* = genital, agenital, anal, and adanal setae, respectively; *iad* = adanal lyrifissure; *po* = preanal organ. *Legs: ω, φ, σ* = solenidia; *e* = famulus; *d, l, v, bv, ev, ft, tc, it, p, u, a, s, pv* = setae; *pa* = porose area.

**Notes** — References to the original descriptions of taxa in the List of identified taxa section are not included in the References section. Paired structures are described in the singular, unless otherwise noted.

## List of identified taxa

Distribution: mostly from Subías (2022, online version 2024). Unidentified species are not included. The number of the collected specimens is in parentheses.

The list includes 70 species belonging to 54 genera and 33 families. Eight species (*Annectacarus insculptus*, *Neoliodes terrestris*, *Zetorchestes ornatus*, *Oxyoppia (Oxyoppiella) bituberculata*, *Eupelops forsslundi*, *Protoribates ziemowiti*, *Transoribates agricola*, *Benoibates rugosus*), five genera (*Annectacarus*, *Zetorchestes*, *Transoribates*, *Benoibates*, *Punctoribates*) and two families (*Zetorchestidae*, *Oripodidae*) are reported in Ethiopia for the first time. One species (*Punctoribates zachvatkini*) is reported for the first time in the Afrotropical region. Seventeen species are known only from Ethiopia, 27 species are known from the Afrotropical region and 26 species (including six cosmopolitan/semicosmopolitan species) have broad distributions (more than one geographical region).

### Lohmanniidae

*Annectacarus insculptus* Wallwork, 1962: location 19 (3 exs.). Distribution: Ghana. New record of the genus and species in Ethiopia.

### Cosmochthoniidae

*Cosmochthonius lanatus* (Michael, 1885): location 15 (1 ex.). Distribution: Semicos-mopolitan.

### Sphaerochthoniidae

*Sphaerochthonius splendidus* (Berlese, 1904): locations 17 (5 exs.), 18 (1 ex.). Distribution: Tropical and Subtropical regions.

### Epilohmanniidae

*Epilohmannia pallida* Wallwork, 1962: locations 18 (1 ex.), 19 (4 exs.). Distribution: Tropical and Subtropical regions.

### Nanhermanniidae

*Masthermannia mammillaris* (Berlese, 1904): location 12 (1 ex.). Distribution: Tropical and Subtropical regions.

### Nothridae

*Nothrus crassisetus* Mahunka, 1982: locations 18 (1 ex.), 19 (3 exs.). Distribution: Afrotropical region.

### Malaconothridae

*Malaconothrus ensifer* Mahunka, 1982: locations 12 (1 ex.), 18 (1 ex.), 19 (3 exs.). Distribution: Afrotropical region.

### Hermanniellidae

*Hermanniella dubiosa* Mahunka & Mahunka-Papp, 2007: locations 16 (9 exs.), 17 (8 exs.), 18 (2 exs.), 19 (7 exs.). Distribution: Afrotropical region.

### Plasmobatidae

*Plasmobates foveolatus* Ermilov, Sidorchuk & Rybalov, 2010: location 12 (1 ex.). Distribution: Afrotropical region.

### Neoliiodidae

*Neoliodes terrestris* (Wallwork, 1963): locations 15 (1 ex.), 18 (1 ex.). Distribution: Tropical and southern Palaearctic regions. New record of the species in Ethiopia.
**Aleurodamaeidae**

*Aleurodamaeus recenfesevpi* Ermilov and Rybalov, 2012: location 15 (3 exs.). Distribution: Ethiopia.

**Damaeidae**

*Metabelba (Pateribelba) glabriseta* Mahunka, 1982: locations 16 (5 exs.), 18 (1 ex.), 19 (1 ex.). Distribution: Afrotropical region.

**Microzetidae**

*Berlesezetes ornatissimus* (Berlese, 1913): locations 15 (4 exs.), 18 (4 exs.), 19 (4 exs.). Distribution: Tropical and Subtropical regions.

*Microzetes (Megazetes) eckeri* (Mahunka, 1982): location 16 (1 ex.). Distribution: Ethiopia.

**Basilobelbidae**

*Basilobelba gigantea* Ermilov, Sidorchuk & Rybalov, 2011: locations 12 (1 ex.), 14 (1 ex.), 18 (5 exs.), 19 (4 exs.). Distribution: Ethiopia.

**Damaeolidae**

*Fosseremus laciniatus* (Berlese, 1905): locations 12 (1 ex.), 19 (7 exs.). Distribution: Cosmopolitan.

**Eremobelbidae**

*Eremobelba tuberculata* Mahunka, 1982: location 14 (6 exs.). Distribution: Ethiopia.

**Eremulidae**

*Eremulus flagellifer* Berlese, 1908: locations 15 (1 ex.), 19 (4 exs.). Distribution: Cosmopolitan.

**Zetorchestidae**

*Zetorchestes ornatus* Mahunka, 1985: locations 16 (5 exs.), 18 (4 exs.). Distribution: Afrotropical region. New record of the family, genus and species in Ethiopia.

**Carabodidae**

*Austrocarabodes (Austrocarabodes) glabrus* Mahunka, 1982: location 18 (12 exs.). Distribution: Ethiopia.

*Austrocarabodes (Austrocarabodes) heterosetosus* Ermilov, Sidorchuk and Rybalov, 2010: location 16 (3 exs.). Distribution: Afrotropical region.

*Austrocarabodes (Uluguroides) arboreus* Ermilov, Sidorchuk and Rybalov, 2010: location 15 (3 exs.). Distribution: Ethiopia.

*Carabodes (Klapperiches) paradilatatus* Ermilov **n. sp.**: locations 18 (4 exs.), 19 (2 exs.). Distribution: Ethiopia.

*Congocepheus setiformis* Ermilov **n. sp.**: location 18 (7 exs.). Distribution: Ethiopia.

**Dampfiellidae**

*Beckiella opposita* Mahunka, 1982: location 12 (1 ex.). Distribution: Ethiopia.

**Machadobelbidae**

*Machadobelba shtanchaevae* Ermilov, Sidorchuk & Rybalov, 2010: location 12 (5 exs.). Distribution: Ethiopia.

**Oppiidae**

*Antennoppia capilligera* (Berlese, 1916): locations 16 (1 ex.), 17 (4 exs.), 19 (3 exs.). Distribution: Afrotropical region.

*Afroppia brevipila* (Mahunka, 1982): location 12 (7 exs.). Distribution: Ethiopia.

*Arcoppia rugosa* (Mahunka, 1974): location 14 (11 exs.). Distribution: Afrotropical region, Hawaii.

*Neoamerioppia polygonata* (Mahunka, 1982): locations 12 (21 exs.), 13 (13 exs.), 14 (16 exs.), 16 (3 exs.). Distribution: Afrotropical region.

*Oppia kuehnelti* Csiszár, 1961: location 18 (3 exs.). Distribution: Tropical and southern Palaearctic regions.

*Oxyoppia (Oxyoppiella) bituberculata* (Balogh, 1958): location 19 (2 exs.). Distribution: Afrotropical and Sub-Antarctic regions. New record of the species in Ethiopia.

*Pseudoamerioppia barrancensis* (Hammer, 1961): location 15 (1 ex.). Distribution: Neotropical, Oriental and Afrotropical regions, Canary Is.
*Ramusella* (*Insculptoppia*) *seniczakae* Hugo-Coetzee, 2016: locations 15 (6 exs.), 17 (1 ex.), 19 (10 exs.). Distribution: South Africa.

**Suctobelbidae**

*Suctobelbella* (*Flagrosuctobelba*) *kontschani* (Mahunka & Mahunka-Papp, 2007): locations 12 (2 exs.), 17 (1 ex.), 19 (3 exs.). Distribution: Afrotropical region.

*Suctobelbella* (*Ussuribata*) *spirochaeta* Mahunka, 1983: locations 12 (1 ex.), 15 (1 ex.). Distribution: Afrotropical region.

**Tectocepheidae**

*Tectocepheus sarekensis* Trägårdh, 1910: locations 14 (5 exs.), 17 (24 exs.), 19 (11 exs.). Distribution: Cosmopolitan.

*Tegeozetes tunicatus* Berlese, 1913: location 16 (1 ex.). Distribution: Tropical region, Hungary.

**Phenopelopidae**

*Eupelops forsslundi* (Balogh, 1959): location 15 (9 exs.). Distribution: Afrotropical region, Vietnam. New record of the species in Ethiopia.

**Caloppiidae**

*Zetorchella arsiensis* Ermilov, 2023: location 16 (1 ex.). Distribution: Ethiopia.

*Zetorchella nortoni* Ermilov, Sidorchuk & Rybalov, 2010: location 15 (2 exs.). Distribution: Ethiopia.

**Haplozetidae**

*Pilobatella dhatiensis* Ermilov, 2019: location 14 (3 exs.). Distribution: Ethiopia.

*Protoribates aethiopicus* Ermilov & Rybalov, 2013: locations 16 (4 exs.), 17 (1 ex.), 18 (5 exs.), 19 (8 exs.). Distribution: Afrotropical region.

*Protoribates paracapucinus* (Mahunka, 1988): locations 18 (1 ex.), 19 (7 exs.). Distribution: Tropical and Subtropical regions.

*Protoribates ziemowiti* Ermilov & Starý, 2020: locations 16 (4 exs.), 17 (2 exs.), 18 (7 exs.), 19 (4 exs.). Distribution: Madagascar. New record of the species in Ethiopia.

*Transoribates agricola* (Nakamura & Aoki, 1989): location 16 (7 exs.). Distribution: eastern Palaearctic region, Vietnam, South Africa. New record of the genus and species in Ethiopia.

**Mochlozetidae**

*Unguizetes atypicus* (Mahunka, 1982): location 12 (1 ex.). Distribution: Afrotropical region.

**Oripodidae**

*Benoibates rugosus* Mahunka, 2001: location 16 (3 exs.). Distribution: Kenya. New record of the family, genus and species in Ethiopia.

**Scheloribatidae**

*Muliercula walalensis* Ermilov, 2019: location 16 (4 exs.). Distribution: Ethiopia.

*Scheloribates aethiopicus* Mahunka, 1982: locations 12 (5 exs.), 14 (47 exs.). Distribution: Afrotropical region, Canary Islands.

*Scheloribates fimbriatus* Thor, 1930: location 14 (1 ex.). Distribution: Pantropical and Subtropical regions.

*Scheloribates leleupi* Balogh, 1959: location 12 (50 exs.). Distribution: Afrotropical region.

*Scheloribates pallidulus* (Koch, 1841): location 13 (3 exs.). Distribution: Cosmopolitan.

*Scheloribates perisi* Pérez-Íñigo, 1982: location 14 (21 exs.). Distribution: Afrotropical region.

*Scheloribates praeincisus* (Berlese, 1910): locations 12 (1 ex.), 16 (10 exs.). Distribution: Tropical and Subtropical regions.

*Scheloribates* (*Bischeloribates*) *munesaensis* Ermilov, 2016: location 15 (4 exs.). Distribution: Afrotropical region.

*Scheloribates* (*Perscheloribates*) *paratranslamellatus* (Ermilov & Rybalov, 2014): location 15 (1 ex.). Distribution: Ethiopia.
**Punctoribatidae**

*Afroleius valerieae* Coetzee, 2014: location 18 (16 exs.). Distribution: Afrotropical region.

*Lamellobates molecula* (Berlese, 1916): locations 16 (6 exs.), 17 (5 exs.), 18 (9 exs.). Distribution: Tropical and Subtropical regions.

*Punctoribates zachvatkini* Shaldybina, 1969: location 18 (2 exs.). Distribution: southern Palaearctic region. New record of the species in the Afrotropical region. New record of the genus in Ethiopia.

**Galumnidae**

*Allogalumna machadoi* (Balogh, 1960): location 12 (1 ex.). Distribution: Paleotropical region.

*Galumna flabellifera* Hammer, 1958: locations 12 (1 ex.), 16 (45 exs.), 17 (17 exs.), 18 (15 exs.), 19 (12 exs.). Distribution: Tropical and Subtropical regions.

*Galumna incisa* Mahunka, 1982: location 12 (7 exs.). Distribution: Afrotropical region.

*Pergalumna pocsi* Mahunka, 1984: locations 16 (11 exs.), 17 (10 exs.), 18 (1 ex.). Distribution: Afrotropical region.

*Pilizetes anufrievi* Ermilov, Sidorchuk & Rybalov, 2010: location 12 (1 ex.). Distribution: Ethiopia.

*Trichogalumna africana* Ermilov, Sidorchuk & Rybalov, 2011: locations 12 (8 exs.), 13 (12 exs.), 14 (33 exs.). Distribution: Afrotropical region.

*Trichogalumna nipponica* (Aoki, 1966): location 15 (18 exs.). Distribution: Semicommopolitan.

**Galumnellidae**

*Galumnella apiculata* Mahunka, 1992: location 12 (1 ex.). Distribution: Afrotropical region.

*Galumnella subareolata* Mahunka, 1969: location 15 (7 exs.). Distribution: Afrotropical region.

*Galumnopsis ruginervis* Balogh, 1962: location 15 (1 ex.). Distribution: Afrotropical region.

## Descriptions

**Carabodes (Klapperiches) paradilatatus** Ermilov *n. sp.*

Zoobank: 1FEEB9AE-E513-459C-9B27-D2CFB0F9C746

(Figures 2, 3)

**Type material** — Holotype (female) and three paratypes (three females): Southern Ethiopia, South Ethiopia Regional State, Gamo Zone, 6°01'36.7"N, 37°35'42.7"E, 1170 m a.s.l., 5 km East from the town Arba Minch, 1 km from the lake Abai, litter in a mixed forest on the terrace (first layer of forest: *Celtis* sp., *Ficus sycomorus*, *Schefflera* sp., *Acacia* sp.; second layer of forest: *Lemon* sp., *Coffea arabica*) on brown (black-brown) soil with light-medium loam soil texture, 7.XII.2018, dry season (L.B. Rybalov). Two paratypes (two females): same as the holotype, but mineral soil (0–5 cm).

The holotype is deposited in the collection of the Senckenberg Museum of Natural History, Görlitz, Germany; five 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: 300–345. Prodorsum between lamellae with unevenly spaced tubercles, frequently fused and forming short tuberculate ridges. Lamella triangular distally; translamella absent. Rostral and lamellar setae short, setiform, slightly barbed; interlamellar seta comparatively short, phylliform, dilated mediodistally, barbed; bothridial seta long, fusiform, with barbed head. Notogaster with unevenly spaced tubercles (dorsocentral part with some areas without tubercles). All notogastral setae comparatively short (posterior setae shorter than others), phylliform, dilated mediodistally, barbed. Subcapitular mentum and epimal
<figure><img src="image_4.png" /><figcaption>**Figure 2** *Carabodes (Klapperiches) paradilatatus* Ermilov *n. sp*., adult: a - dorsal view; b - anterior part of prodorsum, anterior view; c - ventral view (gnathosoma and legs omitted); d - right lateral view (legs omitted); e - posterior view (part of left half omitted). Scale bar 50 µm.</figcaption></figure>
<figure><img src="image_5.png" /><figcaption>**Figure 3** *Carabodes (Klapperiches) paradilatatus* Ermilov *n. sp*., adult: a - subcapitulum, ventral view; b - palp, left, paraxial view; c - chelicera, left, paraxial view; d - leg I, right, antiaxial view; e - leg II (tarsus omitted), right, antiaxial view; f - leg III (tarsus omitted), left, antiaxial view; g - leg IV, left, antiaxial view. Scale bars 20 µm (a, c-g), 10 µm (b).</figcaption></figure>
region without sculpture; epimere I with one pair of setae; all epimeral setae short, simple. Anogenital region with slightly developed unevenly spaced tuberculate ridges, simple ridges and some small depressions. Genital, aggenital and anal setae short, simple; all adanal setae short, slightly dilated and barbed mediodistally. Leg tarsus IV with 12 setae.

**Description** — *Measurements*. Body length: 345 (holotype), 300–345 (female paratypes); notogaster width: 180 (holotype), 165–180 (female paratypes). Body ratio (length/width): ≈ 1.8–1.9.

*Integument* (Figs 2(a–e), 3(e, d)) — Body color light brown. Body covered by thin layer of gel-like cerotegument. Prodorsum between lamellae with unevenly spaced tubercles (diameter up to 7), frequently fused and forming short tuberculate ridges; lateral part of prodorsum partially with foveolae (diameter up to 5); notogaster with unevenly spaced tubercles (diameter up to 7; dorsocentral part with some areas without tubercles); anogenital region with slightly developed unevenly spaced tuberculate ridges, simple ridges and some depressions; subcapitular mentum, epimeral region, genital and anal plates without sculpture; antiaxial side of leg femora I, II partially with large, poorly developed foveolae.

*Prodorsum* (Figs 2(a, b, d)) — Rostrum broadly rounded. Lamella long and broad, triangular distally; translamella absent; tutrorium short, ridge-like; lateral part of prodorsum with slight ridge (*lpr*) directed to end of lamella and bearing indistinct teeth. Rostral (13–15) and lamellar (17–19) setae setiform, slightly barbed; *ro* inserted dorsally on rostrum; *le* inserted dorsomedially in distal part of lamella; interlamellar seta (26–30) phylliform, dilated mediodistally, barbed; bothridial seta (45–55) fusiform, with long, roughened stalk and short, barbed head; exobothridial seta not observed. Bothridium interrupted ventrally.

*Notogaster* (Figs 2(a, d, e)) — Anterior notogastral margin slightly convex medially. Humeral shoulder small, visible in dorsolateral and lateral aspects. Ten pairs of notogastral setae (*p*₁, *p*₂, *p*₃, *h*₃: 13–15; others: 22–30) phylliform, dilated mediodistally, barbed. Opisthonotal gland opening and lyrifissures *ia*, *im* visible; lyrifissures *ip*, *ih* and *ips* not observed.

*Gnathosoma* (Figs 3(a–c)) — Subcapitulum size: 75–77 × 60–64; subcapitular seta *a* (11) setiform, roughened; *m* (9–11) and *h* (7) setiform, nearly smooth. Palp length: 37; setation: 0-2-1-2-7(+ω); setae *acm* and *v*' on tarsus and *l*'' on tibia absent; postpalpal seta (4) spiniform, smooth. Chelicera length: 82–86; seta *cha* (22–26) setiform, barbed; seta *chb* (11–13) setiform, roughened.

*Epimeral and lateral podosomal regions* (Figs 2(c, d)) — Epimeral setation: 1-1-3-3; all setae (7) setiform, nearly smooth. Discidium tubercle-like.

*Anogenital region* (Figs 2(c–e)) — Anogenital formula: 4-1-2-3; genital (7), aggenital (11) and anal (11) setae setiform, nearly smooth; all adanal setae (11) slightly dilated and barbed mediodistally. Adanal lyrifissure not observed.

*Legs* (Figs 3(d–g)) — Claw of each leg strong, slightly roughened dorsally, with ventrobasal indistinct microtubercle. Dorsoparaxial porose area on femora I–IV elongate oval. Formulas of leg setation and solenidia: I (1-4-3-4-16) [1-2-2], II (1-4-3-2-15) [1-1-2], III (2-3-1-2-15) [1-1-0], IV (1-2-2-2-12) [0-1-0]; homology of setae and solenidia indicated in Table 1; seta *s* of tarsus I spiniform; phylliform seta absent on segments; solenidion φ₁ of tibia I long, subflagellate versus other solenidia medium-sized or short, thickened, rounded distally.

**Remarks** — *Carabodes* (*Klapperiches*) *paradilatatus* Ermilov **n. sp.** is most similar to *C. (K.) dilatatus* Ermilov, Sidorchuk and Rybalov, 2010 from Ethiopia in having tuberculate notogaster and dorsal part of prodorsum, long bothridial seta with well-developed head, and short, phylliform interlamellar and notogastral setae. However, the new species differs from the latter by smaller body size (length: 300–345 versus 365–464), the morphology of lamellar, bothridial and adanal setae (*le* setiform; *bs* fusiform, with barbed head; *ad*₁–*ad*₃ dilated mediodistally versus *le* dilated in median part; *bs* clavate, nearly smooth; *ad*₁–*ad*₃ setiform), the density of arrangement of the tubercles in dorsocentral part of the notogaster (unevenly spaced, with some comparatively large areas without tubercles versus evenly and densely spaced, without large free areas between tubercles), and the absence (versus presence) of foveolae on the subcapitular mentum.
<table><caption>**Table 1** Leg setation and solenidia of *Carabodes* (Klapperiches) *paradilatatus* Ermilov *n. sp.*.</caption><thead><tr><th>Leg</th><th>Tr</th><th>Fe</th><th>Ge</th><th>Ti</th><th>Ta</th></tr></thead><tbody><tr><td>I</td><td>v'</td><td>d, (l), bv''</td><td>(l), v', σ</td><td>(l), (v), φ₁, φ₂</td><td>(ft), (tc), (it), (p), (u), (a), s, (pv), e, ω₁, ω₂</td></tr><tr><td>II</td><td>v'</td><td>d, (l), bv''</td><td>(l), v', σ</td><td>(v), φ</td><td>(ft), (tc), (it), (p), (u), (a), s, (pv), ω₁, ω₂</td></tr><tr><td>III</td><td>l', v'</td><td>d, l', ev'</td><td>l', σ</td><td>(v), φ</td><td>(ft), (tc), (it), (p), (u), (a), s, (pv)</td></tr><tr><td>IV</td><td>v'</td><td>d, ev'</td><td>d, l'</td><td>(v), φ</td><td>ft'', (tc), (p), (u), (a), s, (pv)</td></tr></tbody></table>

Note: *Tr*, *Fe*, *Ge*, *Ti*, *Ta* = trochanter, femur, genu, tibia, and tarsus, respectively; 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.

**Etymology** — The species name *paradilatatus* refers to the similarity of *C. (K.) paradilatatus* to *C. (K.) dilatatus*.

**Congocephalus setiformis** Ermilov *n. sp.*

Zoobank: 1B2C9206-2CCE-4309-AF17-5D9B43984037

(Figures 4, 5)

**Type material** — Holotype (male) and six paratypes (three males and three females): Southern Ethiopia, South Ethiopia Regional State, Gamo Zone, 6°01'36.7"N, 37°35'42.7"E, 1170 m a.s.l., 5 km East from the town Arba Minch, 1 km from the lake Abai, litter in a mixed forest on the terrace (first layer of forest: *Celtis* sp., *Ficus sycomorus*, *Schefflera* sp., *Acacia* sp.; second layer of forest: *Lemon* sp., *Coffea arabica*) on brown (black-brown) soil with light-medium loam soil texture, 7.XII.2018, dry season (L.B. Rybalov).

The holotype is deposited in the collection of the Senckenberg Museum of Natural History, Görlitz, Germany; six 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: 375–450. Notogaster with specific system of comparatively sparse ridges and depressions; two anteromedial ridges fused distally. Lamella rounded distally; translamella present. Dorsocentral prodorsal region distinctly hump-like; mediobasal prodorsal part with large depression. Rostral seta medium-sized, phylliform, barbed, inserted on translamella; lamellar seta medium-sized phylliform, covered by strong spines, inserted in ventrodistal part of lamella; interlamellar seta long, phylliform, barbed; bothridial seta medium-sized, setiform, heavily barbed. Fourteen pairs of phylliform, barbed notogastral setae: *c*₂, *dp* longest; *p*₁, *p*₂, *p*₃, *h*₃ shortest; other setae medium-sized; *c*₂ directed forward. Epimeral setae *la*, *lc*, *2a*, *3a* and anal setae short, spiniform; genital setae short, setiform, nearly smooth; other epimeral setae well as aggenital and adanal setae short, setiform, slightly barbed; seta *ad*₃ in preanal position. Leg tarsus IV with 12 setae; seta *l''* of genua I, II and *l'* of genua III, IV phylliform.

**Description** — *Measurements*. Body length: 375 (holotype), 375–450 (male and female paratypes); notogaster width: 210 (holotype), 210–255 (male and female paratypes). Body ratio (length/width): ≈ 1.7–1.9. No difference between males and females in size.

*Integument* (Figs 4(a–e)) — Body color light grey to dark brown. Body surface nearly smooth, covered by thin layer of dense microgranulate cerotegument. Notogaster and anogenital region with specific system of ridges and depressions; two oblique anteromedial notogastral ridges fused distally.

*Prodorsum* (Figs 4(a, b, d)) — Rostrum broadly rounded. Lamella long and broad, rounded distally; translamella present (observed in dorsal aspect); tutrorium long (slightly shorter than lamella), ridge-like. Dorsocentral region distinctly hump-like; mediobasal part with large depression. Rostral seta (37–41) phylliform, dilated in median part, barbed, inserted on distinct tubercle (tubercle located on translamella); lamellar seta (37–41) phylliform, dilated in median
<figure><img src="image_6.png" /><figcaption>**Figure 4** *Congocepheus setiformis* Ermilov *n. sp*., adult: a - dorsal view; b - anterior part of prodorsum, anterior view; c - ventral view (gnathosoma and legs omitted); d - right lateral view (legs omitted); e - posterior view (part of left half omitted). Scale bar 100 µm.</figcaption></figure>
<figure><img src="image_7.png" /><figcaption>**Figure 5** *Congocepheus setiformis* Ermilov *n. sp.*, adult: a -subcapitulum, ventral view; b -palp, left, paraxial view; c -chelicera, left, paraxial view; d -leg I, right, antiaxial view; e -leg II (tarsus omitted), right, antiaxial view; f -leg III (tarsus omitted), left, antiaxial view; g -leg IV, left, antiaxial view. Scale bars 20 µm (a, c-g), 10 µm (b).</figcaption></figure>
part, covered by strong spines, inserted in ventrodistal part of lamella; interlamellar seta (75–82) phylliform, dilated mediobasally or in median part, barbed; bothridial seta (52–56) setiform, heavily barbed, curved mediodistally; exobothridial seta not observed. Bothridium interrupted ventrally.

*Notogaster* (Figs 4(a, d, e)) — Anterior margin slightly convex medially; lateral and posterior parts slightly depressed. Humeral shoulder small, well visible. Fourteen pairs of notogastral setae ($c_3$ absent; $c_2$, dp: 82–90; $p_1$, $p_2$, $p_3$, $h_3$: 37–41; others: 67–71) phylliform, dilated mediobasally or in median part, barbed; $c_2$ directed forward; $c_1$ located medial to $c_2$. Opisthonotal gland opening and all lyrifissures visible.

*Gnathosoma* (Figs 5(a–c)) — Subcapitulum size: 94–97 × 75–79; subcapitular seta *h* (9–11) setiform, slightly barbed; *a* (11–15) and *m* (11–15) narrowly phylliform, slightly dilated in median part, slightly barbed. Palp length: 60–64; setation: 0-2-1-3-9(+ω); postpalpal seta (7) spiniform, roughened. Chelicera length: 109–112; seta *cha* (30–34) setiform, barbed; seta *chb* (15–17) setiform, roughened.

*Epimeral and lateral podosomal regions* (Figs 4(c, d)) — Epimeral setation: 3-1-3-3; setae *1a*, *1c*, *2a*, *3a* (4) spiniform, smooth; other setae (*1b*: 30; others: 19–22) setiform, slightly barbed. Discidium tubercle-like.

*Anogenital region* (Figs 4(c–e)) — Anogenital formula: 4-1-2-3; all genital setae (11–15) setiform, nearly smooth; aggenital and adanal setae (19–22) setiform, slightly barbed; seta $ad_3$ in preanal position, posterior to aggenital seta; both anal setae (4) spiniform, smooth. Adanal lyrifissure distanced from anal plate.

*Legs* (Figs 5(d–g)) — Claw of each leg strong, barbed dorsally, with small ventrobasal tooth. Dorsoparaxial porose area on femora I–IV elongate oval. Formulas of leg setation and solenidia: I (1-4-3-4-16) [1-2-2], II (1-4-3-3-15) [1-1-2], III (2-3-1-2-15) [1-1-0], IV (1-2-2-2-11) [0-1-0]; homology of setae and solenidia indicated in Table 2; seta *s* of tarsus I eupathidial; seta *l''* of genua I, II and *l'* of genua III, IV phylliform, dilated in median part; solenidion $\phi_1$ of tibia I long, subflagellate; $\omega_2$ long, rod-like; other solenidia medium-sized or short, thickened, rounded distally.

**Remarks** — *Congocepheus setiformis* Ermilov *n. sp.* is most similar to *C. rwandensis* Fernández, Theron and Leiva, 2016 from the Afrotropical region in having setiform bothridial seta and the presence of one pair of long notogastral setae of *c*-row directed forward. However, the new species differs from the latter by the seta $c_2$ (versus $c_1$) directed forward and a significantly smaller number of ridges and depressions on the notogaster.

**Etymology** — The species name *setiformis* refers to the morphology of the bothridial seta (setiform).

## General remark

This section discusses taxonomic status of some supraspecies taxa within Carabodidae based on the direction of notogastral setae in anterior part of the notogaster. The subgenus *Austrocarabodes* (*Austroflexa*) was proposed by Subias (2019), with *Austrocarabodes brasiliensis* Ermilov and Tolstikov, 2015 as type species. The genus *Flexa* was proposed by Kulijev (1977),

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

Note: See Table 1 for explanations.
with *Carabodes dubius* Kulijev, 1968 as type species; Subías (2004) considered *Flexa* as the subgenus within *Carabodes* Koch, 1835. *Tanzaniacepheus* was proposed by Fernández *et al.* (2017), with *Tanzaniacepheus ornatus* (Mahunka 1983) (in Mahunka 1983a) as type species; Subías (2022) considered *Tanzaniacepheus* as a subgenus within *Congocepheus*. *Austrocarabodes* (*Austroflexa*), *Flexa* and *Tanzaniacepheus* are morphologically similar to *Austrocarabodes* (*Austrocarabodes*), *Carabodes* (*Carabodes*) and *Congocepheus*, respectively, but differs from them mainly by the direction of notogastral setae of the *c*-row (usually one or two setae directed forward).

In our opinion, the direction of notogastral setae of the *c*-row in the cases listed above is a specific morphological character and is sufficient only to distinguish groups of species within the genera (see Balogh 1988; Ermilov *et al.* 2015), but not enough to support the generic or subgeneric status. Moreover, other important morphological characters are ignored in justifying these genera (for example, body sculpture/ornamentation; presence/absence, size and deep of notogastral depressions). Therefore, we consider *Austrocarabodes* (*Austroflexa*), *Flexa* and *Tanzaniacepheus* as junior subjective synonyms of *Austrocarabodes* (*Austrocarabodes*), *Carabodes* (*Carabodes*) and *Congocepheus*, respectively.

## Acknowledgements

We thank Dr. Gezahegn Degefe for supporting our field studies and organizing laboratory work, Dr. Oleg G. Gorbunov for sampling assistance and two anonymous reviewers for valuable comments. 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. The work was performed within the framework of the Joint Russian-Ethiopian Biological Expedition, financially supported by the Russian Academy of Sciences.

## ORCID

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

Leonid B. Rybalov https://orcid.org/0000-0002-1121-1183

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