# Two new species of Cavernocepheus (Acari, Oribatida, Otocepheidae) from Malawi

Sergey Ermilov, Josef Starý

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Sergey Ermilov, Josef Starý. Two new species of Cavernocepheus (Acari, Oribatida, Otocepheidae) from Malawi. Acarologia, 2022, 62 (2), pp.396-403. 10.24349/eyzw-dpwh. hal-03643307

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# Two new species of *Cavernocepheus* (Acari, Oribatida, Otocepheidae) from Malawi

Sergey G. Ermilov IDª, Josef Starý IDᵇ

## Original research

### ABSTRACT

The oribatid mite genus *Cavernocepheus* (Oribatida, Otocepheidae) is recorded for the first time in Malawi. Two new species of the subgenus *Cavernocepheus* (*Paracavernocepheus*) — *C. (P.) hlavaci* n. sp. and *C. (P.) mulanjensis* n. sp. — are described from forest litter. Morphological comparison between known species of *Cavernocepheus* (*Paracavernocepheus*) and their distribution are presented.

**Keywords** otocepheid mites; taxonomy; new species; morphology; distribution; Afrotropical region

**Zoobank** http://zoobank.org/A717BB51-A40A-4336-A048-5310E5FC9614

## Introduction

The oribatid mite genus *Cavernocepheus* (Acari, Oribatida, Otocepheidae) was proposed by Balogh and Mahunka, 1969 with *Cavernocepheus monstruosus* Balogh and Mahunka, 1969 as type species. At present, the genus comprises two subgenera and nine species (*Cavernocepheus* (*Cavernocepheus*) Balogh and Mahunka, 1969 — seven species; *C. (Paracavernocepheus)* Ermilov and Starý, 2018 — two species), which are distributed in the Afrotropical (*C. (Paracavernocepheus*)*) and Neotropical (*C. (Cavernocepheus*)*) regions (Subías 2004, updated 2021). The revised generic and subgeneric diagnoses and an identification key to known species of the genus were presented by Ermilov and Starý (2018).

Among the oribatid mite materials (all specimens were kindly provided by the Moravian Museum, Brno, Czech Republic) collected from Malawi, we found two new species of *C. (Paracavernocepheus)*. They are the first representatives of the genus recorded in this country. The main goal of the paper is to describe these two new species. Additionally, the morphological comparison between the known (four) species of the subgenus and their distribution are given.

## Methods

**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”.

<figure><img src="image_6.png" /></figure>

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

<footnote>ᵇBiology Centre v.v.i., Czech Academy of Sciences, Institute of Soil Biology, Ceské Budejovice, Czech Republic.</footnote>
**Terminology** — Morphological terminology partially used in this paper follows that of Aoki (1965, 1967), Norton (1977), Ermilov and Starý (2018), and Ermilov and Khaustov (2020).

**Abbreviations** — Prodorsum: *cos* = costula; *ro*, *le*, *in*, *bs*, *ex* = rostral, lamellar, interlamellar, bothridial, and exobothridial seta, respectively; *cpm*, *cpl* = medial and lateral prodorsal condyles, respectively; *con* = concavity. Notogaster: *cnl* = lateral notogastral condyles; *cr* = crista; *c*, *la*, *lm*, *lp*, *h*, *p* = notogastral setae; *ia*, *im*, *ip*, *ih*, *ips* = notogastral lyrifissures; *gla* = opisthonotal gland opening. Gnathosoma: *a*, *m*, *h* = subcapitular setae. Epimeral and lateral podosomal regions: *1a–1c*, *2a*, *3a–3c*, *4a–3c* = epimeral setae; *PdI*, *PdII* = pedotectum I, II, respectively; *dis* = discidium. Anogenital region: *g*, *ag*, *an*, *ad* = genital, aggenital, anal, and adanal seta, respectively; *vr* = ventral ridge; *iag*, *iad* = aggenital and adanal lyrifissure, respectively; *cvr* = circumventral ridge; *po* = preanal organ. Legs: *Tr*, *Fe*, *Ge*, *Ti*, *Ta* leg trochanter, femur, genu, tibia, and tarsus, respectively; *pa* = porose area; *ω*, *φ*, *σ* = solenidia; *ε* = famulus; *d*, *l*, *v*, *bv*, *ev*, *ft*, *tc*, *it*, *p*, *u*, *a*, *s*, *pv* = leg setae.

# Taxonomy

## *Cavernocepheus (Paracavernocepheus) hlavaci* n. sp.

Zoobank: D4EF017C-EF51-4699-BFC6-A7012C72A447

(Figures 1a-c, 2a-d)

**Diagnosis** — Body length: 730–813. Rostral and lamellar setae long, setiform, barbed; interlamellar seta long, rod-like, barbed; bothridial seta spindleform, with long setiform apex, slightly barbed; exobothridial seta present. Lateral and medial prodorsal condyles present; lateral notogastral condyle quadrangular, top slightly oblique. All notogastral setae long, rod-like, barbed. Epimeral setae setiform, slightly barbed. Genital setae setiform, roughened; other anogenital setae setiform, barbed.

**Description of adult** — Measurements – Body length: 813 (holotype, male), 730, 796 (two paratype males); body width: 365 (holotype), 315, 348 (two paratypes). Length/width ratio of body: 2.2–2.3.

Integument – Body color light brown. Body surface densely microgranulate and partially sparsely foveolate (diameter of foveola up to 8, poorly visible). Lateral part of body between bothridium and acetabula I–III densely tuberculate (diameter of tubercle up to 8).

Prodorsum – Rostrum broadly rounded. Costula long, reaching bothridium basally and insertion of lamellar seta distally. Rostral (102–106) and lamellar (110–114) setae setiform, barbed. Interlamellar seta (106–114) rod-like, barbed. Bothridial seta (135–147) spindleform, with long setiform apex, slightly barbed. Exobothridial seta (32–41) setiform, thin, slightly barbed. Tutorium absent. Paired lateral and medial prodorsal condyles tubercle-like, located separately.

Notogaster – Concavity of dorsosejugal region large. Lateral notogastral condyle broadly quadrangular, top slightly oblique. Crista (extending from inner part of *cnl*) long, almost reaching the level of insertions of setae *lm*. Ten pairs of rod-like, barbed setae (86–98). Lyrifissures distinct, *ia* located lateral to *c*, *im* lateral to *lm*, *ip* between *p*₂ and *p*₃, *ips* between *h*₃ and *p*₃, *ih* anterior to *h*₃. Opisthonotal gland opening located close to *im*.

Gnathosoma – Morphology of subcapitulum, palp and chelicera similar to the other representatives of *Cavernocepheus* (*Paracavernocepheus*) (see Ermilov and Starý 2018). Subcapitulum size: 147–159 × 102–114. Subcapitular setae (*a*: 20; *m*: 49–53; *h*: 57–61) setiform, barbed. Palp (102–110) setation: 0–2–1–3–8(+ω). Solenidion of palptarsus connected with setae *ul'* and *ul''* mediodistally. Postpalpal seta (6) spiniform, smooth. Chelicera (159–168) with two setiform, barbed setae (*cha*: 61; *chb*: 28).

Epimeral and lateral podosomal regions – Apodemes I, II and sejugal apodeme well-developed; apodeme III also present, but poorly visible. Epimeral formula: 3-1-3-3. All setae
<figure><img src="image_7.png" /><figcaption>**Figure 1** *Cavernocepheus (Paracavernocepheus) hlavaci* n. sp., adult: a – dorsal view (legs omitted); b – ventral view (legs omitted); c – lateral view (legs omitted). Scale bar 100 µm.</figcaption></figure>
<figure><img src="image_8.png" /><figcaption>**Figure 2** *Cavernocepheus (Paracavernocepheus) hlavaci* 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 50 µm.</figcaption></figure>
(1a, 2a, 3a, 4b: 24–32; 1b: 73–82; 3b: 61–69; 3c: 32–41; others: 57–65) setiform, slightly barbed. Pedotectum I represented by large lamina. Discidium subtriangular.

Anogenital region – Aggenital lyrifissure located close and anterior to genital aperture. Genital setae (20–24) setiform, roughened. Aggenital (49–53), adanal (61–69) and anal (41–49) setae setiform, barbed. Distance *ad*<sub>3</sub>–*ad*<sub>3</sub> larger than *ad*<sub>2</sub>–*ad*<sub>2</sub> and *ag*–*ag*. Adanal lyrifissure located close and parallel to anal plate. Circumventral ridge slightly developed.

Legs – Claw of each leg strong, barbed on dorsal side. Dorsal side of all tarsi without teeth. Genua III, IV and femora I, II with one pair of teeth ventrodistally versus genua I, II and femora III, IV without teeth. Porose areas on all femora and on trochanters III, IV well visible. 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 (1–2–2–2–12) [0–1–0]; homology of setae and solenidia indicated in Table 1. Seta *u* setiform on tarsus I versus thorn-like on tarsi II-IV. Seta *l'* on genu II and *pv'* on tibia IV phylliform.

**Material examined** — Holotype (male) and two paratypes (all males): sample #12, Malawi, Zomba, Mt. Zomba, 15°20'37.2"S, 35°16'40.2"E, 1846 m a.s.l., sifting forest litter, Winkler extraction, 27.XI.2012 (leg. P. Baňař & P. Hlaváč).

**Type deposition** — The holotype and two paratypes are deposited in the collection of the Tyumen State University Museum of Zoology, Tyumen, Russia.

**Etymology** — The species name is dedicated to our colleague, Dr. Peter Hlaváč (Prague, Czech Republic) renowned entomologist, specialist on Coleoptera, collector of large material of soil samples from Malawi.

**Remarks** — The new species clearly differs from all species of the subgenus in the presence of spindleform (versus clavate/fusiform) bothridial seta having long, setiform apex (versus without long, setiform apex).

**Cavernocephus (Paracavernocephus) mulanjensis** n. sp.

Zoobank: 4093E79F-CC1C-474D-A8D8-B7D027526014

(Figures 3a-c)

**Diagnosis** — Body length: 630–747. Rostral and lamellar setae long, setiform, barbed; interlamellar seta long, rod-like, barbed; bothridial seta clavate, slightly barbed; exobothridial seta present. Lateral and medial prodorsal condyles present; lateral notogastral condyle quadrangular, concave medially. Posterior notogastral setae setiform, with attenuate and flexible tip, barbed; others rod-like, barbed. Epimeral setae setiform, slightly barbed. Genital setae setiform, roughened; other anogenital setae setiform, barbed.

**Description of adult** — Measurements – Body length: 747 (holotype, male), 630, 747 (two paratype males); body width: 340 (holotype), 282, 340 (two paratypes). Length/width ratio of body: 2.2.

Integument – Body color light brown. Body surface densely microgranulate and partially sparsely foveolate (diameter of foveola up to 8). Lateral part of body between bothridium and acetabula I–III densely tuberculate (diameter of tubercle up to 8).

<table><caption>**Table 1** Leg setation and solenidia of adult *Cavernocephus (Paracavernocephus) hlavaci* n. sp. and *C. (P.) mulanjensis* 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), ε, 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>l', (v), 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>v', l'</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>
<figure><img src="image_9.png" /><figcaption>**Figure 3** *Cavernocepheus (Paracavernocepheus) mulanjensis* textbfn. sp., adult: a – dorsal view (legs omitted); b – ventral view (legs omitted); c – lateral view (legs omitted). Scale bar 100 µm.</figcaption></figure>
Prodorsum – Rostrum broadly rounded. Costula long, reaching bothridium basally and insertion of lamellar seta distally. Rostral (86–98) and lamellar (98–110) setae setiform, barbed. Interlamellar seta (61–69) rod-like, barbed. Bothridial seta (102–114) clavate, slightly barbed. Exobothridial seta (24–28) setiform, thin, slightly barbed. Tutorium absent. Paired lateral and medial prodorsal condyles tubercle-like, located separately.

Notogaster – Concavity of dorsosejugal region large. Lateral notogastral condyle broadly quadrangular, concave medially. Crista (extending from inner part of cnl) long, almost reaching the level of insertions of setae *lm*. Ten pairs of notogastral setae (57–65); posterior setae ($h_3, p_1-p_3$) setiform, with attenuate and flexible tip, barbed; other setae rod-like, barbed. Lyrifissures distinct, *ia* located lateral to *c*, *im* lateral to *lm*, *ip* between $p_2$ and $p_3$, *ips* between $h_3$ and $p_3$, *ih* anterior to $h_3$. Opisthonotal gland opening located close to *im*.

Gnathosoma – Morphology of subcapitulum, palp and chelicera similar to the other representatives of *Cavernocepheus* (*Paracavernocepheus*) (see Ermilov and Starý 2018). Subcapitulum size: 139–147 × 86–94. Subcapitular setae (*a*: 20; *m*: 45–53; *h*: 53–61) setiform, barbed. Palp (86–94) setation: 0–2–1–3–8 (+ω). Solenidion of palptarsus connected with setae *ul'* and *ul''* mediodistally. Postpalpal seta (6) spiniform, smooth. Chelicera (151–159) with two setiform, barbed setae (*cha*: 57; *chb*: 28).

Epimeral and lateral podosomal regions – Apodemes I, II and sejugal apodeme well-developed; apodeme III also present, but poorly visible. Epimeral formula: 3-1-3-3. All setae (*1a*, *2a*, *3a*, *4b*: 24–28; *1b*: 53–61; others: 41–49) setiform, slightly barbed. Pedotectum I represented by large lamina. Discidium subtriangular.

Anogenital region – Aggenital lyrifissure located close and anterior to genital aperture. Genital setae (16–20) setiform, roughened. Aggenital (41–49), adanal (57–65) and anal (41–49) setae setiform, barbed. Distance $ad_3-ad_3$ larger than $ad_2-ad_2$ and *ag-ag*. Adanal lyrifissure located close and parallel to anal plate. Circumventral ridge slightly developed.

Legs – Generally, similar to *Cavernocepheus* (*Paracavernocepheus*) *hlavaci n. sp.*, but: 1) all genua and femora without teeth ventrodistally; 2) seta *l'* on genu II and *pv''* on tibia IV setiform.

**Material examined** — Holotype (male) and two paratypes (all males): sample #9, Malawi, Maloza, Mulanje Mts., Maloza stream valley, 16°0′3.9″S, 35°32′36.6″E, 972 m a.s.l., sifting litter, Winkler extraction, 24.XI.2012 (leg. P. Baňař).

**Type deposition** — The holotype and two paratypes are deposited in the collection of the Tyumen State University Museum of Zoology, Tyumen, Russia.

**Etymology** — The species name refers to the place of origin, Mulanje Mts., Malawi.

**Remarks** — The new species clearly differs from all species of the subgenus in the presence of posterior notogastral setae ($h_3, p_1-p_3$) having attenuate and flexible tip (versus posterior notogastral setae rod-like).

## Morphological comparison

The known species of *Cavernocepheus* (*Paracavernocepheus*) can be distinguished from each other by the morphology of bothridial seta, lateral notogastral condyle, notogastral and adanal setae, leg tarsi, genua and femora, and leg seta *l'* on genu II and *pv''* on tibia IV (see Table 2).

## Distribution of *Cavernocepheus* (*Paracavernocepheus*)

Species of the subgenus are known from the Afrotropical region only and have a highly circumscribed geographic distribution, i.e. are endemic to a single country.

*Cavernocepheus (Paracavernocepheus) nortonroyi* and *C. (P.) concavus* are known from Cameroon: forest litter and soil in Korup National Park (300 m a.s.l.), Rengo Camp, about 8 km NW of Mundemba South-West Province. *Cavernocepheus (Paracavernocepheus) hlavaci* and *C. (P.) mulanjensis* are known from Malawi: *C. (P.) hlavaci* was described from forest litter on Mt. Zomba (1846 m a.s.l.); *C. (P.) mulanjensis* from forest litter in Maloza stream valley on Mulanje Mts. (972 m a.s.l.).
<table><caption>**Table 2** Morphological differences between *Cavernocepheus* (*Paracavernocepheus*) spp.</caption><thead><tr><th>Character</th><th>C. (P.) nortonroyi Ermilov and Starý, 2018</th><th>C. (P.) concavus Ermilov and Starý, 2018</th><th>C. (P.) hlavaci n. sp.</th><th>C. (P.) mulanjensis n. sp.</th></tr></thead><tbody><tr><td>Bothridial seta</td><td>Clavate, with three or four teeth distally</td><td>Lanceolate, slightly barbed</td><td>Spindleform having setiform apex</td><td>Clavate, slightly barbed</td></tr><tr><td>Lateral notogastral condyle</td><td>Truncate</td><td>Concave medially</td><td>Truncate</td><td>Concave medially</td></tr><tr><td>Notogastral setae</td><td>Setae h<sub>3</sub>, p<sub>1</sub>-p<sub>3</sub> longer than others, all setae rod-like</td><td>All setae similar in length, rod-like</td><td>All setae similar in length, rod-like</td><td>All setae similar in length, h<sub>3</sub>, p<sub>1</sub>-p<sub>3</sub> with attenuate and flexible tip, others rod-like</td></tr><tr><td>Adanal setae</td><td>With flexible tip</td><td>Without flexible tip</td><td>Without flexible tip</td><td>Without flexible tip</td></tr><tr><td>Leg tarsi dorsally</td><td>With small teeth</td><td>With small teeth</td><td>Without teeth</td><td>Without teeth</td></tr><tr><td>Leg genua and femora ventrodistally</td><td>Rounded</td><td>Rounded</td><td>Genua III, IV and femora I, II pointed</td><td>Rounded</td></tr><tr><td>Leg seta l' on genu II and pv" on tibia IV</td><td>Setiform</td><td>Setiform</td><td>Phylliform</td><td>Setiform</td></tr></tbody></table>

## Acknowledgements

We thank to Dr. Julia Baumann and two anonymous reviewers for valuable comments; Drs P. Baňař (Brno, Czech Republic) and P. Hlaváč (Prague, Czech Republic) for long-term cooperation in study of soil invertebrates; and the Moravian Museum in Brno, Czech Republic, which kindly provided material for our study. Samples exportation to Czech Republic was based on permit no. EAD-12-07-086-18-20. This research was supported by Czech Academy of Sciences (Research Plan No. RVO: 60077344) and the cooperative agreement No. FEWZ-2021-0004 from the Russian Ministry of Science and Higher Education.

## References

* Aoki J. 1965. A preliminary revision of the family Otocepheidae (Acari, Cryptostigmata). I. Subfamily Otocepheinae. Bul. Nat. Sci. Mus., Tokyo, 8(3): 259-341.

* Aoki J. 1967. A preliminary revision of the family Otocepheidae (Acari, Cryptostigmata). II. Subfamily Tetracondylinae. Bul. Nat. Sci. Mus., Tokyo, 10(3): 297-359.

* Balogh J., Mahunka S. 1969. The scientific results of the Hungarian soil zoological expeditions to South America. 10. Acari: Oribatids, collected by the second expedition. I. Acta Zool. Acad. Sci. Hung., 15(1-2): 1-21.

* Ermilov S.G., Khaustov A.A. 2020. A new species of *Dolichermaeus* Jacot 1938 (Acari, Oribatida, Otocepheidae) from Zanzibar, with a key to the known species of the genus from the Ethiopian region. Zool. Zh., 99(1): 13-23. https://doi.org/10.31857/S0044513419110047

* Ermilov S.G., Starý J. 2018. Contribution to the knowledge of the oribatid mite genus *Cavernocepheus* (Acari, Oribatida, Otocepheidae). Syst. Appl. Acarol., 23(3): 489-500. https://doi.org/10.11158/saa.23.3.8

* Norton R.A. 1977. A review of F. Grandjean's system of leg chaetotaxy in the Oribatei (Acari) and its application to the family Damaeidae. In: Dindal D.L. (Ed.). Biology of oribatid mites. Syracuse: SUNY College of Environmental Science and Forestry. pp. 33-61.

* Subías L.S. 2004. Listado sistemático, sinonímico y biogeográfico de los ácaros oribátidos (Acariformes, Oribatida) del mundo (1758-2002). Graellsia, 60 (número extraordinario): 3-305. https://doi.org/10.3989/graellsia.2004.v60.iExtra.218

* Subías L.S. 2021. Listado sistemático, sinonímico y biogeográfico de los Ácaros Oribátidos (Acariformes: Oribatida) del mundo (excepto fósiles), 16ª actualización. 532 pp. Available from: http://bba.bioucm.es/cont/docs/RO_1.pdf (accessed March 2021).