Frolov, A.V. 2014. Revision of the genus Delopleurus Boheman (Coleoptera: Scarabaeidae: Scarabaeinae) with description of new species from Africa. Journal of Natural History, 49: 1-25. 

DOI: 10.1080/00222933.2014.909072

Revision of the genus Delopleurus Boheman (Coleoptera: Scarabaeidae: Scarabaeinae) with description of new species from Africa

A.V. Frolov

Laboratory of Insect Systematics, Zoological Institute, Russian Academy of Sciences, Universitetskaya nab., 1, 199034 St.Petersburg, Russia. E-mail: avfrolov@mail.ru

Abstract

The genus Delople​urus Erichson, 1847, is revised. Five new species are described from Africa: Delopleurus naviauxi sp. nov., Delopleurus krikkeni sp. nov., Delopleurus darrenmanni sp. nov., Delopleurus fossatus sp. nov. and Delopleurus pubescens sp. nov. One new synonymy is established (Delopleurus parvus (Sharp, 1875) = Delopleurus cardoni Paulian, 1934, syn. nov.). Lectotype of Delopleurus pullus Boheman, 1857, is designated. Delopleurus janssensi Frey, 1963, is transferred to the genus Metacatharsius Paulian, 1939. A key to the Delopleurus species and locality maps are given.

Keywords: Coleoptera, Scarabaeidae, Delopleurus, new species, new synonymy, key to species, Africa-Southern Asia disjunction, mycetophagy

Introduction

This paper is the last in a series of the taxonomic revisions of the peculiar scarabaeine "dung beetles" of the Sarophorus group of genera (Sarophorus Erichson, Frankenbergerius Balthasar, Coptorhina Hope, and Delopleurus Erichson). This group was first recognized by Erichson ((1847)) who noted the sinuate epipleura (strongly in Coptorhina Hope and Delopleurus Erichson and not so strongly in Sarophorus Erichson) and correspondingly widened metepisterna. Later the status and position of Frankenbergerius Balthasar within the group was validated ((Frolov & Scholtz, 2005)) and other characters shared by the members of the group were found ((Frolov, 2002; Frolov, Akhmetova, & Scholtz, 2008; Frolov & Scholtz, 2003, 2005)).

   Delopleurus comprises superficially similar, small-sized, dark brown to black, strongly convex beetles distributed in Africa south of the Sahara and is Southern Asia. The genus is commonly classed as the dung beetles, however there are no evidences of its association with dung. Available data suggest that the beetles are associated with higher fungi similar to the related genus Coptorhina Hope, 1830, members of which are obligate feeders of basidiomycetes ((Frolov et al., 2008)). Delopleurus is also peculiar in having a disjunctive range including most of the Afrotropical Region and Indian Peninsula. Distribution of the other genera of Sarophorus group is limited to the Afrotropical region.

Of the four genera of the Sarophorus group, Delopleurus is the rarest in the collections. This may indicate that the beetles have a highly cryptic life style and short mating period and depend on the highly ephemeral and patchy food resources (mushrooms). It is however possible that some species may be locally common as can be seen from a reasonable series of D. darrenmanni  sp. n. collected in Caprivi Park Nova (Namibia).

Before the present contribution Delopleurus comprised 6 nominal species, three described from Africa and three from Asia ((Davis, Frolov, & Scholtz, 2008; Ferreira, 1972; Gillet, 1911; Janssens, 1939)). Examination of the material accumulated in museums after the last taxonomic works on the group showed that the characters used by the previous workers are not sufficient to separate species. Other characters, especially the sculpture of the pygidium, suggested that there are nine species of Delopleurus in the available material, five of which were not yet described. This necessitated examination of the type specimens of all nominal species and a revision of the genus.

Material and methods

The study is based on the material housed in the following institutions:

BMNH  - The Natural History Museum, London,

DMAGD - Durban Museum and Art Gallery, Durban,

IRSNB   - Institut Royal des Sciences Naturelles de Belgique, Brussels,

MNHN  - Muséum National d'Histoire Naturelle, Paris,

MRAC   - Musee Royal de l’Afrique Centrale, Tervuren,

NHMB  - Naturhistorisches Museum, Basel,

NHRS    - Naturhistoriska Riksmuseet, Stockholm,

SAMC   - South African Museum, Cape Town,

SANC    - National Collection of Insects, Plant Protection Research Institute, Pretoria,

TMSA   - Transvaal Museum, Pretoria,

UMO - University Museum, Oxford,

UPSA    - University of Pretoria Insect Collection, Pretoria,

ZIN   - Zoological Institute of Russian Academy of Sciences, Sankt-Petersburg.

Preparation of specimens follows the common technique used in entomological research. Photographs were taken with a Leica MZ9.5 stereo microscope and a Leica DFC290 digital camera from dry specimens except for aedeagi and internal sacs, which were photographed in glycerol. Partially focused serial images were combined in Helicon Focus software (Helicon Soft Ltd.) to produce completely focused images. Distribution maps were prepared with ArcGIS software (ESRI Inc.). As a base map, terrestrial ecoregions of the world map ((Olson et al., 2001)) was used. Co-ordinates of the localities were taken from the specimens labels, if available, or from the NGA GEOnet Names Server (GNS, http://earth-info.nga.mil/gns/html/index.html).

Deloplerus (Erichson, 1847)

Type species: D. pullus (Boheman, 1857).

Delopleurus was established by Erichson ((1847)) for "one small south African species, which differs from Coptorhina in narrower hind legs". Ten years later Boheman ((1857)) gave the name to this species, D. pullus, and thus fixed the type species of the genus. Sharp ((1876)) described Coptorhina parva from Northern India. Arrow ((1931)) moved C. parva to Delopleurus and described the second species from India, D. striatus. Paulian ((1934)) described the third species from India, D. cardoni. Janssens ((1939)) described the second species from Africa, D. gilleti, and gave a diagnostic key to all species known by then. Frey ((1963)) described D. janssensi from Ethiopia. Except for this primary taxonomic literature, members of the genus were listed in the catalogues and monographs ((Balthasar, 1963; Davis et al., 2008; Ferreira, 1972; Gillet, 1911; Péringuey, 1901)) but little additional data was given.

Diagnosis

Delopleurus and closely related Coptorhina share somewhat rectangular metepisternum, widest in its hind part, very convex epipleural margin (Figure 1), and other taxonomic characters of the Sarophorus group of genera ((Frolov et al., 2008)). From Coptorhina it can easily be separated by quadridentate clypeus and smaller body size (3.5–6.1 mm in Delopleurus and 8.0–20.0 mm in Coptorhina).

Delopleurus spp. D. fossatus sp. n., D. pullus, D. naviauxi, female, holotype
Figures 1–4. Delopleurus spp. 1, D. fossatus sp. n., lateral view of body (sinuate epipleuron is arrowed); 2–4, anterior tibiae (2, D. pullus, male, 3, D. pullus, female, 4, D. naviauxi, female, holotype).

Description

Beetles are small-sized (3.5–6.1 mm), strongly convex, black or dark brown, glabrous.

Clypeus distinctly quadridentate in most species. Two medial teeth are almost always acute, lateral ones are acute to right-angled. In some species, lateral clypeal teeth are almost not separated from the lateral margin (Figures 34, 37). Head without carinae on disc, and any horns or tubercles. Small carina present near inner margin of eye. Genae right-angled, indistinctly separated from  clypeus. Frontoclypeal and genal suturae almost indistinct. Eyes small, their dorsal parts small, slit-shaped in some species, ventral parts larger, sub-rectangular. Distance between eye and gula approximately 2 times the width of eye in ventral view. Gula with longitudinal groove (Figure 51). Dorsal surface of clypeus rugose in most species, frons densely punctate to rugose.

Pronotum more or less trapezoidal, about 2 times wider than long. Anterior and lateral margins with distinct border, base with or without border. Pronotum is not excavated, without horns or ridges but with fine short longitudinal groove basally.

Elytra trapezoidal, as wide as long, shiny to opaque, with deep sinuation on lateral margins receiving convex metepisternum (Figure 1). Stria fine but distinct, punctate; striae 1-7 reach base of elytron, stria 8 reaches the sinuation but not the base; striae 9-10 are very close to epipleura and mostly inseparable from each other except apically. Elytral intervals flat to convex, with minute punctuation. Elytra fused along suture. Scutellum not visible from above.

Wings are well developed and feature a number of characters which Delopleurus share with species of the Sarophorus group of genera ((Frolov et al., 2008)).

Anterior tibiae have 3 outer teeth. Margin basad of 3rd tooth feebly or not serrate. Spur of anterior tibia more or less bifurcated in males and apically acute to rounded and curved inwards in females. In one species (D. naviauxi sp. n.), anterior tibia with 1 small acute dent between 1st outer tooth and spur (Figure 4). Outer margins of middle and posterior tibiae without transverse carinae, serrate. Tarsi of all legs well developed, shorter than tibiae. Claws about half the length of 5th tarsal segment.

Pygidium about 2 times wider than long, its sculpture is species-specific and, in some species, differs strongly from that in the other scarabaeine taxa. For descriptive purposes, three parts of the pygidium are recognized: basal border, apical border and disc. In some Delopleurus species basal and apical borders are relatively narrow and the disc occupies most of pygidium (Figure 9). In others, however, the borders are much wider and the disc is almost slit-shaped (Figure 6). Sculpture of pygidium is sex-dependent in some species. 

Aedeagus of typical scarabaeinae shape (Figures 10, 14, 20, 26, 36, 32, 42). Phallobase with two symmetrical tubercles. Parameres symmetrical, without setae apically, slightly distinctive in different species. Internal sac with armature consisting of a few sclerites of complex shape.

Sexual dimorphism

Males can be easily separated from females by having more or less bifurcated spurs of anterior tibiae (Figure 2, 3). Some species differ strongly in the sculpture of the pygidium.

Diagnostic characters

Unlike other scarabaeines, Delopleurus species can be best separated by the sculpture of the pygidium. Although shape of the parameres and internal sac armature shows some species specific features, they are not considered reliable diagnostic characters. However, availability of the other characters, especially that of the pygidium, makes dissection unnecessary in routine identification.

Distribution

Delopleurus has a disjunctive range covering most of Afrotropical Region and Indian Peninsula. The ranges of the different species are rather limited (except for D. gilleti Janssens) although it may be a result of undesampling. Two regional faunas, Afrotropical and Indian, share no common species which may indicate that they have long developed in isolation. The genus range disjunction can be best explained by the contraction of the previously very large range since the late Neogene. It is possible that in the late Miocene, when huge regions of Africa and Eurasia, including the territories of the present-day Sahara and Mediterranean, were covered by the African-type savannas, Delopleurus was widely distributed in the Eurasia. No paleontological data however exist on the past occurrence of Delopleurus in Europe and extratropical Asia, but considering very low taphonomic potential of these soil dwellers, discovery of the fossil Delopleurus seems highly improbable.

Biology

Little information is available about biology of Delopleurus and the premature stages of the genus are unknown. In Ivory Coast, a few specimens of D. gilleti Janssens were collected on mushrooms of Termitomyces cf. schimperi along with more numerous Coptorhina nitidipennis Boheman (Yves Cambefort, pers. comm.). In Namibia (Caprivi, Darren Mann, pers. comm.), beetles of D. darrenmanni sp. n. were collected in mushrooms (Termitomyces sp.) on and around termite mounds, also in association with Captorhina (C. nitidipennis and C. angolensis Arrow). There were apparently Coptorhina burrows under fungal umbrella or a few centimeters away. No smaller burrows of Delopleurus were seen however. It is possible that Delopleurus can utilize mushroom pieces buried by Coptorhina.

Delopleurus beetles have never been found in or under dung pads, and they have never been collected by pitfalls baited with dung or carrion (Philippe Walter collected a few specimens in traps baited with fish (Yves Cambefort, pers. coom.) but these were most probably chance trappings).

Available data suggest that Delopleurus are specialist feeders of agaric mushrooms (Agaricales). They seem to be associated with termitophilous genus Termitomyces, a widespread fungal genus with large fruit bodies. In case of Delopleurus and Coptorhina, feeding on mushrooms can be a secondary adaptation from the putatively generalist saprophagy of their ancestors. Feeding on mushrooms as an adaptation to arid biotopes in beetles was hypothesized by Striganova ((1980)).

All Delopleurus specimens examined have well developed wings and apparently are good fliers. But no specimens have been collected at light, according to the labels and data from the collectors. This, along with small eyes (typical for the members all other Sarophorus group genera), suggests that the beetles have diurnal flight activity.

Key to Delopleurus species

Males

1.  Disc of pygidium more or less densely punctate (Figures 13, 19, 25) or granulate with short brown setae (Figure 9). Species from Africa ............................................ 2

— Disc of pygidium bilobate, almost smooth, without setae (Figure 41). Species from Asia ...............................................................................................................  D. parvus

2.  Disc of pygidium rugose and with sparse brown setae (Figure 9)  D. pubescens sp. n.

— Disc of pygidium with different sculpture, without setae........................................... 3

3.  Border of pygidium relatively narrow; punctation of pygidium disc sparser (Figure 13) ..................................................................................................................  D. pullus

— Border of pygidium thicker (Figures 19, 31, 35), if similar to D. pullus than punctation of pygidium disc coarser (Figure 25) ....................................................... 4

4.  Disc of pygidium almost as long as apical border (Figure 19) D. darrenmanni  sp. n.

— Disc of pygidium 1.5–2 times longer than apical border............................................ 5

5. Disc of pygidium feebly convex, almost in plain with basal border (in lateral view, Fig. 25) ...........................................................................................................  D. gilleti

— Disc of pygidium strongly convex, distinctly separated from basal border by a deep hollow (in lateral view, Fig 31, 35) ............................................................................ 6

6.  Elytra with well-developed humeral umbones, sinuate behind umbones in dorsal view (Figure 30). Basal border of pygidium strongly widened medially (Figure 31) .  D. krikkeni sp. n.

-   Elytra with feebly developed humeral umbones, feebly or not sinuate behind umbones in dorsal view (Figure 34). Basal border of pygidium slightly widened medially (Figure 35) .........................................................................  D. fossatus sp. n.

Females

1.  Disc of pygidium more or less densely punctate, or slit-shaped, or granulate (with short brown setae). Species from Africa .................................................................... 2

— Disc of pygidium almost smooth (Figure 44) or granulate, without setae (Figure 48). Species from Asia ...............................................................................................  7

2. Fore tibia with small acute tooth near apical spur (Figure 4). Pygidium with slit-shaped pubescent disc (Figure 6)...................................................... D. naviauxi sp. n.

-   Fore tibia without tooth near apical spur (Figure 3). Sculpture of pygidium different; if disc of pygidium slit-shaped than it lacks setation (Figure 22).............. 3

3.  Borders of pygidium relatively narrow; disc feebly convex, granulate and pubescent (Figure 16)....................................................................................................... D. pullus

— Borders of pygidium relatively wide, disc without setation ...................................... 4

4.  Disc of pygidium slit-shaped, with transversal carina (Figure 22)..... D. darrenmanni  sp. n.

— Disc of pygidium 1.5–2 times longer than apical border thickness ........................... 5

5. Disc of pygidium feebly convex, almost in plain with basal border (in lateral view, Fig. 28) ..........................................................................................................................

     ......................................................................................................................... D. gilleti

— Disc of pygidium strongly convex, distinctly separated from basal border by a deep hollow (in lateral view, Fig. 38) .......................................................  D. fossatus sp. n.

7.  Elytral striae with larger punctures (Figure 43). Disc of pygidium smooth to indistinctly rugose (Figure 44) .....................................................................  D. parvus

— Elytral striae with smaller punctures (Figure 46). Disc of pygidium granulate (Figure 48)..................................................................................................... D. striatus

Delopleurus naviauxi Frolov et Cambefort, sp. nov.

(Figures 4–7)

Diagnosis

This species can be separated from other Delopleurus species by having deep transversal fossa on pygidium and by the fore tibia with small acute tooth near apical spur.

Description

Holotype, female. Body strongly convex, black, glabrous, length 4.9 mm (Figure 5).

Clypeus quadridentate. All four clypeal teeth acute, two medial relatively slender. Head without carinae on disc, small carinae present near inner margin of eyes. Genae right-angled, indistinctly separated from clypeus. Frontoclypeal and genal suturae indistinct. Eyes small, their dorsal parts slit-shaped, ventral parts sub-rectangular. Clypeus rugose in anterior part and laterally, frons densely punctate with elongate punctures .

Pronotum trapezoidal, about 2 times wider than long. Anterior and lateral margins bordered, base without border. Pronotum relatively densely punctate on disc (punctures separated by 1-2 puncture diameters), punctuation becoming denser laterally.

Elytra trapezoidal, as wide as long, shiny. Stria distinct, with punctures larger than striae. Elytral intervals slightly convex on disc, with minute punctuation.

Anterior tibiae have 3 outer teeth and a small acute tooth between 1st outer tooth and apical spur (Figure 4).

Pygidium with deep transversal slit-shaped fossa with yellowish setae (Figure 6).

Paratype, female, slightly larger than the holotype (6.0 mm). The specimen has malformed or worn, reduced anterior tibiae without outer teeth and tarsi, and reduced clypeal teeth. It also differs from the holotype in having less shiny dorsal side.

Male unknown.

Type material

Holotype, female with the labels "Kenya Colony Makuyu iii-v 1937 C.D.Knight", "Imp. Inst. Entom." (BMNH) and paratype, female with the labels "Kenya Maji ya Chumvi 30.4.78 R.N.", "coll. R. Naviaux", "Delopleurus n. sp? Y.Cambefort det" (MNHN).

Distribution

The species is known from two localities in Northern Acacia-Commiphora bushlands and thickets ecoregion in East Africa (Figure 7).

Etymology

The species is named after Roger Naviaux, a French collector.

Remark

Yves Cambefort (MNHN) examined the Delopleurus specimen from Maji ya Chumvi in 1980s and noted that it was distinct from other described species of this genus. However, he did not describe it at that time. In the course of the revision of the genus, I examined the specimen from Makuyu which is housed in the BMNH. After comparison of the two specimens we concluded that they are conspecific and belong to a new species described above.

Delopleurus spp. D. naviauxi sp. n. D. pubescens sp. n.
Figures 5–11. Delopleurus spp. 5, 8, habitus; 6, 9, pygidium; aedeagus in lateral view and internal sac; 7, 11, locality maps. 5–7, D. naviauxi sp. n.; 8–11, D. pubescens sp. n.

Delopleurus pubescens Frolov, sp. nov.

(Figures 8–11)

Diagnosis

Males of this species can be separated from other Delopleurus species by the disc of pygidium being rugose, somewhat granulate and pubescent with sparse brown setae (Figure 9). Such a sculpture is similar to that of females of D. pullus (Figure 16) but it is not rugose in the latter. Despite females of D. pubescens  sp. n. are not yet known and despite similarity of the male D. pubescens  sp. n. and female D. pullus in the pygidium sculpture, I believe that the females with a pubescent granulate disc of pygidium are conspecific with D. pullus and not with D. pubescens sp. n.

Description

Holotype, male. Body strongly convex, black, glabrous, length 5.0 mm (Figure 8).

Clypeus quadridentate, two medial tooth are acute, lateral ones are somewhat angulate. Genae right-angled, indistinctly separated from  clypeus. Frontoclypeal and genal suturae almost indistinct. Clypeus rugose in anterior part and laterally, frons densely punctate (punctures separated by about 1 puncture diameter).

Pronotum more or less trapezoidal, about 2 times wider than long. All margins with distinct border. Disc and base punctate with punctures separated by 2–3 puncture diameters, sides and especially anterior angles more densely punctate.

Elytra trapezoidal, as wide as long, somewhat opaque. Stria deep, with punctures slightly larger than striae width. Elytral intervals slightly convex (more distinctly on disc).

Anterior tibiae have 3 outer teeth, without a small acute tooth between 1st outer tooth and apical spur.

Pygidium with relatively slender borders and convex disc. Basal border slender and almost parallel-sided except in the middle. Apical border about 2 times thicker in the middle than the basal border, becoming slenderer laterally. Disc with irregular, rugose and granulate sculpture, pubescent with short yellowish setae (Figure 9).

Parameres with a small protrudence ventroapically (visible in lateral view, Fig. 10).

Female unknown.

Paratypes. Body length 4.3–5.5 mm.

Distribution

The species is known from a few localities in Congolian forest-savanna and Miombo woodlands (Figure 11).

Type material

Holotype, male with the label "Salisbury Mashonaland Feb. 1906 G.A.Marshall" (BMNH). Paratypes: 3 males with the same label as the holotype (BMNH); 1 male with the label “Salisbury Mashonaland G.A.Marshall 189[4]” (SAMC); 2 males with the labels “COLL. MUS. CONGO Mayidi -1945 R. P.Van Eyen” (MRAC); 1 male with the label “Musee du Congo Lulua: r. Kapelekese 17.XI.1933 G.F.Overlaet” (MRAC); 1 male with the label “Coll. I.R.Sc.N.B. Rhodesie du Sud  Salisbury Dec. 1900 G.A.K.Marshall Coll. J.J.Gillet” (IRSNB); 1 male with the label “Salisbury 14/1/15” (DMAGD).

Delopleurus pullus Boheman

(Figures 12-17)

Delopleurus pullus (Boheman, 1857); (Péringuey, 1901); (Ferreira, 1972); (Gillet, 1911); (Janssens, 1939).

Diagnosis

This species is most similar to D. gilleti but can be separated from it in having the pygidium with relatively larger disc punctate with smaller and sparser punctures in males and granulate and pubescent in females, and in having relatively sparsely punctate disc of pronotum.

Description

Beetles are small-sized (4.5-6.1 mm), strongly convex, black or dark brown, upper surface glabrous.

Male (Figure 12). Clypeus quadridentate, two medial teeth acute, lateral ones right-angled to acute-angled. Clypeus rugose in anterior part and laterally, frons densely punctate (punctures separated by less than a puncture diameter).

Pronotum more or less trapezoidal, about 2 times wider than long. Anterior and lateral margins bordered, base not bordered. Surface relatively sparsely punctate on disc (punctures separated by 2–3 puncture diameters) and more densely laterally.

Elytra trapezoidal, as wide as long, shiny. Stria relatively deep, with punctures slightly larger than striae width. Elytral intervals feebly convex to almost flat, with minute punctuation.

Anterior tibiae have 3 outer teeth, without a small acute tooth between 1st outer tooth and apical spur.

Pygidium with relatively slender borders and convex disc. Basal border slender and almost parallel-sided except in the middle. Apical border about 2 times thicker in the middle than the basal border, becoming slenderer laterally. Disc glabrous, punctate with punctures separated by 1–2 puncture diameters (Figure 13).

Parameres with a small protrudence ventroapically (visible in lateral view, Fig. 14).

Female (Figure 15) can be separated from male in having not bifurcated spurs of anterior tibiae and disc of pygidium with small granules and setae (Figure 16).

Delopleurus pullus.
Figures 12–17. Delopleurus pullus. 12, 15, habitus (12, male, 15, female); 13, 16, pygidium (13, male, 16, female); 14, aedeagus in lateral view and internal sac; 17, locality map.

Type material

Lectotype (here designated), male with the labels "Caffraria", "J. Wahlb[erg].", "Type", "pullus Boh. det J.Ferrer" (NHRS). Paralectotypes, male and female with the labels "Caffraria", "J. Wahlb[erg]." (NHRS).

Additional material

ANGOLA: Namakunde, II.1923, 1 male, 2 females (SAMC). ZIMBABWE: Mutare [Umtali], 1903, A.Bodong leg., 1 male (SAMC); Empandeni, 1911, J.O'Neil leg., 2 males, 2 females (SAMC); Salisbury, 1894, G.A.Marshall leg., 3 males (SAMC); Salisbury, XII.1897, G.A.Marshall leg, 1 male (BMNH); Salisbury, XII.1900, G.A.K.Marshall leg., 1 male (IRSNB); 7 mi S of Matimba, 5.I.1972, Bornemissza & Kirk leg., 1 male (SANC); Bulawayo, XII.1903, G.A.K.Marshall leg., 1 female (BMNH). NAMIBIA: Otjiwarongo district, Abachaus, XII.1951, G.Hobohm leg., 1 male (TMSA); Otjiwarongo district, Abachaus, 16.I.1956, G.Hobohm leg., 1 male, 2 females (NHMB); Kamanyab [Kamanjab], 1 male, 4 females (SAMC); Mafa, II.1923, 2 females (SAMC); Nagusib [Farm 25 SE Namutoni], I.1922, 1 spm. (SAMC); Kaokoveld, 44 km NW Ohopoho, 7.II.1975, from under stones, leg. Endrody-Younga leg., 2 females (TMSA). MOZAMBIQUE: Chimoio, XII.1929, P.Lesne leg., 2 females (MNHN). SOUTH AFRICA: D'Nyala Nature Reserve, Ellisras District, 18-20.XII.1987, Grobbelaar leg., 1 male, 1 female (SANC); Lydenburg District, 1896, P.A.Krantz leg., 1 female (TMSA); Vryburg, 1918, J.Brown leg., 1 male (SAMC); Vryburg, 1893, E.Simoni leg., 1 male, 1 female (MNHN); Malta, 1928, G.van Son leg., 2 males (TMSA); Shiluvane, 1902, 1 female (SAMC); Shiluvane, H.A. Janod leg., 1 female, 2 spm. (MNHN); Ben Alberts Nature Reserve, Thabazimbi, 24-28.XI.1980, S.J. van Tonder leg., 1 male (SANC); Moorddrift, XII.1914, C.J.Swierstra leg., 1 male (IRSNB); Durban, 1846, Boheman, 1 female (MNHN); Legonyane, 11.I.2011, from mushroom, C. Deschodt leg., 1 male, 4 females (UPSA); "Natal", 1 male (MNHN).

Distribution

The species is rather widely distributed in the northern part of Southern Africa (Figure 17).

Delopleurus darrenmanni Frolov, sp. nov.

(Figures 18–23)

Diagnosis

This species can be separated from other Delopleurus species by the sculpture of its pygidium. Males have punctate, glabrous, narrow disc of pygidium which is almost as long as apical and basal border in the middle (Figure 19). Females have characteristic slit-shaped disc with a transverse carina in the middle (Figure 22).

Description

Male, holotype. Body strongly convex, black, glabrous, its length 6.0 mm (Figure 19).

Clypeus quadridentate, medial teeth acute (right tooth malformed), lateral ones acute-angled. Clypeal margin laterad of lateral teeth somewhat angulate so clypeus looks hexadentate. Clypeal surface rugose in anterior part and laterally, disc of clypeus and frons densely punctate (punctures separated by less than a puncture diameter).

Pronotum more or less trapezoidal, about 2 times wider than long. Anterior and lateral margins distinctly bordered, base feebly bordered. Surface relatively sparsely punctate on disc (punctures separated by 2–3 puncture diameters) and more densely laterally.

Elytra trapezoidal, as wide as long, shiny. Stria relatively deep, with punctures slightly larger than striae width. Elytral intervals feebly convex, with minute punctuation.

Anterior tibiae have 3 outer teeth, without a small acute tooth between 1st outer tooth and apical spur. Anterior tibial spur bifurcated apically.

Pygidium with relatively thick borders and convex disc. Apical border almost as long as disc in the middle. Disc glabrous, punctate with punctures separated by 2–4 puncture diameters (Figure 13).

Parameres rounded in lateral view (Figure 20).

Female can be separated from male in having not bifurcated spurs of anterior tibiae and in sculpture of pygidium, which has very wide borders and very narrow disc with transverse keel (Figure 22).

Variability. Body length of the paratypes 4.0–6.2 mm.

Delopleurus darrenmanni
Figures 18–23. Delopleurus darrenmanni sp. n. 18, 21, habitus (18, male, 21, female); 19, 22, pygidium (19, male, 22, female); 20, aedeagus in lateral view and internal sac; 23, locality map.

Distribution

The species is known from three localities in a relatively small area between Zambezi and Okavango rivers in Zambezian woodland region (Figure 23).

Ethymology

The species is named after Darren Mann (UMO), who collected the type series.

Type material

Holotype, male with the labels “Namibia W. Caprivi Park Nova, 5 km N of Okavango River S18°09'56" E21°44'31" 19.xii.1999. In fungi. Coll. Mann, Marais & Newman”, “D.J.Mann colln. OX.UNI.MUS.NAT.HIST (OUMNH) OUMNH-2006-093”. Paratypes, 17 males and 6 females from Namibia, Zambia, and Botswana. Namibia: same data as the holotype, 13 males and 5 females (OUMNH), 2 males and 1 female (ZIN). BOTSWANA: 1 male with the label “Savuti Botswana S18.5651 E24.0629 Deschodt Tshikai 07.12.05” (UPSA). ZAMBIA: 1 male with the label “Kabula Loge, Zambia S17.041314 E024.013017 26-31.xi.2003 Deschodt & Groenewald” (UPSA).

Delopleurus gilleti Janssens, 1939

(Figures 24–29)

Delopleurus gilleti (Janssens, 1939)

Diagnosis

This species is most similar to male D. pullus but can be separated from it in having the pygidium with smaller disc which is punctate with bigger punctures (Figures 25, 28) and in having relatively densely punctate disc of pronotum (Figures 24, 27). This species does not show sexual dimorphism in the sculpture of the pygidium.

Description

Beetles are small-sized (3.5–5.0 mm), strongly convex, black or dark brown, glabrous.

Male (Figure 24). Clypeus quadridentate, medial teeth acute, lateral ones acute-angled. Clypeal margin laterad of lateral teeth somewhat angulate so clypeus looks hexadentate. Clypeal surface rugose in anterior part and laterally, frons densely punctate (punctures separated by less than puncture diameters).

Pronotum about 2 times wider than long. Anterior and lateral margins distinctly bordered, base not bordered. Pronotum densely almost uniformly punctate with punctures separated by less than a puncture diameter.

Elytra trapezoidal, as wide as long, shiny. Stria relatively deep, with punctures slightly larger than striae width. Elytral intervals convex, with minute punctuation.

Anterior tibiae have 3 outer teeth, without a small acute tooth between 1st outer tooth and apical spur. Anterior tibial spur bifurcated apically.

Pygidium with relatively narrow borders and feebly convex disc. Disc glabrous, coarsely punctate with punctures separated by about 1 puncture diameter (Figure 25).

Parameres in lateral view somewhat abruptly truncate, with distinct processes  (Figure 26).

Female (Figure 27) can be separated from male in having not bifurcated spurs of anterior tibiae. Sculpture of pygidium is similar to that of male (Figure 28).

Delopleurus gilleti
Figures 24–29. Delopleurus gilleti. 24, 27, habitus (24, male, 27, female); 25, 28, pygidium (25, male, 28, female); 26, aedeagus in lateral view and internal sac; 29, locality map.

Type material

Holotype, female with the labels. "Coll. R.I.Sc.N.B Togo Togo Coll. J.J.Gillet", "A. Janssens det., 1939 Delopleurus gilleti n. sp. type", "cf. Expl. P.N.A. G.F. de Witte (1933–1935) fasc 29, 1939, p. 130", "TYPE", and "A. Janssens det., 1941 gilleti A. Janssens" (IRSNB). Paratypes: male and female with the same locality data as the holotype (IRSNB); female, CONGO (Kinshasa), Kasai, forêt de Luebo, G. Babault (IRSNB).

Additional material

 CÔTE D’IVOIRE: National Park De La Comoe, Quango, Fitini, IX.1980, Y.Cambefort leg., 1 female (MNHN); Lamto-Savane, Girard leg., I.1968, 1 spm. (MNHN); Lamto, Girard leg., X.1968, 1 male (MNHN); Touba, Biemasso, 1 female (UPSA). MALI: Kita, 1 female (MNHN). NIGERIA: Yankari, 25.VII.1974, G.F.Bornemissza leg., 1 female (SANC); 10 km E of Kontagora, 19.VII.1974, G.F.Bornemissza leg., 14 females (SANC); Samaru, 20.VII.74 G.F.Bornemissza leg., 1 male (MNHN). MALAWI: Mlanje, S.A.Neave: 21.I.1913, 1 female, 6.II.1913, 1 female, 27.XII.1912, 1 male (BMNH); between Ft. Mangoche and Chikala Boma, 20-25.III.1910, S.A.Neave, 1 female (BMNH). NAMIBIA: Kaross, II.1925, 1 spm. (SAMC). TANZANIA: Kigonsera, 1 female (MNHN). ZAMBIA: Mwengwa, 1919, H.C.Dollman leg., 3 females (BMNH). MOZAMBIQUE: Canxixe, 1926, J. Surcouf leg., 1 male (MNHN).

Distribution

The species is widely distributed in the savannas both south and north from the equator (Figure 29).

Delopleurus krikkeni Frolov, sp. nov.

(Figures 30–33)

Diagnosis

This species can be easily separated from other Delopleurus species by sparsely punctate to impunctate disc and base of pronotum. Its pygidium is similar to that of D. fossatus but differs in having convex basal border distinctly widened medially (Figure 31). From D. fossatus it also differs in having larger eyes (in dorsal view).

Description

Holotype, male. Body strongly convex, black, glabrous, length 4.6 mm (Figure 30).

Clypeus quadridentate. Two medial teeth acute, lateral ones right-angled. Clypeal surface coarsely punctate in anterior part and laterally, frons densely punctate (punctures separated by 1–2 puncture diameters).

Pronotum about 2 times wider than long. Anterior and lateral margins distinctly bordered, base feebly bordered laterally. Disc and base of pronotum impunctate, sides relatively densely punctate.

Elytra trapezoidal, as wide as long, shiny. Stria relatively deep, with punctures 1.5–2 times larger than striae width. Elytral intervals convex, with minute punctuation.

Anterior tibiae have 3 outer teeth, without a small acute tooth between 1st outer tooth and apical spur. Anterior tibial spur bifurcated apically.

Pygidium with relatively strongly convex disc separate by deep circular fossa from border. The apical border wider medially and somewhat tapering laterally; apical border somewhat tuberculate in the middle (Figure 31).

Parameres in lateral view with small processes (Figure 32).

Paratype, male. Differs from the holotype in being slightly larger (length 4.8 mm) and having only anteromedial part of pronotal disc impunctate.

Female unknown.

Delopleurus krikkeni
Figures 30–33. Delopleurus krikkeni sp. n., holotype. 30, habitus; 31, pygidium; 32, aedeagus in lateral view and internal sac; 33, locality map.

Distribution

The locality where the holotype was collected is uncertain (Adrian Davis, pers. comm.). In 1896, "Samburu" might refer to a large area to the northwest of Mount Kenya and to the south of Lake Turkana, that was occupied by the Samburu people. But it might also be the railway station [S 3°46' E 39°16'] on the railroad Mombassa–Nairobi. This latter locality is some 100 km ENE of the locality where the paratype was collected, Mkomazi Game Reserve (Figure 33).

Ethymology

The species is named after Jan Krikken (Naturalis, Leiden).

Type material

Holotype, male with labels "Samburu [KENYA] 30.X to 20.XI.[18]96", "B. E. Africa C. S. Bretton 98-12", "diff. from the others Krikken 1975" (BMNH). Paratype, male: TANZANIA: Mkomazi GR, Ibaya Camp, unburnt grassland, pitfall traps, 7-8.IV.1995, J.G. Davies leg. (BMNH).

Remark

Two new species are described in this work from the Eastern Arc Region, D. krikkeni sp. n. and D. naviauxi sp. n. The former species is known from two males and the latter from two females only. I believe that these specimens are not conspecific because the male specimens lack a characteristic small tooth apicad of the first major outer tooth, which is found in the D. naviauxi holotype. Sex-dependent modifications of the legs are common in the Scarabaeinae but found in the males only. I thinks that this additional tibial tooth is sex-independent and should be present in the males of D. naviauxi.

Delopleurus fossatus Frolov, sp. n.

(Figures 34–39)

Diagnosis

This species is similar to D. krikkeni in having pygidium with deep circular fossa, but can be separated from it by having basal border of pygidium not or indistinctly widened medially, punctate disc of pronotum and smaller, slit-shaped dorsal parts of eyes.

Description

Holotype, male. Body strongly convex, black, glabrous, its length 5.3 mm (Figure 34).

Clypeus quadridentate. Two medial teeth acute, lateral ones obtuse. Clypeus rugose in anterior part and laterally, frons densely punctate (punctures separated by 1-3 puncture diameters).

Pronotum about 2 times wider than long. Anterior and lateral margins distinctly bordered, base feebly bordered. Pronotum densely punctate (punctures separated by 1–2 puncture diameters on disc, becoming denser laterally).

Elytra trapezoidal, as wide as long, shiny. Stria relatively deep, with punctures 1.5–2 times larger than striae width. Elytral intervals convex, with minute punctuation.

Anterior tibiae have 3 outer teeth, without a small acute tooth between 1st outer tooth and apical spur. Anterior tibial spur bifurcated apically.

Pygidium with strongly convex disc separated by deep circular fossa from borders. Apical and basal borders about the same thickness, not significantly thicker in the middle. Disc glabrous and coarsely punctate (Figure 35).

Parameres with small processes in lateral view (Figure 36).

Female can be separated from male in having not bifurcated spurs of anterior tibiae. Sculpture of pygidium is similar to that of male but the pygidium is 1/8–1/9 shorter (Figure 38).

Paratypes. Body length varies from 4.5 mm to 5.5 mm, otherwise the paratypes are similar to the holotype.

 Delopleurus fossatus
Figures 34–39. Delopleurus fossatus sp. n. 34, 37, habitus (34, male, 37, female); 35, 38, pygidium (35, male, 38, female); 36, aedeagus in lateral view and internal sac; 39, locality map.

Type material

Holotype, male with label "Nyassaland Mlanje [MALAWI, Mulanje] Feb. 11, 1913 S.A.Neave 1913-140" (BMNH). Paratypes: 1 male with the same data as the holotype; 13 specimens with the same data as the holotype except for the date of collecting: 27.II.1913, 1 female (BMNH), 24.II.1913, 1 female (BMNH), 14.II.1913, 1 female (ZIN), 7.III.1913, 1 male (BMNH), 19.XII.1912, 2 males (BMNH), 7.I.1913, 1 male (BMNH), 20.II.1913, 1 male (BMNH), 5.III.1913, 1 male (BMNH), 12.I.1913, 1 male (ЗИН), 1.II.1913, 1 female (BMNH), 8.II.1913, 1 female (BMNH), 21.I.1913, 1 female (BMNH).

Distribution

The species is known from Mlanje, Malawi (Figure 39) in southern Rift Valley.

Ethymology

The epithet is a Latin word denoting the sculpture of the pygidium.

Delopleurus parvus (Sharp, 1876)

(Figures 40–45)

Coptorhina parva ((Sharp, 1876)).

Delopleurus cardoni (Paulian, 1934), syn. n.

Diagnosis

This species differs from the other Delopleurus species in having bilobate glabrous disc of pygidium with very thick borders similar in both sexes (Figures 14, 44), and elytral striae with large punctures (Figures 40, 43).

Description

Beetles are small-sized (4.5–5.2 mm), strongly convex, black or dark brown, glabrous.

Male (Figure 40).Clypeus quadridentate, medial teeth acute, lateral ones right-angled. Clypeal surface rugose in anterior part and laterally, frons densely punctate (punctures separated by less than puncture diameter).

Pronotum about 2 times wider than long. Anterior and lateral margins distinctly bordered, base not or feebly bordered laterally. Surface of pronotum densely punctate anterolaterally, sparsely punctate with smaller punctures basally and anteromedially.  

Elytra trapezoidal, as wide as long, shiny. Stria fine, with punctures much larger than striae width. Elytral intervals slightly convex to almost flat, with minute punctuation.

Anterior tibiae have 3 outer teeth, without a small acute tooth between 1st outer tooth and apical spur. Anterior tibial spur bifurcated apically.

Pygidium with wide borders. Disc relatively small, somewhat bilobate, smooth and glabrous (Figure 44).

Parameres rounded in lateral view, without processes (Figure 42).

Female (Figure 43) can be separated from male in having not bifurcated spurs of anterior tibiae. Sculpture of pygidium is similar to that of male (Figure 44).

Delopleurus parvus
Figures 40–45. Delopleurus parvus. 40, 43, habitus (40, male, 43, female); 41, 44, pygidium (41, male, 44, female); 42, aedeagus in lateral view and internal sac; 45, locality map.

Type material

Delopleurus parvus — holotype, female with the labels "Ind. Bor.", "Laforte 5821", "ex Mus. D.Sharp 1890", "Museum Paris 1952 Coll. R. Obertür", "Coptorhina parva Type D.S." (MNHN).

Delopleurus cardoni — 3 syntypes with the label "Chota-Nagpore Palkot R.P.Cardon VII-VIII 1897" (MNHN).

Additional material

INDIA: Chota Nagpore Nowatoli R.P.Cardon VIII-IX 1896, 3 females, 6 spm. (IRSNB); Chota-Nagpore Nawatoli R.P.Cardon X.1897, 2 spm. (MNHN); Chota-Nagpore Nawatoli R.P.Cardon 1898, 3 spm. (MNHN); Chota-Nagpore Barway R.P.Cardon VI-VII.1897, 1 spm. (MNHN); Bangalore Chikmagalur Tabourel 1900, 1 spm. (MNHN); Pulney Hills R.P.Castets 1898, 3 spm. (MNHN). SRI LANKA: Coll.R.I.Sc.N.B Ceylan coll. Doherty Collection E.Candeze, 1 female (IRSNB).

Distribution

The species is known from a few localities in Indian Peninsula — south eastern Deccan Plateau and Chota Nagpur Plateau. Locality map (Figure 45) suggests that it can be associated with transitional zones between moist and dry deciduous forest ecoregions.

Remark

Paulian ((1934)) described Delopleurus cardoni from 9 specimens from Northern India ("Chota-Nagpore Nawatoli Palkot, R.P.Cardon VII-VIII.1897"). He separated D. cardoni from D. parvus on the basis of the punctate disk of pronotum in the former species and smooth in the latter. Comparison of the types of the both nominal species showed, however, that the both have relatively densely punctate disk of pronotum and do not differ significantly in other characters. The new synonymy is therefore established.

Delopleurus striatus Arrow, 1931

(Figures 46–49)

Delopleurus striatus (Arrow, 1931), syn. n.

Diagnosis

This species (known from a single female) differs from other Delopleurus species in having characteristic granulate disc of pygidium (Figure 48).

Description

Female. Body strongly convex, black, glabrous, length 5.1 mm (Figure 46).

Clypeus appear bidentate with two medial teeth acute (Figure 47). Clypeus rugose in anterior part and laterally, frons densely punctate (punctures separated by 1–3 puncture diameters).

Pronotum about 2 times wider than long. Anterior and lateral margins distinctly bordered, base bordered except in the middle. Pronotum densely punctate anterolaterally, sparsely punctate on the base and anterior part medially.

Elytra trapezoidal, as wide as long, shiny. Stria relatively deep, with punctures 1.5–2 times larger than striae width. Elytral intervals flat, with minute punctuation.

Anterior tibiae have 3 outer teeth, without a small acute tooth between 1st outer tooth and apical spur. Anterior tibial spur rounded apically.

Pygidium with relatively thick borders. Disc granulate, without setae (Figure 48).

Male unknown

Delopleurus spp., D. striatus, D. fossatus, Metacatharsius janssensi
Figures 46–51. Delopleurus spp. and Metacatharsius janssensi. 40, 50, habitus (46, D. striatus, 50, M. janssensi); 47, D. striatus, head and pronotum in dorsal view; 48, D. striatus, pygidium; 47, D. striatus, locality map; 51, D. fossatus sp. n., ventral view of head showing gula with distinct longitudinal groove.

Type material

Holotype, female with the labels "India. United Provs. Dehra Dun. 2,000 ft. viii.1927 H.G.Champion", "India H.G.Champion B.M. 1931-8", "Delopleurus striatus type Arrow", "Type", "SYNTYPE", "Delopleurus striatus Arrow M.E.Bacchus det 1975 SYNTYPE" (BMNH).

Distribution

The species is known from the single locality in northern periphery of the Indo-Gangetic Plain (Figure 49).

Metacatharsius janssensi (Frey, 1963), comb. nov.

(Figure 50)

Delopleurus janssensi Frey, 1963

Delopleurus janssensi was described from 2 specimens from Warder, Somali region of Ethiopia. The holotype, female (Figure 50), proved to be a member of the genus Metacatharsius Paulian. The paratype is similar to and apparently conspecific with the holotype. In the original description, the author compared this species with D. pullus and D. gilleti but did not provide characters specific for the genus Delopleurus. However the original description and locality data agree well with the type specimens. There is no doubt that Delopleurus janssensi was described from the misidentified Metacatharsius specimens and the new generic combination is therefore established here.

It is possible that M. janssensi is conspecific with another described species of the genus. There are up to 18 species of this genus known from adjacent Somalia ((Ferreira, 1972)) and the genus is currently a subject of taxonomic revision (François Génier, personal communication). It is out of the scope of the present work to specify the taxonomic status of M. janssensi more exactly.

Type material

Holotype, female with the labels: "Wardere Somalia", "Holotype", "TYPE", "Brit. Mus 1963-277", "Delopleurus janssensi det. G.Frey 1963, n. sp. Typus" (BMNH) and paratype with the labels: 'Wardere Somalia Oct. 1943, T.H.E.Jackson", "Delopleurus janssensi det. G.Frey 1963, p. Typus", "Cotype" (NHMB).

Acknowledgments

I am thankful to the colleagues who provided material and data for this work: Adrian Davis (UPSA), Alain Drumont (IRSNB), Bert Viklund (NHRS), Christian Deschodt (UPSA), Darren Mann (UMO), Eva Sprecher-Uebersax (NHMB), Kirstin Williams (DMAGD), Marc De Meyer (MRAC), Margie Cochrane (SAMC), Max Barclay (BMNH), Olivier Montreuil (MNHN), Riaan Stals (SANC), and Ruth Müller (TMSA). Special thanks are due to Marc De Meyer who hosted me at MRAC on two long-term occasions.

Funding

Russian Foundation for Basic Research [grants 10-04-00539-a, 13-04-01002-a]; Belgian Science Policy [postdoctoral fellowship].

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