Frolov, A.V., Scholtz, C.H. 2005. Revision of the southern African genus Frankenbergerius Balthasar with description of new taxa (Coleoptera: Scarabaeidae: Scarabaeinae). Journal of Natural History, 39, 2355-2377.

DOI: 10.1080/00222930500101829

Revision of the southern African genus Frankenbergerius Balthasar with description of new taxa (Coleoptera: Scarabaeidae: Scarabaeinae)

A.V. Frolov1, 2* & C.H. Scholtz1

1Department of Zoology and Entomology, University of Pretoria, Pretoria, 0002 South Africa.

2Laboratory of Insect Systematics, Zoological Institute, Russian Academy of Sciences, Universitetskaya nab., 1, St. Petersburg 199034, Russia.

*To whom correspondence should be addressed. Laboratory of Insect Systematics, Zoological Institute RAS, Universitetskaya nab., 1, St. Petersburg 199034, Russia. E-mail: afrolov@zin.ru

Abstract

The southern African genus Frankenbergerius Balthasar is revised. Two new species, F. opacus sp. n. and F. nitidus sp. n. (South Africa, Western Cape), and one new subspecies, F. armatus tuberculatus ssp. n. (South Africa, Mpumalanga), are described. F. imitativus (Péringuey) is considered a junior synonym of F. forcipatus (Harold). A key to the species and notes on biology are given.

Key words: Coleoptera, Scarabaeidae, Frankenbergerius, new species, new synonymy, key to species, South Africa.

Introduction

Southern Africa has a rich Scarabaeinae fauna with many groups retaining ancient Gondwanaland affiliations and relict distribution on the subcontinent (Halffter and Matthews, 1966; Cambefort, 1991; Davis, 1997). In the Scarabaeinae, two 'old' Gondwanaland tribes are recognized, the Canthonini and the Dichotomiini (Cambefort, 1991), however the latter is probably polyphyletic (Génier, 1996, Montreuil, 1998). The major deterrent to phylogenetic, biogeographic, ecological studies of the group is the lack of taxonomic data. While the African taxa of Canthonini are relatively well known (Scholtz and Howden, 1987a, b; Howden and Scholtz, 1987; Davis et al., 1999), many of the nominal genera currently placed in the Dichotomiini are still in need of taxonomic treatment. One of these poorly known groups is the genus Frankenbergerius Balthasar which was long considered a junior synonym of Coptorhina Hope.

The generic name Frankenbergerius was proposed by Balthasar (1938: 212) for one species, F. mirabilis Balthasar, described in the same paper from 'Columbien, Chiriguana'. The genus was, however, inadequately described from apparently mislabeled material and one year later Paulian (1939: 37) remarked that the description of F. mirabilis belongs to the African species Coptorhina armata (Boheman, 1857). Janssens (1939: 31) followed Paulian and listed Frankenbergerius as a junior synonym of Coptorhina.

Ferreira (1954: 7) erected a new genus, Pseudocoptorhina (type species C. armata (Boheman)), to accommodate five species previously placed in Coptorhina and a new one, P. gomesi, which differ from other Coptorhina in having triangular metepisterna and a bifurcated spur on the anterior tibiae in males. In a later publication, Ferreira (1972: 359) restored Frankenbergerius as a valid name since she recognized that Frankenbergerius and Pseudocoptorhina are based on the same type species and therefore the latter is a junior synonym of the former.

According to the original description the types of F. mirabilis were deposited in the author's collection, most of which is now housed in the National Museum of Natural History, Prague, although the older specimens may be deposited elsewhere (S. Bílý, pers. comm.). We were unable to trace the types. Since there is no evidence that F. mirabilis does occur in South America and because no statement in the original description of this speciescontradicts with characters of C. armata, we follow Paulian (1939) and Ferreira (1972) and consider Frankenbergerius a valid name for the taxon in question.

The genus differs from Coptorhina in a number of characters apart from those indicated by Ferreira (see below) and generic rank of this group is well supported. Both genera share a numbers of apomorphies with Delopleurus Erichsonand Sarophorus Erichson; Frankenbergerius is a putative sister group to the clade Coptorhina+Delopleurus.

The genus currently comprises seven species. Two new species and one new subspecies are described below; F. imitativus (Péringuey) is placed in synonymy. Although the peculiar shape of the clypeus can readily distinguish some of the species, others are still poorly described and inadequately keyed. Most of the species have not been illustrated, nor have their aedeagi been described or figured.

Material and methods

Little material was available to past researchers because representatives of the genus were very rare in collections. However, currently large numbers of specimens of some species are available for study in South Africa, most of which were collected over the past few decades by the staff of the Transvaal Museum, Pretoria (TMSA), chiefly by means of pitfall trapping. Other institutions from which material was borrowed for this study are: Albany Museum, Grahamstown (AMSA), The Natural History Museum, London (BMNH), Durban Museum, Durban (DMSA), Institut Royal des Sciences Naturelles de Belgique, Brussels (IRSNB), Musee Royal de l'Afrique Centrale, Tervuren (MRAC), Muséum National d'Histoire Naturelle, Paris (MNHN), National Collection of Insects, Plant Protection Research Institute, Pretoria (SANC), National Museum, Bloemfontein (BMSA), Naturhistoriska Riksmuseet, Stockholm (NHRS), South African Museum, Cape Town (SAMC), University of Pretoria Insect Collection, Pretoria (UPSA).

The distribution maps were generated with ArcView GIS 3.0 software (ESRI, Inc.). Co-ordinates were taken from the specimens' labels, if available, otherwise from the Alexandria Digital Gazetteer (fat-albert.alexandria.ucsb.edu:8827/gazetteer/).

Aedeagi were prepared according to the common technique used in entomological research and photographed in glycerol. Scanning electron micrographs were taken with a JEOL 840 electron microscope from uncoated specimens at low accelerating voltage (figures 9 and 10) and from the specimens coated with gold (figures 25 and 26). Outline figures are not to scale. Authors' comments are in square brackets.

Frankenbergerius Balthasar

Frankenbergerius Balthasar, 1938. Type species: F. mirabilis Balthasar, 1938: 212, by monotypy. Paulian, 1939: 37 (as synonym of Coptorhina); Janssens, 1939: 31 (as synonym of Coptorhina); Ferreira, 1972: 359.

Pseudocoptorhina Ferreira, 1954: 7; synonymy by Ferreira, 1972: 359.

Diagnosis.

Males with less developed clypeal processes and females of some species of Frankenbergerius, especially F. barratti (Waterhouse) with smooth elytra, are similar to species of Coptorhina. The two genera can immediately be distinguished, however, by the shape of the metepisternon. That of Frankenbergerius is triangular, widest in its anterior part, with slightly convex epipleural margin (figure 1), whiles Coptorhina's is somewhat rectangular, widest in its hind part, with secondary suture and very convex epipleural margin (figure 2). Elytra of Frankenbergerius are not fused along sutural margins, as opposed to Coptorhina, and have complex sculpture in most species. The two genera differ strongly in secondary sexual characters. In Coptorhina, sexual dimorphism is very weak; males differ from females in having the last abdominal sternite slightly convex. In Frankenbergerius,sexes can easily be separated by the shape of the spur of the anterior tibiae, which is simple, acute in females and bifurcated in males. Males of some species also differ from females in having curious horn-like clypeal processes. However the size of these horns, similar to other head processes in different groups of scarab beetles, is subject to much variation among individuals.

Fiigs. 1-12. Frankenbergerius spp. 1 – Frankenbergerius sp.; 2 – Coptorhina sp.; 3, 6, 8, 9, 11 – F. nanus; 4, 7, 10, 12 – F. opacus sp. n., holotype; 5 – F. gomesi. 1, 2 –scheme of thoracic pleurites (ms – metepisternon, s – secondary suture, ep – epipleuron); 3, 4 – pronoptum in lateral view; 5, 6 – anterior tibia of ♂; 7, 8 – habitus of ♂; 9, 10 – abdomen.

Figs. 1-12. Frankenbergerius spp. 1 – Frankenbergerius sp.; 2 – Coptorhina sp.; 3, 6, 8, 9, 11 – F. nanus; 4, 7, 10, 12 – F. opacus sp. n., holotype; 5 – F. gomesi. 1, 2 –scheme of thoracic pleurites (ms – metepisternon, s – secondary suture, ep – epipleuron); 3, 4 – pronoptum in lateral view; 5, 6 – anterior tibia of ♂; 7, 8 – habitus of ♂; 9, 10 – abdomen; 11, 12 – aedeagus in dorsal and lateral view.

Description.

Small to medium-sized beetles (length 4,5-16 mm, width 3.1-16 mm). Colour monotonous black to dark brown, sometimes anterior part of clypeus, legs or elytra slightly paler. Head and pronotum densely punctate and pubescent (except for F. barratti), in some species most of pronotum rugose. Each puncture, except for those of elytral striae, bears a short yellowish seta. Setae sometimes abraded on disc of pronotum and head.

Clypeus deeply sinuate in middle, angulate to dentate at sides in females and with more or less developed horns in males of most species. In some species anterior angles of clypeus with short carina directed proximally. Genae rectangular to rounded. Frontoclypeal suture interrupted on disc. Eyes small, the dorsal part slit-shaped, ventral part sub-rectangular. Distance between eye and gula approximately 2 times the width of eye in ventral view. Gula with longitudinal groove.

Pronotum more or less parallel-sided, wider than long. Anterior margin and base not bordered in most species; lateral margins bordered.

Elytra not fused, with humeral umbones, sinuate laterally near base. Elytral intervals flat to convex in apical part, with tubercles in most species. Scutellum not visible from above. Wings fully developed.

Anterior tibiae have 3 outer teeth with smaller intermediate teeth between (except for F. nanus, F. opacus sp. n. and F. nitidus sp. n.). Outer margins of middle and posterior tibiae without transverse keels, serrate.

Pygidium punctate on disc, bordered; with small longitudinal keel in F. opacus sp. n.

Parameres symmetrical, their apices strongly to feebly sclerotized, without setae. Armature of internal sac of aedeagus is similar in most species.

The immature stages are unknown.

Diagnostic characters.

The most important diagnostic character of Frankenbergerius species is the sculpture of the dorsal surface of the body and especially the elytra. The shape of the parameres is species-specific but in some individuals the character may be ambiguous. The shape of the clypeus in males is distinctive for some species (except for F. nanus, F. opacus sp. n. and F. nitidus sp. n.) in which clypeal processes are strongly developed.

Biology

Little is known of Frankenbergerius biology although inferences can be made from information recorded with museum specimens. The beetles have been collected most frequently in dense vegetation in association with litter and decomposing plant matter. Several records imply a close association with mushrooms. As opposed to Coptorhina, which has been studied in more detail and is known to be an obligatory basidial mushroom eater (Tribe, 1976), Frankenbergerius presumably retained a more ancestral life style with no strict preference to mushrooms but rather to any rotten plant matter. The largest series of the beetles (F. armatus, 31 specimens) was collected in Silaka Forest Reserve (Eastern Cape Province) in rotten Cussonia fruit.

Frankenbergerius specimens have been collected by means of pitfall traps baited with different types of bait (banana, meat, and faeces) but this does not necessarily imply that the specimens were attracted to the baits. Very long trap exposures (up to 68 days) suggest that the beetles might be captured occasionally along with other litter dwellers. The only indication that Frankenbergerius specimens might be attracted to dung is one specimen of the rare species F. nanus collected near Darling (Western Cape Province) in a pitfall trap with fresh cattle dung which was exposed for 24 hours. However, in some 15 months of trapping in the area (10 traps on 3 occasions per month), this was the only Frankenbergerius specimen recorded so, this was probably a chance trapping (Adrian Davis, pers. comm.). No specimens have been collected in dung pads, nor are there direct indications that specimens are attracted to carrion.

Mycetophagy has been hypothesized to be the ancestral feeding type of the Scarabaeinae, with a change to dung in most taxa evolving much later (Scholtz and Chown, 1995). However, feeding on the higher fungi, the mushrooms, is more likely to be a later change from humus or dung to a more nutritious substance. In the lineage to which the genus Frankenbergerius belongs, the genera Coptorhina and Delopleurus, the obligatory mushroom eaters, represent the most derived taxa with the greatest number of apomorphies, while the genus Sarophorus Erichson, which feeds on dung, humus, and carrion, largely retained plesiomorphic character states (Frolov and Scholtz, unpublished).

Nesting behavior of Frankenbergerius has not been studied but it can be assumed that it is similar to related taxa. Coptorhina specimens have been observed feeding on mushrooms with two types of fruit-body. In the case of  "puff-ball" mushrooms the adults burrow into the fruit-body, detach pieces and drag them into their burrows. In "parasol" mushrooms, the beetles climb the stalk and detach pieces of the gills, which they then drag into their burrows. Brood balls are constructed from the macerated pieces, eggs are laid and the balls coated with soil. Because males of some species of Frankenbergerius have long clypeal horns, they are probably unable to burrow into a fresh mushroom or detach pieces of it and probably do not take part in brood ball construction. Field observations are needed to clarify the genus' nesting behavior.

Frankenbergerius specimens are apparently not attracted to light and are presumably day-fliers.

Figs. 13-22. Frankenbergerius spp. 13 – F. nitidus sp. n., holotype; 14-16, 22 – F. armatus armatus; 17-21 – F. armatus tuberculatus ssp. n. 13, 21, 22 – habitus (13 - ♀, 21, 22 - ♂; 21 – holotype); 14-20 – head (14, 15, 17-19 – ♂, 16, 20 – ♀).

Figs. 13-22. Frankenbergerius spp. 13 – F. nitidus sp. n., holotype; 14-16, 22 – F. armatus armatus; 17-21 – F. armatus tuberculatus ssp. n. 13, 21, 22 – habitus (13 - ♀, 21, 22 - ♂; 21 – holotype); 14-20 – head (14, 15, 17-19 – ♂, 16, 20 – ♀).

Key to Frankenbergerius species

1. Disc of pronotum densely punctate. Body smaller (4,5-10 mm) ............ 2

― Disc of pronotum smooth. Body larger (8.5-16 mm) ...... F. barratti (Waterhouse)

2. Anterior tibiae with smaller teeth between major outer teeth (figure 5). Anterior margin of pronotum usually distinctly sinuate (very feebly or not sinuate in F. gomesi). Eastern Cape, KwaZulu-Natal, Mpumalanga, Northern provinces ....... 3

― Anterior tibiae without distinct teeth between major outer teeth (figure 6). Anterior margin of pronotum not sinuate. Western Cape Province ...... 5

3. Elytral interval 8 with long carina occupying more than basal half of elytron. Length 4.0-7.3 mm ...... F. gomesi (Ferreira)

― Elytral interval 8 with a row of cariniform tubercles or with 2 carinae widely separated in the middle of elytron ...... 4

4. Elytral intervals 2 to 7 with smaller to indistinct tubercles (figure 28). Apices of parameres widened in lateral view (figure 37). Length 8.0-10.2 mm ...... F. forcipatus (Harold)

― Elytral intervals laterad of 1st or 2nd usually with distinct tubercles (figure 21, 22). Apices of parameres not widened in lateral view (figure 23, 24). Length 4.6-11.0 mm. ...... F. armatus (Boheman)

Figs. 23-27. Frankenbergerius armatus. 23, 25 – F. armatus armatus, 24, 26 - F. armatus tuberculatus ssp. n. (23, 24 - aedeagus in dorsal and lateral view, 25, 26 – elytron); 25 – distribution map of F. armatus.

Figs. 23-27. Frankenbergerius armatus. 23, 25 – F. armatus armatus, 24, 26 - F. armatus tuberculatus ssp. n. (23, 24 - aedeagus in dorsal and lateral view, 25, 26 – elytron); 25 – distribution map of F. armatus.

5. Pygidium without longitudinal carina in basal part. Disc of meso- and metasternum and dorsal surface of femora smooth to finely punctate. Base of pronotum not concave in lateral view (figure 3) ..... 6

― Pygidium with small longitudinal carina in basal part (figure 10). Disc of meso- and metasternum and dorsal surface of femora densely punctate. Base of pronotum slightly concave in lateral view (figure 4). Length 5.8-7.1 mm. ...... F. opacus sp. n.

6. Last abdominal sternite with transverse convexity near pygidium (figure 9). Elytral intervals matt with shiny, elongated tubercles (figure 8). Length 5.5-6.5 mm. ...... F. nanus (Péringuey)

― Last abdominal sternite flat. Elytral intervals shiny with round to elongated tubercles (figure 13). Length 7.5 mm ..... F. nitidus sp. n.

Frankenbergerius barratti (Waterhouse)

(Figures 29–32, 38, 43)

Figs. 28-38. Frankenbergerius spp. 28, 33-37 – F. forcipatus; 29-32, 38 – F. barratti. 28, 29 – habitus of ♂; 30-36 – head (30, 31, 33-35 – ♂, 32, 36 – ♀), 37, 38 - aedeagus in dorsal and lateral view.
Figs. 28-38. Frankenbergerius spp. 28, 33-37 – F. forcipatus; 29-32, 38 – F. barratti. 28, 29 – habitus of ♂; 30-36 – head (30, 31, 33-35 – ♂, 32, 36 – ♀), 37, 38 - aedeagus in dorsal and lateral view.

Coptorhina barratti Waterhouse 1876, p 22; Péringuey 1901, p 278, 293.

Pseudocoptorhina barratti (Waterhouse): Ferreira 1954, p 8.

Frankenbergerius barratti (Waterhouse): Ferreira 1972, p 360, 363.

Diagnosis.

This species can easily be separated by its large size (8.5-16 mm) and almost smooth upper side of body. Males usually have long and slender clypeal horns (figure 30).

Description.

Black, shiny beetle (figure 29). Body length of males 8.5-16 mm, width 7.5-13 mm, of females 8.6-15 mm and 6.7-11 mm. Dorsal surface without visible setae.

Head. Clypeus of males with two long, acute horns curved upwards; horns 1-0.5 times the length of the head (figures 30, 31). In females, clypeus with protruding anterior angles separated by deep sinuation (figure 32). Genae right-angled, finely bordered. Genal sutures distinct. Lateral margin of clypeus distinctly sinuate near genal suture in males but without sinuation in females. Frontal suture obsolete. Head densely punctate except for disc which is smooth and slightly convex. Eyes small, almost completely divided by canthus, dorsal parts slit-shaped.

Pronotum with subparallel sides, 2 times wider than long. Anterior margin sinuate medially, anterior angles slightly sinuate laterally. Disc smooth, anterior and lateral parts densely punctate (puncture separated by 1-2 puncture diameters).

Elytra. Striae distinct, punctate (punctures separated by 3-4 puncture diameter). Intervals slightly convex, smooth, without tubercles.

Underside. Pygidium with deep border, its disc densely punctate. Abdominal and thoracic sternites coarsely punctate except for disc of metasternum.

Legs. Anterior tibiae with smaller teeth between major outer teeth in most specimens. Intermediate teeth can be obsolete in older individuals. Spur of anterior tibia bifurcated in males and simple, acute in females.

Aedeagus. Parameres with acute apices in lateral view (figure 36).

Variability. Shape and length of clypeal processes in males vary (figures 30, 31) otherwise variation among examined specimens is very slight. Females differ from males in not having bifurcated spur of anterior tibiae and in the shape of the clypeus (figure 32).

Distribution. This rare species is known from a few localities in South Africa and Lesotho (figure 43).

Type material examined. Holotype ♂ with labels 'Type', 'Transvaal', and 'Coptorhina Barratti, C. Waterh. Type' (BMNH).

Additional material examined. Mpumalanga: Uitsoek, Grootkloof Indigenous Forest, 25°15'S 30°33'E, 15.XII.1986, pitfall traps, 1 ♂, S. Endrödy-Younga leg. (TMSA); Road Barbeton to Bulembu near Swaziland border [25°55'S 31°10'E], 20.I.2000, 1 ♀ (TMSA). Gauteng: Pretoria [25°45'S 28°10'E], 1 ♀ (BMNH). KwaZulu-Natal: Durban [29°50'S 31°01'E], 1 ♂ (BMNH); Farm Boschberg, 50 km N of Ladysmith 28°12'S 29°48'E, 9-11.I.2001, A. Davis leg., 1 ♀ (UPSA). Lesotho: Maleata [not traced], III.1944, H. K. Munro leg., 3 ♂ and 1 ♀ (TMSA), 3 ♂ and 2 ♀ (BMNH). 'Transvaal”, 1 ♂, (BMNH), 1 ♂ and 1 ♀ (IRSNB).

Frankenbergerius gomesi (Ferreira)

(figures 39-43)

Figs. 39-43. Frankenbergerius spp. 39-42 – F. gomesi (39 – habitus of ♂; 40 – aedeagus in dorsal and lateral view; 41, 42 – head of ♂ and ♀ respectively); 43 – distribution map of Frankenbergerius.

Figs. 39-43. Frankenbergerius spp. 39-42 – F. gomesi (39 – habitus of ♂; 40 – aedeagus in dorsal and lateral view; 41, 42 – head of ♂ and ♀ respectively); 43 – distribution map of Frankenbergerius.

Pseudocoptorhina gomesi Ferreira, 1954: 10.

Frankenbergerius gomesi (Ferreira): Ferreira, 1972: 362.

Diagnosis.

This species can be distinguished by elytral interval 8 with a long carina occupying 2/3 of the length of the elytron and by its small body size.

Description.

Black to dark brown, shiny beetle (figure 39). Body length of males 4.6-7.2 mm, width 2.5-3.2 mm, of females 4-7.3 mm and 2.4-3.9 mm. Dorsal surface with minute setae which may be abraded on most surfaces.

Head. Clypeus with two horn-like processes; processes slightly longer in males (figures 41, 42). Genae obtuse, finely bordered. Genal sutures distinct. Lateral margin of clypeus with deep sinuation near genal suture. Frontal suture obsolete medially. Dorsal surface of horn-like processes granular, rest of head densely punctate except for middle area of clypeus which is smooth. Eyes small, almost completely divided by canthus, their dorsal parts slit-shaped.

Pronotum narrower in anterior part, 1.6 times wider than long. Anterior margin not sinuate medially, or with minute unclear sinuation. Lateral margins and anterior angles finely bordered, anterior margin and base not bordered. Surface coarsely punctate with more or less elongated punctures.

Elytra. Striae distinct, punctate (punctures separated by 3-4 puncture diameter). Intervals indistinctly punctate on disc, with elongated tubercles. Interval 8 with long carina occupying 2/3 of elytron.

Underside. Pygidium with deep border, its disc densely punctate. Abdominal and thoracic sternites coarsely punctate except for disc of metasternum.

Legs. Anterior tibiae with smaller teeth between major outer teeth in most specimens. Spur of anterior tibia bifurcated in males and simple, acute in females.

Aedeagus. Parameres with angulate apices in lateral view (figure 40).

Variability. Except for body size, the specimens examined show very little variation. Females differ from males in not having bifurcated spur of anterior tibiae and clypeus with somewhat shorter processes with smaller sinuation between (figure 41, 42).

Distribution. This species is known from a number of localities in Limpopo, Mpumalanga, and KwaZulu-Natal provinces.

Type material examined. Holotype ♂ with labels 'Uitzicht, Zoutpansberg [Soutpansberg], Nov[ember] 1924, Heske [leg.]' and 'Holotypus Pseudocoptorhina gomesi sp. n. M.C.Ferreira 1954' (TMSA).

Additional material examined. Mpumalanga: Nelspruit Nature Reserve, 25°29'S 30°55'E, 18.XII.1986, dry valley, pitfall traps, 53 days, 4 ♂ and 5 ♀; rivulent valley, pitfall traps, 4 ♂ and 2 ♀; litter, riverine bush, 2 ♂ and 1 ♀; Koppie, 2 ♂ and 3 ♀; 9.II.1987, dry valley, pitfall traps, 58 days, 13 ♂ and 24 ♀; rivulent valley, 3 ♂ and 1 ♀; 25.X.1986, rivulent valley, pitfall traps, 34 days, 1 ♂ and 3 ♀; dry valley, pitfall traps, 34 days, 4 ♂ and 1 ♀; 29.XI.1986, dry valley, pitfall traps, 19 days, 5 ♂ and 7 ♀, S. Endrödy-Younga leg., (TMSA). 18 km S of Nelspruit, 25°37'S 30°58'E, 10.II.1987, pitfall traps, 57 days, 2 ♂, S. Endrödy-Younga leg. (TMSA). 16 km N of Barbeton, 25°42'S 30°57'E, 24.X.1986, pitfall traps, 31 days, 1 ♀; 30.XI.1986, pitfall traps, 53 days, 1 ♀; pitfall traps, 57 days, 1 ♂ and 2 ♀, S. Endrödy-Younga leg. (TMSA). 17 km NNW of Barbeton, 25°36'S 29°53'E, 10.XI.1980, 1 ♂, S. Endrödy-Younga leg. (TMSA). Nelspruit, botanical garden, 25°31'S 30°32'E, 24.I.1981, pitfall traps, 44 days, 8 ♂ and 8 ♀; 6.XII.1980, pitfall traps, 45 days, 3 ♂ and 3 ♀, S. Endrödy-Younga leg. (TMSA). Nelspruit, I.1939, 1 ♀ (SAMC). Road between Sabie and Sudwala Caves [25°05'S 30°40'E], 25.I.1996, fungi in eucalyptus forest next to road, 1 ♂, I. Pajor leg. (SANC). Hazyview, 25°04'S 31°07'E, 27.I.1996, forest litter, 3 ♀ (TMSA). White river Distr., Farm Lichtfontein [25°19'S 31°01'E], 28.XII. 1992, 2 ♂, R. Müller leg. (TMSA). KwaZulu-Natal: Hluhluwe Game Reserve, 28°05'S 32°04'E, 18.I.1992, intercept trap, open biotope, 2 ♀; 20.I.1992, fungous trunk and litter, 1 ♂, S. Endrödy-Younga leg. (TMSA). Empangeni [28°45'S 31°55'E], X.1976, 1 ♀; XI.1976, 1 ♀; XII.1976, 1 ♀, P. E. Reavel leg. (TMSA).

Frankenbergerius forcipatus (Harold)

(figures 28, 33-37)

Coptorhina forcipata Harold, 1881: 149; Péringuey,1908: 627; Janssens, 1939: 32, 36.

Pseudocoptorhina forcipata (Harold): Ferreira, 1954: 9.

Frankenbergerius forcipatus (Harold): Ferreira, 1972: 360, 363.

Coptorhina imitativa Péringuey, 1901: 288; Janssens, 1939: 32, 36; Ferreira, 1954: 10 (as Pseudocoptorhina); Ferreira, 1972: 362, 363 (as Frankenbergerius); syn. n.

Diagnosis.

This species is similar to F. armatus but can be separated from it by having elytral intervals 1-4 smooth or with only traces of tubercles. It also differs in the shape of the parameres (figure 37).

Description.

Black to dark brown, shiny beetle (figure 28). Body length of males 8.1-10.2 mm, width 5.5-6.1 mm, of females 8-9.5 mm and 5.4-6 mm. Dorsal surface with fine setae.

Head. Clypeus with two horn-like processes; processes sometimes longer in males (figures 33-36). Genae obtuse, finely bordered. Genal sutures distinct. Lateral margin of clypeus slightly sinuate near genal suture or without sinuation. Frontal suture obsolete medially. Dorsal surface of horn-like processes granular, other part of head densely punctate except for middle area of clypeus which is smooth. Eyes small, almost completely divided by canthus, their dorsal parts slit-shaped.

Pronotum narrower in anterior part, 1.8 times wider than length. Anterior margin not sinuate medially, or with minute unclear sinuation. Lateral margins and anterior angles finely bordered, anterior margin and base not bordered. Punctures separated by a puncture diameter on disc, becoming denser and somewhat elongated laterally.

Elytra. Striae distinct, punctate (punctures separated by 3-4 puncture diameter). Intervals 1 and 2 without tubercles; intervals 3-7 with small tubercles; interval 8 with long keel interrupted at middle of elytron; interval 9 with smaller keel situated in the middle of elytron.

Underside. Pygidium with deep border, its disc densely punctate. Abdominal and thoracic sternites coarsely punctate except for disc of meso- and metasternum.

Legs. Anterior tibiae with smaller teeth between major outer teeth in most specimens. Spur of anterior tibia bifurcated in males and simple, acute in females.

Aedeagus. Parameres with widened apices in lateral view (figure 37).

Variability. Except for body size there is some variation in the length of the clypeal processes in males (figures 33-35). Females can be separated by acute, not bifurcated spur of anterior tibiae and in most cases by shorter clypeal processes (figure 36).

Distribution. This species is known from a few distant localities in Magaliesberg (Northwest Province), southern Drakensberg and Ciskei (Eastern Cape Province) (figure 43).

Type material examined. F. forcipatus: holotype ♂ with labels 'Cap.' and 'forcipata type Harold' (MNHN). F. imitativus: holotype ♂ with labels 'Coptorhina imitativa type [18]97' and '474' (SAMC).

Additional material examined. Northwest Province: Rustenburg [25°40'S 27°15'E], 14.I.1895, 1 ♀ (SANC). KwaZulu-Natal Province: Drakensberg, Cathedral Peak, forest, 28°56'S 29°12'E, 13.III.1976, pitfall traps, 7 days, 1 ♂ and 2 ♀; 15.III.1976, pitfall traps, 5 days, 3 ♀, S. Endrödy-Younga leg. (TMSA). Giant's Castle [29°14'S 29°29'E], XII.1979, 1 ♂, C. Scholtz leg. (UPSA). Eastern Cape Province: Pirie Bush 2 ♂ [? Pirie Forest, 27°14'E 32°45'S] (BMNH). "Promontorium Bonae Spei, Transvaal", 2 ♂ (BMNH).

Remark. C. forcipata was unknown to Péringuey when he described C. imitativa and his later record of the former species (Péringuey 1908: 627) is based on its original description. Examination of the type specimens of the two nominal species shows no noticeable differences in the punctuation of the pronotum nor sculpture of the elytra, contrary to the characters given in the keys by Janssens (1939: 32) and Ferreira (1954: 8, 9).

Frankenbergerius armatus (Boheman)

(figures 14-27)

Epirhinus armatus Boheman, 1857: 200.

Coptorhina armata (Boheman): Harold, 1872: 205; Péringuey, 1901: 288, 294; Janssens, 1939: 32, 36.

Frankenbergerius armatus (Boheman): Ferreira, 1972: 362.

Coptorhina granulifera Harold, 1871: 112; synonymy by Harold, 1872: 205.

Frankenbergerius mirabilis Balthasar, 1938: 212; synonymy by Paulian, 1939: 37 (as Coptorhina).

Diagnosis.

This species is most similar to F. forcipatus but can be separated by its smaller size, by having elytral intervals 2-4 with more distinct tubercles and by the parameres narrower in lateral view (figures 23, 24).

Polymorphism.

In the examined material, there are two distinct forms of males: one with clypeal horns curved upwards and slightly backwards (figures 14, 22) with somewhat truncate apices, and another form with acute horns curved upwards but not backwards (figures 17, 21). An additional, although minor difference, is the less tuberculate elytra in the first form (figures 25, 26). The form with truncate apices of clypeal horns was collected in the Eastern Cape and KwaZulu-Natal provinces and the other one in Mpumalanga and Northern Province (figure 27).

The distinctness of the form with acute clypeal horns from 'typical' F. armatus was apparently recognized by M. Ferreira; one such specimen in the TMSA bears the label 'Type ♂ Pseudocoptorhina armata var. tuberculata nov. M.G.Ferreira 1954'. However, the name was not published.

Because of the allopatric distribution and distinct morphological differences between males with well developed horns we treat these two forms as subspecies. The absence of noticeable differences in paramere shape and ambiguous differences in elytral sculpture (characters that are normally distinctive for other Frankenbergerius species) prevent us from assigning them specific rank. However, if further research shows reproductive isolation or sympatric distribution, their rank should be reconsidered.

Remark. Harold (1871: 7) described C. granulifera from Port Natal [Durban]. Later he transferred E. armatus to Coptorhina and wrote that C. granulifera belongs to this species (Harold, 1872: 205). We did not have the opportunity to examine the type of C. granulifera, however the collection locality suggests that it is the nominotypical subspecies of F. armatus, to which the type of C. granulifera belongs.

Frankenbergerius armatus armatus (Boheman)

(figures 14-16, 22, 23, 25, 27)

Diagnosis.

This subspecies differs from F. armatus tuberculatus by the clypeal horns of males that are curved upwards and backwards and truncate at the apices.

Description.

Male (figure 22). Dorsal surface of pronotum, elytra and head with small yellowish setae. Most of setae may be abraded; in most specimens setae absent on disc of head and pronotum.

Head. Clypeus in some individuals with horns curved upwards and slightly backwards (Fig. 14) with somewhat truncate apices. Genae obtuse, finely bordered. Genal sutures visible as fine lines. Lateral margin of clypeus not, or very feebly, sinuate near genal suture. Frontal suture broadly interrupted at middle. Area behind genae slightly concave, coarsely punctate with big adjoining punctures, with distinct margins. Genae and anterior part of clypeus densely punctate; margins of punctures indistinct. Very anterior part of clypeus with sculpture nearly rugose. Disc of head with minute, feebly visible punctuation. Inner (dorsal) surface of clypeal horns finely granular, outer (ventral) surface smooth. Anterior margin of clypeus flattened, deeply sinuate at middle, slightly rugose.

Pronotum. Anterior angles rounded. Lateral margins with fine border, hind and anterior angles and anterior margin with wider border. Border of anterior margin widened and sinuate at middle. Base not bordered. Hind angles obtuse. Dorsal surface densely and coarsely punctate; punctures separated by 0.2-0.3 their diameters on disc, becoming denser laterally.

Elytra. Striae fine on disc becoming slightly wider and deeper apically, shiny, punctate (punctures separated by 3-4 puncture diameter). Intervals flat on disc, feebly convex in apical part, smooth to slightly shagreened and punctate (punctures separated by 1.5-2 puncture diameter). Intervals 2-7 with longitudinal tubercles, less developed than those in F. armatus tuberculatus. Interval 8 with long keel interrupted at middle of elytron. Interval 9 with smaller keel situated at about middle of elytron. Keels, especially hind part of keel on interval 8, serrate in most individuals.

Underside. Pygidium with deep border, its disc densely punctate. Abdominal and thoracic sternites coarsely punctate except for disc of meso- and metasternum.

Legs. Anterior tibiae with smaller teeth between major outer teeth in most specimens. Spur of anterior tibia bifurcated in males and simple, acute in females.

Aedeagus. Similar to F. armatus tuberculatus (figure 23).

Variability. Body length of males 4.6-8.2 mm, width 3.5-5.2 mm, of females 6.2-8.3 mm and 3.9-5.0 mm. Colour of body from black to dark brown. Length of clypeal processes varies, some males have shorter clypeal processes (figure 15). Females can be separated by acute, not bifurcated spur of anterior tibiae and, in some cases, by shorter clypeal processes (figure 16).

Distribution. The subspecies is distributed is Eastern Cape and KwaZulu-Natal provinces up to 29°S in the north (figure 27).

Type material examined. Lectotype (here designated): ♂ with labels 'Caffraria', 'J. Wahlb[erg]', 'armata Boh.', and '3636 E92' (NHRS). Paralectotypes: ♂ with labels 'Caffraria', 'J. Wahlb', 'Type', 'armatus Bhm.', 'Typus', '157 54', '3632 E92'; 3 ♀ with labels 'Caffraria', 'J. Wahlb' and catalogue numbers '3633 E92', '3634 E92', and '3635 E92' (NHRS).

In the original description, Boheman (1857: 200) did not designate the holotype nor did he indicate the number of specimens examined. However the size range provided (length 7.5-8.0 mm, width 5.0-5.2 mm) suggests that a few specimens were studied. The five specimens collected by Wahlberg fit these measurements well and might have all been examined by Boheman. One of these specimens bears a label 'typus', but it has less developed clypeal horns (and, its left horn is broken) to class it unequivocally as the subspecies armatus. The locality label 'Caffraria' suggests that this specimen could have been collected in the natural habitat of either of the two subspecies. To ensure stability of the nomenclature, another male specimen which has the explicit features of F. armatus armatus is here designated as the lectotype.

Additional material examined. KwaZulu-Natal Province: Durban [29°50'S 31°01'E], 4 ♂ and 6 ♀, (BMNH), 4 ♂ and 3 ♀ (IRSNB). uMvoti [29°09'S 30°45'E], 1 ♂, H. Fry leg. (SAMC). Eastern Cape Province: Ntsubane Forest, 31°27'S 29°44'E, 25.XI.1988, pitfall traps, 14 days, 8 ♂ and 12 ♀; fungi and forest litter, 1 ♀; fungous tree trunks, 1 ♀; 26.XI.1988, pitfall traps, 14 days, 1 ♀ and 2 ♂; 1.XII.1988, forest litter, 1 ♀, S. Endrödy-Younga leg. (TMSA). Dwesa Forest Reserve, 32°17'S 28°50'E, 26.II.1985, pitfall traps, 7 days, 12 ♀ and 9 ♂; grassnetting, day, 1 ♀ and 1 ♂; 27.II.1985, sifting, indigenous forest litter, 2 ♂; 11.XII.1979, sifting, forest litter, 2 ♂ and 3 ♀; 12.XII.1979, owl pellet, 1 ♀; 17.XII.1979, 1 ♂, S. Endrödy-Younga leg. (TMSA). Silaka Forest Reserve, 31°33'S 29°30'E, 24.XI.1987, pitfall traps, 8 days, 1 ♂; indigenous forest litter, 1 ♂ and 1 ♀; 24.XI.1988, rotten Cussonia fruit, 14 ♂ and 17 ♀; 24.XI.1988, ground traps, 8 day, 4 ♂ and 2 ♀, S. Endrödy-Younga leg. (TMSA). Amatole, Isidenge Forest Station, 32°41'S 27°15'E, 16.XI.1987, Quercus forest litter, 19 ♂ and 5 ♀; Quercus and Eucalypt forest, fungi, 6 ♂ and 7 ♀; 14.XI.1987, indigenous forest litter, 1 ♀; 12.XI.87, dead Quercus bark, 3 ♂; 18.XI.1987, Pinus forest litter, 1 ♂, S. Endrödy-Younga leg. (TMSA). Alexandria Forest Station, 33°44'S 26°23'E, 5.XII.1987, pitfall traps with banana bait, 2 days, 3 ♀; 6.XII.1987, indigenous forest litter, 1 ♀, S. Endrödy-Younga leg. (TMSA). Grahamstown [33°18'S 26°32'E], II.1978, 1 ♂ and 1 ♀, C. Scholtz leg. (UPSA). Port Saint John's [31°38'S 29°32'E], XI.1923, 1 ♂ and 1 ♀ (BMNH).

There are two series of this subspecies in TMSA with doubtful locality data: 6 ♂ and 3 ♀ from Nelshoogte, Knuckles rocks forest, 25°47'S 30°50'E, 24.X.86, S. Endrödy-Younga leg., and 11 ♂ and 15 ♀ with the label 'Probably Uitsoek'. They were apparently mislabeled and the former series probably originated from Dwesa Forest Reserve (Ruth Müller, pers. comm.).

Frankenbergerius armatus tuberculatus ssp. n.

(figures 17-21, 24, 26, 27)

Diagnosis.

This subspecies differs from the nominotypical one chiefly by the shape of the clypeal horns in males: the horns curve upwards, with acute apices (figure 17, 21).

Description.

Holotype ♂ (figure 21). Body length 9.5 mm, width 5.5 mm. Dorsal surface of pronotum, elytra and head with minute setae.

Head. Clypeus with two horns slightly curved upwards; horns acute at apices. Genae obtuse, finely bordered. Genal sutures visible as fine lines. Lateral margin of clypeus not or very feebly sinuate near genal suture. Frontal suture broadly interrupted at middle. Area behind genae slightly concave, coarsely punctate with big adjoining punctures with distinct margins. Genae and anterior part of clypeus densely punctate; margins of punctures indistinct. Very anterior part of clypeus sculpture nearly rugose. Disc of head with minute, feebly visible punctuation. Inner (dorsal) surface of clypeal horns finely granular; outer (ventral) surface smooth. Anterior margin of clypeus flattened, deeply sinuate at middle, slightly rugose.

Pronotum. Anterior angles rounded. Lateral margins with fine border, hind and anterior angles and anterior margin with wider border. Border of anterior margin widened and sinuate at middle. Base not bordered. Hind angles obtuse. Dorsal surface densely and coarsely punctate; punctures separated by 0.2-0.3 their diameters on disc, becoming denser laterally.

Elytra. Striae fine, becoming slightly wider and deeper apically, shiny, punctate (punctures separated by 3-4 puncture diameter). Intervals flat on disc, feebly convex in apical part, smooth to slightly shagreened and punctate (punctures separated by 1.5-2 puncture diameter). Intervals 2-7 with longitudinal tubercles more developed than those in F. armatus armatus. Interval 8 with long keel interrupted at middle of elytron. Interval 9 with smaller keel situated at about middle of elytron. Keels, especially hind part of keel on interval 8, serrate in most individuals.

Underside. Pygidium with deep border, disc densely punctate. Abdominal and thoracic sternites coarsely punctate except for disc of meso- and metasternum.

Legs. Anterior tibiae with smaller teeth between major outer teeth in most specimens. Spur of anterior tibia bifurcated in males and simple, acute in females.

Aedeagus. Similar to F. armatus armatus (Fig. 17).

Paratypes. Body length of males 7.0-11.0 mm, width 4.1-5.2 mm, of females 6.3-8.6 mm and 4.0-5.1 mm. Colour of body from black to dark brown. In most specimens, disc of head and pronotum without setae. Length of clypeal horns varies in males (figures 18, 19). Females can be separated by acute, not bifurcated spur of anterior tibiae and by shorter clypeal processes in some cases (figure 20).

Distribution. This subspecies is distributed is Mpumalanga and Limpopo provinces. The southernmost known locality is Wakkerstroom in southern Mpumalanga (figure 27).

Type material. Limpopo Province: Entabeni Forest Station [23°00'S 30°14'E], XI.1931, 3 ♀, G. van Son leg. (TMSA). Malta [24°10'S 30°14'E], 1.II.1927, 1 ♀ (TMSA); Uitzicht [23°19'S 30°01'E], Soutpansberg Distr., XI.1924, 1 ♂ and 1 ♀ (IRSNB); Brak River [? 29°44'E 22°36'S], X.1927, 1 ♂ (TMSA). Mpumalanga: Nelshoogte, Knuckles Rocks Forest, 25°47'S 30°50'E, 24.X.86, intercept traps, 41 day, 6 ♂ 3 ♀, S. Endrödy-Younga leg. (TMSA). Wakkerstroom [27°21'S 30°08'E], I.1925, 1 ♀, G. van Dam leg. (TMSA). Uitsoek, 25°15'S 30°34'E, 7.II.1987, high altitude grassveld, pitfall traps, 61 days, 9 ♂ and 7 ♀; 5.XII.1986, Grootkloof indigenous forest, 25°15'S 30°33'E, pitfall traps, 53 days, 5 ♂ 6 and ♀, S. Endrödy-Younga leg. (TMSA). Nelshoogte, 11.II.1987, Knuckles grassveld, Agaricaceae fungus, 1 ♂; pitfall traps, 58 days, 2 ♂ and 4 ♀; 4.XII.1986, Knuckles rocks forest, pitfall traps, 68 days, 2 ♂; 8.IV.1987, Knuckles grassveld, grassnetting, 1 ♀, S. Endrödy-Younga leg. (TMSA). Berlin, 24°32'S 30°44'E, 4.II.1987, pitfall traps, 41 days, 1 ♂ and 1 ♀, S. Endrödy-Younga leg. (TMSA). Mariepskop, 24°35'S 30°50'E, 2.V.1981, pitfall traps, 5 days, 1 ♂, S. Endrödy-Younga leg. (TMSA). Road Barbeton to Bulembu near Swaziland border [25°55'S 31°10'E], 20.I.2000, 1 ♂ and 1 ♀ (TMSA). White river Distr., farm Lichtfontein [25°19'S 31°01'E], 28.XII.1992, 1 ♂, R. Müller leg. (TMSA).

Frankenbergerius opacus sp. n.,

(figures 4, 7, 10, 12, 43)

Diagnosis.

This species is similar to F. nanus and F. nitidus sp. n. but can be separated from them by the concave base of the pronotum, small longitudinal keel on the pygidium, and the shape of the parameres.

Description.

Holotype ♂ (figure 7). Body black, shiny, length 7.0 mm, width 4.1 mm. Dorsal surface with minute setae.

Head. Clypeus with anterior angles dentiform, separated by deep sinuation. Genae rounded, finely bordered. Genal sutures almost indistinct. Lateral margins of clypeus not sinuate near genal suture. Frontal suture indistinct on disc. Genae and anterior part of clypeus densely punctate, almost rugose; disc more sparsely punctate, margins of punctures indistinct.

Pronotum. Anterior and hind angles rounded. Lateral and anterior margins bordered, base not bordered. Border of anterior margin widened at middle, without sinuation. Dorsal surface densely and coarsely punctate; punctures almost adjacent. Base of pronotum concave (figure 4).

Elytra. Striae distinct, punctate (punctures as wide as striae, separated by 4-5 puncture diameters on disc); striae 9 and 10 widely separated, with deeper and larger punctures medially. Intervals matt, impunctate, with rounded to elongated shiny tubercles.

Underside. Pygidium with strong border and small longitudinal keel, its disc deeply depressed and densely punctate (figure 10). Mesosternum coarsely punctate; metasternum smooth on disc, otherwise punctate as mesosternum.

Legs. Anterior legs with three outer teeth with indistinct teeth between. Anterior tibial spur bifurcated.

Aedeagus. Parameres with widely rounded apices (figure 12).

Paratypes. Body length varies in males from 5.8 mm to 7.1 mm, width from 4.0 mm to 3.6 mm, in females 7.5-6.1 mm and 4.6-3.9 mm, otherwise variation among specimens is very weak.

Distribution. The only recorded locality for this species is Stellenbosch (Western Cape Province) (figure 43).

Type material. Holotype ♂ with labels 'Stellenbosch [33°56'S 18°51'E] 24.5.22 Ch. K. Brain”, 'Coptorhina sp. Not in B.M. Det. G.E.Bryant', and 'Pres. by Com. Inst. Ent. (BMNH) 1949-114” (BMNH). Paratypes: 2 ♂ and 3 ♀ with the same data but without identification label by Bryant (BMNH). One ♂ paratype lacks right elytron.

Additional material examined. One ♂ from unknown locality with handwritten label '33' (SANC).

Frankenbergerius nanus (Péringuey)

(figures 3, 6, 8, 9, 11, 43)

Coptorhina nana Péringuey, 1888: 95; 1901: 288, 293; Janssens, 1939: 32, 36.

Frankenbergerius nanus (Péringuey): Ferreira, 1972: 362, 364.

Diagnosis.

This species is similar to F. opacus sp. n. and F. nitidus sp. n. but can be separated from them by having distinct transverse convexity on the last abdominal sternite. From the former in can also be separated by the pronotum having a flat base, and from the latter by matt elytral intervals.

Description.

Body black to dark-brown, matt, elytra with shiny tubercles (figure 8). Body length of males 5.8-6.2 mm, width 3.1-3.8 mm, of females 5.5-6.5 mm and 3.6-4.1 mm. Dorsal surface of pronotum with minute setae.

Head. Clypeus with anterior angles dentiform with deep sinuation between. Genae rounded, finely bordered. Genal sutures visible as fine lines. Lateral margins of clypeus very feebly sinuate near genal suture. Frontal suture indistinct. Genae and anterior part of clypeus densely punctate, almost rugose. Disc of head with minute, feebly visible punctures.

Pronotum. Anterior angles rounded. Lateral and anterior margins bordered, base not bordered. Border of anterior margin widened at middle, without sinuation. Hind angles rounded. Dorsal surface densely and coarsely punctate; punctures separated by 0.2-0.3 their diameters on disc, becoming denser laterally. Base not concave (figure 3).

Elytra. Striae distinct, punctate (punctures as wide as striae, separated by 3-4 puncture diameter); striae 9 and 10 very close, with deeper and larger punctures. Intervals matt, impunctate, with elongated tubercles.

Underside. Pygidium with strong border, disc densely punctate. Disc of meso- and metasternum smooth. Last abdominal sternite with distinct transverse convexity (figure 9).

Legs. Anterior legs with three outer teeth without smaller teeth between. Spur of anterior tibia bifurcated in males and simple, acute in females.

Aedeagus. Parameres with slightly sclerotized apices narrower in lateral view than in F. opacus sp. n. (figure 11).

Variability. Except for body size variation indicated above, the specimens examined are very similar.

Distribution. This species is known from a few localities in the vicinity of Cape Town (figure 43).

Type material examined. Syntype, ♂: Constantia [Cape Town], IX.1885 Péringuey leg. (SAMC).

Additional material examined. Western Cape Province: Cape Town, VI.1892, Péringuey leg., 3 ♀ (SANC) and 2 ♂ and 4 ♀ (SAMC); Pearly Beach, Bredasdorp [34°45'S 19°30'E], 1 ♂ (SAMC); Groote Post farm, near Darling, 33º36'S 18º25'E, 19.IX.1988, pitfall trap with cow dung, 1 ♂, A. Davis leg. (UPSA).

Frankenbergerius nitidus sp. n.

(figure 13, 43)

Diagnosis.

This species is similar to F. nanus and F. opacus sp. n. but can be separated from the former by having the last abdominal sternite flat, without a transverse convexity, and by shiny elytral intervals with rounded to elongated tubercles. From F. opacus n. sp. it differs in having the base of the pronotum flat, very close elytral striae 9 and 10, and pygidium without longitudinal keel in basal part.

Description.

Holotype ♀ (figure 13). Body black, shiny, length 7.5 mm, width 4.5 mm. Dorsal surface without setae.

Head. Clypeus with anterior angles dentiform with deep sinuation between. Genae rounded, finely bordered. Genal sutures visible as fine lines. Lateral margins of clypeus very feebly sinuate near genal suture. Frontal suture indistinct. Genae and anterior part of clypeus densely punctate, almost rugose. Disc of head with minute, feebly visible punctures.

Pronotum. Anterior angles rounded. Lateral and anterior margins bordered. Border of anterior margin widened at middle, without distinct sinuation. Base not bordered. Hind angles rounded. Dorsal surface densely and coarsely punctate; punctures separated by 0.2-0.3 their diameters on disc, becoming denser laterally.

Elytra. Striae distinct, punctate (punctures as wide as striae, separated by 3-4 puncture diameter); striae 9 and 10 very close, with deeper and larger punctures. Intervals shiny, impunctate, with rounded to elongated tubercles.

Underside. Pygidium with strong border, its disc densely punctate. Disc of metasternum and mesosternum smooth.

Legs. Anterior legs with three outer teeth without smaller teeth between. Anterior tibial spur acute and curved outwards.

Male unknown.

Distribution. This species is known from one locality in Namaqualand (Northern Cape Province) (figure 43).

Type material. Holotype ♀: RSA, Northern Cape Province, Hoekbaai, 31º11'S 17º47'E, 28.VIII.1979, white mesembr. [Mesembryanthemaceae] flow[er], Endrödy-Younga leg. (TMSA).

The specimen was collected on a flower most probably by chance since no species of related taxa are know to be associated with flowers.

Acknowledgements

The authors thank Margaret Cochrane (SAMC), Marc De Meyer (MRAC), Didier Drugmand (IRSNB), Friedrich Gess (AMSA), James Harrison (TMSA), Malcolm Kerley (BMNH), Olivier Montreuil (MNHN), Ruth Müller, (TMSA), Riaan Stals (SANC), Bert Viklund (NHRS), and Cheryl Whitmore (DMSA) for loan of material. Adrian Davis (UPSA) provided the data obtained during his dung beetle surveys in Western Cape Province. Svatopluk Bílý (National Museum of Natural History, Prague) is acknowledged for his help in tracing the types of F. mirabilis. Frank-Thorsten Krell (BMNH) and Izyaslav Kerzhner (Zoological Institute, St. Petersburg) made valuable comments on an early version of the manuscript. This study was supported by a University of Pretoria Postdoctoral Fellowship and, partly, by a grant no 04-04-49109 from Russian Foundation for Basic Research to AVF and by a National Research Foundation (South Africa) grant to CHS.

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