Frolov, A.V., Scholtz, C.H. 2003. Revision of the Afrotropical dung beetle genus Sarophorus Erichson (Coleoptera, Scarabaeidae). African Entomology, 11, 183-198.

Revision of the Afrotropical dung beetle genus Sarophorus Erichson (Coleoptera, Scarabaeidae)

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

1Department of Zoology and Entomology, Faculty of Natural and Agricultural Sciences, University of Pretoria, Pretoria 0002, Republic of 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. E-mail: avfrolov@zoology.up.ac.za.

Abstract

The genus Sarophorus Erichson is revised. Six new species, S. bidentatus sp. n., S. punctatus sp. n., S. carinatus sp. n., S. striatus sp. n., S. latus sp. n., and S. nitidus sp. n., are described. A key to the species is given. Two species-groups are established.

Key words: Sarophorus, new species, species-groups, key, distribution, southern Africa.

Introduction

Sarophorus Erichson is a small genus of "tunneling" dung beetles distributed mainly in Southern Africa. It shares a few apparently synapomorphic characters with Coptorhina Hope and Delopeurus Erichsonand is usually placed in the tribe Dichotomiini (Cambefort, 1991). Validity of this tribe is debated (Montreuil, 1998), however it is beyond the scope of the present study to specify the position of the genus or improve the tribal system of the Scarabaeinae.

The genus Sarophorus was erected by Erichson (1847) for Pedaria tuberculata Castelnau on the grounds of the elytra being sinuate laterally near the base as opposed to being straight in Pedaria. Subsequently, three more species originally described in Pedaria Castelnau were also placed in this genus: P. cicatricosa Péringuey, P. setulosa Brancsik (Boucomont, 1922), and P. costata Fåhraeus in Boheman (Janssens, 1939). Recently we had the opportunity to examine the type of P. setulosa, deposited in the Field Museum, Chicago. This species was transferred to Sarophorus incorrectly and actually belongs to Pedaria. Consequently, the genus Sarophorus contained three described species up to now.

The genera Sarophorus and Pedaria are superficially similar and are often confused in collections. Péringuey (1901) commented that the sinuation of the elytra was variable in the three Sarophorus species known to him (S. tuberculatus, S. costatus and S. cicatricosus), hence he considered Pedaria and Sarophorus to be congeneric. However, the two genera differ strongly in a number of characters apart from the sinuation of elytra (Table 1).

Table 1. Characters to separate Sarophorus and Pedaria (Figs 1-6):
Character Sarophorus Pedaria
Gular plate

 

with longitudinal groove

 

without longitudinal groove

 

Eyes

 

small: distance between eye and gula approximately twice the width of eye in ventral view

 

large: distance between eye and gula approximately half the width of eye in ventral view

 

Genae

 

more protruding past eyes

 

less protruding past eyes

 

Body shape

 

somewhat rectangular

 

sub-oval

 

Abdominal sternites

 

all sternites visible on disc

 

sternite 6 widened and approaching metacoxae

 

Metacoxae

 

separated from epipleura by abdominal sternite 1

 

contiguous to epipleura

 

Lateral margins of elytra

 

more or less sinuate in basal part

 

straight in basal part

 

Metepisterna

 

widened, with convex lateral margin

 

slender, with straight or slightly concave lateral margin

 

Parameres

 

symmetrical

 

asymmetrical

 

Figs 1-13. Sarophorus

Figs 1-13. 1, 3, 5 - Sarophorus sp.; 2, 4, 6 – Pedaria sp.; 7, 10-12 - S. tuberculatus, ♂; 8, 9, 13 - S. bidentatus sp. n., holotype, ♂. 1, 2 – body shape; 3, 4 – abdomen in ventral view; 5, 6 – pterothorax and abdomen in lateral view; 7, 8 – general view; 9, 10 – head and pronotum; 11 – apex of middle tibia; 12, 13 prothorax in lateral view (concavity for apices of retracted middle tibia in 12 indicated by an arrow).

Large numbers of Sarophorus specimens have been collected in the past few decades and are deposited in South African and European museums. During examination of this material new species were found which are described below.

Institutions in which studied material and types are deposited are listed below together with the curators in charge of the collections:

AMSA - Albany Museum, Grahamstown, South Africa: F. Gess; BMNH - British Museum (Natural History), London, Great Britain: M. Kerley; MRAC - Royal Museum of Central Africa, Tervuren, Belgium: M. De Meyer; SANC - National Collection of Insects, Plant Protection Research Institute, Pretoria, South Africa: R. Stals; SAMC - South African Museum, Cape Town, South Africa: M. Cochrane; TMSA - Transvaal Museum, Pretoria, South Africa: R. Müller, J. Harrison; UPSA - University of Pretoria Insect Collection, Pretoria, South Africa: K. Stamhuis.

Genus Sarophorus Erichson

Sarophorus Erichson, 1847: 761. Type species: S. tuberculatus (Castelnau, 1840: Pedaria), by monotypy.

Pedaria: Péringuey, 1901: 277, p. p.

Description.

Medium-sized beetles (6-10 mm). Colour monotonous black to dark brown without metallic tint; anterior part of clypeus, legs or elytra sometimes slightly paler. Surface of body densely punctate, in some species most of pronotum rugose. Each puncture, except for that of elytral striae, bears a short yellowish seta. Setae sometimes abraded on disc of pronotum and head.

Clypeus sinuate in middle, rounded to angulate at sides (anterior angles of clypeus strongly protruding and curved upwards in S. bidentatus sp. n.). In some species anterior angles of clypeus with short carina directed proximally. Genae obtuse. Frontoclypeal and genal sutures indistinct in most species. Head without horns or tubercles. Eyes small, their dorsal part slit-shaped, ventral part sub-rectangular. Distance between eye and gula approximately twice width of eye in ventral view. Gula with longitudinal groove.

Sides of pronotum more or less parallel, width grater than length. Apical margin not bordered, membranous. Lateral margins with or without border. Base of pronotum not bordered, or only indistinctly bordered with a row of punctures.

Elytra not fused, with humeral umbones, sinuate laterally near base. Elytral intervals flat to convex in apical part, with tubercles in some species. Striae with carinate margins. Scutellum not visible from above.

Wings fully developed.

Anterior tibiae with three outer teeth. Anterior tibial spur usually acute and curved downwards. Outer margins of middle and posterior tibiae without transverse keels, serrate in some species. Apices of middle and posterior tibiae densely, regularly or irregularly setose.

Pygidium densely punctate on disc, bordered, with small longitudinal keel in some species.

Parameres symmetrical, their apices strongly to weakly sclerotized, without setae, dorso-ventrally depressed in most species. Armature of internal sac of aedeagus variyng among species.

Sexual dimorphism apparent in three characters: 1) shape of anterior tibial spur (more strongly curved inwards and downwards in ♂), 2) setation of apices of middle tibiae (denser in ♂), and 3) sculpture of clypeus (anterior part more sparsely punctate, usually shiny in ♀ and as coarsely punctate as remaining part of clypeus, usually covered with indument in ♂). Characters 1 and 2 may occur in the same species.

Diagnostic characters of Sarophorus. Shape of the parameres and the armature of the internal sac of the aedeagus are highly specific and should be considered as the most important diagnostic species characters. Other characters include the sculpture of the pronotum and elytra, shape of the clypeus and the anterior tibial spur, as well as the shape and setation of the apices of the middle tibiae.

Biology. Although the genus is broadly classed as a tunneling dung beetle, the little information available on its biology rather indicates a feeding preference for very old dung or carrion remains which may imply detritus as a preferred food rather than dung. The largest known series (S. costatus, 81 specimens) was collected by J. Harrison in a thin soil layer under an old, dry horse carcass. Beetles can fly but rarely come to light. The preimaginal stages is unknown.

The examined species of Sarophorus constitute two distinct species-groups:

1. Sarophorus (species-group tuberculatus): S. tuberculatus, S. punctatus sp. n., S. carinatus sp. n., and S. bidentatus sp. n. belong to this group. The group is characterized by the following characters: 1) elytral intervals, especially intervals 3, 5 and 7, tuberculate; 2) pygidium with small longitudinal carina; 3) metepisterna relatively narrow with aspect ratio ca. 1:2.0 (Fig. 43); 4) internal sac of aedeagus finely granulate, without armature.

S. bidentatus sp. n., known from only a single male specimen, is provisionally placed in this species-group on the basis of tuberculate elytral intervals and proportions of metepisterna. However, it differs from other members of the group in a number of characters, especially in the absence of concavities on the base of the propleurae and the internal sac of the aedeagus being armed with asymmetrical sclerites. The shape of the clypeus and apices of the middle tibiae may be characters of sexual dimorphism. Additional material is needed to clarify the position of this species.

2. Sarophorus (species-group costatus): S. costatus, S. cicatricosus, S. striatus sp. n., S. latus sp. n., and S. nitidus sp. n. are placed in this group. Species of this group are characterized by the following characters: 1) elytral intervals more or less densely punctate but not tuberculate; 2) pygidium without longitudinal carina; 3) metepisterna relatively wide with an aspect ratio of ca. 1:2.5 (Fig. 44); 4) internal sac of aedeagus armed with symmetrically located spinules.

Key to species of Sarophorus

1. Elytral intervals with more or less developed tubercles, sometimes carinate, with indistinct punctures (Figs 7, 8, 14, 16) ..... (species-group tuberculatus) 2

– Elytral intervals punctate, without tubercles (Figs 15, 19-22) ..... (species-group costatus) 5

2. Elytral intervals carinate (Fig. 16). Pygidium deeply foveate laterraly of medial carina (Fig. 18) ..... S. carinatus sp. n. (♀)

— Elytral intervals with oval tubercles (Figs 7, 8, 14). Pygidium less foveate at sides of medial carina (Fig. 17) ..... 3

3. Pronotum with smooth, elevated areas (Figs 7, 8) ..... 4

— Pronotum densely punctate to rugose, without smooth areas (Fig. 14) ..... S. punctatus sp. n.

4. Clypeus with protruding anterior angles (Fig. 9). Concavities on base of propleura absent (Fig. 13). Apex of middle tibiae with a bunch of setae (Fig. 26)

..... S. bidentatus sp. n. (♂)

— Clypeus with angulate, not protruding anterior angles (Fig. 10). Base of propleura with concavities (Fig. 12). Apex of middle tibiae regularly setose (Fig. 11) ..... S. tuberculatus (Castelnau)

5. Pronotum coarsely and irregularly punctate, with sub-symmetrically located, more sparsely punctate areas (Fig. 15) ..... S. cicatricosus (Péringuey)

— Pronotum more finely, regularly punctate (Fig. 19-22) ..... 6

6. Anterior angles of clypeus angulate. Middle tibiae with relatively sparse apical setae in both sexes. Elytra strongly shiny (Fig. 22) ..... S. nitidus sp. n.

— Anterior angles of clypeus rounded. Middle tibiae with denser apical setae in ♂ (Fig. 25). Elytra matt in most specimens ..... 7

7. Elytral intervals more sparsely punctate on disc (punctures separated by a puncture diameter). Apices of parameres with rounded lateral margins in dorsal view (Fig. 32) ..... S. striatus sp. n.

— Elytral intervals more densely punctate on disc (punctures separated by 0.5 puncture diameter). Parameres of different shape (Figs 30, 31) ..... 8

8. Body wider (Fig. 20). Apices of parameres wider, weakly sclerotized (Fig. 31) ..... S. latus sp. n.

— Body narrower (Fig. 19). Apices of parameres narrower, strongly sclerotized (Fig. 30) ..... S. costatus (Fåhraeus in Boheman)

Figs 14-18. Sarophorus: S. punctatus, S. cicatricosus, S. carinatus

Figs 14-18. 14, 17 - S. punctatus sp. n., holotype, ♂; 15 - S. cicatricosus, holotype, ♂; 16, 18 - S. carinatus sp. n., holotype, ♀. 14-16 - general view; 17, 18 – pygidium.

Sarophorus tuberculatus species-group

Sarophorus (tuberculatus) tuberculatus (Castelnau), Figs 7, 10-12, 27, 45

Pedaria tuberculata: Castelnau, 1840: 88.

Sarophorus tuberculatus: Erichson, 1847: 761. Ferreira, 1969: 338.

Diagnosis.

This species is most similar to S. punctatus sp. n. differing from it by the smooth areas on the pronotum and the shape of parameres.

Description.

Upper side of body black, legs slightly paler. Body length 6.0-8.5 mm.

Male (Fig. 7). Head. Anterior margin of clypeus slightly sinuate and flattened medially, anterior angles angulate, with small keels (Fig. 10). Genae obtuse, separated from clypeal margin by distinct sinuation. Anterior and lateral margins of head with fine border. Frontoclypeal and genal sutures indistinct. Dorsal surface regularly and densely punctate with almost adjoining punctures.

Pronotum. Margins not bordered. Surface very densely punctate in posterior part and rugose on disc and in anterior part. Surface usually with 16 irregular, smooth areas sub- distributed symmetrically.

Elytra. Sutural intervals 3, 5, and 7 with small rounded tubercles on disc and each with elongated tubercle at base. Intervals 6 and 8 with elongated tubercles at apex. Interval 9 with a row of partly fused tubercles. Intervals indistinctly punctate. Striae with carinate margins, indistinctly punctate.

Venter. Base of propleura with small concavities for apices of retracted middle femora (Fig. 12). Disc of pygidium densely punctate and pubescent, with small longitudinal carina from middle of basal margin to middle of disc. Abdominal and prothoracic sternites regularly, coarsely punctate. Disc of metasternum concave. Mesosternum with two small convex areas.

Legs. Spur of anterior tibia curved downwards. Apices of middle and posterior tibiae with relatively sparse, unequal setae (Fig. 11).

Aedeagus. Apices of parameres rounded in lateral aspect (Fig. 27). Internal sac finely granulate, without armature (Fig. 35).

Female differs from male in having anterior part of clypeus smooth and shiny.

Material examined: 99 specimens. SOUTH AFRICA: Eastern Cape Province. Algoa Bay, Brauns leg., 1 spm. (SAMC), 2 spm. (TMSA); Hogsback, II.1978, 1 spm., C. Scholtz leg. (UPSA). Western Cape Province. Ysterfontein, 16 spm. (SAMC); 8 km W of Ysterfontein, 33°15’S 18°11’E, 28.VIII.1983, groundtraps, 70 days, 1 spm., Endrödy-Younga and Penrith leg. (TMSA); Saldanha, IX.1912, 37 spm. (SAMC); 8 km NW of Ysterfontein, Farm Blombos, 5.IX.1978, 7 spm., Aschenborn leg. (SANC); Geelbek, 4.IX.1979, 3 spm., Davis leg. (SANC); Velddrif, 17.X.1971, 2 spm., Bornemissza, Kirk leg. (SANC); 33 km NW Port Alfred, 30.VIII.1977, 1 spm., Davis leg. (SANC); 23 km S of Avontuur, 4.V.1976, 1 spm., Davis, Aschenborn leg. (SANC); 12 km SE of Langebaan, Farm Geelbek, 7.IX.1979, 5 spm., Davis leg. (SANC); Rust en Vrede, Oudtshoorn Distr., X.1951, 14 spm. (SAMC); Bo Kouga, Uniondale Distr., III.1954, 1 spm. (SAMC); Darling, 1905, 2 spm., L. Péringuey leg. (SAMC); Swartberge Hagas Farm, 33°24’S 22°46’E, 2.III.1979, sandy valley, 1 spm., S. Endrödy-Younga leg. (TMSA); Somerset, I.1884, 3 spm. (SAMC); Little Karoo, Raubenheimer dam, 33°25’S 22°19’E, 21.X.1993, donkey dung, 2 spm., S. Endrödy-Younga leg. (TMSA).

Sarophorus (tuberculatus) punctatus sp. n., Figs 14, 17, 29, 36, 45

Diagnosis.

This species is most similar to S. tuberculatus but differs from it in having the pronotum without smooth areas and in the widened and depressed apices of parameres.

Description.

Male, holotype (Fig. 14). Upper side of body black, legs and lateral parts of head slightly paler. Body length 7.2 mm.

Head. Anterior margin of clypeus sinuate at medially, anterior angles obtuse. Genae obtuse, separated from clypeal margin by weak sinuation. Anterior and lateral margins of head with very fine border. Frontoclypeal and genal sutures indistinct. Dorsal surface regularly, coarsely punctate with adjoining punctures.

Pronotum. Lateral margins with fine border in posterior part. Surface mostly rugose, coarsely punctate near base, with sub-symmetrically located tubercles; some lateral tubercles cariniform.

Elytra. Sutural intervals convex. Intervals 3, 5, 7, and 9 with elongated tubercles. Intervals 6 and 8 with elongated tubercles at base and apex. Intervals indistinctly punctate. Striae 1-3 more or less distinctly punctate in posterior half, with carinate margins. Punctation of other striae indistinct.

Venter. Base of propleura with concavities for apices of retracted middle femora. Disc of pygidium densely, regularly punctate and pubescent, with short carina near middle of basal margin (Fig. 17). Abdominal and prothoracic sternites coarsely and almost regularly punctate. Disc of metasternum concave. Mesosternum with 2 small convex areas.

Legs. Spurs of anterior tibiae very short, with rounded apex. Apices of middle and posterior tibiae with relatively sparse, unequal setae.

Aedeagus. Parameres with strongly sclerotized, widened and flattened apices (Fig. 29). Internal sac granulate, without armature (Fig. 36).

Female differs from male in having anterior part of clypeus smooth and shiny.

Paratypes. Body length 7,0 – 7,5 mm.

Type material: 8 spm. Holotype ♂: SOUTH AFRICA, Western Cape Province: Keurboomstrand, 34°00’S 23°27’E, 8.XII.1976, groundtraps, 8 days, S. Endrödy-Younga leg. 7 paratypes (♂ and 6 ♀) with the same data. Type material is deposited in TMSA.

Sarophorus (tuberculatus) bidentatus sp. n., Figs 8, 9, 13, 26, 28, 37, 45

Diagnosis.

This species can be separated from other Sarophorus species by the protruding anterior angles of the clypeus with relatively long setae on the dorsal surface, the absence of the concavities on the base of the propleura, the bunch of apical setae on the ventral side of the middle tibia, and the internal sac of the aedeagus being armed with asymmetrical sclerites.

Description.

Male, holotype (Fig. 8). Upper side of body dark brown, pubescent with short yellowish setae. Body length 7,5 mm.

Head. Anterior angles of clypeus strongly protruding and curved upwards (Fig. 9), with long setae on dorsal side. Genae auriculate, separated from clypeal margin by sinuation. Frontoclypeal suture absent. Dorsal surface regularly and densely punctate with almost adjoining punctures.

Pronotum. Margins not bordered. Surface very densely punctate (punctures longitudinally elongated on anterior part), with 14 small, elevated, smooth areas located sub-symmetrically.

Elytra. Sutural intervals convex. Intervals 3, 5, and 7 with small elongated tubercles on disc and base. Intervals 6 and 8 with elongated tubercles at base and apex. Interval 9 with a row of partly fused tubercles. Striae with carinate margins, punctate.

Venter. Base of propleura without concavities for apices of retracted middle femora (Fig. 13). Disc of pygidium densely punctate and pubescent. Abdominal and prothoracic sternites almost regularly, coarsely punctate. Disc of metasternum concave.

Legs. Spur of anterior tibia short and slightly curved downwards. Apices of middle tibiae irregularly setose, with bunch of setae in the middle of ventral side (Fig. 26).

Aedeagus. Apices of parameres rounded in lateral aspect. Base of parameres with distinct angle in lateral aspect (Fig. 28). Internal sac armed with asymmetrical sclerites (Fig. 37).

Type material. Holotype ♂: SOUTH AFRICA, Northern Cape Province, Namaqualand, Kamieskroon, 30°12'S 17°56'E, III.1930 (SAMC).

Sarophorus (tuberculatus) carinatus sp. n., Figs 16, 18, 45

Diagnosis.

This species is similar to S. tuberculatus and S. punctatus sp. n. but differs from them by having the elytra with cariniform tubercles, and a pygidium that is deeply foveate on each side of the middle carina.

Description.

Female, holotype (Fig. 16). Upper side of body dark brown, elytra and legs slightly paler. Body length 6.2 mm.

Head. Anterior margin of clypeus sinuate medially and angulate laterally. Genae rounded, separated from clypeal margin by distinct sinuation. Anterior and lateral margins of head with very fine border. Frontoclypeal suture absent. Anterior part of clypeus smooth, remaining part of head regularly punctate with adjoining punctures.

Pronotum. Lateral margins with fine border, anterior margin and base not bordered. Surface irregularly and coarsely punctate with punctures becoming larger towards base of pronotum. Punctures on disc more or less round, elongated on sides. Pronotum with 17 elevated, irregular, smooth areas sub-symmetrically located, including one near anterior margin.

Elytra. Sutural intervals convex. Intervals 3, 5, 7 and 9 with long, cariniform tubercles on disc. Intervals 6 and 8 with shorter tubercles at base and apex. Intervals indistinctly punctate. Striae with carinate margins, punctate in apical part.

Venter. Base of propleura with feebly visible concavities for apices of retracted middle femora. Disc of pygidium coarsely punctate, with strong carina almost reaching apical border, deeply foveate on each side of the carina (Fig. 18). Abdominal and prothoracic sternites with almost regular, coarse punctation.

Legs. Spur of anterior tibia short and slightly curved downwards. Apices of middle and posterior tibiae with relatively sparse, unequal setae.

Type material. Holotype ♀: SOUTH AFRICA, Mpumalanga, Lydenburg Distr., Ohrigstad, 24°45'S 30°34'E, III.1962, humus, N. Leleup leg. (TMSA).

Sarophorus costatus species-group

Sarophorus (costatus) costatus (Fåhraeus in Boheman), Figs 19, 30, 38, 45

Pedaria costata: Fåhraeus in Boheman, 1857: 204.

Sarophorus costatus: Janssens, 1939: 11, 12. Ferreira, 1969: 338.

Figs 19-24. Sarophorus, S. costatus, S. latus, S. striatus, S. nitidus

Figs 19-24. 19 - S. costatus; 20 - S. latus sp. n., holotype; 21, 23, 24 - S. striatus sp. n. (21, 23 – holotype, 24 – paratype); 22 - S. nitidus sp. n., holotype. 19-22 - general view of ♂; 23, 24 – anterior tibia (23 – ♂, 24 – ♀).

Diagnosis.

This species is most similar to S. striatus sp. n. and S. latus sp. n. but differs from the first in having more coarsely punctate elytral intervals and striae, the presence of a keel at the apex of the elytra and parameres with angulate lateral margins. It differs from the second species in its narrower body, the lateral margins of the pronotum being not bordered, and the presence of narrower, strongly sclerotized apices of the parameres.

Description.

Upper side of body black, legs slightly paler. Body length 7.0-10.0 mm.

Male (Fig. 19). Head. Anterior margin of clypeus sinuate medially, anterior angles rounded. Genae obtuse. Anterior and lateral margins of clypeus with very fine border. Frontoclypeal and genal sutures indistinct. Dorsal surface regularly and densely punctate (punctures separated by 0.5-0.3 puncture diameter).

Pronotum. Anterior angles with fine border, other margins not bordered. Surface densely punctate with almost adjoining punctures.

Elytra. Intervals flat on disc, becoming slightly convex toward apex, coarsely punctate with almost adjoining punctures. Striae shiny, with carinate margins (each stria appears as 3 fine lines), punctate (punctures separated by 2-3 puncture diameters). Intervals 5 to 7 fused at apex forming a distinct keel.

Venter. Base of propleura with concavities for apices of retracted middle femora. Disc of pygidium densely punctate. 6th abdominal sternite sparsely punctate on disc and more densely at sides. Sternites 2-5 punctate with a row of punctures along anterior margin on disc and regularly punctate with almost adjoining punctures at sides. Disc of metasternum concave and coarsely punctate (punctures separated by 0.5 puncture diameter).

Legs. Spur of anterior tibia strongly curved downwards and inwards. Apices of middle and posterior tibiae with dense, unequal setae.

Aedeagus. Apices of parameres flattened and widened, their lateral margins angulate in dorsal view (Fig. 30). Internal sac of aedeagus armed with symmetrically located spinules (Fig. 38).

Female can be separated from male by anterior tibial spur not curved inwards and by sparsely setose apices of middle and posterior tibiae.

Material examined: 283 spm. SOUTH AFRICA: Limpopo Province. 15 km E Louis Trichardt, Farm Welgevonden, 15-17.I.1995, horse carcass, 81 spm., J. Harrison leg. (TMSA); Amatola, Scott farm, 22°56’S 29°23’E, 26.I.1998, dung, 1 spm., R. Müller leg. (TMSA); Blouberg, 3-5.I.1955, 5 spm. (TMSA); Blyderiver Canyon, 24°35’S 30°49’E, 5.V.1981, groundtrap with faeces, 1 spm., sifted gallery forest litter, 1 spm., S. Endrödy-Younga leg. (TMSA); Guernsey, 24°29’S 31°06’E, 12-26.XII.1985, 3 spm., C. Scholtz leg. (UPSA); Happy Rest Nature Reserve, 23°02’S 29°40’E, 1-5.IV.1976, from under stones, 1 spm., Proz.-Schulze leg. (TMSA); Klaserie, X.1981, 1 spm., E. Holm leg. (UPSA); Kruger NP, Nwashtsumbe, 22°47’S 31°17’E, 5.II.1994, buffalo dung, 2 spm., S. Endrödy-Younga leg. (TMSA); Kruger NP, Punda Maria sands, 22°38’S 30°59’E, 11.II.1994, elephant dung, 3 spm., S. Endrödy-Younga leg. (TMSA); Letaba, 1978, 1 spm., C. Scholtz leg. (UPSA); Messina, Hart’s farm, 21.X.1978, 4 spm., C. R. Owen leg. (TMSA); Mmafete, 24°11’S 30°06’E, 28.XI.1985, 6 spm., A. Evans and C. Bellamy leg. (UPSA); Sand River Mt., 24°32’S 27°39’E, 18-19.XII.1985, 1 spm., C. Bellamy and D. d’Hotman leg. (UPSA); Zoutpansberg Distr., Thabina, 13.XI.1905, 1 spm., C. Swierstra leg. (TMSA); Waterberg, 1898-99, 1 spm., V. Jutrzencka leg. (TMSA); Waterberg, Farm 223, 24°11’S 27°50’E, 11.II.1976, groundtraps, 3 spm., UV light, 7 spm., A. Strydom leg. (TMSA); Wildtuin, 22°55’S 31°26’E, 25.IV.1989, 1 spm., L. Le Roux leg. (UPSA); 10 km W of Magoebaskloof, 23°46’S 29°57’E, 10.II.1971, 1 spm., Bornemissza, Ascherborn leg. (SANC); 15 km SW of Kuruman, 27°30’S 23°17’E, 19.XII.1975, 5 spm., Ascherborn leg. (SANC); 30 km SE of Phalaborwa, 24°04’S 30°55’E, 17.I.1973, 2 spm., Davis leg. (SANC); 4 km NW of Mandadzidzi, Kruger National Park, 22°41’S 31°09’E, 25.III.1977, 1 spm., Cambefort, Davis leg. (SANC); Mkluhlu, Kruger National Park, 24°59’S 31°46’E, 8.III.1972, 1 spm., Kirk leg. (SANC); Numbi Gate, Kruger National Park, 25°08’S 31°19’E, 4.II.1971, 3 spm., Bornemissza leg. (SANC); 42 km E of Pietersburg, 23°56’S 29°53’E, 18.I.1973, 2 spm., Davis leg. (SANC); 6 km W of Wylies Poort, 22°53’S 29°52’E, 18.III.1972, 1 spm., Kirk leg. (SANC); Ben Alberts Nature Reserve, Thabazimbi, 24°37’S 27°23’E, 24-28.XI.1980, 3 spm., van Tonder leg. (SANC); Mogol Nature Reserve, Ellisras Distr., 23°58’S 27°45’E, 19-23.XI.1979, 10 spm., van Tonder leg. (SANC). Nort-West Province. 30 km SE of Zeerust, II.1970, 7 spm., L. Schulze and R. Jones leg. (TMSA); 9 km NE of Zeerust, 25°31’S 26°09’E, 27.II.1973, 1 spm., Bornemissza, Temby leg. (SANC); Harte, 25°45’S 27°55’E, 11.III.1995, 1 spm., D. de Beer leg. (UPSA); Potchefstroom, IV.1903, 2 spm. (TMSA); Rustenburg, 12.IV.1936, 1 spm.; 27.II.1999, 1 spm., A. Wagemaker leg. (UPSA); 22 km W of Rustenburg, 25°38’S 27°02’E, 9.I.1973, 1 spm., Davis leg. (SANC); Saartjiesnek, 25°46’S 27°54’E, XII.1985, 2 spm., E. Holm leg. (UPSA); Schoemansville, XII.1932, 1 spm., G. Kobrow leg. (TMSA); Kalkheuvel, X.1908, 2 spm., C. Swierstra leg. (TMSA); Castle Gorge, 66 km W of Pretoria, 25°45’S 27°33’E, 02.I.1970, 4 spm., Bornemissza leg. (SANC). Gauteng. Boekenhoutskloof, 25°33’S 28°29’E, 25.III.1994, blesbok skin, 11 spm., J. Harrison leg. (TMSA); Boekenhoutskloof, 30 km NE of Pretoria, 5.XI.1976, 10 spm., Stickler leg. (SANC); 15 km W of Rust de Winter, 25°19’S 28°28’E, 10.II.1973, 1 spm., Bornemissza leg. (SANC); Hartebeespoort Dam, 25°44’S 27°51’E, 25.I.1971, 1 spm., Bornemissza leg. (SANC); Ditholo, 25°29’S 28°16’E, 2.III.1996, 1 spm., A. Schoeman leg. (UPSA); Mdala, 25°14’S 28°52’E, 14.XII.1997, 1 spm., C. Sole leg. (UPSA); Meyerton, 1-7.XII.1957, 1 spm., J. Heyns leg. (TMSA); Pretoria, X.1949, 1 spm., G. van Son leg. (TMSA); Soutpan, Pretoria, 25°25’S 28°05’E, 8.II.1973, cow dung, 1 spm., S. Endrödy-Younga leg. (TMSA); Vereniging, 26°43’S 28°67’E, 1.III.1997, 1 spm., I. Russo leg. (UPSA); Wagendrift, 25°31’S 28°32’E, 26.II.1990, 1 spm., J. Warrens leg. (UPSA); 5 km S of Soutpan, Pretoria, 3.II.1968, 2 spm., L. and R. Jones leg. (TMSA); Johannesburg, X.1934, 1 spm., G. Kobrow leg. (TMSA). Mpumalanga. 18 km S of Nelspruit, 25°37’S 30°58’E, 30.XI.1986, groundtrap with faeces, 53 days, 1 spm., S. Endrödy-Younga leg. (TMSA); 20 km W of Nelspruit, 25°25’S 30°47’E, 31.X.1972, 1 spm., Davis leg. (SANC); De Kuilen, Lydenburg Distr., 25°10’S 30°32’E, 12.II.1981, 5 spm., Moolman, Harrop leg. (SANC); Mahushe Shongwe Nature Reserve, near Mzinti, 25°45’S 31°44’E, 4-5.II.1995, 41 spm., Chown, Stals, Browne leg. (SANC); 20 km NE of Skukuza, 24°53’S 31°45’E, 3.XI.1984, 2 spm., C. Scholtz leg. (UPSA); Kaapmuiden, 5.IV.1983, 1 spm., S. Chown leg. (UPSA); Kruger NP, 1 km NW of Skukuza, 24°59’S 31°37’E, 22.I.1995, groundtraps with faeces, 1 spm., S. Endrödy-Younga and Ch. Bellamy leg. (TMSA); Kruger NP, Pumbe sands, 24°13’S 31°56’E, 24.I.1995, groundtraps with faeces, 1 spm., S. Endrödy-Younga and Ch. Bellamy leg. (TMSA); Loskop Nature Reserve, 4.III.1995, 1 spm., E. Westhuizen leg. (UPSA); Loskopdam, 25°25’S 29°20’E, 3.III.1995, 2 spm., A. Nel leg. (UPSA); Lydenburg, 1896, 1 spm., P. A. Krantz leg. (TMSA); Nascott Game Reserve, 25°54’S 29°40’E, 3.III.1986, groundtrap with faeces, 45 days, 1 spm., G. du Plessis leg. (TMSA); Nelspruit Distr., Farm de Hoop, 20-21.XI.1972, 1 spm., A. Strydom leg. (TMSA); Nelspruit Nature Reserve, Koppie, 25°29’S 30°55’E, 29.XI.1986, groundtraps with meat and faeces, 13 spm., S. Endrödy-Younga leg. (TMSA); Pretoriuskop, 25°10’S 31°16’E, 3.XII.1984, 1 spm., C. Scholtz leg. (UPSA); Watervalriverpass, 24°54’S 30°21’E, 5-6.XI.1980, cattle dung, 4 spm., S. Endrödy-Younga leg. (TMSA); Waterval-Onder, 22.XI.1910, 1 spm., A. Janse leg. (TMSA). Free State. Cornelia, 28°47’S 27°16’E, 17.III.1987, 2 spm., S. Gothe leg. (UPSA). KwaZulu-Natal. Gwaliweni, Ingwavuma Distr., 7-14.IV.1961, 1 spm., D.W. Rorke leg. (TMSA); Hluhluwe, XII.1960, 12 spm., G. van Son leg. (TMSA); Hluhluwe Game Reserve, 28°05’S 32°05’E, 19-28.XI.1992, buffalo dung, 46 spm., S. Endrödy-Younga leg. (TMSA); Hluhluwe, 28°01’S 32°17’E, 1-7.X.1970, 1 spm., Bornemissza, Ascherborn leg. (SANC); Mkuzi, IV.1950, 2 spm., Koch leg. (TMSA); Mkuze Game Reserve, Msinga Hide, 27°39’S 32°14’E, 27.III.1972, 1 spm., Olsen leg. (SANC); Ndumu, 7.I.1971, 16 spm., de Moor leg. (TMSA); Ndumu Game Reserve, 26°54’S 32°17’E, 1.XII.1992, white rhino dung, 1 spm., S. Endrödy-Younga leg. (TMSA); Ndumu, XII.1960, 2 spm., G. van Son leg. (TMSA); Ndumu Game Reserve, 26°53’S 32°16’E, 9.X.1972, 2 spm., Olsen leg. (SANC); New Hanover, XII.1954, 1 spm., M. B. Baver leg. (TMSA); Pietermaritzburg, X.1956, 1 spm., M. C. Rees leg. (TMSA); Umfolozi Game Reserve, 28°03’S 31°32’E, 6.IV.1974, cattle dung, 8 spm., S. Endrödy-Younga leg. (TMSA); Umfolozi, 28°29’S 32°10’E, 1-7.X.1970, 10 spm., Bornemissza, Ascherborn leg. (SANC); Umgeni Valley, 27.IV.1959, 1 spm., C. Dickson leg. (TMSA); 1 km E of Jozini, 27°25’S 32°05’E, 15.X.1978, 2 spm., Bornemissza, Ascherborn leg. (SANC); 5 km SE of Kokstad, 30°34’S 29°27’E, 18.I.1976, 1 spm., Dewhurst leg. (SANC). Northern Cape Province. Smythe Farm, near Olifantshoek, 27°53’S 22°55’E, X.1992, 8 spm., Steenkamp leg. (SANC). Eastern Cape Province. Dwesa, settlement, 32°15’S 28°49’E, 25.II.1985, horse dung, 1 spm., S. Endrödy-Younga leg. (TMSA); Ntsubane Forest Station, 31°27’S 29°44’E, 25.XI.1987, cattle dung, forest, 1 spm., S. Endrödy-Younga leg. (TMSA); Port St Jonh's, Silaka Bay, 31°33’S 29°30’E, 2.XII.1987, UV light, 7 spm., S. Endrödy-Younga leg. (TMSA); 16 km E of Mount Frere, 30°51’S 29°06’E, 9.XI.1974, 1 spm., Temby leg. (SANC). ZIMBABWE: Bulawayo, 30.I.1928, 1 spm., R. Stevenson leg. (TMSA); 40 mi W of Birchenough Bridge, 19°58’S 32°20’E, 4.I.1972, 1 spm., Bornemissza, Kirk leg. (SANC); Kyle Game Reserve, 20°11’S 31°01’E, 2.I.1972, 2 spm., Bornemissza, Kirk leg. (SANC). BOTSWANA: 4 km N of Lobatsi, 14.IV.1973, 2 spm., Davis leg. (SANC).

Figs 25-34. Sarophorus, parameres

Figs 25-34. 25, 31 - S. latus sp. n., holotype; 26, 28 - S. bidentatus sp. n., holotype; 27 - S. tuberculatus; 29 - S. punctatus sp. n., holotype; 30 - S. costatus; 32 - S. striatus sp. n., holotype; 33 - S. nitidus sp. n., holotype; 34 - S. cicatricosus, holotype. 25, 26 – middle tibiae of ♂; 27-34 – parameres in dorsal and lateral view.

Figs 35-44. Sarophorus, internal sacs

Figs 35-44. 35 - S. tuberculatus; 36 - S. punctatus sp. n., holotype; 37 - S. bidentatus sp. n., holotype; 38 - S. costatus; 39 - S. nitidus sp. n., holotype; 40 - S. striatus sp. n., holotype; 41 - S. cicatricosus; 42 - S. latus sp. n., holotype; 43 – S. (tuberculatus) sp.; 44 – S. (costatus) sp. 35-42 – internal sacs of aedeagi; 43, 44 – shape of metepisternum (MSt – metepisternum, EPl – elytral epipleuron).

Sarophorus (costatus) cicatricosus (Péringuey) Figs 15, 34, 41, 45

Pedaria cicatricosa:Péringuey, 1901: 280, 281.

Sarophorus cicatricosus: Boucomont, 1922: 490. Ferreira, 1969: 338.

Diagnosis.

This species can be separated from similar species (S. costatus, S. striatus sp. n., S. latus sp. n. and S. nitidus sp. n.) by the irregularly punctate pronotum with sparsely punctate, slightly elevated areas and by the apices of the parameres being rounded in lateral aspect.

Description.

Upper side of body black, legs slightly paler. Body length 7.5-8.0 mm.

Male (Fig. 15). Head. Anterior margin of clypeus deeply and triangularly sinuate, anterior angles angulate. Genae obtuse. Anterior and lateral margins of clypeus with very fine border. Frontoclypeal and genal sutures indistinct. Dorsal surface regularly and densely punctate with adjacent punctures.

Pronotum. Anterior angles with fine border, other margins not bordered. Surface densely and irregularly punctate, with a few feebly elevated sparsely punctate areas.

Elytra. Intervals almost flat on disc becoming slightly convex toward apex, coarsely punctate with almost adjoining punctures. Striae with carinate margins, punctate (punctures separated by 2-4 puncture diameters). Intervals 5 to 7 fused at apex forming a keel.

Venter. Base of propleura with concavities for apices of retracted middle femora. Disc of pygidium densely punctate. Abdominal sternites densely and regularly punctate. Disc of metasternum concave and coarsely punctate with adjoining punctures.

Legs. Spur of anterior tibia short and strongly curved downwards. Apices of middle and posterior tibiae with relatively sparse, unequal setae. The holotype has the right anterior tibia with bifurcated spur. Although indicated as diagnostic for this species in the original description, this character is abnormal; spur of left tibia is not bifurcated, of typical shape for the genus.

Aedeagus. Apices of parameres weakly sclerotized, rounded in lateral aspect (Fig. 34). Internal sac with fine, sub-symmetrically distributed spinules (Fig. 41).

Female unknown.

Type material examined. We could not find an exemplar labeled as the type of the P. cicatricosa in SAMC, where it should be deposited according to Péringuey (1901), but we found two specimens labeled “Mashunaland, Salisbury [Harare: ZIMBABWE], G. A. Marshall, 1894” [printed], “Pedaria vulnerata typ, 1897[?]” [Péringuey’s handwriting]. The name “vulnerata” has never been published in combination with either Pedaria or Sarophorus. Because the two specimens agree very well with the description and originate from the type locality of S. cicatricosus we suspect that these two specimens are the only available types of this species. The holotype was not properly designated in the publication, but it can be inferred from the text that the description was largely based on the specimen with an abnormally bifurcated spur of the anterior tibia. This specimen should be considered as the holotype. The paratype comprises anterior (head and pronotum) and posterior parts from two individuals that differ considerably in size.

Additional material examined: 12 spm. ZIMBABWE: Harare, XI.1897, 2 spm., I.1091, 7 spm., Marshall leg. (BMNH); Matopo Mountains, I.1903, 1 spm. (BMNH). ZAMBIA: Serenje District, 12.XII.1907, 1 spm. (BMNH). DEMOCRATIC REPUBLIC OF CONGO: Kundelungus [mountains?], 10°0'S 28°0'E, Mme Tinant, 1 spm. (MRAC).

Fig. 45. Distribution map, Sarophorus

Fig. 45. Distribution map of Sarophorus species.

Sarophorus (costatus) nitidus sp. n. Figs 22, 33, 39, 45

Sarophorus cicatricosus: Balthasar, 1960: 61, 62 (misidentification).

Diagnosis.

This species is most similar to S. costatus but differs from it in its strongly shiny dorsal surface, the absence of a distinct keel on the apices of the elytra, sparsely setose apices of middle and posterior tibiae in both sexes, and the shape of the apices of the parameres.

Description.

Male, holotype (Fig. 22). Upper side of body black, strongly shiny. Body length 7.6 mm.

Head. Anterior margin of clypeus sinuate medially and angulate laterally. Genae obtuse. Anterior and lateral margins of clypeus with very fine border. Frontoclypeal and genal sutures indistinct. Dorsal surface regularly and densely punctate with almost adjoining punctures.

Pronotum. Anterior angles with fine border, other margins not bordered. Surface densely, regularly punctate.

Elytra. Intervals almost flat on disc, becoming slightly convex toward apex, densely punctate. Striae shiny, with slightly carinate margins, punctate (punctures separated by 2-3 puncture diameters). Apices of elytra without keel.

Venter. Base of propleura with concavities for apices of retracted middle femora. Disc of pygidium densely punctate. Abdominal sternite 6 sparsely punctate on disc and more densely so at sides. Sternites 2-5 punctate with a row of punctures along anterior margin on disc and regularly punctate with almost adjoining punctures at sides. Disc of metasternum concave and coarsely punctate (punctures separated by 0.5 puncture diameter).

Legs. Spur of anterior tibia curved strongly downwards and slightly inwards. Apices of middle and posterior tibiae sparsely setose.

Aedeagus. Apices of parameres acute in lateral view, their lateral margins angulate in dorsal view (Fig. 33). Internal sac with sub-symmetrically located spinules (Fig. 39).

Female differs from male in the more sparsely punctate anterior part of clypeus and the more slender and less inwardly curved spur of anterior tibiae. Body length 7,4 mm.

Type material. Holotype ♂: TANZANIA, Tanganyika Terr., Longido, Masai Distr., 1500 m., 17-20.IV.1957, Basilewsky and Leleup leg. Paratype ♀ with the same data. Type material is deposited in MRAC.

Sarophorus (costatus) striatus sp. n. Figs 21, 23, 24, 32, 40, 45

Diagnosis.

This species is most similar to S. costatus and S. latus sp. n. and differs from them in its more finely punctate elytral intervals, the absence of a distinct keel at apex of elytra, the apices of the parameres being rounded lateral margins, and the paler, brown colour of the body.

Description.

Male, holotype (Fig. 21). Upper side of body brown, disc of head and pronotum slightly darker. Body length 8.5 mm.

Head. Anterior margin of clypeus slightly sinuate medially, rounded at sides of sinuation. Genae obtuse. Anterior and lateral margins of clypeus with very fine border. Frontoclypeal and genal sutures absent. Dorsal surface regularly, densely punctate (punctures separated by 0.3-0.5 puncture diameter).

Pronotum. Anterior angles with fine border, other margins not bordered. Surface densely punctate with two sub-symmetrically located, small, smooth areas.

Elytra. Intervals almost flat on disc becoming slightly convex toward apex, shagreened, punctate (punctures separated by a puncture diameter on disc becoming indistinct at apex). Striae fine, shiny, with slightly carinate margins, punctate (punctures separated by 2-3 puncture diameters on 1st stria and about 5 puncture diameters on other striae).

Venter. Base of propleura with concavities for apices of retracted middle femora. Disc of pygidium densely punctate. 6th abdominal sternite sparsely punctate on disc and more densely so at sides. Sternites 2-5 with a row of punctures along anterior margin on disc and regularly punctate with almost adjoining punctures at sides. Disc of metasternum concave and coarsely punctate (punctures separated by 0.5 puncture diameter).

Legs. Spur of anterior tibia strongly curved downwards and inwards (Fig. 23). Apices of middle and posterior tibiae with dense, unequal setae.

Aedeagus. Apices of parameres flattened, their lateral margins rounded in dorsal aspect (Fig. 32). Internal sac with sub-symmetrically located spinules (Fig. 40).

Female. Separated from male by the anterior tibial spur what is not curved inwards (Fig. 24) and by the sparsely setose apices of middle and posterior tibiae.

Paratypes. Body length 8.0-9.0 mm.

Type material: 33 spm. SOUTH AFRICA, Holotype ♂: Western Cape Province, 23 km S of Avontuur, 4.V.1976, Davis and Aschenborn leg. (SANC). Paratypes: Eastern Cape Province, 11 mi S of Grahamstown, 17.V.1971, ♂ and ♀, Aschenborn leg. (SANC); Grahamstown, Strowan, 25.I.1969, 2 ♂ and ♀; 18.II.1969, ♂; 22.X.1968, ♂; 11.XII.1968, ♂; Burntkraal, 13.III.1969, ♀; 13.III.1969, ♀; 1.X.1969 ♀, Gess leg. (AMSA); 7 mi W of Paterson, 6.III.1971, ♀, Aschenborn leg. (SANC); 7 km N of Paterson, 6.IV.1974, ♂, Temby leg. (SANC); Demiskraal, 28.I.1977, ♀, Bornemissza leg. (SANC); Addo Park, 16-20.XII.1996, 3 ♂, Wolmarans leg. (TMSA); 9 km E of Hankey, 28.I.1977, 1 ♀, Bornemissza leg. (SANC); Western Cape Province, Uniondale Distr., Bo Kouga, III.1954, 5 ♂ and 9 ♀ (SAMC).

Sarophorus (costatus) latus sp. n. Figs 20, 25, 31, 42, 45

Diagnosis.

This species is most similar to S. striatus sp. n. and S. latus sp. n. and differs from them in its wider body, the distinctly bordered lateral margins of the pronotum and the weakly sclerotized, widened and downwardly curved apices of the parameres.

Description.

Male, holotype (Fig. 20). Body relatively wide. Upper side of body black to dark brown, legs slightly paler. Body length 8.5 mm.

Head. Anterior margin of clypeus slightly sinuate medially, anterior angles rounded. Genae obtuse. Anterior and lateral margins of clypeus with very fine border. Frontoclypeal and genal sutures absent. Dorsal surface regularly and densely punctate (punctures separated by 0.5-0.3 puncture diameter).

Pronotum. Anterior angles and lateral margins bordered, other margins not bordered. Surface densely and regularly punctate with almost adjoining punctures.

Elytra. Intervals flat on disc becoming slightly convex toward apex, coarsely punctate with almost adjoining punctures. Striae shiny, with carinate margins, punctate (punctures separated by 2-3 puncture diameters). Intervals 5 to 7 fused at apex of elytron forming a distinct keel.

Venter. Base of propleura with concavities for apices of retracted middle femora. Disc of pygidium densely punctate. 6th abdominal sternite sparsely punctate medially and more densely laterally. Sternites 2-5 punctate with a row of punctures along anterior margin on disc and regularly punctate laterally. Disc of metasternum concave and coarsely punctate (punctures separated by 0.5 puncture diameter).

Legs. Spur of anterior tibia strongly curved downwards and inwards. Apices of middle and posterior tibiae with dense, unequal setae (Fig. 25).

Aedeagus. Apices of parameres widened and curved downwards (Fig. 31). Internal sac with sub-symmetrically located spinules (Fig. 42).

Female. Separated from male by the anterior tibial spur, which is not curved inwards, and by the sparsely setose apices of middle and posterior tibiae.

Paratypes. Body length 8.0-9.0 mm.

Type material: 20 spm. SOUTH AFRICA. Holotype ♂: Gauteng, Farm Rhenosterpoort, 25º43’S 28º56’E, 27.XII.1975, bird carcass, L. Schulze leg. Paratypes: 3 ♂ and 4 ♀ with the same data; 4 spm. with the same data but collected in horse dung (♂) and baboon faeces (2 ♂ and ♀); 3 ♂ and ♀ with the same data but collected 3.XII.1975 in horse dung; 2 ♂ with the same data but collected 6.I.1979 and 17.XII.1980; ♂: Pretoria, Waterkloof [other data lacking; the specimen was presumably collected by S. Endrödy-Younga (R. Müller, pers. comm.)]; ♂: Limpopo Province, Happy Rest NR, 23.02’S 29.40’E, 1-5.IV.1976, from under stones, Prozesky-Schulze. Type material is deposited in TMSA.

Acknowledgements

 

The authors thank the curators of collections for providing material for this study and the two anonymous reviewers for valuable comments on the manuscript. Staff of the Laboratory of Microscopy and Microanalysis, University of Pretoria, are acknowledged for providing access to digital photography facilities and technical support. This work has been supported by a University of Pretoria Postdoctoral Fellowship to A. F.

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