Giant Dung Beetles Of The Genus Heliocopris

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S VAT O P L U K P O K O R N ¯, J I ¤ Í Z Í D E K , K A R L W E R N E R

GIANT DUNG BEETLES OF THE GENUS HELIOCOPRIS

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28. Clypeal apex distinctly emarginate or excised. Lateral pronotal margin strongly concave behind anterior angles, which are directed distinctly outward in both sexes. Anterior steep part of pronotum covered by large punctae that become more confluent mesially. Elytral intervals finely punctate or impuctate. .............................................................................. 29 -- Clypeal apex truncate or rounded. Lateral pronotal margin obliquely cut, either not concave behind anterior angles or steep anterior part of pronotum covered by irregular granules or reticulate. Pronotal crest of major and intermediate males formed by a wide truncate process whose sides extend into laterally projecting sharp teeth posteriorly bordered by concavities. ................................................................................................................. 30 29. Frontal carina straight or weakly emarginate medially, on either side with divergent horns or teeth. Punctation of steep anterior part of pronotum very confluent, on disc very dense and asperate, in center of base changes to granules. Major male: Frontal carina between horns effaced, horns very long and pointed. Base of pronotal crest very wide, laterally excavate, narrowed toward truncate or weakly emarginate apex that is narrower than vertex. Minor male: Pronotal carina more developed, lateral horns reduced to teeth. Pronotal crest medially angular, weakly excavate. Median part bordered by excavations nearly as wide as head. Female: Frontal carina straight, without lateral teeth. Median convex part of pronotal carina narrower than head. faunus Boheman Length 33-40 mm, width 19-24 mm. (page 70)

-- Major male: Frons with a wide open V-shaped carina laterally extending into dorsally flattened and curved branches that reach beyond genal margins. Punctation of steep anterior part of pronotum very sparse, on disc very irregular, in center of base changes to ill-defined granules. Pronotal disc broadly elevate, anteriorly truncate and forming a tridentate process wider than vertex. Minor male: Frontal carina narrower than distance between eyes, laterally bears divergent teeth. Median pronotal process reduced to a minor triangle. Female: Frontal carina similar to that of minor male. Pronotal carina medially swollen, spans nearly entire width of pronotum. Length 32-37 mm, width 20-23 mm. anadematus Gillet (page 32)

30. Steep anterior part of pronotum covered by transversely prolonged granules more-or-less arranged in irregular lines, but never forming a network of wrinkles (both sexes). Frons of males armed with a conical horn or a sharp carina. ...................................................... 31 -- Steep anterior part of pronotum covered by large, simple punctae that near anterior margin merge into confluent asperities and form wrinkles mixed with strong reticulation (both sexes). Frons of males always markedly swollen, laterally with horns. Clypeal apex nearly semicircular. .................................................................................................................... 32 31. Genae of males truncate laterally, genal margin of major males emarginate, that of minor males straight or weakly convex. Pronotum granulose in median part of disc and toward base, remaining surface reticulate. Clypeal apex truncate. Major male: Frons with a frontal blunt, broadly based horn. Lateral teeth of pronotal crest very sharp but short. Minor male: Frons with a wide carina laterally delimited by weak teeth. Lateral teeth of pronotal crest blunt or truncate. Female: Genae laterally extended in front of eyes. Frontal carina wekly curved, without lateral teeth. Pronotal carina spanning nearly entire pronotal width, slightly deflected but not sinuous. Length 36-45 mm, width 23-27 mm. densissa Roth (page 56)

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-- Genae of both sexes markedly extended laterally in front of eyes. Clypeus nearly semicircular, anteriorly weakly truncate. Pronotum completely granulose and punctate, some punctae rimmed. Major male: Frons with a high, wide, sharp and straight carina laterally terminating in straight, conical horns. Lateral teeth of pronotal crest blunt or truncate at apex. Minor male: Lateral horns of frontal carina reduced to minor teeth. Pronotal crest shaped as a half-octahedron with transversely excised lateral margins. Female: Pronotal carina anteriorly convex, medially more arcuate and as wide as frontal carina. Length 36-52 mm, width 22-26 mm. biimpressus Kolbe (page 42)

32. Posterior pronotal surface granulose, granules often in rows, farther forward may be replaced by rimmed and merging punctae. Elytral intervals with many fine, deep punctae visible to naked eye. Major male: Frons markedly swollen between lateral horns, which are laterally compressed and connected by a strong convex-forward carina separated from frontal swelling by a trough. Pronotal crest truncate or shallowly emarginate medially, its lateral processes short and wide, obliquely cut at apex. Minor male: Frontal carina effaced, lateral horns reduced. Lateral pronotal processes less developed. Female: Apex of clypeus slightly emarginate, its lateral margins sinuous. Frontal carina straight and markedly wider than anterior emargination of clypeus. Pronotal carina with midsection more pronounced and narrower than frontal cephalic carina. Length 32-44 mm, width 20-27 mm. atropos Boheman (page 38)

-- Posterior pronotal surface almost completely reticulate, with granules only near base. Elytral intervals densely punctate, punctae shallow and minute, barely discernible to naked eye. Frontal cephalic horns of males never connected by a carina but only by a swelling. Females uncertain or unknown. …................................................................................................. 33 33. Canthus forms evenly excised hind edge of genae. Lateral teeth of pronotal crest short and blunt or truncate. Length 35-45 mm, width 22.5-26 mm. beccarii Harold (page 40)

-- Genae extended laterally in front of eyes. Lateral teeth of pronotal crest long, acutely triangular. Female unknown. Length 40-42 mm, width 25-26 mm. alatus Felsche (page 30)

34. Median tubercle of frontal cephalic carina in both sexes always lower than lateral tubercles. Major male: Genae straight or nearly so, parallel, their lateral margins not extended or barely extended in front of eyes. Apex of pronotal crest wide and rounded, on either side bordered by two teeth. Steep anterior surface of pronotum without a median longitudinal carina. Minor male: Genae oblique, bluntly extended in front of eyes. Apex of pronotal crest straight, laterally bordered by weak, angular tubercles. Female: Frontal cephalic carina at most as wide as anterior truncate part of clypeus, occupies one-third of head width. Pronotal carina evenly curved. Length 40-55 mm, width 24-33 mm. japetus Klug (page 90)

-- Median tubercle of frontal cephalic carina in both sexes always higher than lateral tubercles. Major male: Genae laterally broadly and deeply emarginate, their posterior angles distinctly but bluntly extended in front of eyes. Apex of pronotal crest truncate and narrower than head. Steep anterior surface of pronotum with a short median longitudinal carina. 23

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Heliocopris gigas (Linné)

DESCRIPTION

Scarabaeus gigas Linné 1758: 348 [nec Olivier]; 1764: 16. Heliocopris gigas (Linné): Felsche 1907: 279; Gillet 1911: 66 [as putative syn. of H. isidis]; Arrow 1931: 86 [in part]; Boucomont 1933: 6; Balthasar 1935: 60; Janssens 1939: 79, 97; Ferreira 1972: 231.

Length 37-60 mm, width 20-32 mm. Dark brown to black, matte to semi-matte. Head of both sexes polygonal, with anterior edge of clypeus deeply emarginate to excised (in unworn specimens). Major males with a deep, round impression in center of clypeus and long, sharp, divergent but distally incurved horns in anterolateral corners of genae; frons unarmed. In minor males genal horns reduced to teeth, clypeal impression shallow and ill-defined, frons bears a short carina with lateral teeth. In females genal teeth and clypeal impression absent, frons bears a sinusoidal, weakly tricuspid carina with median cusp situated slightly posterior of lateral cusps. Genae posteriorly divergent in both sexes, narrower in major males than in minor males and females; in major males posterolateral corners of genae sharp and situated directly lateral to eyes, in minor males and females corners blunt and situated at obtuse angles in front of eyes. Pronotum with steep anterior surface granulose and covered by long, rusty setae. Disc of major males anteriorly drawn into a broadly based, forward-tapering medial horn whose narrowly spatulate and excised tip reaches over frons of head. In minor males anterior extent of horn reduced, but tip remains excised and much narrower than cephalic carina. In females anterior edge of disc forms a sinusoidal, mesially convex-forward carina excised at midline and laterally terminating in low, angular processes. Sculpture of disc granulose, with effaced lateral areas. Anterior angles cuspidate in all but weakest males and females. Lateral margins drawn into forward-directed or slightly divergent sharp teeth long in major males, variably reduced in minor males, and becoming mere angulations in weakest males and females. Posterior margin rimmed and straight. Elytra coarsely punctate, punctae irregularly shaped and confluent, giving surface leathery appearance. Basal nodes absent in both sexes. Fifth interval markedly widened at base, nearly as wide as sixth and seventh intervals combined and much wider than fourth interval. Pseudoepipleuron extends laterally, exposing epipleural carina in dorsal view. Pygidium rimmed except a brief interruption at apex.

SYNONYMS Copris isidis Latreille 1819: 249; Laporte 1840: 75; Gillet 1911b: 66 [as valid sp.].

DISTRIBUTION

COMPARISON

Egypt, Eritrea, Ethiopia, Oman, Qatar, Saudi Arabia, Sudan, Yemen.

H. gigas belongs to the gigas group and is close to H. andersoni and H. midas, with which it shares the leathery elytra and armament of the head and pronotum. These three species are rather similar, but can be unequivocally identified by the features given in the key and descriptions. The group is widely distributed in Asia (H. dominus, H. midas) and Africa, where H. andersoni is confined to the Afrotropics whereas H. gigas is northern Afrotropical and southern Palearctic (northeastern Africa, Arabian peninsula).

REMARKS Although the name emphasizes the large size, this is not the largest species of the genus. The types of Linné (UZIU) and Latreille (MNHN) are from Egypt. Museum collections contain only small numbers of old and poorly labeled specimens, creating the impression of a species that has declined due to a Holocene retreat of the elephant to the south. Recent captures of H. gigas in Yemen, Oman and Qatar negate that scenario, however. If a strong dependence on elephant dung ever existed, then H. gigas was able to make the nutritional switch to wild and domesticated ruminants without a noticeable reduction in body size and appears to strive in suitable niches throughout the Arabian peninsula, where it may be presently more common than in Ethiopia, Eritrea or Sudan. 76

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Fig. 130 - Assuan Lake, Egypt, III-1992, photo R. Lízler. Fig. 131 - H. gigas,
minor, Cairo, no coll. data. Fig. 132 - Aedeagus of H. gigas in dorsal view (top) and in left lateral view (bottom).

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Fig. 133 - H. gigas,
major, Egypt, Assuan. Fig. 134 - Head and pronotum of
major (middle),
minor (left) and  (right). Fig. 135 - H. gigas, , Egypt, Gov. di Sharkia, dint Shalya, XI-1992.

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GIANT DUNG BEETLES OF THE GENUS HELIOCOPRIS (SCARABAEIDAE)

Heliocopris gigas (Linné)

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Heliocopris hamadryas (Fabricius)

DESCRIPTION

Copris hamadryas Fabricius 1775: 22; 1801: 36; Olivier 1789: 98; 1790: 152; Laporte 1840: 76. Heliocopris hamadryas (Fabricius): Péringuey 1901: 313, 318; Gillet 1911b: 65; Boucomont 1933: 6; Paulian 1937: 8; Janssens 1939: 65, 89; Ferreira 1962: 99; 1967a: 10; 1967b: 82; 1967c: 1155; 1968: 278, 290; 1972: 231.

Length 39-53 mm, width 23-33 mm. Dark brown to black, glossy. Head transversely rugate, with anterior edge of clypeus broadly truncate. Male frons with a high, transverse carina mesially emarginate and laterally drawn into teeth; area behind carina markedly swollen. Female frontal carina lower and weakly tricuspid. Genae of both sexes extend laterally in front of eyes at obtuse angles. Pronotum with steep anterior surface lacking setae and granulose, granules arranged longitudinally. Disc of major males anteriorly drawn into a tridentate crest with teeth broadly triangular, median tooth reaching over head vertex and low lateral teeth bordered by excavations that extend forward near anterior angles. In minor males configuration of crest similar but subdued. In females disc anteriorly terminates in a long, low, convex-forward carina. Sculpture of disc granulose. Lateral margins in major males parallel to slightly convergent toward base, anterior angles sharp and curved out; in minor males and females margins rounded and anterior angles obtuse. Posterior margin rimmed and straight. Elytra coarsely punctate, basal nodes present only in males. Females with diagonal impressions behind humeral angles. Fifth interval markedly widened at base, nearly as wide as sixth and seventh intervals combined and much wider than fourth interval. Pseudoepipleuron extends laterally, exposing epipleural carina in dorsal view. Pygidium with apex not rimmed.

SYNONYMS None.

DISTRIBUTION Angola, DRC, Kenya, Malawi, Mozambique, PRC, Rwanda, RSA, Somalia, Tanzania, Uganda, Zambia, Zimbabwe.

REMARKS The type of Fabricius (BMNH) is from “Habitat ad Cap B.S. Mus. Dom. Banks”. H. hamadryas occurs from the Tropic of Cancer down to the Cape provinces of RSA and is well represented in collections. K. Werner and R. Lízler collected this species at lights in various RSA locations, and in December 1998 excavated specimens from underneath cattle dung near Vwawa, Mbeya province, Tanzania.

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COMPARISON H. hamadryas belongs to the hamadryas group. The closest species is H. myrmidon, which differs in having a parabolical, anteriorly narrowing and rounded clypeus, lateral teeth of cephalic carina divergent, anterior pronotal angles truncate and not curved out in major males, and calluses present in lateral pronotal excavations.

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Fig. 136 - Habitat of H. hamadryas, Zambia, Ifufa, XII-2002, photo R. Lízler. Fig. 137 - H. hamadryas,
minor, Tanzania, Mbeya Pr., Vwawa, 10.XII.1998, K. Werner & R. Lízler leg. Fig. 138 - Aedeagus of H. hamadryas in dorsal view (top) and in left lateral view (bottom).

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Fig. 139 - H. hamadryas,
major, Zimbabwe, Mt. Silinda, II-1989, R.C. Owen leg. Fig. 140 - Head and pronotum of
major (middle),
minor (left) and  (right). Fig. 141 - H. hamadryas, , Zimbabwe, Mt. Silinda, II-1989, R.C. Owen leg.

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GIANT DUNG BEETLES OF THE GENUS HELIOCOPRIS (SCARABAEIDAE)

Heliocopris hamadr yas (Fabricius)

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Heliocopris hamifer Harold

DESCRIPTION

Harold 1878: 39; Péringuey 1901: 313, 322; Gillet 1911b: 65; Arrow 1928: 74; Janssens 1939: 57, 85; Ferreira 1962: 98; 1967b: 81; 1968: 277, 291; 1972: 232.

Length 30-48 mm, width 17-23 mm. Dark brown to black, semi-matte. Head with clypeus polygonal, anteriorly truncate or weakly emarginate. Frons of both sexes bears a variably long transverse carina with small lateral teeth. Genae of both sexes extend laterally in front of eyes at obtuse angles. Pronotum with steep anterior surface transversely rugate, lacking setae. Disc of major males anteriorly drawn into a long median horn with low basal angulations and narrowly truncate slender tip that reaches over clypeus and on its underside bears small reclined, hook-shaped teeth. In male intermorphs (“H. jupiter”, “H. sirius”) horn progressively shorter, wider, with apex emarginate or excised; in minor males horn reduced to a high, forward-inclined, broadly emarginate carina. In females anterior edge of disc forms a long, low carina with convex-forward central portion approximately as long as cephalic carina. Sculpture of disc vermicular, without granules. Lateral margins vary from posteriorly parallel to rounded throughout, anterior angles tooth-like and usually divergent (except some minor males). Posterior margin rimmed and weakly angular at midline. Elytra coarsely but shallowly punctate, basal nodes present in most males, absent in females and some minor males. Fifth interval markedly widened at base, nearly as wide as sixth and seventh intervals combined and much wider than fourth interval. Pseudoepipleuron folded beneath elytral margin, concealing epipleural carina in dorsal view. Pygidium rimmed except an interruption at apex.

SYNONYMS Heliocopris jupiter Harold 1878: 40; Gillet 1911b: 65; Janssens 1939: 85; Ferreira 1968: 291; 1972: 232. Heliocopris selousi Péringuey 1896: 153; Gillet 1911b: 65; Janssens 1939: 85; Ferreira 1968: 291; 1972: 232. Heliocopris sirius Gillet 1925: 359; Janssens 1939: 85; Ferreira 1968: 291; 1972: 232. Heliocopris longirostris Müller 1941: 338; Ferreira 1968: 291; 1972: 232.

COMPARISON H. hamifer appears to be most conveniently placed in the sensu stricto subgroup of the eryx group, but probably is closely related to the neptunus subgroup. Its pronotal horn facilitates identification, because in all species of the neptunus subgroup it is tridentate.

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DISTRIBUTION Ethiopia, Malawi, Mozambique, RSA, Tanzania, Uganda, Zambia, Zimbabwe. REMARKS The holotype males of Harold’s H. hamifer and H. jupiter (both ZMHB) are from “Nyassa”; that of H. sirius Gillet (BMNH) is from “E. Africa”; that of H. longirostris Müller (MSNT) is from Gondar, Ethiopia; and the syntypes of H. selousi Péringuey (SAMC) are from Mozambique (Manica) and Zimbabwe (Mazoe). Multiple male intermorphs make H. hamifer the most variable species of the genus and are responsible for the synonyms. K. Werner captured several specimens at lights in an area of Ethiopian highlands where no large game occurs and cattle dung is the only available source of food for these beetles. 80

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Fig. Fig. Fig. Fig.

142 143 144 145

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Ethiopia, Kaffa Province, Jimma - Soddo, 9.V.2002, photo K. Werner. Head and pronotum of
“jupiter” (left) and
“sirius” (right). H. hamifer,
minor, Ethiopia, Kaffa Prov., Soddo - Waka,, V-2002, K. Werner leg. Aedeagus of H. hamifer in dorsal view (top) and in left lateral view (bottom).

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Fig. 146 - H. hamifer,
major, D. ost-Afrika [Tanzania], Kigon Sera. Fig. 147 - Head and pronotum of
major (middle),
minor (left) and  (right). Fig. 148 - H. hamifer, , Zambia, Kaful, I-2000.

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GIANT DUNG BEETLES OF THE GENUS HELIOCOPRIS (SCARABAEIDAE)

Heliocopris hamifer Harold

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Heliocopris haroldi Kolbe

DESCRIPTION

Kolbe 1893: 194; Gillet 1907a: 595; Gillet 1911b: 65; Janssens 1939: 76, 94; Ferreira 1967a: 10; 1967c: 1155; 1972: 233.

Length 39-52 mm, width 23-31 mm. Dark brown to black, very glossy. Head transversely wrinkled, anteriorly truncate or weakly emarginate in both sexes. Frontal horn of major males flattened, with emarginate termination and base situated in front of line connecting anterior angles of genae. Frontal process of small males formed by a strong and high carina with small lateral teeth, narrower than one-third of head width. Frontal carina of females distinctly narrower than weakly emarginate apex of clypeus, with weak lateral teeth. Anterior angles of genae strongly and sharply extended outward, posterior angles not extended laterally in front of eyes in major males. Genae weakly extended in front of eyes in minor males, extention stronger in females. Pronotum with steep anterior surface covered by numerous rusty setae in both sexes. Anterior pronotal angles of major males drawn forward into more-or-less strong and sharp teeth; in small specimens anterior angles more-or-less rounded and maximum width shifted forward of mid-length. Pronotal crest of major males with a median process whose excised tip forms a pair of sharp teeth underneath equipped with denticles; space between crest and anterior angles bears strong, rectangular processes. Pronotal crest of minor males weakly convex medially, with small lateral teeth. Pronotum of females with a wide, medially convex carina. Sculpture of disc vermicular, without granules. Posterior margin rimmed, distinctly angular at midline. Elytra very glossy, striate and weakly punctate; fifth interval very wide at base, nearly as wide as sixth and seventh intervals combined and much wider than fourth interval. Elytral base without nodes in both sexes. Pseudoepipleuron extends laterally, exposing epipleural carina from above. Pygidium with apex not rimmed. Protibiae markedly longer in males than in females. Terminations of female meso- and metatibiae underneath smooth and impunctate.

SYNONYMS None.

COMPARISON

DISTRIBUTION Angola, Burundi, Cameroon, DRC, Ethiopia, Guinea, Kenya, PRC, Tanzania.

REMARKS The holotype male of H. haroldi (ZMHB) is from Cameroon. This species is well represented in collections.

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H. haroldi belongs to the bucephalus group and is very similar to H. pirmal and H. samson. Prior to 1939 these three taxa were confused and often regarded as only races of H. pirmal, but Janssens (1939) recognized them as distinct species because major males are easily distinguishable. However, separation of minor males and females of these species is difficult and requires attention to the characters hereby given in the key and descriptions. Provenance helps to separate H. pirmal, which is southern. H. haroldi occurs in eastern, central and western Africa, H. samson in central and western Africa, and these two species co-occur in DRC.

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Fig. 149 - El Sod, south of Yabello, Ethiopia, X-2008, photo R. Filipsk˘. Fig. 150 - H. haroldi,
minor, Ethiopia, Malca Wakana, II-1988. Fig. 151 - Aedeagus of H. haroldi in dorsal view (top) and in left lateral view (bottom).

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Fig. 152 - H. haroldi,
major, W-Afrika. Fig. 153 - Head and pronotum of
major (middle),
minor (left) and  (right). Fig. 154 - H. haroldi, , Ethiopia, Malca Wakana, II-1988.

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GIANT DUNG BEETLES OF THE GENUS HELIOCOPRIS (SCARABAEIDAE)

Heliocopris haroldi Kolbe

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Heliocopris hermes Gillet

DESCRIPTION

Gillet 1911a: 310; 1911b: 65; Janssens 1939: 54, 83; Ferreira 1972: 233.

Length 28-39 mm, width 18-24 mm. Dark brown to black, rather glossy. Head with clypeus polygonal, in front truncate or weakly emarginate. Frons of major males with a horn whose apex is excised and wider than apex of excised medial tooth of pronotal crest. Frontal horn of minor males reduced to a narrow carina incised at apex. Frons of females with a convex carina that is narrower than one-third of head width and bears weak lateral teeth. Genae of males strongly and widely emarginate laterally in front of eyes, so that their lateral margins form distinct angles; in females these angles less developed. Pronotum with steep anterior surface lacking setae in both sexes. Pronotal crest of males with three horns, medial horn weakly incised at apex, lateral ones long and sharp, almost parallel and shorter than medial horn. Horns of major males long, reaching over head base; those of minor males rather reduced. Pronotal carina of females strongly bulged medially, bulge narrower than frontal carina. Steep anterior surface transversely wrinkled, on disc sculpture vermicular, without any granules. Lateral margins emarginate behind anterior angles that are somewhat extended outward. Posterior margin rimmed and weakly angular at midline. Elytra rather glossy, striate, intervals punctate, fifth interval markedly widened at base, nearly as wide as sixth and seventh intervals combined and much wider than fourth interval. Basal nodes present only in males. Pseudoepipleuron folded beneath elytral margin, concealing epipleural carina in dorsal view. Pygidium with apex rimmed.

SYNONYMS None.

COMPARISON H. hermes belongs to the neptunus subgroup of the eryx group. The species of this subgroup are mutually very similar and their separation is reliable only for major males, whose characters are sufficiently well developed. Determination of minor males and females is exceedingly difficult and requires comparisons of multiple specimens collected together at one locality, and preferably also examination of male genitalia.

DISTRIBUTION Cameroon, Eritrea, Ethiopia, Kenya, Tanzania.

REMARKS The holotype male (ISNB) is from Ethiopia. This species is not rare in collections. It has been found occasionally at lights and also excavated from burrows near cattle dung in both highlands and lowlands of Ethiopia (K. Werner).

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Fig. 159 - H. hermes, 씹 minor, Ethiopia, Kaffa Province, Gojeb, Jimma, 8.V.02, photo K. Werner. Fig. 160 - H. hermes, 씹 minor, Ethiopia, Gamu Gofa prov., Ch’en’cha 2500 m, VI-1996 K.Werner leg. Fig. 161 - Aedeagus of H. hermes in dorsal view (top) and in left lateral view (bottom).

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Fig. 162 - H. hermes, 씹 major, Ethiopia, Awash N. P., VI.1994, K. Werner leg. Fig. 163 - Head and pronotum of 씹 major (middle), 씹 minor (left) and 씸 (right). Fig. 164 - H. hermes, 씸, Ethiopia, Awash N. P., VI.1994, K. Werner leg..

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GIANT DUNG BEETLES OF THE GENUS HELIOCOPRIS (SCARABAEIDAE)

Heliocopris hermes Gillet

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Heliocopris hunteri Waterhouse

DESCRIPTION

Waterhouse 1891: 508; Fairmaire 1894: 389; Gillet 1911b: 66; Paulian 1937: 8; Janssens 1939: 73, 93; Ferreira 1967c: 1155; 1972: 233.

Length 34-44 mm, width 21-37 mm. Dark brown to black, glossy. Head transversely rugate, with anterior edge of clypeus weakly emarginate. Frons of major males with a sharp median horn. Frons of minor males with an obtuse denticle, that of females with a laterally dentate carina considerably shorter than distance between eyes. Genae of major males laterally drawn into sharp, upward-directed horns; these horns reduced to denticles in minor males and altogether absent in females. Pronotum with steep anterior surface transversely rugate and densely setose. Anterior pronotal edge of males strongly emarginate behind eyes; in major males emargination posteriorly bordered by a deep, glossy fovea. Disc of major males drawn medially into a long and slender horn whose truncate or weakly bifurcate tip reaches over head and base lacks any lateral angulations. In minor males horn reduced to crest. In females disc bears an arcuate median process, anteriorly rounded and laterally angular. Sculpture of disc vermicular and reticulate, without granules. Posterior margin rimmed and weakly angular at midline. Elytra highly glossy, finely punctate, with basal nodes present only in major males. Fifth interval markedly widened at base, nearly as wide as sixth and seventh intervals combined and much wider than fourth interval. Pseudoepipleuron folded beneath elytral margin, concealing epipleural carina in dorsal view. Pygidium with apex not rimmed. Protibiae markedly longer and narrower in males than in females.

SYNONYMS Heliocopris trilobus Kolbe 1893: 196; 1895: 338; Gillet 1911b: 66; Janssens 1939: 93; Ferreira 1972: 233.

DISTRIBUTION Angola, Kenya, Tanzania.

REMARKS The holotype males of both H. hunteri Waterhouse (BMNH) and H. trilobus Kolbe (ZMHB) are from the Tanzanian side of Kilimanjaro. This species is rare in collections.

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COMPARISON H. hunteri belongs to the hunteri group and is close to H. felschei, from which it differs in having the pronotal horn more slender, without any lateral angulations at base; the genae laterally drawn into sharp, upward-directed horns; and the anterior pronotal edge strongly emarginate behind eyes. Females of these two species differ only in details given in the key.

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170

Fig. 165 - Kenya, Mt. Kenya, II-2009, photo R. Filipsk˘. Fig. 166 - H. hunteri, 씹, no data. Fig. 167 - Aedeagus of H. hunteri in dorsal view (top) and in left lateral view (bottom).

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Fig. 168 - H. hunteri, 씹 major, Kenya, Mt. Kenya, 5.X.2001, local collector leg. Fig. 169 - Head and pronotum of 씹 major (left) and 씸 (right). Fig. 170 - H. hunteri, 씸, Kenya, Mt. Kenya, 5.X.2001, local collector leg.

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GIANT DUNG BEETLES OF THE GENUS HELIOCOPRIS (SCARABAEIDAE)

Heliocopris hunteri Waterhouse

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Heliocopris japetus Klug

DESCRIPTION

Klug 1855: 655; Péringuey 1901: 312, 317; 1908: 629; Gillet 1911b: 66; Janssens 1939: 70, 91; Gomes Alves 1956: 15; Ferreira 1962: 99; 1967b: 82; 1967c: 1155; 1968: 278, 293; 1972: 234.

Length 40-55 mm, width 24-33 mm. Dark brown to black, glossy. Head transversely rugate, anterior margin of clypeus broadly rounded or truncate. Frontal carina in both sexes trituberculate, median tubercle considerably lower. Genae of major males parallel, their posterior angles not extended in front of eyes. In minor males and females posterior angles of genae obtusely extended in front of eyes. Pronotum with steep anterior surface transversely rugate, without setae in either sex, in males bears deep and smooth lateral excavations posteriorly bordered by semicircular carinae with teeth at both ends. Median part of pronotal crest rather broad (nearly as wide as head), weakly convex, laterally terminating in teeth that are slightly offset toward base and directed upward. Transverse carina of females long, broadly flexed forward medially and shallowly emarginate near lateral ends. Median part of disc near base granulose. Posterior margin rimmed and straight. Elytra punctate, without basal nodes in either sex. Fifth interval slightly wider at base than fourth, and much narrower than sixth and seventh intervals combined. In males fifth interval near base with a short (ca. 4 mm) carina adhering to fourth stria, in females carina absent. Pseudoepipleuron extends laterally, exposing epipleural carina in dorsal view. Pygidium rimmed all around, granulose at base, shallowly punctate at apex.

COMPARISON

SYNONYMS Heliocopris bicarinulatus Boheman 1860: 108; Gillet 1911b: 66; Janssens 1939: 91; Ferreira 1968: 293; 1972: 234.

DISTRIBUTION Angola, Botswana, DRC, Kenya, Mozambique, Namibia, RSA, Rwanda, Tanzania, Zambia, Zimbabwe.

REMARKS The holotype of Klug (ZMHB) is from Mozambique, and the holotype of H. bicarinulatus Boheman (NHRS) is from “Ngamisee” [N’Gami Lake], Botswana. This is a common species frequently encountered at lights. K. Werner and R. Lízler collected specimens in proximity of national parks in Kenya, Tanzania, Zambia and Botswana. The presence of elephants near most of the localities suggests their dung as the primary food.

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H. japetus belongs to the hamadryas group. Males are easily distinguished from other species on presence of the short carina near base of the fifth elytral interval. The only other species that possesses this carina is H. antenor, which differs in the shape of head and of the pronotal process (males), and in the frontal cephalic carina (both sexes) that has the median tubercle always higher than the lateral ones.

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Fig. 171 - H. japetus, Zambia, 30 km N Livingstone, 13.XII.01, photo K. Werner. Fig. 172 - H. japetus, 씹 minor, RSA, Kombat, II-1990, C. R. Owen leg. Fig. 173 - Aedeagus of H. japetus in dorsal view (top) and in left lateral view (bottom).

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Pokorn˘, Zídek, Werner

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Fig. 174 - H. japetus, 씹 major, Kenya, Voi, XII-1996, M. SníÏek leg. Fig. 175 - Head and pronotum of 씹 major (middle), 씹 minor (left) and 씸 (right). Fig. 176 - H. japetus, 씸, Kenya, Voi, XII-1996, M. SníÏek leg.

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GIANT DUNG BEETLES OF THE GENUS HELIOCOPRIS (SCARABAEIDAE)

Heliocopris japetus Klug

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