GIANT DUNG BEETLES OF THE GENUS HELIOCOPRIS

SVATOPLUK POKORN¯, J I¤ Í Z ÍDEK, KARL WERNER

G I A N T D U N G B E E T L E SOF 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.

Length 33-40 mm, width 19-24 mm. faunus Boheman

-- 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

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 70)

(page 32)

(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

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

-- 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

-- 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

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

-- 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.

(page 42)

(page 38)

(page 40)

(page 30)

(page 90)

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Heliocopris gigas (Linné)

SYNONYMS

Copris isidis Latreille 1819: 249; Laporte 1840: 75;

Gillet 1911b: 66 [as valid sp.].

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.

DISTRIBUTION

Egypt, Eritrea, Ethiopia,

Oman, Qatar, Saudi

Arabia, Sudan, Yemen.

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.

DESCRIPTION

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.

COMPARISON

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).

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133131 135

132

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).

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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Pokorn˘, Zídek, Werner GIANT DUNG BEETLES OF THE GENUS HELIOCOPRIS (SCARABAEIDAE) Heliocopris gigas (Linné)

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Heliocopris hamadryas (Fabricius)

SYNONYMS

None.

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.

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.

DESCRIPTION

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.

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.

136


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139137 141

138

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).

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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Pokorn˘, Zídek, Werner GIANT DUNG BEETLES OF THE GENUS HELIOCOPRIS (SCARABAEIDAE) Heliocopris hamadryas (Fabricius)

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Heliocopris hamifer Harold

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.

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.

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.

DESCRIPTION

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.

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.

143

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146144 148

145

Fig. 142 - Ethiopia, Kaffa Province, Jimma - Soddo, 9.V.2002, photo K. Werner.Fig. 143 - Head and pronotum of � “jupiter” (left) and � “sirius” (right).Fig. 144 - H. hamifer, � minor, Ethiopia, Kaffa Prov., Soddo - Waka,, V-2002, K. Werner leg.Fig. 145 - Aedeagus of H. hamifer in dorsal view (top) and in left lateral view (bottom).

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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Pokorn˘, Zídek, Werner GIANT DUNG BEETLES OF THE GENUS HELIOCOPRIS (SCARABAEIDAE) Heliocopris hamifer Harold

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Heliocopris haroldi Kolbe

SYNONYMS

None.

Kolbe 1893: 194; Gillet 1907a: 595; Gillet 1911b:

65; Janssens 1939: 76, 94; Ferreira 1967a: 10; 1967c:

1155; 1972: 233.

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.

DESCRIPTION

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.


Protibiae markedly longer in males than in females. Terminations of female

meso- and metatibiae underneath smooth and impunctate.

COMPARISON

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.

149


83

152150 154

151

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).

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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Pokorn˘, Zídek, Werner GIANT DUNG BEETLES OF THE GENUS HELIOCOPRIS (SCARABAEIDAE) Heliocopris haroldi Kolbe

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Heliocopris hermes Gillet

SYNONYMS

None.

Gillet 1911a: 310; 1911b: 65; Janssens 1939: 54, 83;

Ferreira 1972: 233.

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).

DESCRIPTION

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.

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.

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162160 164

161

Fig. 159 - H. hermes, � minor, Ethiopia, Kaffa Province, Gojeb, Jimma, 8.V.02, photo K. Werner.Fig. 160 - H. hermes, � 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).

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

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Pokorn˘, Zídek, Werner GIANT DUNG BEETLES OF THE GENUS HELIOCOPRIS (SCARABAEIDAE) Heliocopris hermes Gillet

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Heliocopris hunteri Waterhouse

SYNONYMS

Heliocopris trilobus Kolbe 1893: 196; 1895: 338;

Gillet 1911b: 66; Janssens 1939: 93; Ferreira 1972:

233.

Waterhouse 1891: 508; Fairmaire 1894: 389; Gillet

1911b: 66; Paulian 1937: 8; Janssens 1939: 73, 93;

Ferreira 1967c: 1155; 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.

DESCRIPTION


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.


Protibiae markedly longer and narrower in males than in females.

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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168166 170

167

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

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

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Pokorn˘, Zídek, Werner GIANT DUNG BEETLES OF THE GENUS HELIOCOPRIS (SCARABAEIDAE) Heliocopris hunteri Waterhouse

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Heliocopris japetus Klug

SYNONYMS

Heliocopris bicarinulatus Boheman 1860: 108; Gillet

1911b: 66; Janssens 1939: 91; Ferreira 1968: 293;

1972: 234.

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.

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.

DESCRIPTION


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

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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174172 176

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Fig. 171 - H. japetus, Zambia, 30 km N Livingstone, 13.XII.01, photo K. Werner.Fig. 172 - H. japetus, � 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).

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

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Pokorn˘, Zídek, Werner GIANT DUNG BEETLES OF THE GENUS HELIOCOPRIS (SCARABAEIDAE) Heliocopris japetus Klug

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GIANT DUNG BEETLES OF THE GENUS HELIOCOPRIS

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