helioz Cover 5/12/09 6:14 PM Page 1

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
Helios 1 4/23/09 12:00 PM Page 22

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
(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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Helios 1 4/23/09 12:00 PM Page 23

-- 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)
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.
SYNONYMS
Copris isidis Latreille 1819: 249; Laporte 1840: 75;
Gillet 1911b: 66 [as valid sp.].
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.
76
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).
130
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131 133 135

  
Fig. 130 - Assuan Lake, Egypt, III-1992, photo R. Lízler. Fig. 133 - H. gigas,
major, Egypt, Assuan.
Fig. 131 - H. gigas,
minor, Cairo, no coll. data. Fig. 134 - Head and pronotum of
major (middle),
minor (left) and  (right).
Fig. 132 - Aedeagus of H. gigas in dorsal view (top) and in left lateral view (bottom). Fig. 135 - H. gigas, , Egypt, Gov. di Sharkia, dint Shalya, XI-1992.

132 134

Pokorn˘, Zídek, Werner GIANT DUNG BEETLES OF THE GENUS HELIOCOPRIS (SCARABAEIDAE) Heliocopris gigas (Linné) 77
Helios Part II 4/23/09 12:02 PM Page 78

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.

COMPARISON
SYNONYMS

H. hamadryas belongs to the hamadryas group. The closest species is H. myrmidon,

None.
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.
DISTRIBUTION

Angola, DRC, Kenya,

Malawi, Mozambique,
PRC, Rwanda, RSA,
Somalia, Tanzania,
136
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.

78
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137 139 141

  
Fig. 136 - Habitat of H. hamadryas, Zambia, Ifufa, XII-2002, photo R. Lízler. Fig. 139 - H. hamadryas,
major, Zimbabwe, Mt. Silinda, II-1989, R.C. Owen leg.
Fig. 137 - H. hamadryas,
minor, Tanzania, Mbeya Pr., Vwawa, 10.XII.1998, K. Werner & R. Lízler leg. Fig. 140 - Head and pronotum of
major (middle),
minor (left) and  (right).
Fig. 138 - Aedeagus of H. hamadryas in dorsal view (top) and in left lateral view (bottom). Fig. 141 - H. hamadryas, , Zimbabwe, Mt. Silinda, II-1989, R.C. Owen leg.

138 140

Pokorn˘, Zídek, Werner GIANT DUNG BEETLES OF THE GENUS HELIOCOPRIS (SCARABAEIDAE) Heliocopris hamadr yas (Fabricius) 79
Helios Part II 4/23/09 12:02 PM Page 80

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.
SYNONYMS
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
Heliocopris jupiter Harold 1878: 40; Gillet 1911b: 65;
Janssens 1939: 85; Ferreira 1968: 291; 1972: 232.
H. hamifer appears to be most conveniently placed in the sensu stricto subgroup
Heliocopris selousi Péringuey 1896: 153; Gillet 1911b:
of the eryx group, but probably is closely related to the neptunus subgroup. Its pronotal horn
65; Janssens 1939: 85; Ferreira 1968: 291; 1972: 232.
facilitates identification, because in all species of the neptunus subgroup it is tridentate.
Heliocopris sirius Gillet 1925: 359; Janssens 1939:
85; Ferreira 1968: 291; 1972: 232.
Heliocopris longirostris Müller 1941: 338; Ferreira
142
1968: 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
143
highlands where no large game occurs and cattle dung
is the only available source of food for these beetles.

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144 146 148

  
Fig. 142 - Ethiopia, Kaffa Province, Jimma - Soddo, 9.V.2002, photo K. Werner. Fig. 146 - H. hamifer,
major, D. ost-Afrika [Tanzania], Kigon Sera.
  
Fig. 143 - Head and pronotum of
“jupiter” (left) and
“sirius” (right). Fig. 147 - Head and pronotum of
major (middle),
minor (left) and  (right).
Fig. 144 - H. hamifer,
minor, Ethiopia, Kaffa Prov., Soddo - Waka,, V-2002, K. Werner leg. Fig. 148 - H. hamifer, , Zambia, Kaful, I-2000.
Fig. 145 - Aedeagus of H. hamifer in dorsal view (top) and in left lateral view (bottom).

145 147

Pokorn˘, Zídek, Werner GIANT DUNG BEETLES OF THE GENUS HELIOCOPRIS (SCARABAEIDAE) Heliocopris hamifer Harold 81
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Heliocopris haroldi Kolbe DESCRIPTION

Kolbe 1893: 194; Gillet 1907a: 595; Gillet 1911b: Length 39-52 mm, width 23-31 mm. Dark brown to black, very glossy.
65; Janssens 1939: 76, 94; Ferreira 1967a: 10; 1967c: Head transversely wrinkled, anteriorly truncate or weakly emarginate in both sexes. Frontal
1155; 1972: 233. 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
SYNONYMS base without nodes in both sexes. Pseudoepipleuron extends laterally, exposing epipleural
carina from above.
None. Pygidium with apex not rimmed.
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

DISTRIBUTION 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
Angola, Burundi, easily distinguishable. However, separation of minor males and females of these species is
Cameroon, DRC, Ethiopia, difficult and requires attention to the characters hereby given in the key and descriptions.
Guinea, Kenya, PRC, Provenance helps to separate H. pirmal, which is southern. H. haroldi occurs in eastern,
Tanzania. central and western Africa, H. samson in central and western Africa, and these two species
co-occur in DRC.

REMARKS
149
The holotype male of H. haroldi (ZMHB)
is from Cameroon. This species is well represented
in collections.

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150 152 154

  
Fig. 149 - El Sod, south of Yabello, Ethiopia, X-2008, photo R. Filipsk˘. Fig. 152 - H. haroldi,
major, W-Afrika.
Fig. 150 - H. haroldi,
minor, Ethiopia, Malca Wakana, II-1988. Fig. 153 - Head and pronotum of
major (middle),
minor (left) and  (right).
Fig. 151 - Aedeagus of H. haroldi in dorsal view (top) and in left lateral view (bottom). Fig. 154 - H. haroldi, , Ethiopia, Malca Wakana, II-1988.

151 153

Pokorn˘, Zídek, Werner GIANT DUNG BEETLES OF THE GENUS HELIOCOPRIS (SCARABAEIDAE) Heliocopris haroldi Kolbe 83
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Heliocopris hermes Gillet DESCRIPTION

Gillet 1911a: 310; 1911b: 65; Janssens 1939: 54, 83; Length 28-39 mm, width 18-24 mm. Dark brown to black, rather glossy.
Ferreira 1972: 233. 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

COMPARISON
None.

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
159

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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160 162 164

  
Fig. 159 - H. hermes, 씹 minor, Ethiopia, Kaffa Province, Gojeb, Jimma, 8.V.02, photo K. Werner. Fig. 162 - H. hermes, 씹 major, Ethiopia, Awash N. P., VI.1994, K. Werner leg.
Fig. 160 - H. hermes, 씹 minor, Ethiopia, Gamu Gofa prov., Ch’en’cha 2500 m, VI-1996 K.Werner leg. Fig. 163 - Head and pronotum of 씹 major (middle), 씹 minor (left) and 씸 (right).
Fig. 161 - Aedeagus of H. hermes in dorsal view (top) and in left lateral view (bottom). Fig. 164 - H. hermes, 씸, Ethiopia, Awash N. P., VI.1994, K. Werner leg..

161 163

Pokorn˘, Zídek, Werner GIANT DUNG BEETLES OF THE GENUS HELIOCOPRIS (SCARABAEIDAE) Heliocopris hermes Gillet 87
Helios Part III 4/23/09 12:06 PM Page 88
Heliocopris hunteri Waterhouse
Waterhouse 1891: 508; Fairmaire 1894: 389; Gillet
1911b: 66; Paulian 1937: 8; Janssens 1939: 73, 93;
Ferreira 1967c: 1155; 1972: 233.
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.
88
DESCRIPTION
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.
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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166 168 170

  
Fig. 165 - Kenya, Mt. Kenya, II-2009, photo R. Filipsk˘. Fig. 168 - H. hunteri, 씹 major, Kenya, Mt. Kenya, 5.X.2001, local collector leg.
Fig. 166 - H. hunteri, 씹, no data. Fig. 169 - Head and pronotum of 씹 major (left) and 씸 (right).
Fig. 167 - Aedeagus of H. hunteri in dorsal view (top) and in left lateral view (bottom). Fig. 170 - H. hunteri, 씸, Kenya, Mt. Kenya, 5.X.2001, local collector leg.

167 169

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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.
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.

171
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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172 174 176

  
Fig. 171 - H. japetus, Zambia, 30 km N Livingstone, 13.XII.01, photo K. Werner. Fig. 174 - H. japetus, 씹 major, Kenya, Voi, XII-1996, M. SníÏek leg.
Fig. 172 - H. japetus, 씹 minor, RSA, Kombat, II-1990, C. R. Owen leg. Fig. 175 - Head and pronotum of 씹 major (middle), 씹 minor (left) and 씸 (right).
Fig. 173 - Aedeagus of H. japetus in dorsal view (top) and in left lateral view (bottom). Fig. 176 - H. japetus, 씸, Kenya, Voi, XII-1996, M. SníÏek leg.

173 175

Pokorn˘, Zídek, Werner GIANT DUNG BEETLES OF THE GENUS HELIOCOPRIS (SCARABAEIDAE) Heliocopris japetus Klug 91