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Revision of Sceliages Westwood, a millipede-eating genus of southern African dung beetles (Coleoptera : Scarabaeidae)

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The genus Sceliages Westwood (Scarabaeinae : Scarabaeini) from southern Africa is revised. Seven species are recognised: Sc. granulatus, sp. nov. (Botswana, South Africa), Sc. augias Gillet (Angola, Democratic Republic of Congo, Zambia), Sc. adamastor (Le Peletier de Saint-Fargeau & Serville) (South Africa), Sc. brittoni zur Strassen (South Africa), Sc. difficilis zur Strassen (South Africa, Zimbabwe), Sc. gagates Shipp (South Africa, Moçambique) and Sc. hippias Westwood (South Africa). The new species is described and the others are redescribed. Neotypes are assigned to Sc. adamastor and Sc. gagates. A key to the species is provided and male genitalia and other diagnostic characters are illustrated. A phylogenetic analysis of the genus is presented. Distribution maps of all species are provided. Mature larvae of Sc. hippias are described, the first for the genus. They can be distinguished from other Scarabaeinae larvae by a markedly reduced torma on the epipharynx and complete absence of hypopharyngeal sclerites (oncyli). Millipede relocation and burial behaviour of the adults of Sc. hippias and Sc. adamastor are described. We also provide descriptions of the brood chamber and brood balls of Sc. hippias.
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Invertebrate Systematics
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Invertebrate Taxonomy
Volume 16, 2002
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© CSIRO 2003 10.1071/IT01025 1445-5226/02/060931
Invertebrate Systematics, 2002, 16, 931–955
(Scarabaeidae) from AfricaS. A. Forgie
Revision of Sceliages Westwood, a millipede-eating genus of southern
African dung beetles (Coleoptera:Scarabaeidae)
Shaun A. Forgi eA,B, Va s i l y V. GrebennikovA and Clarke H. ScholtzA
ADepartment of Zoology and Entomology, University of Pretoria, Pretoria 0002, Republic of South Africa.
BTo whom correspondence should be addressed. Email:
Abstract. The genus Sceliages Westwood (Scarabaeinae : Scarabaeini) from southern Africa is revised. Seven
species are recognised: Sc. granulatus, sp. nov. (Botswana, South Africa), Sc. augias Gillet (Angola, Democratic
Republic of Congo, Zambia), Sc. adamastor (Le Peletier de Saint-Fargeau & Serville) (South Africa), Sc. brittoni
zur Strassen (South Africa), Sc. difficilis zur Strassen (South Africa, Zimbabwe), Sc. gagates Shipp (South Africa,
Moçambique) and Sc. hippias Westwood (South Africa). The new species is described and the others are
redescribed. Neotypes are assigned to Sc. adamastor and Sc. gagates. A key to the species is provided and male
genitalia and other diagnostic characters are illustrated. A phylogenetic analysis of the genus is presented.
Distribution maps of all species are provided. Mature larvae of Sc. hippias are described, the first for the genus.
They can be distinguished from other Scarabaeinae larvae by a markedly reduced torma on the epipharynx and
complete absence of hypopharyngeal sclerites (oncyli). Millipede relocation and burial behaviour of the adults of
Sc. hippias and Sc. adamastor are described. We also provide descriptions of the brood chamber and brood balls of
Sc. hippias.
Additional keywords: biology, larva, new species, phylogeny.
The tribe Scarabaeini contains the genera Scarabaeus
Linnaeus, 1758, Sceliages Westwood, 1837, Drepanopodus
Janssens, 1940, Kheper Janssens, 1940, Pachylomerus
Kirby, 1828 and the subgenus Scarabaeolus Balthasar, 1965,
comprising 146 species. Many of the species exhibit distinct
morphological and biological variability and possess either
facultative or obligate feeding strategies including
necrophagy. Some of the most specialised members of this
tribe belong to the genus Sceliages, which exclusively utilise
millipedes (Diplopoda) for food and reproduction. Millipede
necrophagy has long been known in the Scarabaeinae
(Halffter and Matthews 1966: 25–34). Facultative
opportunistic use of millipede carcasses by Scarabaeus
(Neateuchus (syn.)) proboscideus Guérin, 1844, S. satyrus
(Boheman, 1860) and S. (Scarabaeolus) flavicornis
(Boheman, 1860), has been observed by some of us
(S. A. Forgie and C. H. Scholtz, unpublished data).
Necrophagy of millipedes has also been recorded in several
species in two other tribes. In the Onthophagini, several
species of Onthophagus Latreille, 1802, including
O.bicavifrons d’Orbigny, 1902 and O. latigibber d’Orbigny,
1902, were attracted to fresh millipede carcasses (Krell et al.
1997; Krell 1999). Neotropical canthonines, Canthon
cyanellus cyanellus Le Conte, 1859 and C. morsei Howden,
1966, utilise both live injured and dead diplopods (Villalobos
et al. 1998), whereas Deltochilum kolbei Paulian, 1938
(Halffter and Matthews 1966) and D. valgum acropyge
Bates, 1877 (Cano 1998) are known to actively prey on live
In southern Africa, many animals prey on millipedes (see
Lawrence 1987: 82, 89–90). For example, adult and nymphal
reduviid bugs (Hemiptera : Reduviidae), such as Ectricodia
crux (Thunberg, 1783), Cleptria cinctiventris Stål, 1855 and
nymphs of the genus Glymmatophora Stål, 1853, frequently
specialise in preying on Doratogonus Attems, 1914, a
spirostreptid, but they have never been recorded preying on
species of the genus Centrobolus Cook, 1897 (Lawrence
1987). Various quinone-based defensive allomones are
secreted particularly in spirobolid and spirostreptid
millipedes to repel attack by predators (Krell et al. 1998).
932 S. A. Forgie et al.
Two species of the orders Spirostreptida and Julida were
found to use quinonous defensive secretions as pheromones
(Haacker 1974), which is likely to be a secondary function
for many species of millipedes using these secretions.
Necrophagous onthophagine scarabaeids are reported to be
attracted to millipede secretions used as repellents (Krell et
al. 1997, 1998; Krell 1999) and are also likely to be attracted
to the quinonous secretions used as pheromones by
millipedes during copulation (Kon et al. 1998). Positive
chemotaxis to the defensive secretions of millipedes by
Sceliages has not been tested prior to this study. Live (Krell
1999; S. A. Forgie, unpublished data), injured and freshly
dead millipedes all attract Sceliages, suggesting
quinone-based secretions play a role in attracting these
beetles. Sceliages have also been collected by
Endrödy-Younga in traps containing meat/carrion, horse
dung and fruit, and observed rolling antelope dung pellets
(Mostert and Scholtz 1986:10). However, the observation
recorded by Mostert and Scholtz (1986) is best described as
aberrant behaviour for the genus or more likely the product
of misidentification of the beetle responsible. Likewise, the
records of Sceliages trapped in long-term ground traps baited
with various ingredients by Endrödy-Younga are possibly
misleading. For example, in this case, Sc. brittoni may have
become trapped inadvertently after being attracted to
millipedes that might have stumbled into the traps.
With the description of the new species herein, there are
now seven species in the genus Sceliages, all restricted to
southern Africa. Members of the genus are rarely
encountered in the wild and are likely to be mistaken for
Scarabaeus L. Furthermore, specimens of Sceliages are rare
in collections and often misidentified or unidentified. The
biology of Sceliages has, to date, not been studied. Zur
Strassen’s (1965) revision of the genus is the precedent for
this study. It was based on relatively few specimens held in
several museums in Europe and southern Africa. In his
introduction, zur Strassen mentioned that encounters of
generic misidentifications of a number of specimens in
museum collections reflected that the genus Sceliages was
not well known and the descriptions of the oldest species
were very deficient. Zur Strassen (1965) also pointed out
that the situation had been exacerbated by later authors, who
had described known species as new because they were
unaware of the already described species. In addition, and in
accordance with zur Strassen (1965), the holotypes of Sc.
Adamastor and Sc. gagates could not be located and may be
considered non-existent. As a result, we have assigned a
neotype for each of these species.
In this paper we report the results of our review, present a
phylogenetic analysis of the genus and describe a new
species from the semi-arid western parts of the region.
Moreover, we provide for the first time a larval description
of one species and give details of the remarkable biology of
members of the genus feeding on Diplopoda.
Taxonomy and biology
Materials and methods
Adult material examined
A complete list of the material examined is available as
‘Accessory Material’ on the Invertebrate Systematics web page
The institutions to which the specimens belong are abbreviated as
BMNH The Natural History Museum, Department of Entomology,
Cromwell Road, London SW7 5BD, UK.
DMSA Durban Museum, PO Box 4085, Durban 4000, South
HECO Hope Entomological Collections, Oxford University
Museum of Natural History, Parks Road, Oxford OX1 3PW,
ISNB Institut Royal des Sciences Naturelles de Belgique,
Département d’Entomologie, Rue Vautier 29, B–1000
Bruxelles, Belgium.
SAMC South African Museum, PO Box 61, Cape Town 8000,
South Africa.
SANC The National Collection of Insects, Plant Protection
Research Institute, Private Bag X134, Pretoria 0001, South
TMSA Museum of Natural History (Transvaal Museum), Northern
Flagship Institution, PO Box 413, Pretoria 0001, South
UPSA University of Pretoria, Department of Zoology and
Entomology, Pretoria 0002, South Africa.
Latitude and longitude coordinates for all specimens where full
information was available were utilised for the distribution maps of
each species. Locality information in parentheses represents the current
recognised localities and also information not listed on the specimen
collection data labels. A question mark in parentheses immediately
precedes a locality data item that could not be interpreted. Inverted
commas surround exact wording taken from specimen collection data
labels. Some latitude and longitude coordinates were obtained from
material examined sections of zur Strassen (1965) for Sceliages augias
Gillet, 1908 and are included in the distribution map (Fig. 82) for this
Distribution maps. Map coordinates were obtained either directly
from specimen collection data labels or by submitting specimen
localities into GeoName™ digital gazetteer (GDE Systems, Inc., software. These
coordinates were converted to decimal degrees and plotted as
distribution maps using ArcView® GIS software (Environmental
Systems Research Institute, Inc., Redlands CA, USA).
Male genitalia. Aedeagi were removed from 21 specimens (Sc.
granulatus: 1 holotype SANC, 4 paratypes SANC, 1 paratype UPSA;
Sc. hippias: 1 TMSA, 1 SANC; Sc. augias: 1 BMNH; Sc. adamastor: 1
TMSA; Sc. brittoni: 1 SANC, 1 TMSA; Sc. difficilis: 3 SANC, 1
UPSA, 3 TMSA; Sc. gagates: 2 SANC), and soaked in warm 10% KOH
for c. 15 minutes. The internal sacs were extracted, stretched out and
allowed to soak in warm 10% KOH for a further 5 minutes. Sacs were
soaked successively in distilled H2O, 70% EtOH, distilled H2O prior to
their preservation in glycerine. Virgular sclerites (Matthews 1974) were
dissected from the internal sacs and placed in drops of glycerine on
glass slides for examination under a stereomicroscope.
Larval material examined
Five mature larvae of Sceliages hippias originated from brood balls
collected together with females. Two larvae with one female were
collected on 17 December 2000 at the Rustenburg Nature Reserve
Millipede-eating Sceliages (Scarabaeidae) from Africa 933
(25°40S 27°12E), NW Province, Republic of South Africa by S. Forgie
and V. Grebennikov. Three more larvae with one female were collected
on 12 January 2001 from the same locality and by the same collectors.
Voucher larvae and females are deposited in UPSA and BMNH.
Sceliages larvae were preserved in Bouin’s liquid for a week and
then transferred into 70% ethanol. Two larvae were disarticulated as
follows. The head, left legs, mandibles and the labio-maxillar complex
were separated and cleaned in a hot water solution of KOH. Separated
parts were transferred into glycerol and studied under dissecting and
compound microscopes. Morphological drawings were prepared using
camera lucida. The morphological terms utilised in this description are
those explained by Böving (1936), Ritcher (1966) and Lawrence
(1991). Exception is made for terms applicable to the secondary
thoracic and abdominal subdivisions; instead of ‘prescutum’, ‘scutum’
and ‘scutellum’, we use ‘dorsal lobes’ as explained in Baker (1968: 13).
For comparative purposes, one larva of each of the following
Scarabaeinae genera was studied as described in the ‘Materials and
methods’ section: Circellium bacchus Fabricius, 1785; Heliocopris
andersoni Bates, 1868; Kheper nigroaeneus (Boheman, 1857);
Scarabaeus galenus (Westwood, 1847); Scarabaeus (Pachy soma )
gariepinus (Ferreira, 1953); S. (Pachysoma) striatus (Castelnau, 1840);
Synapsis tmolus (Fischer, 1821); and Tragiscus dimidiatus Klug, 1855.
Biology and nidification
Field study of Sceliages hippias was carried out at the Rustenberg
Nature Reserve and Sc. adamastor was observed at the De Hoop Nature
Reserve (34°25S 20°24), Western Cape Province, South Africa.
Beetles were attracted to a series of pitfall traps baited with freshly
killed millipedes unless otherwise stated. Traps were placed into the
field before 0900 hours and left for no longer than one hour to minimise
stress of captured beetles.
Genus Sceliages Westwoo d
Sceliages Westwood, 1837: 12. – Lacordaire, 1856: 66; Shipp, 1895:
37; Péringuey, 1901: 16, 22, 62, 63; Gillet, 1911a: 16; Ferreira,
1961: 63; Ferreira, 1967: 59–63; Ferreira, 1972: 74–78; zur
Strassen, 1965: 220; Halffter & Edmonds, 1982: 138; Scholtz &
Holm, 1985: 220; Mostert & Scholtz, 1986: 1, 8, 10, 11, 16, 22,
23; Hanski & Cambefort, 1991: 167, 472; Krell, 1999: 287.
Parascarabaeus Balthasar, 1961: 174. – Ferreira, 1972: 76; Mostert
& Scholtz, 1986: 10.
Type species of Sceliages: Sceliages iopas Westwood, 1837 (= S.
adamastor (Le Peletier de Saint-Fargeau & Serville, 1828)), by
Type species of Parascarabaeus: Parascarabaeus tonkineus
Balthasar, 1961, by original designation.
Unique morphological characters diagnosing the genus
Sceliages are as follows. Shape and arrangement of the four
up-turned clypeal teeth (two frontal medial teeth narrow and
protrude further than lateral teeth; rounded suture separates
frontal medial teeth from each other; lateral teeth broad,
angulate, skewed, and separated from frontal medial teeth by
a sharply angled suture). Apex of mesotibia has two markedly
developed spurs. Basal tarsomeres are distally flared
appearing triangulate. Unique behaviour within Scarabaeini:
utilisation of millipedes for feeding and breeding.
Redescription of adults
Body shape. Body mostly hunched, reminiscent of species
of the Scarabaeus (Scarabaeolus) ebenus (Klug, 1885)
group (zur Strassen 1965).
Head (Fig. 2). Clypeal margin with four up-turned
teeth. Two frontal medial teeth narrow, protruding further
than lateral teeth. Rounded suture separating frontal medial
teeth from each other; lateral teeth broad, angulate, skewed
and separated from frontal medial teeth by sharply angled
suture. Genal epistomae more pronounced than lateral teeth
of clypeus. Anterior lateral corner of gena tooth-like and
separated from clypeus by sharply angled suture. Posterior
margin of gena obtusely rounded. Geno-clypeal suture
laterally present with obvious groove at its basal terminus.
Surface texture of genae and clypeus shagreened and rugose
with dense, often deep, punctations. Punctations simplified
and less dense on frons and vertex.
Antennae. Antennal furcle consists of three segments.
First segment bowl shaped; second segment much smaller,
less bowl shaped, fits into first segment. Third segment sits
on top of second segment.
Pronotum. Surface smooth with fine shagreen texture
and covered with minute, regularly spaced punctations.
Sternites. Surface of mesobasisternum shagreened with
complex punctuation of varying density and size. Margins of
punctations smooth. Distal halves of metasternum and
adjacent metepisternites with simple sparse punctation.
Mesosternal process markedly broad and pronounced.
Markedly developed facet present between each inner
margin of mesocoxal cavity and lateral carinae of
mesosternellum. Width of mesosternellum between closest
point separating mesocoxae greater than width of mesocoxal
Legs. Apical (fourth) denticle of protibia sickle-shaped
(Figs 16–19). Antero-ventral margin of profemora adjacent
to protrochanter ridged in all species and armed with at least
one spur-like projection in most species (Figs 3–5). Spur
present in males and females and blunted in aged specimens.
Antero-ventral margin of protrochanter well defined and
ridged in most species (Figs 3–5). Meso- and metatibia
truncated distally (Figs 16–24). Two markedly developed
mesotibial spurs present, outer spur larger than inner spur
(Figs 16–19). Outer spur curved, spatulate and pointed. Inner
spur evenly tapered to point. In males, dorsal truncation of
distal portion of metatibia exaggerated, short and acutely
fading towards medial region of tibia (Figs 14, 21, 23). Three
clumps of setae present at base of metatibial truncation
(Fig. 14). First clump forms row closest to truncation,
comprising dense row of short, equal length setae, aligned to
angle of truncation across dorsal surface of metatibia.
Second clump of setae longer than first clump and
positioned along metatibial margin in row transecting
terminus of first clump of setae. Third clump consisting of
934 S. A. Forgie et al.
short setae positioned basally from first clump along the
outer dorsal margin of metatibia. In females, metatibia broad,
rectilinear, without truncation and dorsal surface lacking
clumps of setae (Fig. 15). Both sexes possess uniform row of
setae running uninterrupted medially along length of inner
metatibial surface (Fig. 20). Setae often arise from medial
longitudinal carina or margin that defines the setal row. Basal
tarsomeres flared distally appearing triangular in dorsal and
ventral perspectives.
Male genitalia (Figs 6–13). Ventral structure, or handle,
of virgular sclerite completely fused with primary sclerotised
circular ring. Apical region of handle upturned with varying
degree and twisted so that apex is approximately
perpendicular to plane of basal region.
Ateuchus adamastor was first described by Le Peletier de
Saint-Fargeau and Serville in 1828. In 1837, Westwood
described the genus Sceliages (based on the species
Sc.iopas) to differentiate species of Scarabaeus, including
those of the genus Ateuchus Weber, which possess several
‘structural peculiarities’ including two mesotibial spurs.
Both Westwood (1837) and Lacordaire (1856) thought
A.adamastor should belong to the genus Sceliages,
presumably without realising A. adamastor was conspecific
with Sc. iopas. Synonymy of both species under the new
combination Sceliages adamastor (Le Peletier de
Saint-Fargeau & Serville, 1828) was formalised by Shipp
(1895). The genus Ateuchus was described by Weber in 1801
without the designation of a type species and in 1901
Péringuey designated it a synonym of Scarabaeus. Currently,
only Ateuchus appears as a new world genus within the tribe
Coprini (Halffter and Edmonds 1982).
As far as we are aware, Ferreira (1972: 76) is the first
author to cite Parascarabaeus Balthasar, 1961 as a synonym
of Sceliages. However, Mostert and Scholtz (1986: 10–11)
Fig. 1. Sceliages granulatus, habitus.
Millipede-eating Sceliages (Scarabaeidae) from Africa 935
later proposed that the type species of Parascarabaeus
tonkineus is likely to be a mislabelled specimen of Sceliages
since all other specimens of Sceliages have been collected
only in the southern half of the African continent.
Nonetheless, its status remains as the only full generic
synonym of Sceliages to date (Mostert and Scholtz 1986).
Regarding the morphological differentiation of the genus,
a second medial/inner mesotibial spur is also present in the
Scarabaeus subgenus Scarabaeolus Balthasar, 1965, but this
differs from Sceliages in being vestigial and difficult to
locate (e.g. zur Strassen 1967: 130).
Key to the species of the genus Sceliages Westwood
1. Protibia slightly and evenlly increasing in width distally; with
slight to no inward angulation on medial facet at level of
second external denticle (angulation less apparent in
females) (Figs 25–32, 42–45) . . . . . . . . . . . . . . . . . . . . . . . 2
Protibia abruptly increasing in width distally at level between
second and third external denticle; with markedly developed
inward angulation of medial facet at level from third external
denticle to between third and halfway to second external
denticle (angulation less apparent in females) (Figs 33–41)
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5
2(1). Elytra surface complex, obvious corrugation or granulation;
course shagreen texture; waxy indumentum present; general
matt appearance; striae well defined, bordered with
microcarinae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
Elytra surface plain, without obvious surface protuberances; f ine
shagreen texture, waxy indumentum usually absent, general
glossy appearance; striae fine, narrow and grooved . . . . . 4
3(2). Elytra surface covered in dense, raised granulations. . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . Sc. granulatus, sp. nov.
Elytra surface longitudinally corrugated; tops of corrugations
smooth, glossy, providing a striped or ribbed appearance. .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Sc. augias Gillet
4(3) Antennae yellow–orange; posterior facet of mesofemora armed
with single row of long setae closely paralleling ventral
margin (Fig. 47); medial facet of mesotibia straight (Fig. 18)
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Sc. hippias Westwo o d
Antennae brown–black; posterior facet of mesofemora armed
with two to three rows of setae (Figs 54, 55); first row closely
paralleling ventral margin; second and third row (if present)
with reduced setation, from a few to several setae; medial
facet of mesotibia slightly curved inwards . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .Sc. gagates Shipp
5(1). Obtuse inward angulation of medial facet of protibia from third
external denticle (Figs 33–36); protibial width (in dorsal
perspective) broadens abruptly in apical quarter; elytra
surface glossy; striae on elytra fine, narrow grooved . . . . 6
Slight inward angulation of medial facet of protibia from
between third and half way to second external denticle (Figs
37–41); protibial width (in dorsal perspective) slightly
increased in apical quarter; elytra surface matt with thin
waxy indumentum; striae on elytra well defined, often
bordered with microcarinae (more apparent in teneral adults)
. . . . . . . . . . . . . . . . . . . . . . . . . . . . Sc. difficilis zur Strassen
6(5). Setation red; mesotibia slightly bowed (Fig. 16); outer
mesotibial spur elongate to approximately 1/2 length of
Fig. 2–13. 2, Head plates of Sceliages, contour. 3–5, Profemora and protrochanter; development of basal region of antero-ventral margin (ventral
perspective). Scale bars: 1 mm. 3, Sceliages hippias; 4, Sc. augias; 5, Sc. brittoni. 6–13, Male genitalia of Sceliages; virgular and circular sclerites
of the internal sac, contour. Scale bars: 0.2 mm. 6, Sceliages granulatus (Sekoma, Botswana); 7, Sc. granulatus (Kimberley, RSA); 8, Sc. hippias;
9, Sc. augias; 10, Sc. brittoni; 11, Sc. adamastor; 12, Sc. difficilis; 13, Sc. gagates.
936 S. A. Forgie et al.
mesotibia (Fig. 16); medial facet of metatibia (in dorsal
perspective) relatively straight (Fig. 23) . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . Sc. brittoni zur Strassen
Setation black; mesotibia obtusely bowed (Fig. 17); outer
mesotibial spur 1/5 to 1/4 length of mesotibia (Fig. 17);
entire metatibia bowed inwards (Figs 21, 22) . . . . . . . . . . . .
. . . Sc. adamastor (Le Peletier de Saint-Fargeau & Serville)
Sceliages adamastor (Le Peletier de Saint-Fargeau &
(Figs 11, 17, 21, 22, 35, 36, 50, 51, 83)
Ateuchus adamastor Le Peletier de Saint-Fargeau & Serville, 1828:
351. – Westwood, 1837: 12; Lacordaire, 1856: 66; Shipp, 1895:
Sceliages adamastor (Le Peletier de Saint-Fargeau & Serville). –
Shipp, 1895: 38; Péringuey, 1901: 63; Felsche, 1910: 339;
Gillet, 1911a: 16; Ferreira, 1961: 63; Ferreira, 1967: 60;
Ferreira, 1972: 77; zur Strassen, 1965: 220, 228, 229, 231;
Halffter & Edmonds, 1982: 138; Scholtz & Holm, 1985: 220;
Krell, 1999: 288.
Sceliages iopas Westwood, 1837: 12. – Lacordaire, 1856: 66; Shipp,
1895: 38.
Sceliages jopas [Sic!] Westwood, 1837. – zur Strassen, 1965: 220,
228–230; Ferreira, 1972: 77.
Sceliages joppas [Sic!] Westwood, 1837. – Ferreira, 1972: 76.
Sceliages curvipes Gillet, 1911b: 310. – Gillet, 1911a: 16; zur
Strassen, 1965: 228.
Material examined
Sceliages adamastor
Neotype., ‘South Africa, WC, De Hoop Nature Res., 34°25S
20°24E, 2.xi.2000, millipede baited p/f trap, S. A. Forgie’, SANC.
(Here designated.)
Sceliages iopas
Holotype., ‘South Africa; Sceliages iopas Westwood, 1837.
Type coll.428’, HECO.
Paratype .1, ‘South Africa; Sceliages iopas Westwood, 1837.
Type coll.428’, HECO.
Sceliages curvipes
Holotype., ‘South Africa; Sceliages curvipes Gillet, 1911’,
Other material examined. 17 and 4 . South Africa, Wes t e rn
Cape Province: De Hoop Nature Reserve (34°25S 20°24E),(1
UPSA); De Hoop Nature Reserve, nr Koppie Alleen, 34°28S 20°28E
(2 SAMC); De Hoop Nature Reserve, Cape Agulhas, 60 km NE
34°25S 20°24E (2 ,2 BMNH; 2 ,1 SANC; 1 ,1 UPSA);
Klippe Rugt Farm, 34°42S 20°12E (1 TMSA). Northern Cape:
Klipvlei, Garies, Namaqualand (30°25S 17°54E) (2 SAMC);
Victoria West, ‘Cape-Karoo’, 31°24S 23°07E (4 TMSA). Eastern
Cape: Papiesfontein, Gamtoos River Mouth (33°57S 25°04E) (3
SAMC). (?)Orange Free State (Free State): Modder River, Brandfort
(1 SAMC).
Sceliages adamastor possesses obtusely curved inward
angulation of the mesotibiae. Presence of an obvious inward
curvature of the metatibia is apparent in both males and
females. The length of the outer mesotibial spur relative to
the length of the mesotibia is markedly reduced. Setation
generally black.
Length 12–22 mm.
Head. Surface smooth to slightly rugose on genal
epistomae. Genae and clypeus densely covered with large
punctations. Geno-clypeal suture well defined.
Sternites. Surface of mesobasisternum coarsely
shagreened with markedly spaced, shallow, feebly developed
crescent-shaped punctations. Dorsal margins of punctations
smooth. Setation generally absent.
Legs. Medial (inner) facet of protibia abruptly angled
inwards from third (penultimate distal) external denticle
(Figs 35, 36). Protibial width (in dorsal perspective) in distal
quarter broadens markedly to apex. Mesotibia obtusely
curved inwards, more rounded than rectilinear and truncate
in apical third (Fig. 17). Inner mesotibial spur less than 1/2
length and thickness of outer mesotibial spur. Inner spur
offset from outer spur by 15–20 degrees. Outer mesotibial
spur 1/5 to 1/4 length of mesotibia (Fig. 17). Mesotarsi
approximately half length of mesotibia. Mesofemora armed
with many rows of dense long black setae on posterior facet
(Fig. 51). Metatibia evenly curved inwards (Figs 21, 22).
Male genitalia (Fig. 11). Handle of virgular sclerite
broad, steep semi-circular concavity along dorso-basal
margin; evenly concave along entire ventral margin. Dorsal
margin notched at terminus of union with circular sclerite
prior to obvious swelling to terminus of apical region of
handle. Secondary sclerotisation of handle reduced between
dorsal and ventral corners of apex, forming slight saddle.
Geographical distribution
Sceliages adamastor is known only from South Africa
(Fig. 84).
Horn et al. (1990) provided no information regarding the
Coleoptera collections of Le Peletier or Audinet-Serville. It
therefore seems to be improbable that their locations,
including the type(s) of Ateuchus (=Sceliages) adamastor,
will ever be found. Moreover, Yves Cambefort (personal
communication) of the Paris Museum of Natural History,
home to portions of Le Peletier’s collections, states the types
described by both Le Peletier and Audinet-Serville are
apparently unknown and are likely to be lost. In accordance
with zur Strassen (1965) and those who have helped us in
attempting to locate the type(s), Sc. adamastor type(s) are
considered as missing or non-existent. We have therefore
assigned a neotype for this species.
Redescription and identification of this species was
therefore based on the descriptions of Ateuchus adamastor
by Le Peletier de Saint-Fargeau and Serville (1828),
examination of the conspecific types of Sceliages iopas and
Sc. curvipes, the description of Sc. iopas by Westwood
(1837), description of Sc. adamastor by zur Strassen (1965)
and identified non-type material.
Millipede-eating Sceliages (Scarabaeidae) from Africa 937
It is worth noting the species labels of the two type
specimens of Sceliages iopas held at the Hope
Entomological Collections, University of Oxford, are hand
written in a manner likely to be misinterpreted as Sc. jopas.
The locality data of the Free State specimen is not considered
accurate and is not included in the distribution map (Fig. 84).
We also examined 12 specimens from the ISNB. Of these,
two are misidentified specimens of Sc. gagates and two are
misidentified specimens of Sc. difficilis. The remaining eight
specimens of Sc. adamastor include minimal to no original
collection data to be of any use in distribution maps.
The body size of Sc. adamastor is approximately as large
as Sc. brittoni and smaller adult specimens may also be
confused initially with large specimens of Sc. difficilis. The
posterior surface of the mesofemora of Sc. adamastor
specimens from De Hoop is armed with a single row of long
black setae closely paralleling the ventral margin and a
second, less dense, incomplete row inset from the posterior
margin (Fig. 50). All other specimens of this species are
heavily setose (Fig. 51). The mesotibia of males is only
slightly more obtusely bowed and the inner facet of the
protibia is more abruptly angled inwards than in females.
Sceliages augias Gillet
(Figs 4, 9, 31, 32, 48, 49, 81)
Sceliages augias Gillet, 1908: 64. – Felsche, 1910: 339; Gillet,
1911a: 16; Ferreira, 1961: 64; Ferreira, 1972: 67, 68; zur
Strassen, 1965: 220; Mostert & Scholtz, 1986: 10.
Sceliages sulcipennis Felsche, 1910: 339. – Gillet, 1911a: 16.
? Scarabaeus delaunay-larivierei Paulian, 1934: 58. – Ferreira,
1961: 64.
Parascarabaeus tonkineus Balthasar, 1961: 174 (mislabelled
specimen, Mostert and Scholtz 1986).
Material examined
Holotype., ‘Angola, Benguela; F. C. Wallman Leg.’, ISNB
Other material examined.2 , 2 . Angola: Casonda (7°23S
20°54) (1 TMSA). Democratic Republic of Congo: Mukana(1
ISNB). Zambia: Abercorn (Mbala) (8°50S 31°23E) (1 ISNB);
Algoa (Kabwe), Broken Hill, 180 km E (14°26S 30°18) (1 BMNH);
Mpika, Muchinga Mtns (11°42S 27°10E) (1 ISNB);
Mweru-Wantipa (1 ISNB); Serenje District (1 BMNH); Serenje
(13°10S 30°47) (1 BMNH). Tan zania: Mpwapwa (6°21S
36°29E) (1 BMNH).
Figs 14–24. 14–15, Metatibia of Sceliages; sexual variation of distal apices (dorsal perspective), contours. Scale bars: 1 mm. 14, Sceliages
hippias, ; 15, Sc. hippias, . 16–19, Mesotibia and mesotibial spurs of Sceliages (dorsal perspective), contours. Scale bars: 1 mm. 16, Sceliages
brittoni; 17, Sc. adamastor; 18, Sc. hippias; 19, Sc. difficilis. 20, Metatibia of Sceliages gagates; medial facet with medio-longitudinal
setation/carina (dorso-lateral perspective), contour. Scale bar: 1 mm. 21–24, Metatibia of Sceliages; inward curvation (dorsal perspective), contours.
Scale bars: 1 mm. 21, Sceliages adamastor, ; 22, Sc. adamastor, ; 23, Sc. brittoni, ; 24, Sc. granulatus, .
938 S. A. Forgie et al.
Sceliages augias is easily differentiated from the other
species of Sceliages by the appearance of its elytra:
pronounced longitudinal ridges/corrugations filled with a
matt grey indumentum and the tops shiny black, providing a
striped appearance.
Length 10–18 mm.
Head. Surface rugosely punctated on genae and
clypeus. Distal halves of geno-clypeal sutures obscured by
rugose surface. Frons and vertex with less dense and smaller
Pronotum. Obtusely rounded with curvature in the
posterior third of lateral margins. Thickness and angulation
of lateral margins unvaried.
Elytra. Pronounced longitudinal carina or corrugations
positioned medially on surface between each stria. Surface
texture coarsely shagreened. Surfaces between carinae
covered with indumentum and appearing matt grey. Carinae
shiny black without waxy indumentum. Elytra appearing
striped or ribbed.
Sternites. Surface of mesobasisternum coarsely
shagreened with well-spaced, crescent-shaped, facetted
punctations. Punctations raised forming protrusions or
turbercle-like structures. Dorsal margins of protrusions
smooth. Protrusions each armed with single long seta.
Legs. Spur-like projection markedly pronounced on
anterior ventral ridges of both profemora and protrochanter
(Fig. 4). Inner face of protibia slightly angled inwards from
second external protibial denticle (Figs 31, 32). Mesofemora
armed with few (Fig. 49) to many (Fig. 48) rows of long setae
on the posterior facet. Inner mesotibial spur 2/3 length of
outer spur and 1/2 its thickness; angle is offset from outer
spur by 30 degrees. Mesotarsus between 1/2 and 2/3 length
of mesotibia.
Male genitalia (Fig. 9). Handle of virgular sclerite broad
in width, steeply concave along dorsal margin of basal region
and evenly concave along ventral margin. Dorsal margin
notched immediately after distal terminus of union with
circular sclerite. Dorsal margin angles abruptly upwards to
apex of handle. Apical region tapers to its widest thickness at
apex. Apex angulate and width nearly as broad as length of
dorsal margin from apex to notch. Secondary sclerotisation
Figs 25–45. Protibiae of Sceliages (dorsal perspective), contours. Scale bars: 1 mm. 25, Sceliages granulatus, (Kimberley, RSA); 26, Sc.
granulatus, (Kimberley, RSA); 27, Sc. granulatus, (Kang, Botswana); 28, Sc. granulatus, (Kang, Botswana); 29, Sc. hippias, ; 30, Sc.
hippias, ; 31, Sc. augias, ; 32, Sc. augias, ; 33, Sc. brittoni, ; 34, Sc. brittoni, ; 35, Sc. adamastor, ; 36, Sc. adamastor, ; 37, Sc.
difficilis, (Rhenosterpoort farm, RSA); 38, Sc. difficilis, (Rhenosterpoort farm, RSA); 39, Sc. difficilis, (Umtali, Zimbab.); 40, Sc. difficilis,
(Umtali, Zimbab.); 41, Sc. difficilis, (holotype, BMNH); 42, Sc. gagates, (Muzi Area, RSA); 43, Sc. gagates, (Muzi Area, RSA); 44, Sc.
gagates, (Delagoa B., Moçamb.); 45, Sc. gagates, (Delagoa B., Moçamb.).
Millipede-eating Sceliages (Scarabaeidae) from Africa 939
of handle reduced in ventral corner of apical region and dorsal
corner of basal region. Handle has small baso-ventral
extension also with reduction in secondary sclerotisation.
Geographical distribution
We saw specimens of Sc. augias from the Dominican
Republic of Congo, Angola, Zambia and a single specimen
labelled ‘Mpwapwa’ with no other collection information
provided. The only locality fitting this name occurs in far
eastern Tanzania. If this locality is correct, the distribution of
the species is extended all the way across central Africa. Its
distribution point however has not been included in Fig. 82.
A further record cited by zur Strassen (1965: 221) from the
Upemba National Park, DRC, is incorporated in Fig. 82.
We are not aware who formally synonymised Scarabaeus
delaunay-larivierei with Sc. augias; we follow Ferreira
(1961: 64), who is the first author known to us to use this
Intra-specific variation in the degree of setation or
pubescence on the mesofemoral posterior surface is apparent
among specimens of Sc. augias. Specimens from coastal
Casonda, Angola possess dense setation similar to that of
Sc.brittoni (Fig. 48) compared with inland specimens (e.g.
Fig. 49).
Sceliages brittoni zur Strassen
(Figs 5, 10, 16, 23, 33, 34, 48, 83)
Sceliages brittoni zur Strassen, 1965: 230. – Ferreira, 1972: 77.
Material examined
Holotype., ‘South Africa: SW Cape (Western Cape Province);
Leipoldtville, Eland’s Bay (32°13S 18°29E); October 1947, museum
exped.’, SAMC.
Paratypes.1 , ‘South Africa: SW Cape (Western Cape
Province); Darling (33°23S 18°23E); October 1906, L.Péringuey’,
SAMC; 1 , ‘Eland’s Bay; October 1947, museum exped.’, SAMC; 1
, ‘Saldanha Bay (33°03S 18°00E); September 1960 ‘S.A.M.’,
Other material examined.South Africa, Wes t ern Cap e P rov. :
Namaqualand Kommandokraal farm, 31°30S 18°13E (1 TMSA);
Langebaan, Geelbek, 12 km SE (33°06S 18°02E) (3 , 2 SANC);
Langebaan, Geelbek, 12 km SE (1 SANC), Nortier farm, 32°02S
18°20E (1 TMSA); Seweputs coast, 31°39S 18°17E (1 TMSA).
Northern Cape Prov.: Hondeklipbaai, 30°19S 17°16E; (1 SANC);
Hondeklipbaai, 12 km E, 30°21S 17°25E (11 ,4 TMSA);
Kotzesrus, 30°57S 17°50E (2 TMSA); Quaggafontein, 30°13S
17°33E (1 TMSA); Vlakte farm, Gemsbok, 30°30S 17°29E (2
Sceliages brittoni is easily diagnosed with the following
morphological characters: red setation; markedly elongate
outer mesotibial spur relative to the length of the mesotibia;
and a large body size. The distribution of Sc. brittoni is
restricted to the west coastal regions of South Africa.
Length 17–25 mm.
Sternites. Surface of mesobasisternum texture coarsely
shagreened. Punctations crescent-shaped and facetted.
Punctations raised forming protrusion or turbercle-like
structures; dense but generally unlinked radiating
anterior-laterally from centre of mesosternal process.
Setation generally absent.
Legs. Medial (inner) facet of protibia abruptly angled
inwards from third external denticle (Figs 33, 34). Protibial
width (in dorsal perspective) in distal quarter broadens
abruptly to apex. Profemoral spur-like projection
pronounced with reduced tooth-like serrations on remainder
of anterior ventral ridge of both profemora and protrochanter
(Fig. 5). Spur-like projection may be present but reduced on
anterior ventral ridge of protrochanter. Mesofemora armed
with many rows of dense, obvious long red/brown setae on
the posterior facet (Fig. 48). Outer mesotibial spur markedly
elongate; approximately half length of mesotibia (Fig. 16).
Inner mesotibial spur less than 1/3 length and width of outer
spur; offset from outer spur by 30 to 45 degrees. Mesotarsus
approximately 1/3 length of mesotibia. Minimal to no inward
curvature of metatibia (Fig. 23).
Male genitalia (Fig. 10). Handle of virgular sclerite
narrow and relatively constant thickness through its length;
widening slightly at each end. Handle evenly concave along
majority of dorsal margin to an abrupt outward angulation
near terminus of apical region; ventral margin slightly
concave to angulate. Dorsal margin unnotched at terminus of
union with circular sclerite. Secondary sclerotisation of
handle reduced between dorsal and ventral corners of apex
forming an obvious saddle. Dorsal and ventral corners of
apical region of handle appear as protruding points.
Baso-ventral corner forms a slight protruding extension with
reduction in secondary sclerotisation from its apex to
baso-dorsal union with circular sclerite.
Biological observations
A single specimen of Sceliages brittoni in Namaqualand,
South Africa, was observed displacing reduviid nymphs
(species unknown) that were attacking a large harpagophorid
millipede, Zinophora sp. (Diplopoda : Spirostreptida).
Sceliages brittoni then relocated the millipede while it was
still alive (J. Colville, personal communication).
Geographical distribution
Sceliages brittoni is known from South Africa only (Fig. 84).
The majority of the external morphological features of
Sc.brittoni closely resemble those of Sc. adamastor.
940 S. A. Forgie et al.
Sceliages difficilis zur Strassen
(Figs 12, 19, 37–41, 52, 53, 82)
Sceliages difficilis zur Strassen, 1965: 224. – Ferreira, 1972: 77.
Material examined
Holotype., ‘South Africa: Eastern Cape Province; Grahams
town; ex. coll. Fry 1905–100’, BMNH.
Paratypes.1 and 1 , ‘South Africa: Natal (Kwazulu-Natal);
Krantzkop (Kranskop) (28°58S 30°51E); November 1917, K.H.
Barnard’, SAMC. 1 , ‘Natal (Kwazulu-Natal); pt; ‘58.13’’, BMNH; 1
, ‘Natal (Kwazulu-Natal); ex. coll. Pascoe’, BMNH; 1 , ‘Natal
(Kwazulu-Natal), Durban; A.E.Miller’, SAMC; 1 , ‘Zimbabwe:
Salisbury (Harare), Mashunaland (17°50S 31°03E); 1893, G.A.
Marshall’, SAMC; 1 , ‘Inyanyadzi R. (Nyanyadzi), Gazaland
(19°45S 32°25E); November 1901, G.A.K. Marshall’, BMNH; 1,
‘Africa, austr.; Coll. J.J.Gillet’, ISNB.
Other material examined.South Africa, Gauteng:
Boekenhoutskloof, Pretoria, 30 km NE (25°31S 28°25E) (6 ,3
SANC); Johannesburg (26°12S 28°05E) (1 ,1 TMSA);
Rhenosterpoort Farm, 25°43S 28°56E (2 ,2 TMSA); Valhalla,
Pretoria (25°49S 28°08E) (1 TMSA); Mpumalanga, Groenvaly
farm, nr Badplaas, 50 km N (25°50S 30°45E) (3 ,1 UPSA);
Lydenburg District, Kopies Kraal, 25°06S 30°12E (2 TMSA).
Northern Province: Motlakeng, Blouberg (23°05S 29°01E) (1
TMSA); Nylsvley Nature Reserve, 24°39S 28°42E (1 UPSA);
Nylsvley, Smith Farm, 24°40S 28°42E (1 TMSA); Pietersburg
(23°50S 29°20E) (1 TMSA); Sand River, Thabazimbi 24°32S
27°39E (1 UPSA); Iron Crown, 2 km W, Wolkberge, Haenertsberg
Distr. (24°00S 29°57E) (1 TMSA). Kwazulu-Natal: ‘ZuluLand’ (1
BMNH; Sceliages iopas W.); ‘Natal’ (1 BMNH); (Free State),
Glen (1 SANC). Zimbabwe: Harare, Mashunaland[sic] (1
BMNH); Peak Mine, Selukwe (Shurugwi) (19°40S 30°00E) (1
SAMC); Chipinga (Chipinge) (20°12S 32°37E) (1 TMSA); Upper
Búzi River (19°51S 32°48E) (1 BMNH 1908-212); Upper Búzi
River, Gazaland (Melsetter Distr) (1 BMNH 1931-138); Mt
Chirinda, ‘Gaza Ld.’ (19°14S 32°14E) (1 BMNH); Mt Chirinda,
Gazaland (1 BMNH; 1931–138); Chirinda Forest (1 TMSA);
Umtali (Mutare) (18°58S 32°40E), (1 SAMC; specimen v. poor
cond.); Umtali (Mutare), Mashonaland (3 , 1 BMNH 1904-206);
‘Rhodésie’, Plumtree, Rhodesia (1 ISNB; Sc. adamastor 10.640);
‘Rhodésie’, Salisbury (Harare), Mashonaland (1 ISNB; Sc.
Sceliages difficilis can be diagnosed using the angulation of
the protibiae (the medial facet of the protibia is slightly
angled inwards from between the third external denticle to
half-way to the second external denticle; protibial width is
even from the base to parallel with the third external denticle
where the width broadens slightly to the apex), and the
protruding extension of the baso-ventral corner of the
virgular sclerite handle.
Length 10–18 mm.
Pronotum. Lateral margins evenly rounded, narrow and
Elytra. Surface smooth with flat to slightly raised,
minute granulations (more apparent in teneral adults). Stria
markedly developed (more apparent in teneral adults)
comprising a longitudinal pair of microcarina. Minimal to no
longitudinal groove bordered by microcarinae.
Sternites. Surface of mesobasisternum coarsely
shagreened, large crescent-shaped punctations, densely
arranged and often linked, radiating anterior-laterally from
centre of mesosternal process. Punctations generally armed
with single very short brown/black setae.
Legs. Medial facet of protibia slightly angled inwards
from between third external denticle to half-way to second
external denticle (Figs 37–41). Even protibial width from
base to parallel with third external denticle, where broadens
slightly to apex. Protibial angulation and start of angulation
more subtle in females (Figs 38, 40, 41). Mesofemora armed
with three (Fig. 53) to four (Fig. 52) rows of long thick
black/brown setae on posterior facet. Most rows
approximately equal in length. Second row from ventral
marginal row often reduced setation. Mesotibia slightly
curved inwards (Fig. 19). Outer mesotibial spur 1/3 length of
mesotibia. Inner spur 2/3 length and 1/3 width of outer spur.
Inner mesotibial spur offset from the outer spur by 10 to 20
degrees. Mesotarsus 2/3 length of mesotibia.
Male genitalia (Fig. 12). Handle of virgular sclerite of
relatively constant thickness through most its length,
widening slightly at each end. Handle evenly concave along
majority of dorsal and ventral margin. Dorsal margin
unnotched at terminus of union with circular sclerite. Apical
margin of handle rectangular. Basal margin between dorsal
and ventral corners slightly concave. Secondary
sclerotisation of handle reduced in apical ventral corner and
apical dorsal corner. Baso-ventral corner forms protruding
extension with reduction in secondary sclerotisation.
Geographical distribution
Sceliages difficilis is known from South Africa and
Zimbabwe (Fig. 83).
The etymology of the species name difficilis could be based
on the difficulty to accurately diagnose this species.
Sceliages difficilis closely resembles Sc. gagates and Sc.
adamastor. Geographic distribution (Sc. gagates is restricted
to the eastern coastal regions), differences in width
curvature/angulation of the protibiae and pronotum lateral
margins (more evenly rounded in Sc. difficilis and more
obtusely rounded/flared in Sc. gagates) are the principal
characters for separating the two species. Some specimens of
Sc. difficilis have been misidentified as Sc. adamastor. This
may be due to the curvature of mesotibia reminiscent of this
strong diagnostic feature in Sc. adamastor (Fig. 17). Note,
however, on Sc. adamastor, the outer mesotibial spur is
markedly shorter than the length of mesotibia (Fig. 17)
compared with Sc. difficilis (Fig. 19) and the mesotarsi of
Millipede-eating Sceliages (Scarabaeidae) from Africa 941
Sc.difficilis are 2/3 the length of the mesotibiae (cf.
mesotarsi of Sc. adamastor are 1/2 the length of the
Sceliages gagates Shipp
(Figs 13, 20, 42–45, 54, 55, 82)
Sceliages gagates Shipp, 1895: 38. – Gillet, 1911a: 16; Ferreira,
1961: 64; Ferreira, 1967: 60; Ferreira, 1972: 78; zur Strassen,
1965: 226–229.
Material examined
Neotype., ‘Sordwana Bay, Natal (3 km from camp); 15.x.1978,
Bornemissza & Aschenborn’, SANC, database No: COLS 00045.
(Here designated.)
Other material examined.South Africa, Kwazulu-Natal: Ko zi
Bay (26°52S 32°50E) (1 UPSA); Muzi, Tongaland (Ingwavuma
Distr.) 26°52S 32°29E (2 , 1 SANC); Muzi Area (Ubombo Distr.)
‘N. Natal’(27°29S 32°23E) (5 ,2 SANC); Sodwana Bay (27°32S
32°41E) (2 SANC); St Lucia Estuary (28°21S 32°29E) (1 ,1
SANC); Tembe Elephant Pk, ‘N. Kwazulu Natal’ 26°51S 32°24E (1
UPSA). North-west Province: (?)Sand R. Mtn, 24°32S 27°39E (1
UPSA). Moçambique: Delagoa Bay (Maputo, Baía de) (25°48S
32°51E) (6 , 6 BMNH): ‘Delagoa’ (1 BMNH; 3 ISNB);
‘Dela. B., Mozamb.’ (1 BMNH): Inhambane (23°52S 35°23E) (1
SAMC): Lourenço Marques (Maputo) (2 SAMC): Nyaka, ‘P E.
Afr.’(30°9S 29°46E) (1 SAMC).
Diagnostic morphological characters that distinguish
Sc.gagates from Sc. difficilis are: the lateral margins of the
pronotum (obtusely rounded; lateral margins flared medially
and narrow posteriorly; dorsal edge of margin upturned
slightly); and the differences in protibia angulation (medial
facet of protibia slightly angled inwards from second
protibial denticle; even and slight increase of protibial width
from base to apex). The distribution of Sc. gagates is
restricted to the coastal regions of north-eastern South Africa
and Moçambique and is regarded as a key diagnostic
character for the species.
Length 10–18 mm.
Pronotum. Lateral margins obtusely rounded, flared
medially and narrow posteriorly. Dorsal edge of margin
upturned slightly.
Elytra. Surface smooth with flat, minute granulations
(more apparent in teneral adults). Stria comprising a groove
without longitudinal microcarina bordering either side.
Sternites. Surface of mesobasisternum with
pronounced crescent-shaped, posteriorly facetted
punctations. Punctations raised forming protrusion or
turbercle-like structures; dense but generally unlinked,
radiating antero-laterally from centre of mesosternal
process. Each punctation armed with a single red–brown
Legs. Medial facet of protibia slightly angled inwards
from second protibial denticle (Figs 42–45). Even and slight
increase of protibial width (in dorsal perspective) from base
to apex. Mesofemora armed with two (Fig. 54) to three
(Fig. 55) rows of red/brown setae on posterior facet. First
row dense and mostly uniform on postero-ventral margin.
Second row with setation well spaced and reduced in number
from first row. Third row further reduced to absent. Outer
Figs 46–55. Mesofemora of Sceliages; setation/pubescence on posterior facet. Left margins are dorsal (lateral perspective). Scale bars: 1 mm. 46,
Sceliages granulatus; 47, Sc. hippias; 48, Sc. brittoni and Sc. augias (Casonda, Angola); 49, Sc. augias (Mpwapura); 50, Sc. adamastor (De Hoop
Nat. Res., RSA); 51, Sc. adamastor; 52, Sc. difficilis (Badplaas, RSA); 53, Sc. difficilis (Boekenhoutskloof, RSA); 54, Sc. gagates (Muzi Area,
RSA); 55, Sc. gagates (Muzi Area, RSA).
942 S. A. Forgie et al.
mesotibial spur 1/3 length of mesotibia. Inner spur 2/3 length
and 1/3 width of outer spur. Inner mesotibial spur offset from
outer spur by 20 to 30 degrees. Mesotibia slightly curved
inwards. Mesotarsi greater than 2/3 length of mesotibia.
Male genitalia (Fig. 13). Handle of virgular sclerite
relatively thin medially widening at each end. Handle evenly
concave along dorsal margin of medio-basal region of handle
and majority of ventral margin. Dorsal margin extended
horizontally outward from terminus of distal union with
circular sclerite prior to upward angulation to apex. Apical
margin of handle roughly triangular. Basal margin between
dorsal and ventral corners straight. Secondary sclerotisation
of handle heavily reduced in apical ventral corner and
extending baso-ventrally in apical region. Baso-ventral
corner forms a pronounced downward protruding extension
with reduction in secondary sclerotisation throughout and
extending to baso-dorsal union with circular sclerite.
Geographical distribution
Sceliages gagates is known from the lowland coastal regions
of north-eastern South Africa and southern Moçambique
(Fig. 83).
Type specimens could not be located for this revision, nor
could their existence be confirmed (Horn et al. 1990). Only
a few types from the original Shipp collection were retained
by the Hope Entomological Collections, Oxford University
Museum of Natural History. Some of the collection was
purchased by P. M. Bright, but that part contains no
scarabaeines (D. Mann, personal communication). The
location of the location of most of Shipp’s types remains a
mystery. According to zur Strassen (1965), a [holo]type of Sc.
gagates with the locality ‘limpopo’ was apparently lodged
with the BMNH. All specimens of Sceliages were loaned
from the BMNH for this revision and no types of Sc.gagates
were present. Zur Strassen (1965) was met with the same
circumstances and also questioned the existence of types,
most especially, the supposed type specimen. Zur Strassen
(1965) also argued that the original description by Shipp was
inaccurate and could easily have been based on a specimen
of Sc. hippias lacking its yellow apical antennomeres, or Sc.
difficilis, an undescribed species in 1895.
With an absence of types, our description of this species
was initially based on the original description by Shipp
(1895) and, in accordance with zur Strassen (1965), found
Shipp’s description more akin to that of Sc. difficilis. We
therefore relied on the description and key by zur Strassen
(1965) and examination of as many specimens we could
obtain. We noticed an obvious lack inzur Strassen’s work of
the geographic localities of the specimens he examined and
subsequently identified either as Sc. gagates or Sc. dificilis,
which made their differentiation rather ambiguous. This lead
us to redescribe the species based on lowland coastal
specimens of Sc. gagates or mislabelled Sc. difficilis, which
possess uniformity in the morphological characters we
regard as descriptive for Sc. gagates.
Based on the current redescription of Sc. gagates the
[holo]type with the locality ‘limpopo’ is likely to be correct
only if the specimen was collected in the eastern lowland
coastal region of Moçambique through which the Limpopo
River flows. However, its location, or that of any other type
for this species, remains a mystery. General agreement by
those who have helped us try to locate missing types is that
they should be treated as non-existent and neotypes should
be assigned. This we have done.
Subtle intra-specific variations or external wear on
diagnostic characters of both Sc. gagates and Sc. difficilis
can easily lead to misidentification for either species. There
are relatively few consistently sound diagnostic characters
that could be described to separate the two species.
Ambiguous male specimens of Sc. gagates that could be
misidentified as Sc. difficilis do, however, show reasonably
consistent similarities in their virgular sclerites.
Sceliages granulatus Forgie & Scholtz, sp. nov.
(Figs 1, 6, 7, 24, 25–28, 46, 80)
Material examined
Holotype., ‘Botswana: Kang, 35 km SE, millipede-baited
pitfall trap; 23 January 1978, A.L.V. Davis’ (23°46S 22°51E), SANC.
Paratypes.3 , 1 , same data as holotype, SANC; 1 , 2 ,
‘Botswana: Sekoma, 26 km E, millipede-baited pitfall trap, 24–25
January 1978, A.L.V. Davis’ (24°24S 23°53), 1 in UPSA, 2 in
Other material examined.South Africa: 1 , ‘Northern Cape
Province; Olifantshoek, 45 km SW, 28 February 1973, Bornemissza
and Temby’ (27°56S 22°44), SANC; 1 , ‘Vryburg’ (26°57S
24°44), SAMC; 2 , 1 , ‘South Africa: ‘Cape Province’, Kimberley
28°44S 24°46, October–November, 1980, S. Erasmus’, SANC.
Sceliages granulatus shares a similar overall appearance
with small teneral adults of Sc. difficilis except for the
following differences present in Sc. granulatus: an obvious
indumentum covering the elytra; granulations on elytra more
obvious and pronounced; striae on the elytra are more
pronounced; pronotum lateral margins are obtusely rounded
(similar to Sc. gagates).
Length 11–13 mm.
Head. Surface rugose, densely punctated on genae and
clypeus. Punctations on marginal regions of clypeus and
gena form prolonged ripple-like ridges. Geno-clypeal suture
obscured along apical half by punctations and roughness of
surface near geno-clypeal margins. Antennae armed with
dense array of pale minute sensilla on apical margins and
dorsal surfaces on antennal club antennomeres.
Millipede-eating Sceliages (Scarabaeidae) from Africa 943
Pronotum. Surface coarsely shagreened, covered in
minute punctations and fine, flat granulations. Lateral
margins of pronotum obtusely rounded. Width of pronotal
lateral margins constant in posterior half. Angulation of
lateral margin without variation posteriorly and minimal to
none medially. Entire dorsal lateral margin well defined.
Elytra. Surface coarsely shagreened, covered in well
developed granulations. Stria bordered with 2 evenly spaced
carinae; granulation absent in strips running parallel with
outer margins of each carina. Elytra covered with thin layer
of indumentum providing dull matt appearance.
Sternites. Surface of mesobasisternum finely
shagreened, covered with large punctations often linked in
ripple-like chains of four or more. Punctations
crescent-shaped and slightly facetted. Facets between inner
margins of mesocoxal cavities and lateral carinae of
mesosternellum undermined posteriorly and straightened to
vertical at deepest point. Each facet outwardly curved
twisting to horizontal anteriorly. Carina curved at union with
mesobasisternal–mesepisternal suture.
Legs. Medial (inner) facet of protibia poorly curved
inwards. In females, curvature even (Figs 26, 27). In males,
curvature subtly more apparent from second external
protibial denticle (Figs 25, 28). Medial facet of protibia
twisted by 45 degrees from vertical in proximal half to
horizontal position in distal half. In males, medial carina of
keel of first (most basal) protibial denticle reaches dorsal
longitudinal carina. Antero-ventral margin of profemora
adjacent to protrochanter armed with one spur-like
projection. Posterior facet of mesofemora armed with three
rows of setae running parallel to dorsal and ventral margins
(Fig. 46). Rows of variable length. Setae of variable density.
Vagrant setae located sparsely between rows. Mesotibiae
poorly curved inwards. Outer mesotibial spur approximately
1/3 length of mesotibia. Inner mesotibial spur more than 2/3
Length of outer spur. Inner mesotibial spur offset from outer
spur by 15 degrees. Mesotarsus approximately 2/3 length of
mesotibia. Metatibia (in dorsal perspective) without inward
curvature (Fig. 24).
Male genitalia (Figs 6, 7). Handle of virgular sclerite
broadly concave along medial region; dorsal margins steeply
curved upwards at both ends. Apical region tapers to its
widest at apex. Apex rounded at dorsal corner; angled at
ventral corner. Secondary sclerotisation reduced transversely
in ventral corner of apex of handle. Basal region of handle
has reduction of secondary sclerotisation at dorsal corner.
Baso-ventral corner of handle has pronounced rectangulate
Geographical distribution
Sceliages granulatus is known from semi-arid regions of
southern Botswana and the northern part of central South
Africa (Fig. 81) in areas considered to represent Kalahari
The virgular sclerite of Sc. granulatus from Kimberley
(Fig. 7) differs slightly from the Botswana specimens (Fig. 6)
in possessing a slightly thinner, less tapered apical region
without angulation at the apex. Generally, virgular sclerites
show little to no intraspecific variation and are therefore
regarded as species specific and highly conserved. Only two
male specimens have been collected from Kimberley, which
prevents testing whether or not these differences are
consistent. Although no external differences are apparent
between the South African and Botswana specimens, the
former have not been included in the type series.
Specific epithet
The species name is derived from the small but pronounced
granulations on the elytra.
Sceliages hippias Westwood
(Figs 3, 8, 14, 15, 18, 29, 30, 47, 56–79, 80)
Sceliages hippias Westwood, 1844: 100. – Shipp, 1895: 39;
Péringuey, 1901: 64; Gillet, 1911a: 16; Ferreira, 1961: 65;
Ferreira, 1972: 78; zur Strassen, 1965: 221, 222, 224, 226–228.
Ateuchus microcephalus Boheman, 1857: 176. – Shipp, 1895: 38.
Material examined
Holotype., ‘?type’ ‘Int. S. Afr.’, BMNH 54-76.
Other material examined.South Africa: ‘Cap’ (1 ISNB).
Mpumalanga: Watervalriverpass, 24°54S 30°21E (1 TMSA);
Lydenburg, Waterval Pass, 11 km NE (1 TMSA); Lydenberg District
(1 , 1 SAMC); Schmoemansville (25°01S 30°31E) (1 TMSA).
Kwazulu-Natal: ‘Burn’ (1 SAMC); Tugela Ferry (28°44S 30°27E)
(1 TMSA); Weenen (28°51S 30°04E) (2 BMNH); Tugela R.,
Wee n e n ( 1 , 1 DMSA). Northern Province: Blouberg (23°05S
29°01) (1 TMSA); Blouberg, 23°05S 29°01 (1 SANC);
Nylstroom 24°40S 28°15E (1 UPSA); Nylsvlei, Smith Farm
24°40S 28°42E (1 TMSA); Pietersburg, 24°1440′′S 29°1530′′E (1
, 2 UPSA); Rhenosterpoort Farm (24°45S 28°23E) (2 TMSA);
Vic. Mmafete 24°11S 30°06E (1 UPSA); Warmbad (24°50S
28°20E) (1 UPSA). North-west Province: Rustenberg Nature
Reserve 25°40S 27°12E (1 , 1 SANC; 1 , 8 ) UPSA);
Tonguani Gorge, Pretoria, 81 km W (25°50S 27°30E) (1 SANC);
Retiefskloof, Rustenberg, 15 km SE (25°49S 27°16E) (1 SANC);
Thambazimbi, 30 km W, SE (24°35S 27°20E) (1 UPSA). Gauteng:
Johannesburg (26°12S 28°05E) (3 TMSA; 1 SAMC);
Kameeldrift, Pretoria, 24 km NE (25°39S 28°21E) (1 SANC): ‘nr
Johannesburg’ (1 BMNH 1906-29); Roodeplaat Farm, Pretoria, 20
km NE (25°34S 28°22E) (1 SANC); Hennopsriver, Pretoria, 32 km
W (25°50S 27°58E) (1 SANC); Pretoria (3 SANC; 1 SAMC;
1 , 1 UPSA); Swartkops, nr Jhb (26°05S 27°45E) (1 BMNH);
Welgedacht, Pretoria, 50 km N (25°20S 28°20E) (1 UPSA).
Sceliages hippias is easily identifiable with the
yellow/orange colour of the antennomeres, especially in live
specimens. Should this character be ambiguous or missing in
dead pinned specimens, then correct diagnosis of the species
944 S. A. Forgie et al.
is assured with the unique setation on the posterior facet of
the mesofemora and mesotibiae are relatively short, evenly
tapering to the distal apex and without any curvature.
Length 12–16 mm.
Head. Surface of frons and vertex smooth with smaller,
fewer punctations than in geno-clypeal region. Geno-clypeal
suture is obscured by dense punctations and rough surface
along its apical half. Antennal club yellow~orange.
Pronotum. Lateral margins of pronotum obtusely
rounded. Angulation of lateral margins of pronotum
unvaried. Lateral margins of even width in posterior half.
Elytra. Surface smooth, finely punctated and without
waxy indumentum. Striae faint, lineal grooves infrequently
bordered by minute carinae, and interrupted by punctations.
Sternites. Surface of mesobasisternum smooth with
large, evenly spaced, scalloped punctations radiating in
vague linear fashion, antero-laterally from centre of
mesosternal process. Mesosternum loosely covered in
minute punctations, most armed with a single minute pale
Legs. Medial facet of protibia angled slightly, if at all,
inwards from second protibial denticle (Figs 29, 30). Protibia
width increasing evenly and slightly from base to apex.
Antero-ventral margin of profemora adjacent to trochanter
an obvious ridge, rounded, without serration; spur-like
projection absent (Fig. 3). Mesofemora armed with single
row of setae closely paralleling ventral margin in basal half
of posterior facet (Fig. 47). Inner facet of mesotibia straight
(Fig. 18). Outer mesotibial spur approximately 1/4 length of
mesotibia. Inner mesotibial spur equal in length to outer spur
and 1/2 its thickness. Angle of inner mesotibial spur is
generally offset from outer spur by 10 degrees. Mesotarsus
approximately 2/3 length of mesotibia.
Male genitalia (Fig. 8). Handle of virgular sclerite thin
in width, unevenly concave along baso-dorsal and ventral
margins, and of relatively constant thickness. Dorsal margin
angle down or notched at distal terminus of union with circular
sclerite. Apical region of handle abruptly curved upward
giving handle an overall sickle-shape appearance. Apical
region is slightly thicker than basal region and rectilinear at
its apex. Both dorsal and ventral corners of apex have
approximately equal reduction in secondary sclerotisation.
Geographical distribution
Sceliages hippias is known only from the north-eastern
corner of South Africa (Fig. 81).
Much of the Westwood collection, including many of the
types, was housed in the Zoological Society of London’s
collection, which was later incorporated into the main series
collections of the BMNH before 1900. According to
D. Mann (personal communication), the Westwood type of
Sceliages hippias, if extant, would have been in the series we
received from the BMNH for this revision. D. Mann warned
that the specimen (if present) may not be labelled as a
Westwood species, as the labelling of that period was scant
to say the least. We examined a specimen labelled ‘?type’
and ‘Int. S. Afr.’ from the BMNH series with an accession
number, 54-76. The type label may well have been added by
zur Strassen during his revision of the genus in which he
states this specimen, along with the female Ateuchus
microcephalus paratype labelled ‘Caffraria’ in The Natural
History Museum of Stockholm, are dubious. Zur Strassen
(1965) concluded in his discussion of Sc. hippias that it is
likely that the holotype and paratype(s) are lost and should be
regarded as non-existent. However, further examination of
the BMNH ‘?type’ specimen labels, and cross referencing of
the museum accession number with the accessions registrar
in the entomology library of the BMNH, reveals this
specimen to be the actual holotype for Sceliages hippias. The
specimen in question has subsequently been correctly
labelled as the holotype by D. Mann and remains housed in
the BMNH.
Redescription and identification of this species was
therefore based on the descriptions by Westwood (1844) and
zur Strassen (1965), identified material and a diagnostic key
of the genus by zur Strassen (1965).
NB. If some or all of the setae on the posterior facet of the
mesofemora are absent, then the setal sockets should be
evident enough to identify this species.
Description of mature larvae of Sceliages hippias
The appearance of mature Sc. hippias larvae is typical of
those of the subfamily Scarabaeinae (Edmonds and Halffter
1978). They differ, however, from other larvae of the
subfamily in possessing the following unique (*) or rarely
found characters: (1*) tormae and epitormae are markedly
reduced, close to absent; (2) dorsal surface of the stipes is
without an irregular row of conical (=‘stridulatory’) teeth
along the basal margin; (3*) hypopharyngeal area is without
two dissimilar sclerites (oncyli); (4) antennae with three
segments; (5) venter of the last abdominal segment lacks any
rows or patches of short setae; and (6) raster absent.
Body-shape typical for Scarabaeinae larvae: whitish body
strongly bent at about middle with markedly developed
secondary dorsal folds. Head capsule (Figs 58–61): width c.
3.0 mm. Each hypostomal ridge is subdivided into two short
sub-elements. Epipharynx (Fig. 57): tormae markedly
reduced and nearly absent. Antenna (Figs 71, 72): three
segments; basal segment with five pores and no setae;
middle segment with flat sensorium, two pores and four
Millipede-eating Sceliages (Scarabaeidae) from Africa 945
setae; distal segment with three apical conical sensillae and
nine pores on ventral surface. Mandible (Figs 62–67): nearly
symmetrical; scissorial part on right mandible markedly
shorter than that on left one; each mandible with three groups
of short setae in molar part and with one long seta in
proximal third of lateral surface. Maxilla (Fig. 69): dorsal
surface of stipes without irregular row of conical teeth along
basal margin. Labium and hypopharynx (Fig. 69):
hypopharyngeal bracon as on Fig. 69; hypopharynx without
oncyli. Thorax (Figs 77): pro- and metathorax not
subdivided by folds; mesothorax weakly subdivided dorsally.
Tergum of prothorax with anterior process on each side.
Mesothoracic spiracles not found. Legs (Figs 73–76):
two-segmented, with weak additional dorsal fold between
presumably trochanter and femur. Abdomen (Fig. 56):
segments 1–5 subdivided dorsally in three dorsal lobes;
segment 6 subdivided in two dorsal lobes; segments 7–10 not
subdivided. Anal opening transverse (Fig. 79). Raster absent.
Remarks and discussion
Sceliages hippias larvae resemble those of dung-feeding
Scarabaeinae groups. They share a number of morphological
characters outlined by Edmonds and Halffter (1978; with a
few exceptions, see description). In the current paper we
outline some of the major characters of Sc. hippias larvae
and compare them to the larvae of other taxa of the
Scarabaeinae. Putative phylogenetic trends are discussed
(1) Tormae and their associated sclerites (epitorma,
dexiotorma, laeotorma, pternotorma) are markedly
reduced or absent. These structures are normally present
throughout larvae of the Scarabaeoidea (Ritcher 1966;
Edmonds and Halffter 1978). In some cases tormae are
not united mesally or are somewhat reduced (Ritcher
1966; Schuster and Reyes-Castillo 1981). As far as we
are aware, the degree of reduction of the tormae in larval
Sc. hippias is the most extreme within the subfamily.
(2) The dorsal surface of the stipes is without an irregular
row of conical teeth along the basal margin. These teeth
are present throughout Scarabaeinae (except Sisyphus;
see Edmonds and Halffter 1978) and many other groups
in the Scarabaeoidea. We are not aware of a study
demonstrating homology in these structures between the
different lineages of the Scarabaeoidea. Occasionally
these teeth are referred to as stridulatory teeth (Ritcher
1966: 26). This assumption has been doubted by
Edmonds and Halffter (1978: 313). Moreover,
Hirschberger and Rohrseitz (1995) were not able to
detect any sound pattern from Aphodius larvae
possessing these teeth (P. Hirschberger, personal
(3) The hypopharyngeal area is without oncyli.
Hypopharyngeal sclerotisation is present in all known
larvae of the subfamilies Scarabaeinae and Aphodiinae.
Hypopharyngeal sclerotisation is also present in the
larvae of the families Geoptrupidae and Lucanidae. As
with the presence of ‘stridulatory’ teeth on the stipes, no
proof has been found to demonstrate the homology of
these structures in the different groups of the
Scarabaeoidea. The absence of these sclerites in
Figs 56–57. Mature larva of Sceliages hippias. 56, Habitus, lateral view, scale bar: 5 mm; 57, epipharynx, scale bar: 0.2 mm.
946 S. A. Forgie et al.
Sc.hippias larvae appears to be unique within the
(4) The antennae of Sc. hippias larvae consist of three
antennomeres. In contrast, all other described larvae
within the Scarabaeinae possess four antennomeres,
even when the subdivision between the two basal
segments is poorly distinguished. In Sc. hippias, this
subdivision is absent and, consequently, the two basal
antennomeres are fused to form a single one. A
three-segmented antenna is a character often utilised to
separate Geotrupidae larvae from those of Scarabaeinae
possessing four-segmented antennae. Three-segmented
antennae have also been found in one unidentified
Scarabaeus larva from South Africa (Grebennikov and
Scholtz, unpublished data).
(5) The raster is absent from the ventral surface of the last
abdominal segment of Sc. hippias. This structure is
often inconspicuous, but still present in larvae of the
Scarabaeinae (except larvae of the genus Sisyphus;
Edmonds and Halffter 1978). We failed to see any
setation on the median part of the tenth ventrite, even
under high magnification.
Biology and nidification
Three relatively abundant species of millipedes from the
order Spirostreptida are utilised by Sc. hippias at the
Rustenberg Nature Reserve. Two species belong to the
family Spirostreptidae: Doratogonus rugifrons (Attems
1922) and D. levigatus (Attems 1928). The former is a large
black species approximately 12–14 cm long and 4–11 mm
wide. The latter species is approximately 8–11 cm long and
5–7 mm wide. The third large orange/brown-banded species,
Zinophora robusta (Attems, 1928), with similar dimensions
to D. levigatus, belongs to the family Harpagophoridae.
Attraction to millipedes
It was important to confirm whether attraction to millipedes
by Sceliages beetles is primarily due to a positive chemotaxic
response to the quinone-based secretions produced by the
millipedes (Krell et al.1998). To provisionally test this,
healthy, uninjured Z. robusta and D. rugrifrons millipedes
were wrapped in pieces of tissue paper and agitated to collect
their quinone-based secretions (after Krell et al. 1997).
Figs 58–65. Mature larva of Sceliages hippias, details. Scale bars: 1 mm. 58, Head, dorsal; 59, head, ventral; 60, left mandible, dorsal;
61, left mandible, mesal; 62, left mandible, ventral; 63, right mandible, dorsal; 64, right mandible, mesal; 65, right mandible, ventral.
Millipede-eating Sceliages (Scarabaeidae) from Africa 947
These pieces of tissue (including a control piece of tissue
paper not containing secretions) were then suspended above
three pitfall traps and left for approximately 1/2 an hour at
1000 hours. Only the ‘quinone’ traps collected several adult
Sceliages. A healthy, uninjured millipede suspended above a
pitfall trap also attracted four adults of Sceliages within 1/2
an hour. Other species of beetles known to utilise millipede
carcasses in the research area were not attracted to the traps.
Millipede relocation
We tested the assumption that Sceliages make balls from the
internal tissues of a freshly crushed millipede and roll the
ball backwards using its forelegs for locomotion. We crushed
several millipedes and left them for observation. Beetles,
instead of making a ball, utilised the freshest and most intact
portion of the millipede, which was relocated using their
head and forelegs. Whole millipede corpses and injured
millipedes were relocated in the same way up to a distance of
5 m.
Laboratory trials were set-up using six pairs of Sc.
adamastor to observe male7female co-operation in
millipede burial. Fighting took place when a single millipede
corpse was introduced to each pair. When a second millipede
corpse was added to each pair, relocation and burial were
carried out individually. Similarly, no male–female
cooperation of Sc. hippias was observed in the field.
In Rustenberg Nature Reserve, six females of Sc. hippias
were each given a whole millipede carcass in order to
observe burial behaviour and to obtain larvae. Locations of
each burial site were tagged and recorded using global
Figs 66–67. Mature larva of Sceliages hippias, details. Scale bars: 1 mm. 66, Head, frontal; 67,
head, lateral.
948 S. A. Forgie et al.
positioning system. Two burial techniques were observed.
(1) Sceliages hippias excavates directly in front of the
millipede corpse and draws it into the tunnel as it is
excavated. Initially, this is carried out by undermining the
millipede either directly underneath or at an angle at one end
of the millipede. Tunnels constructed by Sc. hippias are
relatively straight and excavated at a 30° angle. (2) A female
of Sc. adamastor was observed at the De Hoop Nature
reserve excavating a tunnel wide enough for both beetle and
millipede to fit. Once the tunnel was started, the beetle
aligned the millipede lengthways at its entrance and pushed
the millipede in gradually from behind making directional
and postural adjustments of the corpse using its head.
Six burial sites were excavated one month later. Four sites
contained vacant burrows and/or chambers containing the
empty, disarticulated segments of the millipedes presented to
the beetles a month earlier. Two sites contained brood
chambers with balls and an accompanying adult female.
Each brood ball contained a third-instar larva. Chambers
were found at a depth of 7 to 14 cm beneath the surface. The
Figs 68–72. Mature larva of Sceliages, details. 68, Right apical labial palpomere, dorsal, scale bar:
0.01 mm; 69, labium, hypopharynx and right maxilla, dorsal, scale bar: 0.25 mm; 70, right maxillary
palp, dorsal, scale bar: 0.1 mm; 71, left antenna, ventral and 72, left antenna, dorsal, scale bar: 0.1 mm.
Millipede-eating Sceliages (Scarabaeidae) from Africa 949
tunnel leading to each chamber, and the entrance side of the
chambers themselves, were filled with empty, separated
millipede exo-skeletal body segments.
Brood balls
Millipedes of different sizes were measured to see how many
balls a reproducing female of Sc. hippias could make from
them. Beetles presented with millipedes up to 7 mm in
diameter produced a single brood ball. Millipedes around 11
mm in diameter yielded two brood balls. Three millipedes
between 7 and 8 mm in diameter were presented to a single
female to test whether she could utilise the entire resource
and produce more than the standard one or two brood balls
from a single millipede. Four weeks later, the same female
was recovered brooding three balls: two balls containing
third-instar larvae; and a third with a second-instar larva.
Brood balls are pear-shaped. The ball is encapsulated by a
compact protective layer of soil up to 3.5 mm thick. Soil is
also used to make an egg chamber on the side of the ball. The
larval food ball of Sc. hippias is approximately 12 mm in
diameter and constructed from the internal tissues, intestinal
dung and remnant chitinous pieces of the millipede.
Addition of a compacted soil substrate occurs within the
brood chamber and is thought to prevent desiccation and
protection against pathogens (Halffter and Matthews 1966).
Parental brooding by female Sceliages occurs in conjunction
with the soil-encrusted balls, a behaviour different to that
described for the majority of the Scarabaeini that were
thought not to brood or encrust balls with soil (Halffter and
Edmonds 1982: 40).
Phylogenetic analysis of the genus Sceliages Westwood
Tax a
A cladistic analysis was performed including all seven species of
Sceliages. Ingroup-taxa character states were polarised against two
species of Scarabaeus, S. zambesianus Péringuey, 1908 and S. rusticus
(Boheman, 1857). Outgroup selection was based on a preliminary
combined morphological and molecular phylogenetic analysis of the
tribe by Forgie, Philips and Scholtz (unpublished data), taking into
consideration arguments by Nixon and Carpenter (1993). Twenty-seven
characters (including three multi-state) were coded from the sclerotised
external structures of teneral adults. The aedeagus and virgular sclerite
of the internal sac were also utilised (see Materials and methods, ‘Male
genitalia’for preparation). Larval characters were not used owing to a
lack of material. Morphological characters and their states were
described using the terminology of Doyen (1966) and Lawrence and
Britton (1991).
Phylogenetic analysis
A character matrix was compiled in DADA, version 1.2.7 (Nixon 1998).
Sceliages hippias has several interesting character states (i.e. character
0/state 1,16/0, 19/0 and 21/0), which it shares with at least one of the
outgroup taxa. These states are either plesiomorphic or were
convergently evolved. Symplesiomorphies are also present in Sc.
Figs 73–76. Mature larva of Sceliages hippias, details. Scale bars: 0.4 mm. 73, Left leg, anterior; 74, left leg, posterior; 75, apex of left leg,
anterior; 76, apex of left leg, posterior.
950 S. A. Forgie et al.
adamastor (26/0) and Sc. brittoni (24/0). Autapomorphic character
states, although info rmative in describing the uniquen ess of each species,
are uninformative in the analyses and bias the consistency and retention
indices by having zero homo plasy. Autapomor phic character states were
therefore avoided during coding and their absence was conf irmed using
the mop-up option in DADA. Character states unique to Sceliages are
informative in differentiating the genus from other genera within the
Scarabaeini, but are generally uninformative in a species-level
phylogenetic study such as this and were therefore not coded.
All characters were spawned in NONA (Goloboff 1993) with 1000
repetitions to ensure all the shortest cladograms were found utilising
branch and bound search options with randomised taxon order in each
run. A single tree found in NONA was submitted to Hennig86, version
1.5 (Farris 1988) and subjected to successive approximations
weighting, hennig tree construction and branch breaker options (xs w,
mh*, and bb* commands respectively). Bremer support (decay index)
was calculated with NONA up to a value of 5, i.e. searching for trees up
to five steps longer in the tree(s) submitted for calculation. Trees were
also calculated in Parsimony and Implied Weights (PIWE), version 2.6
(Goloboff 1993, 1997). Five levels of concavity (0, 1, 2, 3 [default], 4,
5) where applied to the characters using rs0, hold1000, hold/100,
mult*100 commands. High repetitions run in NONA ensured the best
PIWE trees were generated. All DOS-based analysis programmes were
run through WinClada (BETA), version 0.9.9 (Nixon 1999a).
Consistency (CI) and retention indices (RI) (Farris 1989) are
indicated for each character in the list below. A matrix of taxa and
character states is provided in Table 1.
List of characters and their states
00. Apical antennomeres: (0) brown to black; (1) yellow to orange. CI,
0.50; RI, 0.00.
01. Medio-longitudinal hump-like process on clypeus: (0) extending
anteriorly; (1) absent anteriorly (clypeus c. flat). CI, 0.50; RI, 0.50.
02. Geno-clypeal sutures: (0) curved medially; (1) c. straight. CI, 0.50;
RI, 0.50.
03.Punctations on dorsum of head plates: (0) uneven and irregular; (1)
even and regular. CI, 0.50; RI, 0.66.
04. Head plate dorsal surface: (0) rugosissimus; (1) rugulosus. CI,
0.50; RI, 0.66.
Figs 77–79. Mature larva of Sceliages hippias, details. Scale bars: 1 mm. 77, Thorax and two first
abdominal segments, lateral; 78, venter of last abdominal segment; 79, anal opening and anal lobes.
Table 1. Character states of the taxa used in the phylogenetic
analysis of the genus Sceliages Wes two o d
Outgroup taxa are indicated in bold.
1 2
Scarabaeus zambesianus 000000000001000000000000100
Scarabaeus rusticus 100000000111000000000000001
Sceliages adamastor 011110000110101011121111210
Sc. augias 000001111011100010021111211
Sc.brittoni 011110100111011111121111011
Sc. difficilis 000110011010110121121111101
Sc. gagates 000001110100110020010101101
Sc. granulatus 000001111010111020010111101
Sc. hippias 111111100101101100000001201
Millipede-eating Sceliages (Scarabaeidae) from Africa 951
05. Lateral margins of pronotum: (0) evenly rounded; (1) obtusely
rounded. CI, 0.50; RI, 0.66.
06. ‘Epipleura of the lateral margins of pronotum: (0) even thickness
throughout; (1) not so. CI, 0.33; RI, 0.33.
07. Indumentum on surface of elytra: (0) absent (appearing glossy); (1)
present. CI, 0.50; RI, 0.66.
08. Elytra surface: (0) smooth (without raised granulation or
corrugation); (1) complex (with raised granulation or corrugation).
CI, 0.50; RI, 0.50.
09. Striae on elytra: (0) markedly defined with bordering
microcarinae; (1) not so. CI, 0.33; RI, 0.33.
10. Mesobasisternum punctation: (0) facet of punc ture forming a raised
turbercle-like protrusion (punctation usually crescent-shaped and
may be vestigial); (1) not so. CI, 0.50; RI, 0.50.
11. Mesobasisternum surface (due to size, density and arrangement of
punctations): (0) rugose; (1) not so. CI, 0.33; RI, 0.33.
12. Width between sclerotised medial margins of aedeagus paramere
(anterior frontal view): (0) uneven, widening in apical half; (1)
relatively even through length. CI, 0.50; RI, 0.50.
13. Dorsal margin at distal terminus of union between handle of
virgular sclerite and circular sclerite: (0) downward turned (slight
to markedly), discontinuous with remainder of dorsal margin; (1)
not downward turned, forming a continuous uninterrupted curve
with remainder of dorsal margin. CI, 0.33; RI, 0.33.
14. Baso-ventral corner of handle of virgular sclerite: (0) forming a
protruding extension (slight to markedly); (1) no protrusion. CI,
0.33; RI, 0.33.
15. Thickness of handle of virgular sclerite: (0) uneven, thickest at
basal and/or apical termini; (1) even throughout. CI, 0.30; RI, 0.00.
16. Anterior-ventral margin of profemora adjacent to protrochanter:
(0) uniform and unmodified; (1) armed with markedly developed
spur; (2) armed with a vestigial spur. CI, 0.66; RI, 0.75.
17. Width of protibia: (0) progressive increase in width from thinnest
proximally to thickest distally; (1) abrupt increase in width distally
between second and third external denticle. CI, 1.00; RI, 1.00.
18. Inward angulation of medial facet of protibia: (0) between or at
external denticles 1 and 2; (1) between external denticles 2 and 3
or at external denticle 3. CI, 1.00; RI, 1.00.
19. Setae on posterior surface of mesofemora: (0) forming a single
evenly spaced row of setae paralleling ventral margin; (1) forming
two even rows of individually positioned setae (not clustered)
parallel to dorsal and ventral margins with a medial third row
containing fewer setae present or not; (2) three or more uneven
rows of evenly or unevenly spaced setae. CI, 1.00; RI, 1.00.
20. Setation on posterior surface of mesofemora: (0) sparse; (1) dense.
CI, 1.00; RI, 1.00.
21. Shape of mesotibia: (0) uncurved; (1) curved inwards. CI, 1.00; RI,
22. Tapering/truncation through length of mesotibia: (0) even; (1)
uneven, truncation in apical half or third. CI, 1.00; RI, 1.00.
23. Number of mesotibial spurs: (0) one; (1) two. CI, 1.00; RI, 1.00.
24. Length of major (or outer) mesotibial spur: (0) c. 1/2 length of
mesotibia; (1) c. 1/3 length of mesotibia; (2) much less than 1/3
length of mesotibia. CI, 0.40; RI, 0.00.
25. Length of mesotarsi: (0) c. 2/3 length of mesotibia; (1) less than 2/3
length of mesotibia. CI, 0.50; RI, 0.50.
26. Curvature of metatibia: (0) markedly bowed inwards; (1) c.
straight. CI, 0.50; RI, 0.00.
Results and discussion
Unweighted v. weighted trees
A single most parsimonious tree generated with unweighted
data had a length of 58 steps and consistency and retention
indices of 0.51 and 0.57 respectively (Fig. 80). Weighting of
data was applied two ways, successively in Hennig86 (Farris
1988), and in PIWE (Goloboff 1997), with the latter assigned
five levels of weight (1–5) towards character homoplasy (see
Goloboff 1993). Both methods generated single trees with
identical ingroup toplogies. In addition, there were no
differences in the ingroup topologies between weighted and
unweighted analyses. Scarabaeus rusticus always appears as
sister to the Sceliages s. str. clade in all trees except the
successive approximations weighting tree in which a basal
trichotomy occurs.
Tree support
While the data is obviously very stable, low overall Bremer
support in the tree topology is likely to result from a low
number of characters and the relatively high consistency and
retention indices (CI, 0.51; RI, 0.57) generated (Fig. 80).
One of the highest Bremer values in the tree topology
supports the genus, thereby indicating its monophyly. Very
high Bremer support and bootstrap values support the
monophyly of Sceliages in morphological and molecular
phylogenetic studies of the Scarabaeini (Forgie, Philips and
Scholtz and Forgie, Bloomer & Scholtz, unpublished data).
The apical Sc. brittoni and Sc. adamastor clade also shares
greater support than the remaining nodes of the tree. This
close relationship is also well supported in both of the above
phylogenetic studies by Forgie et al. (unpublished data). Four
characters (i.e. 0, 15, 24, 26) had retention indices of zero,
suggesting their states were uninformative and therefore
unable to support the branch topology of the species in the
genus. Their deactivation did not significantly improve the
decay indices but resulted, as would be expected, in the
construction of a slightly more parsimoniously robust tree
(length 46 steps; CI, 0.54; RI, 0.64). The topology of the new
unweighted NONA tree remained unchanged, again
indicating good stability of the data.
Evolutionary trends
Regardless of which method is used to discover
topologies/trees, the same ingroup relationships are found.
The genus Sceliages is differentiated from Scarabaeus L. s.l.
in this phylogeny by four synapomorphic character states
(5/1, 6/1, 12/1, 23/1), of which character state 23/1 is the
only non-homoplasious synapomorphy (Fig. 80). All species
of Sceliages have a second mesotibial spur (23/1) that is
markedly developed. The development of the second spur is
unique within the Scarabaeini and is likely to play a
functional role in the utilisation of millipedes, such as aiding
grip and leverage on the smooth surface of the millipede as
it is being discarticulated. A second spur, however, is not
unique to Sceliages, having evolved polyphyletically at least
two other times within the Scarabaeini (Forgie, Philips and
Scholtz, unpublished data). A second spur is also present
(albeit vestigial) in Scarabaeus subgenus Scarabaeolus s. str.
952 S. A. Forgie et al.
Balthasar, 1965 species, S. silenus (Gray 1832) and
S.(Scarabaeolus) scholtzi (Mostert and Holm 1982) of the
Scarabaeini and also in some members of the Gymnopleurini
(Mostert and Scholtz 1986).
Sceliages adamastor, Sc. brittoni and Sc. difficilis show a
reversal in the lateral margin of the pronotum from an
obtusely rounded condition (5/1) back to an evenly rounded
plesiomorphic condition (5/0), the latter shared by many
species of the Scarabaeini (Forgie, Philips and Scholtz,
unpublished data). Similarly, Sc. adamastor and Sc. difficilis
both show a reversal in the epipleural condition of the
pronotum lateral margin, from a derived uneven thickness
throughout (6/1) to epipleura that are unvaried or even in
thickness (6/0). The shape of the aedeagus of Sceliages is
largely identical to that of the majority of the Scarabaeus
species, a condition thought by Mostert and Scholtz (1986)
to be due to convergence rather than synapomorphy.
However, a single character state coded from the aedaegus
parameres is synapomorphic to the genus; all species of
Sceliages, with the exception of Sc. brittoni, have a relatively
even width between the sclerotised medial margins of the
aedeagus parameres (12/1). Only Sc. brittoni exhibits a
reversal to the ancestral condition (12/0).
Sceliages hippias represents the most basal species for the
genus and is differentiated from the remaining clade by
several apomorphic character states, which it also shares
with one or both outgroup taxa. These include: yellow
antennomeres (0/0); a uniform and unmodified
antero-ventral margin of the profemora adjacent to the
protrochanter (16/0); a single, uniform row of setae
paralleling the ventral margin of the posterior facet of the
mesofemora (19/0); and uncurved mesotibiae (21/0).
Setation on the posterior surface of the mesofemora
evolved from a plesiomorphic condition of one row
(19/0) to two rows with a medial and sparsely
pubescent third row present or not (19/1), to 3 or more
uneven rows of setae (19/2), which is a synapomorphy
for Sc. augias (Casonda, Angola exemplar), Sc.
difficilis, Sc. adamastor and Sc.brittoni. Such a
condition, although variable in development both
intra-and inter-specifically, evolved after the shift to
millipede necrophagy since the plesiomorphic condition
persists in Sc. hippias, the most ancestral member of
the genus. The function of increased pubescence is
unclear when sparsely pubescent species such as Sc.
hippias are as successful in millipede necrophagy as
species like Sc. brittoni that have dense pubescence on
the posterior surface of the mesofemora.
A waxy indumentum on the surface of the elytra (7/1) is
synapomorphic for the clade excluding Sc. hippias, Sc.
adamastor and Sc. brittoni. The latter two species exhibit an
evolutionary reversal back to the ancestral condition where a
waxy indumentum is lacking (7/0). The semi-arid species,
Sc.granulatus, exhibits the highest degree of wax
indumentation in the genus and the coastal species, Sc.
gagates, exhibits the least. Such a condition aids in the
reduction of cuticular transpiration (Ward and Seely 1996)
and, when viewed in a broader phylogenetic sense, is
homoplastic at least within the Scarabaeini (Forgie, Philips
and Scholtz, unpublished data) and Tenebrionidae (e.g. Ward
and Seely 1996).
The dorsal margin at the terminus of the union between
the handle of the virgular sclerite and circular sclerite forms
a continuous, uninterrupted curvature with the remainder of
Scarabaeus zambesianus
Scarabaeus rusticus
Sc. adamasto
Sc. augias
Sc. brittoni
Sc. difficilis
Sc. gagates
Sc. granulatus
Sceliages hippias
Fig. 80. Hypothesised relationships among species of Sceliages (Sc.) Westwood (Coleoptera:Scarabaeidae) represented by single NONA tree
(length 58 steps; CI, 0.51; RI, 0.57). Black squares indicate non-homoplasious changes (synapomorphies or autapomorphies), white squares
indicate homoplasies. Numbers above squares represent character, numbers below squares represent character states (Nixon 1999b). Bremer
support indices (in bold) are provided next to each ingroup node.
Millipede-eating Sceliages (Scarabaeidae) from Africa 953
the dorsal margin (13/1) in the species of Sceliages, with the
exception of Sc. hippias, Sc. augias and Sc. adamastor. This
synapomorphic state is reversed, in the latter two species, to
the ancestral condition in which a downward turn of the
dorsal margin occurs at the terminus of the union between
the handle of the virgular sclerite and circular sclerite (13/0).
The virgular sclerite of the internal sac of the male genetalia
tends to be highly conserved and useful for assigning reliable
apomorphic character states (Medina 2000), but may be
relatively uninformative for phylogenetic analyses. A
consistency value of 0.33 indicates a relatively high level of
homoplasy for this character.
The ancestral lineage of Sceliages was distributed centrally
within the geographical range of the genus from which
subsequent lineages dispersed, morphologically adapting to
the habitats they are currently distributed in (Figs 81–84).
81 82
83 84
Sceliages gagates Shipp, 1895
Sceliages difficilis Zur Strassen, 1965
Sceliages adamastor (Le Peletier de
Saint-Fargeau & Serville, 1828)
Sceliages brittoni Zur Strassen, 1965
Sceliages augias Gillet, 1908
Sceliages granulatus, sp. nov.
Sceliages hippias Westwood, 1844
Figs 81–84. Distribution maps of Sceliages species in southern Africa.
954 S. A. Forgie et al.
The sister relationship of Sc. adamastor + Sc. brittoni is
reflected in their sympatric distributions (Fig. 84). Both
species are adapted to similar habitats and exhibit the most
congruency in derived morphological character states in the
genus. The phylogenetic derivation of the common ancestor
of Sc. adamastor and Sc. brittoni from that leading to Sc.
difficilis is biogeographically feasible, with a break between
a north-eastern extension of the ancestral populations of the
common ancestor of these species. The topological
positioning of the remaining members of the genus as
successive sisters to one another suggests their radiation
from the ancestral lineage, leading to Sc. hippias, occurred
on several independent occasions.
Concluding remarks
The majority of external morphological characters of
Sceliages are shared with the genus Scarabaeus, however, a
few synapomorphic character states are unique to Sceliages
(see ‘Diagnosis’ of the genus). The presence of a second
mesotibial spur in this phylogenetic study appears as the
single unique character state, yet it is present in other
members of the Scarabaeini including Scarabaeus s. str.
However, the degree of development of the second spur in
Sceliages is unique. Moreover, although necrophagy in the
Scarabaeini is not unique to Sceliages, the apparent obligate
utilisation of millipedes is and further strengthens the
monophyly of the genus. Mostert and Scholtz (1986) likened
Sceliages as the genus closest to the hypothetical common
ancestor of the genera of the Scarabaeini, having undergone
the least morphological evolution. In contrast, a current
phylogenetic analysis of the Scarabaeini by Forgie et al.
(unpublished data) suggests the genus Sceliages is among the
more derived members of the tribe. Evidence from both
studies support the monophyly of the genus.
In this paper we were able to describe some behavioural
characteristics of the adult beetles provisioning nests with
millipedes for nidification. Many questions, however, remain
unanswered: We know quinonous secretions of millipedes are
responsible for attracting Sceliages, however, this was tested
by stimulating a defensive reaction by millipedes. In a natural
situation, are Sceliages beetles attracted to these secretions
produced as allomones in response to the millipede being
threatened or injured, and/or to these secretions being used as
pheromones during millipede mate attraction and copulation?
Do Sceliages beetles kill uninjured millipedes they may have
been attracted to, or, must they rely solely on the demise of
injured millipedes? Is Sceliages truly an obligate necrophage
or are other food types also utilised? Are millipedes utilised
for maturation feeding or nuptial courtship? Exactly how is
the millipede disarticulated? A leverage action using the
clypeal teeth and protibial external denticles is inferred
(Villalobos et al. 1998), but has not been witnessed. We hope
that these questions will stimulate further study on the biology
of Sceliages.
The authors would like to make special thanks to Darren
Mann and to Frank Krell for assisting with location of type
specimens, and also Keith Philips and Frank Krell for helpful
comments on the manuscript. Ute Kryger and Sybylle
Gussmann translated parts of zur Strassen’s 1965 revision of
the genus Sceliages into English. Barend Erasmus assisted
with the species distribution maps. Michelle Hamer
identified the species of millipedes utilised by Sceliages
hippias and Sc. adamastor. Claudia Medina helped with
fieldwork of Sc. hippias. Thanks to the museum curators
responsible for issuing loan material: M. Kerley (BMNH); T.
Crouch (DMSA); D. Mann (HECO); D. Drugmand (ISNB);
M. Cochrane (SAMC); R. Stals (SANC); and J. du G.
Harrison (TMSA). Thanks also to Richard Newberry
(North-west Parks and Tourism Board) and Magda Goosen
(Rustenberg Nature Reserve) for permitting research of Sc.
hippias in the Rustenberg Nature Reserve, Andre Olwage for
the habitus illustration of Sc. granulatus and Pam Prowse for
inking the illustrations. This research was funded by
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Manuscript received 26 July 2001; revised and accepted 11 August
... Most dung beetles feed on herbivore dung as wet dung, or sometimes on dry dung pellets (Holter, 2016). Dung beetles feed on variety of food substrate, some feed on dry dung pellets (Holter, 2016), detritus (Holter et al., 2009), millipedes (Forgie et al., 2002) and carion (Forgie et al., 2002); however, most dung beetles feed on wet dung, and that is what we are going to focus on in this study. ...
... Most dung beetles feed on herbivore dung as wet dung, or sometimes on dry dung pellets (Holter, 2016). Dung beetles feed on variety of food substrate, some feed on dry dung pellets (Holter, 2016), detritus (Holter et al., 2009), millipedes (Forgie et al., 2002) and carion (Forgie et al., 2002); however, most dung beetles feed on wet dung, and that is what we are going to focus on in this study. ...
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The life cycle of almost all dung beetles revolve around mammalian dung, the feed on dung, look for mating partners on dung and lay eggs in the dung. We know they feed on dung, but we still do not understand how exactly they filter‐feed on the dung and which particles size range they are ingesting. The aim on this study was to investigate the filter feeding by particle selection by adult dung beetles using Scarabaeus goryi and how that improves the nutrient quality of the ingested particles. We compared the particle sizes and nutrient content of the dung with the ingested material in the foregut, hindgut and the faeces of the dung beetle. Adult dung beetles do select smaller dung particles when feeding, we found the maximum particle size for the ingested particle to be around 1400 μm. The average particle size increased through the gut length. Dung beetles also selected particles with higher nitrogen content when feeding, the nitrogen content increased from about 1.5% in the dung to just over 5% in the foregut which then decreased to the level of the unprocessed dung in the dung beetle faeces. Carbon content did not increase from the unprocessed dung to the foregut but decreased through the gut length. Feeding by particle size selection by dung beetles helps in selecting particles with higher nitrogen content to compensate for the low levels found in dung. For the first time, we measured the particle sizes and nutrient content of the dung in the gut of dung beetles and the dung beetle faeces. We confirmed that dung beetles do select smaller particles in the dung when they are filter feeding. Through this filter feeding, the smaller particles selected have about 5% more nitrogen than the dung they are feeding on.
... Here too, the kill occurs by decapitation using the clypeus as a 'cutting weapon' [11][12][13][14][15]. Other potential cases of predation on diplopods may occur in other Neotropical dung beetles, such as Canthon morsei Howden, 1966 [16,17] [18][19][20]. However, although a clear attraction has been observed for the chemical secretions emitted by diplopods, it has not been observed that the dung beetles kill healthy individuals but rather that they are observed on injured, dying or dead diplopods. ...
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We described, for the first time, a case of predation of a non-arthropod species by a dung beetle species. Canthon chalybaeus Blanchard, 1843 kills healthy individuals of the terrestrial snail Bulimulus apodemetes (D'Orbigny, 1835) showing an evident pattern of physical aggressiveness in the attacks using the dentate clypeus and the anterior tibiae. The description of this predatory behaviour was complemented with the analysis of the chemical secretions of the pygidial glands of C. chalybaeus, highlighting those main chemical compounds that, due to their potential toxicity, could contribute to death of the snail. We observed a high frequency of predatory interactions reinforcing the idea that predation in dung beetles is not accidental and although it is opportunistic it involves a series of behavioural sophistications that suggest an evolutionary pattern within Deltochilini that should not only be better studied from a behavioural point of view but also phylogenetically.
... Although studies with dung beetles feeding on invertebrate carcasses are scarce, this behaviour has been reported in generalist feeders (Halffter & Matthews, 1966;Ebert et al., 2019). Previous studies showed that some dung beetle species predate on arthropods, such as leaf-cutter ants and freshly dead millipedes (Hertel & Colli, 1998;Forgie et al., 2002;Schmitt et al., 2004;Larsen et al., 2009;Karimbumkara and Dharma Rajan, 2016); and use their bodies for brood provision and as food. While some studies have used arthropod carcasses as bait (Larsen et al., 2006Ebert et al., 2019), this is the first study to include arthropod carcasses in a laboratory trophic attraction study. ...
• Trophic ecology of dung beetles has been widely studied because of the important ecological role of these taxa. However, previous studies have focused on a limited number of potential food items (mainly vertebrate dung and carrion) and have used only one approach (either field or laboratory). Moreover, recent studies showed high abundance of dung beetles in defaunated areas with a low abundance of these resources. • In this study, we combined a field and laboratory approach to explore dung beetle trophic attraction to different potential native resources in the Atlantic forest; and we evaluated whether results can explain the high abundance of dung beetles in defaunated areas. Through laboratory olfactometry experiments, we first exposed individuals to vertebrate carrion, omnivorous dung, and decomposing fungi. Then, we exposed species that exhibited a preference for dung to monkey, tapir, and feline dung; and those that preferred carrion and decomposing fungi to chicken, cow meat, and arthropod carcasses. We compared trophic attractions in the field and laboratory conditions with generalised additive models. • We found that coprophagous species preferred monkey dung, and all necrophagous and sapro‐necrophagous species preferred arthropod carcasses. These results suggest that the importance of arthropods carcasses as an important resource for dung beetles has been largely underestimated. • The results of this study might provide an explanation for the high abundance of necrophagous and sapro‐necrophagous dung beetles in defaunated areas. In addition, the use of omnivorous dung and arthropod carcasses could be an effective sampling method for dung beetle assemblages.
... Some dung beetles are known to feed exclusively on invertebrate carrion or predate upon invertebrates. These include some Canthon species in Brazil that utilise the alate queens of leafcutter ants (Atta) for brood provision (Hertel & Colli 1998), the millipede-feeding specialist, Sceliages, in South Africa (Forgie et al. 2002) and the millipede predator, Deltochilum valgum, in Peru (Larsen et al. 2009). In our study, more than 15% of specimens collected at invertebrate carrion belonged to several of the abundant AuEG species (Lepanus NQ9, Aptenocanthon monteithi and L. dichrous) and showed a greater association with invertebrate baits than other species. ...
Dung beetles (Scarabaeinae) are mainly coprophagous. Globally, many species co‐exist with large mammalian fauna in grasslands and savannahs. However, tropical and subtropical rainforests, where large herbivorous mammals are scarce, support numerous dung beetle species. Many rainforest dung beetles have been shown to be generalist saprophages or specialists on non‐dung food resources. In Australian rainforests, observations of native dung beetles have indicated that some species are attracted to other resources such as fruit or fungi, although the extent to which this occurs is not known. To learn more about the diet breadth of Australian native rainforest dung beetles, we assessed their attraction to a range of baits, including two types of dung, four types of carrion from both vertebrates and invertebrates, three types of rotting fruit and rotting mushrooms. We primarily surveyed rainforest sites but included two dry open‐forest sites for comparisons. Of the two groups of Australian native dung beetles (Onthophagini and Australian endemic genera), the latter dominated the rainforest dung beetle fauna and were attracted to a greater variety of baits compared with Onthophagini. The Onthophagini were dominant in open forest and were more likely to be attracted to a particular bait type, primarily dung. Our findings suggest that many of the species belonging to the ‘Australian endemic genera’ are generalist feeders and their ability to utilise a range of food resources contributes to their abundance and diversity in Australian rainforests.
... The basal sclerite is known in the literature as a lateral structure (Barbero et al. 1998), ring sclerite , Reid & Storey 2000, virgular sclerite (Forgie 2002) and circular sclerite , see Table 2). ...
... However, their feeding habits are not limited to this substrate. Many species are known to specialize on carrion, the fruiting bodies of basidiomycote fungi, freshly dead millipedes [5,6], rotting fruit and leaves, or the debris of attine leaf-cutter ants. There are even species that are predators of millipedes [7], reproductive ants, and termites. ...
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The astonishing spectrum of scarabaeine lifestyles makes them an attractive group for studies in entomology and evolutionary biology. As a result of adaptions to specific food substrates and textures, the mouthparts of dung beetles, particularly the mandible, have undergone considerable evolutionary changes and differ distinctly from the presumptive ancestral conditions of the Coleoptera and Polyphaga. The possible functions of dung beetle mouthparts and the evolution of dung feeding have been controversial for decades. In this study, 187 scarabs representing all tribes of the Scarabaeinae and the major lineages within the Scarabaeoidea, along with three major feeding types within the Scarabaeoidea (omnivory, phytophagy and coprophagy), were studied. Based on geometric morphometric and three-dimensional (3D) reconstruction approaches, morphological differences in mandibles among the three feeding types were identified. The ancestral forms of the mandible within the Scarabaeinae were reconstructed and compared with those of modern species. The most recent common ancestor of the Scarabaeinae fed on soft materials, and the ancestor of the Scarabaeinae and the Aphodiinae was in an evolutionary transition between processing more solid and softer substrates. Coprophagy originated from omnivorous ancestors that were very likely saprophagous. Furthermore, phytophagy may also have originated from omnivory. In addition, our study addresses the integration and modularity of geometric morphometric data in a phylogenetic context.
... The basal sclerite is known in the literature as a lateral structure (Barbero et al. 1998), ring sclerite (Reid 2000, Reid & Storey 2000, virgular sclerite (Forgie 2002) and circular sclerite (Medina et al. 2003, see Table 2). ...
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The external and internal male genitalia of 327 species of 11 tribes of the subfamily Scarabaeinae, including species of Deltochilini, Scarabaeini, Gymnopleurini, Ateuchini, and Coprini, among others, were examined. Descriptions of the variations in the genital segment, the aedeagus, the internal sac, and its sclerites and raspules are presented. An exhaustive comparison of structures, names, and terminology used in literature for Scarabaeinae male genitalia are discussed. The internal sac of the aedeagus is divided in areas for an easer comparison of its internal structures; basal, submedial, medial, and apical areas are described in detail and compared. The variation of apical and medial sclerites, as well as the raspules of the submedial area, are described and compared in detail among all the taxa studied.
... In general, percentage similarity between reserves decreases from lower to higher altitude ( Fig. 2) and statistical distance increases (Fig. 3). However, qualitative assessments of axes for regional (altitude, rainfall) and local habitat 1: Davis (1989), 2: Davis (1995), 3: Davis (1996), 4: Davis (1997), 5: Doube (1991b), 6: Forgie et al. (2002), 7: A.L.V. Davis, pers. obs. ...
A quantitative dung beetle survey (Coleoptera: Scarabaeidae: Scarabaeinae) was conducted in six Gauteng nature reserves (Tswaing, Leeuwfontein, Roodeplaat, Ezemvelo, Abe Bailey, Suikerbosrand) representative of the provincial range in environmental conditions. The study provided a provincial species inventory that has been tested for completeness by comparison with museum reference material. It also permitted an analysis of major influences on regional (altitude, annual rainfall) and local (soil and vegetation type) patterns of species abundance. The survey recorded a total of 152 species. Although a further 29 species were represented in reference collections, their absence from the present work was probably due to habitat, food, or temporal specializations. Multivariate analyses (clustering, MDS) of species abundance data formed six clusters, each comprising exclusively the sites in single reserves. This indicates that regional between-reserve faunal differences are greater than local within reserve differences, thus demonstrating the value of each reserve. In a hierarchical analysis of oblique factors, five out of seven statistically defined clusters comprised exclusively the sites in single reserves with Suikerbosrand split in two. These clusters were variously correlated with nine extended factors. Along seven factors, correlations were unique or essentially unique to single reserves and were fairly high, particularly at intermediate altitude. Along the remaining two shared factors, there were opposing trends in that correlations either decreased with higher altitude or decreased with lower altitude. Thus, each cluster shows three coefficients of determination, one representing the proportion of variance due to shared highland faunal influence, one representing that due to shared lowland faunal influence and the third representing that due to unique local faunal composition. The analyses identify the transition from lowland to highland-dominated faunas, the relative faunal distinctiveness of each reserve, and omissions from the provincial reserve system.
... Other recent reductions of long-established Afrotropical genera (Kheper, Sceliages) to subgeneric status within Scarabaeus, (Forgie et al. 2005) may, like Pachysoma, also deserve review and revalidation as full genera on morphological, molecular and behavioral grounds. This applies particularly to Sceliages, which shows modification of the clypeal margin associated with the specialized use of millipede gut contents for breeding (Forgie et al. 2002). ...
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Distribution of the subgenus Scarabaeus (Scarabaeolus) Balthasar 1965 (tribe Scarabaeini) is centred on southern and south central Africa with only three out of 27 species recorded from West and northeast Africa. After taxonomic corrections and descriptions of seven new southern African species this somewhat controversial subgenus now comprises 33 valid species of which one is flightless. In this paper, Scarabaeus (Scarabaeolus) vansoni Ferreira 1958 syn. nov. is synonymized with Scarabaeus (Scarabaeolus) lucidulus (Boheman 1860) and Scarabaeus (Scarabaeolus) xavieri Ferreira 1968 syn. nov. is synonymized with Scarabaeus (Scarabaeolus) andreaei zur Strassen 1963. Scarabaeus (Scarabaeolus) reichei Waterhouse 1890 stat. rev. is removed from synonymy with Scarabaeus (Scarabaeolus) canaliculatus Fairmaire, 1888 and reinstated as a valid species. Distribution maps for S. (S.) reichei, S. (S.) canaliculatus and a third close relative, Scarabaeus (Scarabaeolus) fritschi Harold 1868 are provided. The seven new species comprise: Scarabaeus (Scarabaeolus) soutpansbergensis Deschodt and Davis spec. nov., Scarabaeus (Scarabaeolus) megaparvulus Davis and Deschodt spec. nov., Scarabaeus (Scarabaeolus) niemandi Deschodt and Davis spec. nov., Scarabaeus (Scarabaeolus) carniphilus Davis and Deschodt spec. nov., Scarabaeus (Scarabaeolus) ermienae Deschodt and Davis spec. nov., Scarabaeus (Scarabaeolus) planipennis Davis and Deschodt spec. nov. and Scarabaeus (Scarabaeolus) nitidus Davis and Deschodt spec. nov. A key is provided for all the known winged species together with notes on some of the previously described species.
This chapter deals with the traces produced by extant dung beetles. They had been already described by the ancient Egyptians until Fabre introduced detailed observations in the scientific literature by the end of the nineteenth century. Some of Fabre’s observations are included herein to provide some examples of dung beetle behavior and traces. Those observations are still one of the best reports on dung beetle behavior and neoichnology. Behavior and traces of Aphodiinae, and Geotrupinae are briefly reviewed to focus in the third subfamily of Scarabaeidae, Scarabaeinae, which are the producers of the most common trace fossils in paleosols. The modern classification of nesting patterns and behavioral characters is summarized in a table and analyzed. Types of dung transport, nests, brood balls, brood masses, cakes, linings, egg chambers and other characters of brood balls, and pupation chambers are reviewed. A blueprint-style plate concentrates all the significant traces produced by dung beetles. Color plates on producers and their traces are provided. Some ecological preferences of dung beetles are highlighted as useful tools for paleoenvironmental interpretations.
A comparative phylogenetic approach was used to test the following adaptive hypotheses pertaining to the physiological abilities of the Namib desert tenebrionid beetle genus Onymacris to withstand the hot, dry desert environment: (1) Desert-interior species evolved longer legs (relative to body size) than beetles in the cooler coastal region to facilitate stilting, i.e., elevating their bodies out of the hot boundary layer of air close to the substrate. (2) Wax blooms on the exoskeleton, which reduce evaporative water loss, are more likely to evolve in desert-interior species than in coastal species. (3) The high costs of activity in the extreme climates select for perfect coadaptation of preferred body temperatures (i.e., optimal temperatures for activity) and those they achieve in the field. All three of these adaptive hypotheses were supported by the results of squared-change parsimony and independent-contrasts analyses. Additionally, a parsimony approach suggested that a novel means of obtaining water from periodic fogs, known as fog basking, has evolved independently on two occasions.