Chapter 19
Beetles (Insecta: Coleoptera)
Frank-T. Krell and Wolfgang Schawaller
Abstract Nine fossil beetles and seven fossil brood balls
made by dung beetles are described from Laetoli (Pliocene).
Seven beetles are Tenebrionidae, tribes Tentyriini and Molurini,
one is a June beetle of the tribe Schizonychini (Scarabaeidae:
Melolonthinae) and one a rhinoceros beetle (Scarabaeidae:
Dynastinae) described as Calcitoryctes magnificus sp.n. Seven
fossil dung beetle brood balls are described as Coprinisphaera
laetoliensis ichnosp. n. and C. ndolanyanus ichnosp. n., the
first formally described scarab ichnofossils from Africa. Two
specimens of C. laetoliensis show the largest known traces of
kleptoparasites described as Lazaichnus amplus ichnosp. n.
The fossil beetles and brood balls of Laetoli weakly indicate a
grassland, rather than a dense woodland habitat.
Keywords Tenebrionidae • Scarabaeidae • Ichnofossils
• Coprinisphaera
Introduction
With more than 350,000 described species, the beetles
(Coleoptera) form the largest order of organisms with the
oldest stem-group representatives recorded from the Early
Permian (Ponomarenko 2002; Grimaldi and Engel 2005). As
hard-shelled insects they are well-represented in the fossil
record, often, however, as single elytra or flattened sand-
wiches of several layers of dark cuticle, lacking or hiding
crucial characters (Krell 2000). Apart from amber inclusions,
three-dimensional, undistorted beetle fossils are rare and
known from only a few lagerstätten (Krell 2006), mainly
Oligocene-Miocene Riversleigh in Queensland (Duncan
F.-T. Krell (*)
Denver Museum of Nature & Science,
2001 Colorado Blvd, Denver, CO 80205, USA
e-mail: frank.krell@dmns.org
W. Schawaller
Staatliches Museum für Naturkunde, Rosenstein 1,
D-70191, Stuttgart, Germany
e-mail: wolfgang.schawaller@smns-bw.de
T. Harrison (ed.), Paleontology and Geology of Laetoli: Human Evolution in Context. Volume 2: Fossil Hominins
and the Associated Fauna, Vertebrate Paleobiology and Paleoanthropology, DOI 10.1007/978-90-481-9962-4_19,
© Springer Science+Business Media B.V. 2011
et al. 1998), Miocene Barstow Formation in California (Park
and Downing 2001), Eocene London Clay of Bognor Regis
(Britton 1960), and particularly the Miocene of Rusinga and
Mfangano Island in Lake Victoria, Kenya (Leakey 1952;
Paulian 1976). These fossils give unique insights in the actual
shape of Tertiary insects. The beetle fossils from Laetoli
belong to only two families, Tenebrionidae (darkling beetles)
and Scarabaeidae (scarab beetles). They are mineralized,
filled replicas of the exoskeleton without preservation of the
original cuticle. Despite the fairly detailed preservation, they
lack crucial specific and generic characters, particularly legs,
preventing us from formally describing most of them as new
species, but classification at tribal level is possible.
Additionally, mineralized dung beetle brood balls are pres-
ent, the spherical dung portions covered by a soil layer that
coprophagous scarab beetles form in their underground nests
as food provision for their larvae. All specimens are depos-
ited in the National Museum of Tanzania, Dar es Salaam.
Tenebrionidae (W. Schawaller)
The beetle family Tenebrionidae (Darkling Beetles) is one of
the largest families of Coleoptera worldwide with about
20,000 recent species. The family displays high morphologi-
cal and ecological plasticity with species inhabiting the sea-
shore, sandy and rocky deserts, and woodland habitats up to
the alpine zone above the timberline. In spite of this recent
diversity, fossil records of tenebrionids are poor. The serrate
antenna of one fossil from the Mesozoic Crato Formation in
Brazil might point to the family Tenebrionidae, although
definite tenebrionid family characters cannot be seen. This
would be the only fossil record of a tenebrionid beetle from
the Mesozoic (Wolf-Schwenninger and Schawaller 2007).
Younger fossil tenebrionids are known from Tertiary depos-
its, for example from the Florissant Fossil Beds in Colorado
(Wickham 1914) and from the Messel deposits in Germany
(Hörnschemeyer 1994). Tenebrionids from Baltic Amber are
listed by Spahr (1981), and tenebrionids from Dominican
535
536
Amber have been described by Kaszab and Schawaller
(1984) and Doyen and Poinar (1994).
From Laetoli, seven remnants of beetles have been found
that probably belong to the family Tenebrionidae. They can
be assigned to four species and are described and figured, but
not named. Although the fossils are well preserved, they can
be classified only by the general external morphology, but
not by distinct generic and/or specific characters. Thus, they
are not formally described as new taxa.
Tentyriini Species a (? Genus Tentyria)
EP 734/05 (Fig. 19.1)
Laetoli Locality 3 [localities described by Harrison and
Kweka 2011]: Upper Laetolil Beds, 60–70 cm above Tuff 7.
Fig. 19.1 Tentyriini species A
(?genus Tentyria, Tenebrionidae),
Laetoli. EP 734/05; (a) right side;
(b) ventral; (c) left; (d) dorsal.
Scale in mm
Fig. 19.2 Tentyriini species A
(?genus Tentyria, Tenebrionidae),
Laetoli. EP 1598/04; (a) right
side; (b) ventral; (c) frontal; (d)
left; (e) dorsal. Scale in mm
F.-T. Krell and W. Schawaller
Description. Joint elytra, pronotum, dorsal part of head, and
venter of anterior thorax preserved. Combined elytra of oval
shape, widest before the middle, length of elytra 13.5 mm, max-
imal width of combined elytra 9.0 mm. Elytral surface without
recognizable punctural rows or striae and without recognizable
surface structure. Pronotum as wide as long, widest just before
the middle, surface convex, with dense but not confluent puncta-
tion. Dorsal side of head with similar punctation as on prono-
tum, clypeus without tooth or other modification, eyes somewhat
prominent, kidney-like and only slightly excavated by the genae.
Without prominent prosternal process. Anterior and middle
coxal cavities widely separated, posterior coxal cavities not pre-
served. Last abdominal ventrites not distinguishable.
EP 1598/04 (Fig. 19.2)
Laetoli Locality 12: Upper Laetolil Beds, between Tuffs 5
and 8.
19 Beetles 537

Fig. 19.3 Tentyriini species A (?genus Tentyria, Tenebrionidae), Laetoli. EP 1670/00; (a) right side; (b) ventral; (c) left; (d) dorsal. Scale in mm

Description. Joint elytra without tip, pronotum, head,

venter of thorax and abdominal ventrites preserved.
Combined elytra of oval shape, widest before the middle,
length of elytra 13.0 mm, maximal width of combined elytra
8.0 mm. Elytral surface without recognizable punctural rows
or striae or other recognizable surface structure. Pronotum as
wide as long, widest just before the middle, surface convex,
surface with dense but not confluent punctation. Dorsal side
of head with similar punctation as on pronotum. Clypeus
without tooth or other modification, eyes somewhat promi-
nent and kidney-shaped, only slightly excavated by the
Fig. 19.4 Tentyriini species A (?genus Tentyria, Tenebrionidae), Laetoli.
genae. Without prominent prosternal process. Anterior, mid- EP 669/04; (a) from right; (b) ventral; (c) left; (d) dorsal. Scale in mm
dle and posterior coxal cavities widely separated. Five visible
abdominal ventrites, ventrites 3 and 4 not distinctly shorter
than ventrite 2, last visible ventrite 5 shorter than ventrites 3 surface structure. Anterior coxal cavities not preserved, mid-
and 4 combined. dle and posterior coxal cavities widely separated. Five visible
abdominal ventrites, ventrites 3 and 4 of similar length and
EP 1670/00 (Fig. 19.3)
slightly shorter than ventrite 2, last visible ventrite 5 longer
Laetoli Locality 3: Upper Laetolil Beds, 60–70 cm above
than ventrites 3 and 4 combined.
Tuff 7.
Taxonomy. These four fossils represent the same species
Description. Joint elytra without tip, venter of posterior
because of similar characters. The large body size and the
thorax and basal abdominal ventrites 1–3 preserved. Combined
general shape of pronotum and elytra, the elytra without
elytra of oval shape, widest before the middle, length of elytra
punctural rows or striae, the structure of the head with slightly
15.0 mm, maximal width of combined elytra 9.8 mm. Elytral
prominent eyes and the shape of the eyes, the wide distance
surface without recognizable punctural rows or striae and
of all coxal cavities, and the shape of the abdominal ventrites
without recognizable surface structure. Anterior coxal cavi-
coincide with recent species of the genus Tentyria Latreille,
ties not preserved, middle and posterior coxal cavities widely
1802 (tribe Tentyriini Eschscholtz, 1831, subfamily
separated. Only three basal abdominal ventrites preserved.
Pimeliinae Latreille, 1802). Quite similar is the closely
EP 669/04 (Fig. 19.4) related genus Rhytinota Eschscholtz, 1831. However, the
Laetoli Locality 3: Upper Laetolil Beds, 60–70 cm above recent congeners possess a narrower pronotum, narrower and
Tuff 7. longer elytra and non-prominent eyes.
Description. Joint elytra, venter of posterior thorax and all Zoogeography. The numerous extant species of the genus
abdominal ventrites preserved. Combined elytra of oval shape, Tentyria are distributed mainly in the Mediterranean region,
widest before the middle, length of elytra 14.0 mm, maximal Arabia, and Central Asia. Species of the genus Rhytinota
width of combined elytra 8.5 mm. Elytral surface without occur today in eastern Africa and the Indian subcontinent
recognizable punctural rows or striae or other recognizable (Gebien 1937).
538
Fig. 19.5 ?Tentyriini species B (Tenebrionidae), Laetoli, EP 351/03; (a) from left; (b) ventral; (c) right; (d) dorsal. Scale in mm
Fig. 19.6 ?Tentyriini species C (Tenebrionidae), Laetoli, EP 2777/00; (a) from right; (b) ventral; (c) left; (d) dorsal. Scale in mm
? Tentyriini Species B (? Genus)
EP 351/03 (Fig. 19.5)
Laetoli Locality 3: Upper Laetolil Beds, 60–70 cm above
Tuffs 7.
Description. Joint elytra, venter of complete thorax and
all abdominal ventrites preserved. Combined elytra of oval
shape, widest at anterior third, length of elytra 6.5 mm, max-
imal width of combined elytra 4.0 mm. Elytral surface with
an uncertain number of rows of fine punctures or striae, ely-
tral intervals slightly convex without recognizable surface
structure. Without prominent prosternal process. Anterior
coxal cavities touching each other, middle coxal cavities
slightly separated, posterior coxal cavities widely separated.
5 visible abdominal ventrites, ventrites 3 and 4 of similar
length and distinctly shorter than ventrite 2, last visible ven-
trite 5 longer than ventrites 3 and 4 combined.
F.-T. Krell and W. Schawaller
Taxonomy. From the general shape of the joint elytra, this
fossil might also belong to the tribe Tentyriini. However,
because of the nearly confluent anterior coxal cavities and
the structure of the elytra with rows of fine punctures or striae
and with slightly convex elytral intervals, I feel unable to
assign this and the following fossil specimen to any tenebrionid
genus, but they differ on the species level because of distinctly
different shape of the elytra.
? Tentyriini Species C (? Genus)
EP 2777/00 (Fig. 19.6)
Laetoli Locality 3: Upper Laetoli Beds, 60–70 cm above
Tuff 7.
Description. Joint elytra without tip, pronotum, venter of
complete thorax and all abdominal ventrites preserved.
19 Beetles
Combined elytra of longitudinal shape, widest at anterior
third, length of elytra 11.5 mm, maximal width of combined
elytra 3.5 mm. Elytral surface with at least 6 rows of distinct
punctures without striae, elytral intervals nearly flat without
recognizable surface structure. Pronotum as wide as long,
widest shortly before the middle, surface convex, with dense
but not confluent punctation; without prominent prosternal
process. Anterior coxal cavities touching each other, middle
coxal cavities slightly separated, posterior coxal cavities
widely separated. Five visible abdominal ventrites, basal ven-
trite 1 long, nearly as long as ventrites 2, 3 and 4 combined.
Taxonomy. The general shape of the joint elytra might
point to the tribe Tentyriini. However, as in the previous fos-
sil, the nearly confluent anterior coxal cavities and the struc-
ture of the elytra with rows of distinct punctures prevent a
definitive assignment to any genus.
Molurini Species a (? Genus Arturium)
EP 668/04 (Fig. 19.7)
Laetoli Locality 3: Upper Laetolil Beds, 60–70 cm above
Tuff 7.
Description. Joint elytra, venter of posterior thorax pre-
served. Combined elytra oval shaped, widest at anterior third,
length of elytra 13.0 mm, maximal width of combined elytra
9.0 mm. Elytral disc with two high undulated keels and with
a somewhat lower humeral keel, medial part of elytral
between the humeral keels flat, lateral parts of elytra besides
the humeral keels vertical and not to be seen in dorsal view,
surface between the keels slightly uneven und densely punc-
tured. Middle and posterior coxal cavities widely separated.
Abdominal ventrites not preserved.
Taxonomy. From the larger body size and the dorsal struc-
ture of the elytra the fossil might be assigned to the genus
Arturium Koch, 1951 (tribe Molurini Latreille, 1829, sub-
family Pimeliinae Latreille, 1802), although further characters
cannot be compared.
Fig. 19.7 Molurini species A (?genus Arturium, Tenebrionidae), Laetoli, EP 668/04; (a) from right; (b) ventral; (c) left; (d) dorsal. Scale in mm
539
Zoogeography. The genus Arturium, with few recent
species, is restricted to eastern Africa.
Biology
Recent tenebrionids belong roughly to two ecological groups.
One group includes characteristic dwellers of dry and even
arid habitats. The second, probably larger, group populates
decayed wood and fungi and could be considered as an indi-
cator of mature forests. A compact body with short legs are
characters of the tenebrionid forest dwellers, whereas slen-
der bodies with longer legs are connected with running
behavior in open habitats, including deserts. Unfortunately,
body appendages are not preserved in the Laetoli tenebrion-
ids, so no conclusions can be deduced about their former
habitat based on morphology alone.
The taxonomic assignment of some fossil tenebrionids to
the tribes Tentyriini and Molurini clearly points to an open
habitat during the Pliocene. Nearly all recent species of these
tribes are soil dwellers in steppes, savannahs and deserts, and
today eastern Africa is populated with abundant elements of
these tenebrionid tribes. The fossil tenebrionids from Laetoli
give no hint of woodland or forested habitats.
Scarabaeidae (F.-T. Krell)
With about 31,000 described extant species (Jameson and
Ratcliffe 2002) the Scarabaeoidea are one of the largest super-
families in the Coleoptera. They are distributed world-wide
and comprise such varied groups as dung beetles, stag beetles,
and chafers, ranging from just over 1 mm to 170 mm body
length. In the fossil record they are fairly well represented
with about 230 species described from the Upper Jurassic to
the Pleistocene (Krell 2007). However, apart from fossil
brood balls made by dung beetles, Pliocene scarab fossils are
540 F.-T. Krell and W. Schawaller

rare (Krell 2007) with some extant species recorded, but only Dynastinae: Oryctini/Pentodontini
two extinct species described: the dung beetle Copris kartli-
nus Kabakov, 1988, from the Kisatibi Formation in Georgia, Calcitoryctes Krell, gen.n.
and the dubious Melolonthites laterosinuatus Piton and
Derivatio nominis. From calcite (calcium carbonate) of which
Théobald 1935, from the Mio/Pliocene cinerites of Varennes,
the specimen consists, and Oryctes, the type genus of Oryctini
France, represented by only one elytron.
Mulsant, 1842, to which it might belong. Gender masculine.
Scarabaeoidea is a monophyletic group diagnosed by two
Type species. Calcitoryctes magnificus sp.n.
autapomorphies: antenna with a lamellate club and anterior bor-
Diagnosis. Outer side of mandibles entire or slightly den-
der of hind wings with sclerotized field proximal to a pinch, as
ticulate. Clypeus truncated with rounded angles. Head with
part of a spring folding mechanism (Krell 2006). Both charac-
tubercle. Pronotum without sculpture. Ventrites 1–4 much
ters are rarely preserved in fossils and are missing in the Laetoli
shorter than 5 and 6. Pygidium ca. three times as broad as
scarabs. Most fossil scarab beetles, including the two specimens
long, transversally bulged with strong apical impressions on
described below, were identified on the basis of other characters
both sides. Some extant Oryctes Illiger, 1798, and Cyphonistes
typical for Scarabaeoidea, such as enlarged prothorax adapted
Burmeister, 1847, species have a similar pygidium, but
for digging with short and powerful legs with tibiae toothed
Oryctes is much larger and has a deeply emarginated clypeus
along outer edge, large and narrowly separated to contiguous
and Cyphonistes has never one tubercle on the head (Endrődi
pro- and mesocoxae, and transverse and narrowly separated to
1985). Within Pentodontini Mulsant, 1842, it resembles the
contiguous metacoxae. The body fossils from Laetoli are well-
South/East African genus Pentodontoschema Péringuey,
preserved three-dimensionally, but are lacking finer surface
1901 (Péringuey 1901; Ferreira 1966) from which it differs
structures like punctation. In the melolonthine, even all sutures
by the margined base of the pronotum, the concave apical
are blurred. Moreover, the legs apart from some femora are
part of the pygidium and the probably less denticulated man-
missing, hardly leaving any genus- or species-diagnostic char-
dibles. It differs from Heteroligus Kolbe, 1900, by the miss-
acters. Both specimens are described, but only the exceptionally
ing pronotal tubercles and the concave apical part of the
preserved rhinoceros beetle is named.
pygidium and from Phyllognathus Eschscholtz, 1830, by the
Apart from the two body fossils, seven fossil dung beetle
more slender mandibles (Endrődi 1985).
brood balls were found at Laetoli. Fossil brood balls were
recorded from Laetoli previously (Sands 1987) and also from
Calcitoryctes magnificus Krell, sp.n.
other Pliocene sites in Africa, such as paleo-lake Chad
(Duringer et al. 2000) and Makapansgat Limeworks in South Derivatio nominis. Magnificus (adj.) (post-classical
Africa (Kitching 1980), as well as from the Pleistocene of Latin) = magnificent.
Rutana in Burundi (Basilewsky 1951). However, none of the Holotype. EP 2704/00 (Fig. 19.8), Laetoli Locality 2:
African specimens has been formally described and named Laetolil Beds, upper unit between Tuffs 5 and 7, deposited in
as ichnospecies. the National Museum of Tanzania, Dar es Salaam.

Fig. 19.8 Calcitoryctes magnificus sp.n. (Scarabaeidae, Dynastinae), holotype, Laetoli, EP 2709/00; (a) from frontal; (b) right; (c) dorsal; (d) left;
(e) ventral; (f) caudal. Scale in mm
19 Beetles
Description. Complete body with all femora apart from
the right profemur; length: 18.4 mm; maximum width of
pronotum: 9.5 mm; maximum width of elytra: 10.5 mm.
Body with left profemur and all other femora present.
Mandibles protruding beyond clypeus laterally and anteri-
orly; outer side of mandibles entire or slightly denticulate,
basally broad, slightly emarginated to a relatively sharp tip,
possibly with a denticle in the middle. Labium basally
broadly rectangular, apically with a deep emargination before
the double-convex anterior border, incised in the middle
(Fig. 19.9). Only scape and pedicel of the left antenna pres-
ent, both short as typical for Dynastinae. Clypeus truncated
with rounded angles, anterior margin bluntly triangular, lat-
eral margins slightly emarginated. Head with median conical
tubercle, steeper declined posteriorly than anteriorly; from
the tubercle, a slight bulge extends to the sides. Short ocular
canthus present. Pronotum regularly convex without any
sculpture; lateral margins regularly convex, broadest in the
middle; laterally and basally margined, lateral margin broader
and sharp. Anterior angles protruding and posterior angles
obtuse. Anterior margin without tubercle. Basal margin con-
vex. Scutellum rounded. Elytra broadest just behind the mid-
dle. Sutural interval basally broad, apically tapered and
slightly elevated. Shallow traces of probably three discal
stripes visible. Humeral callus and anteapical callus present.
Epipleura narrow (maximum 0.65 mm), with sharp border,
extending to pygidium. Ventrites 1–4 short, ventrite 5 longer
than 3 + 4 together, ventrite 6 as long as 3–5 together. All
coxae touching. Hind femora much thicker than middle and
slightly thicker than fore femora (ratio length/width f: 2.1;
m: 2.6; h: 1.9). Propydidium not fully covered by elytra, not
produced posteriorly; no stridulatory area visible due to pres-
ervation. Pygidium short (3.1 × as broad as long) as in most
Oryctini. Basal half of pygidium forms a strongly convex
transversal bulge (visible from ventral), apical half impressed
and convex, with strong impressions on both sides, margin of
the tip of the pygidium slightly protruding.
Fig. 19.9 Melolonthinae, Schizonychini species A (Scarabaeidae), Laetoli, EP 2156/03; (a) from right; (b) ventral; (c) frontal; (d) left; (e) dorsal.
Scale in mm
541
Classification. The only character separating the tribes
Pentodontini and Oryctini in the current typological classifi-
cation is the tibial apex. Tibiae are not preserved in this spec-
imen. However, since the tibial apex is variable within tribes
and the tribes themselves are only typologically defined
(Ratcliffe 2003:249), both groups might not survive a phylo-
genetic analysis as equally ranked taxa as one might become
a subgroup of the other. Therefore, it is not a serious short-
coming that Calcitoryctes cannot be unequivocally assigned
to one of these groups.
Melolonthinae: Schizonychini, Species a
EP 2156/03 (Fig. 19.9)
Laetoli Locality 7: Laetolil Beds, upper unit between
Tuffs 5 and 7 [body, one femur]
Description. Length: 12.4 mm; maximum width of pro-
notum: 5.0 mm; maximum width of elytra: 6.1 mm; height
of specimen: 5.1 mm. No microsculpture or punctation vis-
ible due to preservation. Labrum medianly incised,
Prementum bilobate, with a median furrow extending to
mentum. Clypeus seems to be separated from the frons by a
slight transversal bulge. Pronotum: Sides slightly diverging
in basal third, then straight strongly converging to head.
Elytra: Lateral border behind the humeral callus slightly
emarginate, then elytra broadened, widest just behind the
middle. All coxae adjacent.
Classification. The elevated frontoclypeal suture and the
strongly incised labrum are characters of Schizonychini
Burmeister, 1855, and, together with the shape of the elytra,
might even indicate that the specimen belongs to Schizonycha
(Pope 1960; Lacroix 1989), which is a speciose genus
containing abundant African species. However, two crucial
characters for Schizonychini, the enlarged ventrite 6 and the
metepimeron (Lacroix 1989), are not sufficiently preserved
542 F.-T. Krell and W. Schawaller

to allow a reliable tribal classification. Preservation of antennae, thickness of the outer layer is not only an indication of the
claws, mouthparts and setation would be necessary for thickness of a possible soil cover, but also of the amount of
generic classification. dung still in existence at the time when the dung consump-
tion by the larva ended. The longer the larva feeds, the thin-
ner the walls become (Lengerken 1954). Successful
development of the larva and hatching of the beetle results in
Ichnofossils thin walls of the brood ball. Thick walls might indicate that
the development was disturbed or unsuccessful.
A small and a large type of fossil dung beetle brood balls
were found at Laetoli, similar to the abundant brood balls Coprinisphaera laetoliensis Krell, ichnosp. n.
described as Coprinisphaera Sauer, 1955, from South Derivatio nominis. Adjective meaning ‘from Laetoli’, the
America. Genise et al. (2000) have already assigned the type locality.
“structure resembling a dung ball of Heliocopris” mentioned
by Sands (1987: 423) to this ichnogenus and counted Laetoli
in the localities with Coprinisphaera ichnofacies. All for-
mally described scarab ichnofossils are from the Americas
(Krell 2007). The fossil dung beetle brood balls were recently
revised by Genise (2004) and Laza (2006) and classified into
nine ichnospecies. To facilitate the integration of the Laetoli
ichnofossils into the current ichnological classification, they
will be named although ichnospecies-specific characters are
often not clearly visible due to the replacement of original
material and infilling with calcium carbonate.
These balls represent the prepared portions of resources
that dung beetles form from available feces as food provision
for their larvae. The brood balls of large tunnelling dung Fig. 19.10 Coprinisphaera laetoliensis isp.n., fossil dung beetle brood
ball, Laetoli, holotype, EP 224/04. Cut; (a) cut half; (b) counterpart
beetles are generally covered by a layer of soil to prevent
desiccation (Halffter and Edmonds 1982), whereas the sur-
face of brood balls of large rollers such as Scarabaeus L.,
1758 or Kheper Janssens, 1940 is either only smoothened
(Lengerken 1954) or covered by the mother beetle’s feces
(Sato and Imamori 1987).
Whereas the American fossil brood balls are mostly hol-
low spheres with the original soil cover preserved, the Laetoli
fossils are steinkerns of calcium carbonate rendering it
impossible to identify whether the outer layer was originally
formed by soil or dung. I interpret the faint line between the
outer layer and the infilling (Figs. 19.10–19.12) as the border Fig. 19.11 Coprinisphaera laetoliensis isp.n., fossil dung beetle brood
between the original dung portion and larval chamber. The ball, Laetoli, EP 1719/03. Cut; (a) half cut; (b) counterpart

Fig. 19.12 Lazaichnus amplus isp.n., traces of kleptoparasites in fossil dung beetle brood ball, Laetoli. Holotype of L. amplus in paratype of
Coprinisphaera laetoliensis, EP 1719/03; (a) half cut; (b) counterpart; (c) outer opening of L. amplus
19 Beetles
Fig. 19.13 Coprinisphaera laetoliensis isp.n., fossil dung beetle brood
ball, Laetoli, holotype, EP 224/04. Before cut; (a) from top; (b) lateral.
Scale in mm
Holotype. EP 224/04 (Figs. 19.10, 19.13), Laetoli Locality
15: Upper Ndolanya Beds; deposited in the National Museum
of Tanzania, Dar es Salaam.
The specimen is the fossilized empty, abandoned brood
ball, replaced and filled by calcium carbonate. Cut medianly
in half. Maximum height: 53.4 mm; equatorial diameter:
55.5 mm; diameter of upper opening: 22.5 mm. Regular
sphere with wide upper opening surrounded by collar in form
of a flat bulge (11–12 mm wide). Bottom with a small, oval
depression of about 13 mm diameter. upper lateral thickness
of the wall: ca. 4.5 mm, about 10 mm at the bottom.
Biology. Thin wall, broad opening and undamaged outer
surface indicate that the beetle hatched successfully; no signs
of kleptoparasitic or secondary intrusion.
Paratype. EP 1719/03, Specimen B (see Fig. 19.12) (of 3
specimens), Laetoli Locality 15: Upper Ndolanya Beds;
deposited in the National Museum of Tanzania, Dar es
Salaam. Cut medianly in half, some outer cracks fixed by
glue; spherical with upper opening; height: 47.5 mm, equa-
torial diameter: 54 mm; opening: 22 mm; thickness of the
wall: 10.5–15.8 mm, bottom with two deep holes of 9–10 mm
diameter (ventral hole: Fig. 19.12a, b; ventrolateral hole:
Fig. 19.12c).
Biology. The size of the holes in the outer wall indicate
infestation by larger kleptoparasites of genera such as
Onthophagus Latreille, Hyalonthophagus Palestrini or
Pedaria Laporte, being the holotype of Lazaichnus amplus
ichnosp.n. (described below). Kleptoparasitic species invade
and lay their eggs in a dung portion collected by another spe-
cies, generally reducing the reproductive success of the host
significantly (Rougon and Rougon 1980; Gonzáles-Megías
and Sánchez-Piñero 2003). The thick walls show that the
dung material had not been exhausted, probably due to
unsuccessful or abbreviated development of the host larva.
The upper opening might indicate that the host larva might
have hatched.
Additional material. EP 1077/01 (see Fig. 19.14), Laetoli
Locality 15: Upper Ndolanya Beds. Flat sphere with large
543
opening (55.5 mm × 38.8 mm) on one side and a deep small
hole (10 mm diameter) on the other. Height: 48 mm; equato-
rial diameter: 60 mm. Filled with a lighter (replaced dung)
and a darker substance in the area of the small hole and prob-
ably indicating the development space of a kleptoparasite
(paratype of Lazaichnus amplus ichnosp. n., described
below). Due to the large, irregular opening the ichnospecies-
specific collar is missing. Thus, this specimen is not desig-
nated as a paratype of C. laetoliensis.
Biology. The large upper opening (see Fig. 19.14) might
have been caused by a vertebrate predator. Modern exam-
ples of brood balls opened by bat-eared foxes (Otocyon
megalotis (Desmarest, 1822)) show similarly shaped holes
(see Fig. 19.15). These foxes excavate dung beetle brood
balls on a regular basis to prey on the larvae (Nel and Maas
2004).
EP 1719/03, Specimen A (Figs. 19.11, 19.16), Laetoli
Locality 15: Upper Ndolanya Beds, cut medianly in half;
flat pear-shaped; height: 48.5 mm, equatorial diameter:
58.6 mm, maximum width of neck: 27 mm; bottom flat with
central round area of ca. 14 mm diameter (contact zone of
brood ball to ground); lateral thickness of the wall 6–8.5 mm,
4–10 mm at the bottom; no signs of kleptoparasitic or sec-
ondary intrusion. Surface with six dorsoventral ribs which
were possibly caused by filled cracks of the surrounding
soil. Because of the different shape (rather pear-shaped than
spherical), which is similar of the shape of Vondrichnus
planoglobus Duringer et al., 2007 (Fig. 19.10a, d; termite
nest), it is not designated as a paratype. Because a thick
outer wall can be distinguished from the inner chamber, and
because it forms part of a series of three fossil brood balls
likely found together (the other two being typical
Coprinisphaera), it is identified as Coprinisphaera. Since
the three balls were not found in close proximity, I do not
classify them as Quirogaichnus Laza, 2006 which was intro-
duced for clusters of brood balls in a distinct chamber or
cavity.
Biology. The thin walls and undamaged outer surface
indicate a regular development of the larva. However, an
upper opening is not clearly defined. The irregular inter-
nal part of the neck could be the opening (diameter
13 mm).
EP 1719/03 Specimen C, Laetoli Locality 15: Upper
Ndolanya Beds: flat pear-shaped with flat bottom; height:
47.9 mm; equatorial diameter: 60.6 mm; surface with large,
deep cracks; no indication of kleptoparasitic or secondary
intrusion. Because of this suboptimal infilling or incipient
disintegration this specimen was not cut. Due to poor preser-
vation, I do not designate it as a paratype.
Description. Spherical, solid structures with sometimes
flattened underside resulting in a slight pear-like shape;
height: 47.5–53.4 mm; equatorial diameter: 54.0–55.5 mm
(all specimens: 53.4–60.6 mm). Dorsal opening on top with
544 F.-T. Krell and W. Schawaller

Fig. 19.14 Lazaichnus amplus isp.n., traces of kleptoparasites in fossil view in large opening showing the dark internal trace of L. amplus.
dung beetle brood ball, Laetoli. Paratype of L. amplus in C. laetoliensis, Scale (for Fig. 19.14a–c) in mm
EP 1077/01; (a) from top; (b) bottom; (c) lateral; (d) outer opening; (e)

Fig. 19.15 Recent dung beetle brood balls excavated and preyed on by bat-eared foxes (Otocyon megalotis) in the Laetoli area, showing an irregu-
lar large opening. Scale: 30 mm. Specimens deposited in National Museum of Tanzania, Dar es Salaam. Photo: T. Harrison

surrounding bulge-like collar. Outer wall of sphere between Diagnosis. Spherical to slightly pear-shaped structure
4.5 and more than 15 mm thick. Internal (filled) chamber with single chamber and upper opening with neck; differs
opens directly to the upper aperture. from Coprinisphaera kraglievichi (Roselli, 1939) from
19 Beetles 545

Fig. 19.16 Coprinisphaera laetoliensis isp.n., fossil dung beetle brood ball, Laetoli, EP 1719/03. Before cut; (a) from top; (b) bottom; (c) lateral.
Scale in mm

Uruguay and Argentina by the larger size (more than 53 mm

equatorial diameter versus 32–42.4 mm).
Discussion. The size of dung beetle brood balls varies
intraspecifically, but generally stays within 20–30%
(Lengerken 1954; Klemperer and Boulton 1976; Klemperer
1983; Sato and Imamori 1987). A brood ball of 60.6 mm
diameter is unlikely to belong to the same species as one of
32 mm diameter. Although, according to a recently published
convention (Bertling et al. 2006), the possible tracemaker
should not be considered as defining character for ichnospe-
cies, I consider it inappropriate to typologically combine
specimens under one ichnospecific name that are most likely
produced by different tracemaker species.
Trace maker. Sands’s (1987) interpretation of a
Coprinisphaera from Laetoli as resembling a brood ball of
Heliocopris Hope might be right. The large size and the
shape (spherical to slightly pear shaped) of the brood balls
are typical for extant Heliocopris (cf. Klemperer and Boulton
1976), but could also be produced by large Catharsius Hope
species or even by large rollers such as Kheper Janssens
(Sato and Imamori 1987) in which the mother beetle scrapes
the brood balls into a spherical shape during the development
of the larva.
Nomenclatural note. Coprinisphaera Sauer, 1955 is a junior
subjective synonym of Fontanai Roselli, 1939 (Laza 2006), but
is in prevailing usage (Krell 2007). Genise et al. (2006) and
Krell (2008) proposed conservation of Coprinisphaera with
the International Commission on Zoological Nomenclature.
The ICZN (International Commission on Zoological
Nomenclature 2008) has ruled that Coprinisphaera is to be
maintained as the valid name.
Coprinisphaera ndolanyanus Krell, ichnosp. n.
Fig. 19.17 Coprinisphaera ndolanyanus isp. n., fossil dung beetle brood
Derivatio nominis. Adjective meaning “belonging to ball, Laetoli. Holotype, EP 824/01. Before cut; (a) lateral; (b) from top.
Ndolanya”, the stratigraphic unit where it was found. Scale in mm
Holotype. EP 824/01 (Fig. 19.17), Laetoli Locality 18:
Upper Ndolanya Beds; deposited in the National Museum of cut medianly in half. Pear-shaped (sphere with neck on top),
Tanzania, Dar es Salaam; maximum height: 33.7 mm; tip of neck slightly impressed. Homogenous steinkern; no
equatorial diameter: 27.1–27.6 mm, width neck: 10.5–12 mm; internal structures visible.
546 F.-T. Krell and W. Schawaller

holotype of C. kheprii only. The equatorial diameter of the

smallest C. kheprii would fall into the specific size range of
C. ndolanyanus, but the minimum height of C. kheprii is
50 mm, much larger than in the latter.
Trace maker. Pear-shaped brood balls are typical for dung
rollers (Scarabaeini, Gymnopleurini) (Lengerken 1954) that
are abundant in recent Afrotropical coprocenoses.

Lazaichnus amplus Krell, ichnosp. n.

Derivatio nominis. Amplus (Latin) = vast, spacious;
adjective.
Holotype. EP 1719/03, Specimen B (Fig. 19.12) (para-
type of Coprinisphaera laetoliensis ichnosp.n.), Laetoli
Locality 15: Upper Ndolanya Beds; deposited in the National
Museum of Tanzania, Dar es Salaam. Coprinisphaera is cut
Fig. 19.18 Coprinisphaera ndolanyanus isp. n., fossil dung beetle medianly in half, hitting one hole of Lazaichnus amplus
brood ball, Laetoli. EP 3335/00. Scale in mm
(11.4 mm diameter). The other hole has an outer diameter of
8–11 mm.
Paratype. EP 1077/01 (Fig. 19.14) (Coprinisphaera lae-
Additional Material. EP 3335/00 (Fig. 19.18), Laetoli
toliensis), Laetoli Locality 15: Upper Ndolanya Beds; depos-
Locality 18: Upper Ndolanya Beds; maximum height:
ited in the National Museum of Tanzania, Dar es Salaam.
30.0 mm; equatorial diameter: 27.5–28.4 mm; cut medianly
The chamber behind the opening of Lazaichnus is visible in
in half. Sphere, one half with smooth surface, damaged on
the large upper opening in Coprinisphaera. Hole diameter:
the other side (possible neck missing), next to damage with
14–15.3 mm with lateral lighter infilling, diameter of proper
oval depression; no internal structures visible.
hole: 8.5 mm.
Description. Pear-shaped solid structure with upper impres-
Description. Holes in fossil dung beetle brood balls of
sion in long neck, probably indicating a former opening;
8–15.3 mm diameter, with a simple gallery or an expanding
height: 30.0–33.7 mm; equatorial diameter: 27.1–28.4 mm;
chamber extending into the inner core of the brood ball.
neck diameter: 10.5–12.0 mm. No internal structure visible.
Diagnosis. Holes leading to a gallery in fossil dung beetle
Diagnosis. Pear shaped solid structure; differs from
brood balls (Coprinisphaera), differ from the only other spe-
Coprinisphaera kheprii Laza, 2006 (height of holotype
cies of the ichnogenus, Lazaichnus fistulosus Mikulaš and
56.1 mm) from Argentina by the smaller size and the longer,
Genise, 2003, by the larger diameter of the opening and the
more slender upper protuberance. The upper protuberance of
simple structure of the gallery, which forms a simple tube or
C. tonnii Laza, 2006 is thicker and more bulge-like than in
a chamber in L. amplus, but often a more complex system of
C. ndolanyanus, containing an egg chamber. Coprinisphaera
tubes in L. fistulosus.
ndolanyanus differs from the globular C. laetoliensis from
Trace maker. The large diameter of the holes indicates
the same locality by the smaller size and the shape.
kleptoparasites much larger than Cleptocaccobius Cambefort
Discussion. A distinct outer wall of the brood balls is the
or kleptoparasitic Aphodius Illiger (the most abundant repre-
diagnostic character that distinguishes Coprinisphaera Sauer
sentatives of this guild in Africa). Current Afrotropical klep-
(dung beetle brood balls) from Pallichnus Retallack, 1984
toparasites of a matching size belong to Onthophagus,
(supposedly pupal chambers). Neither specimen of C. ndola-
Hyalonthophagus and Pedaria (Scarabaeinae).
nyanus shows this character due to the type of preservation
(original substance replaced by calcium carbonate). However,
the pear-shaped form of the holotype is an unequivocal indi-
cation of scarabaeine brood balls (Halffter and Edmonds
1982) which originally might not have had a different outer Biology
wall structure than C. laetoliensis. Coprinisphaera ndolan-
yanus is much smaller than the holotype of the pear-shaped Schizonychini, Dynastinae, dung beetles and their kleptopar-
C. kheprii, but other known specimens assigned to C. kheprii asites are distributed in all vegetation zones, from tropical
have an equatorial diameter from 28.5 to 60.5 mm. This size rain forest to arid regions. The same is true for the paleoen-
range is much larger than the 20–30% size range of brood viroments of the Coprinisphaera ichnofacies, but most are
balls of a single dung beetle species (see above). Thus, I associated with open grasslands (Genise et al. 2000).
compared the size of C. ndolanyanus with the size of the Schizonychini feed on leaves as adults and on roots as larvae.
19 Beetles
Dynastinae feed on various living and rotting plant material.
Thus, the scarab fossils and ichnofossils do not contradict
the current hypothesis of a savanna-forest ecotone paleoenvi-
ronment at Laetoli (Kingston and Harrison 2007; Su and
Harrison 2007).
Conclusions
Laetoli is one of the few lagerstätten where insect fossils are
preserved three-dimensionally. Whereas reconstructions are
the only way to visualize the body shape of common fossil
imprints, the Laetoli fossils are almost undistorted replicas of
the original bodies which made formal description and naming
of an exceptionally character-rich rhinoceros beetle fossil
possible. External morphology of beetles is often a reliable
indicator of habits, habitats or soil types (e.g., Medvedev 1965).
Adaptive traits are most clearly expressed in the legs, the body
parts interacting most extensively with the physical environ-
ment. However, with legs missing or only partly preserved the
beetle fossils of Laetoli do not indicate a particular habitat.
Extant dung beetles producing brood balls of the size found
at Laetoli are distributed in subtropical and tropical regions. In
the Afrotropics, the highest abundance of dung beetles is in
savannas (Cambefort and Walter 1991), where we also find the
highest density of kleptoparasitic dung beetles (Krell et al.
2003). The existence of large fossil dung beetle brood balls in
Laetoli, some with traces of kleptoparasites, indicates a higher
probability for grassland rather than an arboreal area.
Acknowledgements We are grateful to Terry Harrison, New York
University, for the patient loan of the specimens, to Harry Taylor, Photo
Unit of The Natural History Museum, London, for most of the photo-
graphs, to Brett Ratcliffe, University of Nebraska State Museum,
Lincoln, and Roger-Paul Dechambre, Muséum national d’Histoire
naturelle, Paris, for helpful comments about the fossil dynastine, and to
Jorge Genise, Museo Paleontológico Egidio Feruglio, Trelew, Argentina,
for careful criticism on a former version of the manuscript. Ken
Carpenter and Thomas Garner, DMNS Earth Sciences Department,
helped with cutting the fossil brood balls.
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