Formicdubius Philips & Scholtz from South Africa, a junior synonym of Haroldius Boucomont, and a survey of the trichomes in the African species (Coleoptera, Scarabaeidae, Onthophagini)
Author
Krell, Frank
Denver Museum of Nature & Science, Denver, United States of America
Author
Philips, T.
Western Kentucky University, Bowling Green ,, United States of America
text
ZooKeys
2010
2010-01-28
34
34
41
48
journal article
22582
10.3897/zookeys.34.342
0981ff15-3ede-42a5-9475-980c31a88b2d
1313–2970
576603
African
Haroldius
species
All African species of
Haroldius
appear to have trichomes on the mesepimeron, the pronotal base, and the elytral base. Trichomes appear as distinct and relatively tight clusters of setae while others are looser or even single rows of thick setae such as those on the elytral base. Without microsections we are unable to decide whether those might be proper trichomes associated with glands or mechanoreceptors. However, as further evidence of a glandular association, longitudinal grooves at the pronotal base are also present that may facilitate the spreading of allomone secretions onto the pronotal surface, increasing the attractiveness of these beetles to their host ants.
Formicdubius convexus
(Figs 1–6) (and the very similar or conspecific
Haroldius leleupi
Janssens, 1953
; Figs 11–13) can easily be distinguished from the remaining
Haroldius
species by the presence of a distinct notch between the pronotum and the elytral base accommodating the pronotal trichomes (Figs 2, 12). In
Haroldius ennearthrus
Janssens, 1949
(Figs 14–19), the basal margins of pronotum and elytra are straight with the pronotal margin slightly emarginate close to the edge (Figs 15, 18); a notch is missing, but a slight lateral depression towards the bases of the pronotum and elytra is visible.
H. modestus
Janssens, 1953
shares the general shape of the pronotal and elytral bases with
H. ennearthrus
but the pronotal margin is slightly more strongly emarginate laterally (Figs 8–9).
Figures Ι–6.
Haroldius convexus
:
Ι
Dorsal habitus
2
Trichomes within cleft between prothorax and mesothorax and showing pronotal sulci
3
SEM (scanning electron microscope) view of pronotal trichome
4
SEM view of pronotal trichome showing possibly ant-gnawed tips of the setae
5
SEM view of possible trichomes on the elytral base
6
SEM view of trichome on the mesepisternum and additional setal row proximally.
The trichomes on these species are located on the mesepisternum anterolaterally, the elytra anterolaterally (on the vertical surface facing the pronotum), and on the pronotal base posterolaterally. One should be aware that these setae may be damaged from the effects of gnawing by their host ants and the actual number of setae may vary in number more than we report due to our limited sample sizes. Regardless, the degree of development of these structures in each species is as follows:
Specimens of
H. ennearthrus
possess a large rounded cluster of setae on the mesepisternum (Fig. 19) and no additional setae proximally. They also appear to have two short and relatively thick setae on the elytral base. Three to five thick setae are present on the pronotum (Figs 16–18) as well as a row of similar aligned setae proximal to these, all of which may be trichomes (Fig. 16).
Haroldius modestus
(Figs 7–10) also has a large, rounded cluster of setae on the mesepisternum (Fig. 10) and lacks any thick setae proximally. There are two to three thick setae on the elytral base. On the pronotum a brush-like trichome is well developed, consisting of six to sometimes ten or more, long, thick setae (Figs 8–9).
The third described
Haroldius
species from Africa,
H. leleupi
(Figs 11–13), is strikingly similar to
Formicdubius convexus
(Figs 1–6). They both have a small cluster of stout setae on the mesepisternum (Figs 6, 13) and proximal to this are a few more thick and elongate setae arranged in a transverse row (Fig. 6). These same setae become more slender and slightly more widely separated the further from the mesepisternal trichome. Additionally, there are four to five short, thick setae at the base of the elytra near the lateral margin (Fig. 5). Both species also have a well developed trichome on the pronotal base consisting of 20 or more lobe shaped setae (Figs 3–4, 12–13).
We are unsure whether
H. leleupi
and
F. convexus
are distinct taxa.
H. leleupi
is generally larger; the five specimens studied measure
1.80–2.20 mm
in length (average
2.06 mm
), whereas the 14 sampled
types
of
F. convexus
measure
1.63–1.90 mm
(average
1.78 mm
).
Haroldius leleupi
also has slightly stronger (more clearly defined) punctures on the disk of the pronotum and the elytral intervals and slightly deeper elytral striae, particularly near the suture. Additional material will be required from localities between the
Congo
and
South Africa
to decide to what extent these minor differences indicate taxonomic differentiation.
Synonymy
Haroldius leleupi
and
Formicdubius convexus
are difficult to distinguish at the species level, but there is no doubt that
Formicdubius
is identical with
Haroldius
at the generic level.
Formicdubius
fully matches
Boucomont’s (1914)
original description of
Haroldius
. It also agrees with the extended diagnosis of
Haroldius
recently published by
Krikken and Huijbregts (2006)
with the exception that all African species currently subsumed under
Haroldius
possess trichomes.
Figures 7–Ι3.
Haroldius
species. Figures
7–Ι0.
H. modestus
7
Dorsal habitus
8
Trichomes within cleft between prothorax and mesothorax and showing pronotal sulci
9
Pronotal trichome
Ι0
SEM view of trichomes on the pronotum and mesepisternum. Figures
ΙΙ–Ι3.
Haroldius leleupi
:
ΙΙ
Dorsal habitus
Ι2
Trichomes within cleft between prothorax and mesothorax and showing pronotal sulci
Ι3
SEM view of trichomes on the pronotum and mesepisternum.
We propose the following synonymy:
Haroldius
Boucomont, 1914
(
type
species by subsequent designation by
Arrow, 1931
:
Haroldius rugatulus
Boucomont, 1914
)
=
Formicdubius
Philips & Scholtz, 2000
(
type
species by original designation:
Formicdubius convexus
Philips & Scholtz, 2000
), syn. n.
Consequently, the following new combination is established:
Haroldius convexus
(
Philips & Scholtz, 2000
) Krell & Philips, 2010
,
comb. n.
We refrain from determining the taxonomic status of the African species of the
Haroldius
/
Afroharoldius
group. All African
Haroldius
species have more or less distinct trichomes which could be a synapomorphy justifying a genus or subgenus
Afroharoldius
. However, we neither know if trichomes (vestigial or distinctive) are common in Asian
Haroldius
species, or whether the trichome-bearing species are the sister group of the trichome-less
Haroldius
,
or whether their exclusion from
Haroldius
would leave
Haroldius
or
Afroharoldius
paraphyletic. In
Haroldius brendelli
Krikken & Huijbregts
, antehumeral elytral trichomes are described (
Krikken and Huijbregts 2009
). Based on the illustrations in
Krikken and Huijbregts (2006)
, it appears as though pronotal grooves are present on at least some Asian species and, moreover, trichomes seem to be present in
H. tangkoko
Krikken & Huijbregts
and
H. cambeforti
Krikken & Huijbregts
(see their figs 5 and 6). A revision and phylogenetic analysis of the whole group, including species of
Phaedotrogus
Paulian
and probably
Ponerotrogus
Silvestri
and
Cyclotrogus
Wasmann
(both currently considered to be junior synonyms of
Haroldius
), will probably be necessary to establish a sound genus-level classification. Currently we see no reason to change the status of
Afroharoldius
as junior synonym of
Haroldius
.