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THE AFRICAN JEWEL BEETLES

This monograph is (we hope) the first of a series, intended


to collate our present knowledge of the family Buprestidae
in the Afrotropical Region. The reasons for embarking
on such a venture may be clothed into emotive issues like
ecological planning and documenting the fauna before
it disappears. The fact is, we simply like buprestids and we
like the taxonomic excuse to study them.
The general arranging of information in the book
is aimed at handiness for identification. We therefore list
synonymies, material examined and the bibliography at the end.
Throughout we follow systematic and not alphabetical order.
This is to encourage curators to do the same with the collections
under their care as too many synonyms in large genera went
undetected for decades simply because of alphabetical
ordering. Genera, subgenera and species are therefore
treated in the same sequence as they key out in the various
phylogenetic cladograms. In the case of subspecies, however,
the nominal subspecies is always treated first. Diagnostic
information (comparative notes, diagnoses, detail drawings
and distribution maps) of every species or subspecies is
provided together on one page. The introduction chapters
provide short backgrounds limited to considerations that are
relevant to our taxonomic decisions. They are certainly not
intended as exhaustive treatments of the various themes, as
speculations on historical geography and phylogeny alone
could fill a volume without, however, contributing much to
the taxonomy.
more then 600 color photographs of all species
and subspecies from different localities as well
as photos of behavior and habitats
over 400 pages, luxurious hardcover edition
large format 23x31, text in english
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1.3 ZOOGEOGRAPHY

The known distribution limits of the extant genera and subgenera of the subfamily are given in Figs 8-11. There
are no fossil records that can be assigned to the subfamily.
Scenarios for the possible origins and dispersals of the various groups were discussed in detail by one of
us (Holm 1979a). Since that paper further revisional works and descriptions have changed diversity statistics
somewhat. The fauna of the northern hemisphere (comprising about one half of the extant species) remains
unrevised and there is little to add to the (already excessive) speculations in Holm (1979a).
Aspects that are pertinent to the zoogeography of the various genera are their known distribution; their
phylogenetic relationships; their macro-ecological niches; and the relative species densities, apomorphies and
variabilities in subregions of the distribution limits. These data we briefly summarize below.

Genus AAATA Semenov Tian-Shanskij:


This genus is monotypic and limited to a small area in the Middle East (southeastern Iran and possibly Pakistan, Fig. 8).
It is obviously both a phylogenetic and geographic relic.
Together with Sternocera it forms a sister-group to the rest of the Julodinae.While the biology is unknown,
the relationship and the functional morphology suggest an ecological niche similar to that of Sternocera.
Variation within the single species is very limited. This, together with the large size and predominantly
plesiomorphic character states, suggests a ‘living fossil’ or ‘surviving ancestor’ on the Sternocera lineage. The most
primitive Sternocera occur in semi-arid areas of North East Africa and it is reasonable to suggest that this whole
lineage originated in the Near East.

Genus STERNOCERA Eschscholtz:


The genus has a disjunct distribution in sub-Saharan Africa and the Far East (Fig. 8).
Ecologically species are linked to tree-savannas, from semi-arid to moist climates but not extending into wet
tropical, extremely arid or winter rainfall biomes.
In Africa, variation is highest among species that spread furthest to the west and south, where – at the same
time - the overall species diversity diminishes. It is therefore reasonable to postulate a center of radiation in North East
Africa or even the Near East (see Aaata above), from where both sub-Saharan Africa and the Far East, were colonized.
As neither of these geographical species groups seems to be monophyletic, their isolation (whether by tectonic events
or glaciation) must have happened when species were already diversified. The isolation is now maintained by
the scarcity of suitable
savanna habitats in
the interstitiary region
and/or the well
established presence
A
of Julodis in the
Near East. C

Fig. 8: Approximate distribution of Aaata finchi (Waterhouse) (symbol A, records from literature); distribution of Afrotropical Sternocera spp. and sspp.
(symbol B, exact locality records of all species and subspecies superimposed); approximate distribution of Asian Sternocera spp. and sspp.
(symbol C, records from literature).

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Genus JULODIS Eschscholtz:


As in the case of Sternocera, Julodis has a strikingly disjunct distribution (Fig. 9).
The ecology seems linked to arid shrub biomes, mainly in winter rainfall regions. At present such regions
occur both in North East and South West Africa. In the past an arid corridor repeatedly connected these two areas
during interpluvial episodes (Tinley 1975). The presence in the south of one relic species (J. bennigseni) that fits
the ovipositor type of the northern species-group,
suggests that dispersal along this arid corridor
happened more than once.
Julodis derives from a Julodella-like an-
cestor, but because of the paraphyletic bifur- A
cation between these two groups it is not
possible to sequence this event relative to other
derivations of Julodella. Variation is highest in
the most widespread species in both the sout-
hern and northern groups. In the south, intrusi-
ons in the semi-arid savanna thus seem the most
recent, in the north the invasion of the Medi-
terranean area by one polytypic species (Cobos
1953) is certainly secondary. Species diversity
and variability of the northern fauna is, however,
currently difficult to assess and therefore centers
of origin and radiation remain speculative.
The disjunct distribution in Africa is at
B
present maintained by the lack of suitable habi-
tats and/or competitive exclusion by Sternocera.

Fig. 9: Approximate distribution of northern Julodis spp. and sspp. (symbol A, records from literature); distribution of southern African Julodis spp. and
sspp. (symbol B, exact locality records of all species and subspecies superimposed).

Genus JULODELLA Semenov Tian-Shanskij:


The genus is clearly centered in the Near East (Fig. 10) with limited isolates in Africa. The two southern species
are aberrant relics, probably derived from early expansions along the arid corridor during interpluvials (see
Julodis above).
The ecological and climatic niche is
apparently similar to Julodis, with which the
distribution more or less coincides. As Julodella A
has a significant smaller average body size than
Julodis, competition is possibly avoided by
occupation of different micro-niches.
Phylogenetically Julodella represents
the ‘surviving ancestor’, and conclusions as to its
own derivation cannot be based on its paraphyletic
derivates. The high species diversity in the Near
East, however, suggests an origin in that area.
D

E
B

Fig. 10: Approximate distribution of Palaearctic Julodella spp. and sspp. (symbol A, records from literature); distribution of southern African Julodella
spp. (symbol B: J. bicolor (Obst); symbol C: J. cicatricosa (Germar)); distribution of Amblysterna spp. (symbol D: A. johnstoni Waterhouse;
symbol E: A. natalensis (Boheman)).

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Sternocera castanea boucardii Saunders (Pl. 1.13-16 - pg. 106; Figs 55-69)

DISCUSSION This subspecies includes the largest African Sternocera specimens. Large tomentose spots,
at least along the elytral sides, characterize it.The single basal tomentose spot on the elytron
and the abdominal tomentose spots are well developed, and the elytra are mostly metallic green.The distribution
overlaps widely with S. castanea s. str., sometimes with intermediates (particularly in the west), sometimes without,
suggesting a possible ring cline with partly non-interbreeding populations around the Rift Valley. Populations
transitional to S. castanea s. str. with brown elytra and large lateral tomentose spots on the elytra (as in Pl. 1.12)
occur in the western areas. More in the northern central areas are also populations transitional to S. castanea s. str.;
they have large elytral spots and a green elytral base, but brown apex (rothschildi, Pl. 1.16).This latter form leads
to forms with green elytra and semi-confluent cross bands of tomentose spots (fasciata, Pl. 1.13), or many finer
spots on the disc (multiimpressa, Pl. 1.14), or large lateral but much smaller discal spots (boucardii, fulvoguttata,
microsticta, Pl. 1.15), reaching an extreme reduction in patrizii.

DIAGNOSIS
• Body length 48.6 to 52.8 mm.
• Antennae brown.
• Tibiae and tarsi brown.
• Postero-ventral pronotal margin rounded, without lobe (Fig. 55).
• Lateral pronotal maculae always absent.
• Pronotum with coarse sculpture, often merging into elongate depressions; punctures
and depressions tomentose (Fig. 56); black, or black with metallic sheen.
• Elytron with 1 large, rounded tomentose basal spot, always with additional rounded tomentose
patches (variable in size and number but considerably larger than in S. castanea s. str.)
in apical two thirds (Pl. 1.13-16); large, oval tomentose subhumeral patch present.
• Subhumeral ridge thinly carinate.
• Elytron with fine and regular tomentose punctures; mostly
black with green metallic sheen, rarely with elytral sides and
even apical quarter to one third reddish-brown (e.g. Fig. 61);
some specimens with dark brown elytra (Pl. 1.13-16).
• Venter black, with short setae; abdominal sterna
with tomentose patches (Pl. 1.13-16).
• Meso-metasternal projection pointed to roundedly
pointed, slightly protruding downward (Fig. 55).
• Genitalia as in Fig. 55.

Fig. 56:
Fig. 55: S. c. boucardii Saunders S. c. boucardii Saunders

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S
ETH

t
SOM

UGA
0 KEN

e
DRC
5

r
TAN

n
10
ZAM
Sternocera castanea boucardii Saunders

o
15
15 20 25 30 35 40 45 50

c
e
r
KARL WERNER
Fig. 57: Habitat of Sternocera c. boucardii, Kenya,

a
South Horr, V. 2001

ROBERT LIZLER

Fig. 58:
Sternocera c. boucardii Saunders
f, Tanzania, Uluguru Mts.,
IV.1997, Werner & Lizler leg., KWCG
(1,5x actual size) Fig. 59: Habitat of Sternocera c. boucardii, Kenya, near Isiolo, Shaba N.P., VIII. 1996

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SIBYLLE GUSSMANN

Fig. 60: Habitat of Sternocera c. boucardii, Kenya, Tsavo West N.P., X. 1993

Fig. 61:
Sternocera c. boucardii Saunders
m, Kenya, Wajir to Moyale,
6.V.2001, Werner & Smrz leg., KWCG
(1,5x actual size)
KARL WERNER

Fig. 62: Habitat of Sternocera c. boucardii, Kenya, Hola, V. 1999


SIBYLLE GUSSMANN

Fig. 63: Habitat of Sternocera c. boucardii, Kenya, Tsavo West N.P., X. 1993

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S
t
e
r
n
Fig. 64: Fig. 65: Fig. 66:
Sternocera c. boucardii Saunders Sternocera c. boucardii Saunders Sternocera c. boucardii Saunders
f, Kenya, Wajir to Moyale, f, Kenya, Mado Gashi, f, Kenya, Mado Gashi,
5.V.2001, Werner & Smrz leg., KWCG 1.V.2001, Werner & Smrz leg., KWCG 1.V.2001, Werner & Smrz leg., KWCG

o
(1,5x actual size) (1,5x actual size) (1,5x actual size)

c
e
r
a
Fig. 67: Fig. 68: Fig. 69:
Sternocera c. boucardii Saunders Sternocera c. boucardii Saunders Sternocera c. boucardii Saunders
f, Kenya, Wajir to Marsabit, f, Kenya, Mado Gashi to Wajir, f, Kenya, South Horr,
6.V.2001, Werner & Smrz leg., KWCG 1.V.2001, Werner & Smrz leg., KWCG 8.V.2001, Werner & Smrz leg., KWCG
(1,5x actual size) (1,5x actual size) (1,5x actual size)

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J
DIAGNOSIS
• Body length 19,0 – 37,7 mm.
• Integument extremely variable (Pl. 4.10-12), either dorsum metallic green to blue, venter and legs

u
darker, often with more intense blue or greenish-bronze sheen; or sometimes dorsum dark metallic
green, bluish-green or blue (often with more intense bluish sheen on pronotum) but grading into
castaneous to various degrees on elytron, and venter and legs concolourous with pronotum but
darker; or rarely unicolourous black with weak metallic bronze reflections.
• Pulverulence variable (Pl. 4.10-12), either white to yellowish-white but yellow to red on head

l
(only from between eyes posteriorly across vertex) and on subhumeral setal patch; or sometimes
white to pale-yellow but yellow on pronotum, and orange on head (only from between eyes
posteriorly across vertex) and on setal patches in marginal elytral row.
• Head with densely pulverulent setae from between eyes posteriorly across vertex and less densely

o
pulverulent setae elsewhere.
• Pronotum with foveae medially, antero-paramedially and laterally in anterior half of length merging
into continuous or subcontinuous grooves; with isolated foveae irregular and variable in size
and spacing; grooves and isolated foveae with dense pulverulence.
• Elytron with larger foveae variable in size (sometimes reduced to size of interstitial foveae),

d
relatively regularly spaced, arranged in 5 rows, but with mediobasal, basosutural and mediolateral
fovea always markedly larger; larger foveae with dense, smaller foveae with weaker pulverulence.
• Hypomeron medially with pulverulent setae not forming a tuft.
• Metasternum without, or with weak pulverulence.
• Metepisternum mostly with pulverulence on entire surface.

i
• Metacoxa with postero-internal angle aspinose; median section with glabrous area often limited
to anterior two thirds of metacoxal width; posterior margin unmodified; with pulverulent setae
around glabrous area.
• Legs with tibial setae not longer than mid-tibial width; metatibia dorsally with very densely spaced

s
punctures, these larger and slightly more widely spaced laterally and in proximal one third to
one quarter of tibial length.
• Abdominal sterna with isolated punctures and clusters of punctures, irregular in size and spacing;
each sternum with pulverulence randomly spread over isolated punctures and clusters of punctures
but dense on a larger (but often poorly defined) lateral setal patch and very often along mid-line;
apex of last sternum truncate in male.
• Setation on head erect, approximately as long as half width
of eye, on pronotum erect, shorter
and longer than half width of eye; on elytron recumbent,
shorter than half width of eye,
with additional sparse erect setae near base, majority longer
than half width of eye; on venter
variable in length and density, recumbent to upright.
• Male genitalia as in Fig. 252.

Fig. 252: J. e. egho Gory

Fig. 253:
J. e. egho Gory

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139
14 Julodis egho egho Gory
ANG ZAM
16
MOC
18
ZIM

20 BOT

22

24
NAM
26

28

RSA
30

32

34
12 14 16 18 20 22 24 26 28 30 32 34 36 38 40

Fig. 254: Fig. 255:


Julodis egho egho Gory Julodis egho egho Gory
m, Namibia, 100 km N of Okahandja, f, Namibia, 100 km N of Okahandja,
6.II.1997, K.Werner leg., KWCG 6.II.1997, K.Werner leg., KWCG
(2,5x actual size) (2,5x actual size)

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J
u
l
o
d
i
Fig. 256: Fig. 257: Fig. 258:
Julodis egho egho Gory Julodis egho egho Gory Julodis egho egho Gory
Namibia, Marienfluss, III.1995, Namibia, Marienfluss, III.1995, f, Namibia, Blasskranz, 10.III.1997,
Owen leg., KWCG Owen leg., KWCG Owen leg., KWCG
(2,5x actual size) (2,5x actual size) (2,5x actual size)

Fig. 259: Fig. 260: Fig. 261:


Julodis egho egho Gory Julodis egho egho Gory Julodis egho egho Gory
f, Namibia, Blasskranz, 10.III.1997, Namibia, Kuduberg, III.1986, m, Namibia, Kuduberg, III.1986,
Owen leg., KWCG C.R. Owen leg., KWCG C.R. Owen leg., KWCG
(2,5x actual size) (2,5x actual size) (2,5x actual size)

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J
u
l
o
d
SIBYLLE GUSSMANN

i
Fig. 264: Julodis egho egho Gory, Namibia, Okahandja River, 96 km W Okahandja, sitting on host-plant Lycium sp.,13.IV. 1991

s
SIBYLLE GUSSMANN

Fig. 265: Habitat of Julodis egho egho Gory, Namibia, Windhoek, 14.IV. 1991
SIBYLLE GUSSMANN

SIBYLLE GUSSMANN

Fig. 266: Freshly caught Julodis egho egho Gory specimen, Fig. 267: Freshly caught Julodis egho egho Gory specimen,
Namibia, 45 km NW Giribes Vlakte, 12.IV. 1991 Namibia, Windhoek, 14.IV. 1991

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J
DIAGNOSIS
• Body length 21,8 – 35,9 mm.
• Dorsum usually unicolourous metallic green to blue (sometimes grading to violet on sides),
sometimes elytron with more intense blue sheen than pronotum and head; venter concolourous

u
with elytron or darker; legs usually matt green, greenish-bronze or greenish-blue, rarely black.
• Pulverulence unicolourous yellow (Pl. 5.11, Fig. 348) or orange.
• Head with pulverulent setae from between eyes posteriorly across vertex, sometimes more densely
spaced bilaterally.

l
Pronotum with 5 longitudinal bands of foveae merging into continuous or subcontinuous grooves;
grooves and isolated foveae with dense pulverulence.
• Elytron with larger foveae regularly spaced, rounded (rarely elongated), arranged in 5 rows
and bearing pulverulent setal tufts; with smaller interstitial foveae bearing sparse, usually
non-pulverulent setae; elytral costae sometimes discernibly raised.

o
• Hypomeron medially with pulverulent setal tuft.
• Metasternum without pulverulence.
• Metepisternum with band of pulverulent setae in ventral half of length.
• Metacoxa with postero-internal angle aspinose; median section with usually large glabrous area

d
(rarely limited to anterior two thirds to one third of metacoxal width) barely reaching posterior
margin, the latter unmodified; with pulverulent setal tufts proximad and distad (rarely also posteriad)
of glabrous area.
• Abdominal sterna with irregularly spaced small isolated punctures and clusters of punctures;
each sternum with well-defined lateral pulverulent setal patch, its diameter on sternum 2-4 one third

i
to half of sternum length; apex of last sternum weakly and roundedly concave in male.
• Legs with some tibial setae always longer than mid-tibial diameter; metatibia dorsally with well-spaced
small punctures, these sparser in proximal half to one third of tibial length.
• Setation on head and pronotum erect, majority longer than half width of eye; on elytron either with all

s
setae erect, longer than half width of eye, or with shortest setae erect but grading to recumbent
towards apex, shorter than half width of eye, and with
longest setae erect, longer than half width of eye;
on venter moderately long to long, variably dense.
• Male genitalia as in Fig. 347.

Fig. 347: J. viridipes Laporte


Fig. 348:
J. viridipes Laporte

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16
Julodis viridipes Laporte
18
ZIM
20
NAM

22
BOT MOC

24

26

28

30
RSA
32

34

12 14 16 18 20 22 24 26 28 30 32 34 36

Fig. 349: Fig. 350: Fig. 351:


Julodis viridipes Laporte, Julodis viridipes Laporte, Julodis viridipes Laporte,
f, RSA, Namaqualand, Eksteenfontein, m, RSA, Namaqualand, Eksteenfontein, f, RSA, Namaqualand, near Springbok,
14.X.1999, K.Werner leg., KWCG 14.X.1999, K.Werner leg., KWCG 12. + 17.X.1999, K.Werner leg., KWCG
(2x actual size) (2x actual size) (2x actual size)

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J
u
l
o
d
Fig. 352: Fig. 353: Fig. 354:
Julodis viridipes Laporte, Julodis viridipes Laporte, Julodis viridipes Laporte,

i
f, RSA, Namaqualand, near Springbok, m, RSA, Namaqualand, Steinkopf, f, RSA, Namaqualand, Steinkopf,
12. + 17.X.1999, K.Werner leg., KWCG 12.X.1999, K.Werner leg., KWCG 12.X.1999, K.Werner leg., KWCG
(2x actual size) (2x actual size) (2x actual size)

Fig. 355: Fig. 356: Fig. 357:


Julodis viridipes Laporte, Julodis viridipes Laporte, Julodis viridipes Laporte,
m, RSA, Namaqualand, Steinkopf, f, RSA, Namaqualand, Kamieskroon, m, RSA, Namaqualand, Kamieskroon,
12.X.1999, K.Werner leg., KWCG 11.X.1999, K.Werner leg., KWCG 11.X.1999, K.Werner leg., KWCG
(2x actual size) (2x actual size) (2x actual size)

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Fig. 358: Fig. 359: Fig. 360:


Julodis viridipes Laporte, Julodis viridipes Laporte, Julodis viridipes Laporte,
f, RSA, Cape, Namaqualand, Clanwilliam, m, RSA, Namaqualand, Cedarberg Reg., m, RSA, Namaqualand, near Garies,
9.X.1999, K.Werner leg., KWCG 10.X.1999, K.Werner leg., KWCG 11.X.1999, K.Werner leg., KWCG
(2x actual size) (2x actual size) (2x actual size)
K. & G. WERNER

Fig. 361:
Julodis viridipes Laporte,
m, RSA, Namaqualand, near Garies,
11.X.1999, K.Werner leg., KWCG
(2x actual size) Fig. 362: Julodis viridipes Laporte, RSA, Namaqualand, Steinkopf, 12.X. 1999

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J
u
l
o
K. & G. WERNER

d
Fig. 363: Julodis viridipes Laporte, RSA, Namaqualand,

K. & G. WERNER
near Garies, 11.X. 1999

i
Fig. 364: Julodis viridipes Laporte, RSA, Namaqualand,
Eksteenfontein, 14.X. 1999

s
SIBYLLE GUSSMANN

Fig. 365: Host-plant of Julodis viridipes Laporte, RSA, Clanwiliam, 1.IX. 1990, host-plant Zygophyllum foetidum Schrad. & Wendl.

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SIBYLLE GUSSMANN

Fig. 366: Habitat of Julodis viridipes Laporte, RSA, Spektakelplas, 30.VIII. 1990
SIBYLLE GUSSMANN
SIBYLLE GUSSMANN

Fig. 367: Habitat of Julodis viridipes Laporte, RSA, Fig. 368: Host-plant of Julodis viridipes Laporte, RSA, Clanwiliam,
Spektakelplas, 30.VIII. 1990 1.IX. 1990, host-plant Zygophyllum foetidum Schrad. & Wendl.

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Amblysterna johnstoni Waterhouse (Figs 502-507)

DISCUSSION Despite the parapatric distribution with A. natalensis and the morphological proximity
between the two species, a subspecific relationship between them is ruled out:
the consistent differences in genitalia and the absence of any gradation from one to the other argue against it.
Amblysterna johnstoni occurs in the East African moist savanna (see map). Note an interesting record
from the Seychelles (not illustrated in map), which, in the absence of the species from Madagascar, is probably
due to transportation. The morphology, apart from some colour variation towards blue, and size differences, barely
varies. Three specimens were found in the ZMHB, marked by Kunzen as three different subspecies though they
hardly differ from one another. To our knowledge they were never described. The holotypes of stictica and
johnstoni are identical. Adults of A. johnstoni were mostly collected between April and June, with old records
from December and February. Host-plants and biology are unknown.

DIAGNOSIS
The species A. johnstoni is structurally very similar to A. natalensis, but differs very consistently in
the following:
• Body length 21,5 – 28,2 mm; width 7,8 – 10,5 mm.
• Colour invariably metallic green, while A. natalensis varies from green, purple to black.
• Pronotum without marked pulverulent lateral depressions, irregularly and coarsely sculptured laterally
(Fig. 503).
• Elytra with lateral and sutural pubescent bands absent, instead evenly covered by large punctures with
pulverulent pubescence; these punctures become larger towards the apex (Fig. 503).
• Third and fourth costae not clearly merging near apex, costae less raised than in A. natalensis.
• Pubescence on underside denser and more extensive than in A. natalensis.
• Genitalia as in Fig. 502.

Fig. 503:
Fig. 502: Amblysterna johnstoni Waterhouse Amblysterna johnstoni Waterhouse

T h e J e w e l B e e t l e s o f A f r i c a ( B u p r e s t i d a e : J u l o d i n a e )
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Manuscript3OK.qxd 2.2.2004 14:35 Stránka 245

A
10

m
ETH

5 SOM

b
KEN

l
5
TAN

y
Amblysterna johnstoni Waterhouse

10
10 15 20 25 30 35 40 45 50

s
t
e
r
KARL WERNER

n
Fig. 506: Habitat of Amblysterna johnstoni Waterhouse, Kenya,
NE. Prov., El Wak, 1/3.V. 2001

Fig. 504: Fig. 505:


Amblysterna johnstoni Waterhouse, Amblysterna johnstoni Waterhouse,
m, Kenya, Isiolo, 18.-22.IV.1999, f, Kenya, Isiolo, 18.-22.IV.1999,
K.Werner & R.Lizler leg., KWCG K.Werner & R.Lizler leg., KWCG
(3x actual size)
ROBERT LIZLER

(3x actual size)

Fig. 507: Habitat of Amblysterna johnstoni Waterhouse, Kenya,


Shaba N.P., VIII. 1994

T h e J e w e l B e e t l e s o f A f r i c a ( B u p r e s t i d a e : J u l o d i n a e )
245

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