Julodis Leaflet

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

B

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 ancestor, but because of the paraphyletic bifurA 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 southern and northern groups. In the south, intrusions in the semi-arid savanna thus seem the most recent, in the north the invasion of the Mediterranean 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 habitats 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 A distribution more or less coincides. As Julodella 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

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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: S. c. boucardii Saunders

Fig. 55: S. c. boucardii Saunders

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Fig. 58: Sternocera c. boucardii Saunders f, Tanzania, Uluguru Mts., IV.1997, Werner & Lizler leg., KWCG (1,5x actual size)

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e Fig. 57: Habitat of Sternocera c. boucardii, Kenya, South Horr, V. 2001

Fig. 59: Habitat of Sternocera c. boucardii, Kenya, near Isiolo, Shaba N.P., VIII. 1996

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

KARL WERNER

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

SIBYLLE GUSSMANN

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

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

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S t e r n Fig. 66: Sternocera c. boucardii Saunders f, Kenya, Mado Gashi, 1.V.2001, Werner & Smrz leg., KWCG (1,5x actual size)

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Fig. 65: Sternocera c. boucardii Saunders f, Kenya, Mado Gashi, 1.V.2001, Werner & Smrz leg., KWCG (1,5x actual size)

Fig. 64: Sternocera c. boucardii Saunders f, Kenya, Wajir to Moyale, 5.V.2001, Werner & Smrz leg., KWCG (1,5x actual size)

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Fig. 68: Sternocera c. boucardii Saunders f, Kenya, Mado Gashi to Wajir, 1.V.2001, Werner & Smrz leg., KWCG (1,5x actual size)

Fig. 67: Sternocera c. boucardii Saunders f, Kenya, Wajir to Marsabit, 6.V.2001, Werner & Smrz leg., KWCG (1,5x actual size)

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Fig. 69: Sternocera c. boucardii Saunders f, Kenya, South Horr, 8.V.2001, Werner & Smrz leg., KWCG (1,5x actual size)

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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 (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 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), 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. 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 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.

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

Fig. 252: J. e. egho Gory Fig. 253: J. e. egho Gory

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Fig. 255: Julodis egho egho Gory f, Namibia, 100 km N of Okahandja, 6.II.1997, K.Werner leg., KWCG (2,5x actual size)

Fig. 254: Julodis egho egho Gory m, Namibia, 100 km N of Okahandja, 6.II.1997, K.Werner leg., KWCG (2,5x actual size)

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Fig. 259: Julodis egho egho Gory f, Namibia, Blasskranz, 10.III.1997, Owen leg., KWCG (2,5x actual size)

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Fig. 257: Julodis egho egho Gory Namibia, Marienfluss, III.1995, Owen leg., KWCG (2,5x actual size)

Fig. 258: Julodis egho egho Gory f, Namibia, Blasskranz, 10.III.1997, Owen leg., KWCG (2,5x actual size)

Fig. 260: Julodis egho egho Gory Namibia, Kuduberg, III.1986, C.R. Owen leg., KWCG (2,5x actual size)

Fig. 261: Julodis egho egho Gory m, Namibia, Kuduberg, III.1986, C.R. Owen leg., KWCG (2,5x actual size)

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Fig. 256: Julodis egho egho Gory Namibia, Marienfluss, III.1995, Owen leg., KWCG (2,5x actual size)

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

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

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

Fig. 267: Freshly caught Julodis egho egho Gory specimen, Namibia, Windhoek, 14.IV. 1991

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sometimes elytron with more intense blue sheen than pronotum and head; venter concolourous 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. 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. 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 (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 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 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.

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DIAGNOSIS • Body length 21,8 – 35,9 mm. • Dorsum usually unicolourous metallic green to blue (sometimes grading to violet on sides),

Fig. 347: J. viridipes Laporte Fig. 348: J. viridipes Laporte

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16 Julodis viridipes Laporte 18 ZIM 20 NAM 22

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Fig. 349: Julodis viridipes Laporte, f, RSA, Namaqualand, Eksteenfontein, 14.X.1999, K.Werner leg., KWCG (2x actual size)

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Fig. 350: Julodis viridipes Laporte, m, RSA, Namaqualand, Eksteenfontein, 14.X.1999, K.Werner leg., KWCG (2x actual size)

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Fig. 351: Julodis viridipes Laporte, f, RSA, Namaqualand, near Springbok, 12. + 17.X.1999, K.Werner leg., KWCG (2x actual size)

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J u l o d Fig. 354: Julodis viridipes Laporte, f, RSA, Namaqualand, Steinkopf, 12.X.1999, K.Werner leg., KWCG (2x actual size)

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Fig. 353: Julodis viridipes Laporte, m, RSA, Namaqualand, Steinkopf, 12.X.1999, K.Werner leg., KWCG (2x actual size)

Fig. 352: Julodis viridipes Laporte, f, RSA, Namaqualand, near Springbok, 12. + 17.X.1999, K.Werner leg., KWCG (2x actual size)

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Fig. 356: Julodis viridipes Laporte, f, RSA, Namaqualand, Kamieskroon, 11.X.1999, K.Werner leg., KWCG (2x actual size)

Fig. 355: Julodis viridipes Laporte, m, RSA, Namaqualand, Steinkopf, 12.X.1999, K.Werner leg., KWCG (2x actual size)

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Fig. 358: Julodis viridipes Laporte, f, RSA, Cape, Namaqualand, Clanwilliam, 9.X.1999, K.Werner leg., KWCG (2x actual size)

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Fig. 360: Julodis viridipes Laporte, m, RSA, Namaqualand, near Garies, 11.X.1999, K.Werner leg., KWCG (2x actual size)

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

Fig. 359: Julodis viridipes Laporte, m, RSA, Namaqualand, Cedarberg Reg., 10.X.1999, K.Werner leg., KWCG (2x actual size)

Fig. 362: Julodis viridipes Laporte, RSA, Namaqualand, Steinkopf, 12.X. 1999

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Fig. 363: Julodis viridipes Laporte, RSA, Namaqualand, near Garies, 11.X. 1999

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

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

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

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

Fig. 367: Habitat of Julodis viridipes Laporte, RSA, Spektakelplas, 30.VIII. 1990

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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:

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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: Amblysterna johnstoni Waterhouse

Fig. 502: Amblysterna johnstoni Waterhouse

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KARL WERNER

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

a

Fig. 505: Amblysterna johnstoni Waterhouse, f, Kenya, Isiolo, 18.-22.IV.1999, K.Werner & R.Lizler leg., KWCG (3x actual size)

ROBERT LIZLER

Fig. 504: Amblysterna johnstoni Waterhouse, m, Kenya, Isiolo, 18.-22.IV.1999, K.Werner & R.Lizler leg., KWCG (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 )

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