julodis leaflet

This monograph is (we hope) the first of a series, intendedto collate our present knowledge of the family Buprestidaein the Afrotropical Region. The reasons for embarkingon such a venture may be clothed into emotive issues likeecological planning and documenting the fauna beforeit disappears. The fact is, we simply like buprestids and welike the taxonomic excuse to study them.

The general arranging of information in the bookis aimed at handiness for identification. We therefore listsynonymies, 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 collectionsunder their care as too many synonyms in large genera wentundetected for decades simply because of alphabeticalordering. Genera, subgenera and species are thereforetreated in the same sequence as they key out in the variousphylogenetic cladograms. In the case of subspecies, however,the nominal subspecies is always treated first. Diagnosticinformation (comparative notes, diagnoses, detail drawingsand distribution maps) of every species or subspecies isprovided together on one page. The introduction chaptersprovide short backgrounds limited to considerations that arerelevant to our taxonomic decisions. They are certainly notintended as exhaustive treatments of the various themes, asspeculations on historical geography and phylogeny alonecould fill a volume without, however, contributing much tothe taxonomy.more then 600 color photographs of all speciesand subspecies from different localities as wellas photos of behavior and habitatsover 400 pages, luxurious hardcover editionlarge format 23x31, text in english

T H E A F R I C A N J E W E L B E E T L E S

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 )16

The known distribution limits of the extant genera and subgenera of the subfamily are given in Figs 8-11.Thereare 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 ofus (Holm 1979a). Since that paper further revisional works and descriptions have changed diversity statisticssomewhat. 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; theirphylogenetic relationships; their macro-ecological niches; and the relative species densities, apomorphies andvariabilities in subregions of the distribution limits.These data we briefly summarize below.

1.3 ZOOGEOGRAPHY

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 mostprimitive 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 AAATA Semenov Tian-Shanskij:

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 EastAfrica 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 eventsor glaciation) must have happened when species were already diversified. The isolation is now maintained bythe scarcity of suitablesavanna habitats inthe interstitiary regionand/or the wellestablished presenceof Julodis in theNear East.

Genus STERNOCERA Eschscholtz:

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).

B

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C

Manuscript1OK.qxd 2.2.2004 11:44 Stránka 16


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 areasduring interpluvial episodes (Tinley 1975).The presence in the south of one relic species (J. bennigseni) that fitsthe ovipositor type of the northern species-group,suggests that dispersal along this arid corridorhappened more than once.

Julodis derives from a Julodella-like an-cestor, but because of the paraphyletic bifur-cation between these two groups it is notpossible to sequence this event relative to otherderivations of Julodella. Variation is highest inthe most widespread species in both the sout-hern and northern groups. In the south, intrusi-ons in the semi-arid savanna thus seem the mostrecent, in the north the invasion of the Medi-terranean area by one polytypic species (Cobos1953) is certainly secondary. Species diversityand variability of the northern fauna is, however,currently difficult to assess and therefore centersof origin and radiation remain speculative.

The disjunct distribution in Africa is atpresent maintained by the lack of suitable habi-tats and/or competitive exclusion by Sternocera.

Genus JULODIS Eschscholtz:

The genus is clearly centered in the Near East (Fig. 10) with limited isolates in Africa.The two southern speciesare aberrant relics, probably derived from early expansions along the arid corridor during interpluvials (seeJulodis above).

The ecological and climatic niche isapparently similar to Julodis, with which thedistribution more or less coincides. As Julodellahas a significant smaller average body size thanJulodis, competition is possibly avoided by occupation of different micro-niches.

Phylogenetically Julodella representsthe ‘surviving ancestor’, and conclusions as to itsown derivation cannot be based on its paraphyleticderivates. The high species diversity in the NearEast, however, suggests an origin in that area.

Genus JULODELLA Semenov Tian-Shanskij:

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

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

A

B

A

D

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E


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

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 distributionoverlaps 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. Populationstransitional 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 leadsto forms with green elytra and semi-confluent cross bands of tomentose spots (fasciata, Pl. 1.13), or many finerspots 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.

• 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; puncturesand depressions tomentose (Fig. 56); black, or black with metallic sheen.

• Elytron with 1 large, rounded tomentose basal spot, always with additional rounded tomentosepatches (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 andeven 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 sternawith tomentose patches (Pl. 1.13-16).

• Meso-metasternal projection pointed to roundedlypointed, slightly protruding downward (Fig. 55).

• Genitalia as in Fig. 55.

DISCUSSION

DIAGNOSIS


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


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 )5 1

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Sternocera castanea boucardii Saunders

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

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

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



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Fig. 63: 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)

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

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

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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)

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

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

Fig. 66: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)

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


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 )1 39

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

darker, often with more intense blue or greenish-bronze sheen; or sometimes dorsum dark metallicgreen, bluish-green or blue (often with more intense bluish sheen on pronotum) but grading intocastaneous to various degrees on elytron, and venter and legs concolourous with pronotum butdarker; 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 sometimeswhite to pale-yellow but yellow on pronotum, and orange on head (only from between eyesposteriorly across vertex) and on setal patches in marginal elytral row.

• Head with densely pulverulent setae from between eyes posteriorly across vertex and less denselypulverulent setae elsewhere.

• Pronotum with foveae medially, antero-paramedially and laterally in anterior half of length merginginto continuous or subcontinuous grooves; with isolated foveae irregular and variable in sizeand 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 mediolateralfovea 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 setaearound glabrous area.

• Legs with tibial setae not longer than mid-tibial width; metatibia dorsally with very densely spacedpunctures, these larger and slightly more widely spaced laterally and in proximal one third toone 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 puncturesbut 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 widthof eye, on pronotum erect, shorterand longer than half width of eye; on elytron recumbent,shorter than half width of eye,with additional sparse erect setae near base, majority longerthan half width of eye; on ventervariable in length and density, recumbent to upright.

• Male genitalia as in Fig. 252.

DIAGNOSIS

Fig. 252: J. e. egho Gory

Fig. 253:J. e. egho Gory


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Julodis egho egho Gory

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

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


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

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

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

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

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

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


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

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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. 267: Freshly caught Julodis egho egho Gory specimen,Namibia, Windhoek, 14.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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• 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 concolourouswith 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 denselyspaced 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 rowsand bearing pulverulent setal tufts; with smaller interstitial foveae bearing sparse, usuallynon-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 posteriormargin, 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 thirdto 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-spacedsmall 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 allsetae erect, longer than half width of eye, or with shortest setae erect but grading to recumbenttowards apex, shorter than half width of eye, and withlongest setae erect, longer than half width of eye;on venter moderately long to long, variably dense.

• Male genitalia as in Fig. 347.

DIAGNOSIS

Fig. 347: J. viridipes Laporte

Fig. 348:J. viridipes Laporte



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

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Julodis viridipes Laporte

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

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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Fig. 353:Julodis viridipes Laporte,m, RSA, Namaqualand, Steinkopf,12.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)

Fig. 357:Julodis viridipes Laporte,m, RSA, Namaqualand, Kamieskroon,11.X.1999, K.Werner leg., KWCG(2x actual size)

Fig. 356:Julodis viridipes Laporte,f, RSA, Namaqualand, Kamieskroon,11.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)

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



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

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

Fig. 358:Julodis viridipes Laporte,f, RSA, Cape, Namaqualand, Clanwilliam,9.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. 361:Julodis viridipes Laporte,m, RSA, Namaqualand, near Garies,11.X.1999, K.Werner leg., KWCG(2x actual size)



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

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



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

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

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


Amblysterna johnstoni Waterhouse (Figs 502-507)

Despite the parapatric distribution with A. natalensis and the morphological proximitybetween 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 probablydue to transportation. The morphology, apart from some colour variation towards blue, and size differences, barelyvaries. Three specimens were found in the ZMHB, marked by Kunzen as three different subspecies though theyhardly differ from one another. To our knowledge they were never described. The holotypes of stictica andjohnstoni are identical. Adults of A. johnstoni were mostly collected between April and June, with old recordsfrom December and February. Host-plants and biology are unknown.

The species A. johnstoni is structurally very similar to A. natalensis, but differs very consistently inthe 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 withpulverulent 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.

DISCUSSION

DIAGNOSIS


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



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Amblysterna johnstoni Waterhouse

Fig. 504:Amblysterna johnstoni Waterhouse,m, Kenya, Isiolo, 18.-22.IV.1999,K.Werner & R.Lizler leg., KWCG(3x actual size)

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

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

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Fig. 507: Habitat of Amblysterna johnstoni Waterhouse, Kenya,Shaba N.P., VIII. 1994


julodis leaflet

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