Research Article |
Corresponding author: Renzo Perissinotto ( renzo.perissinotto@nmmu.ac.za ) Academic editor: Andrey Frolov
© 2015 Sébastien Rojkoff, Renzo Perissinotto.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Rojkoff S, Perissinotto R (2015) Review of the genera Anelaphinis Kolbe, 1892 and Atrichelaphinis Kraatz, 1898 (Coleoptera, Scarabaeidae, Cetoniinae). ZooKeys 482: 91-142. https://doi.org/10.3897/zookeys.482.8343
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New material collected recently throughout the Afrotropical region has led to a major reassessment of taxa within the genera Anelaphinis Kolbe, 1892, Atrichelaphinis Kraatz, 1898 and other closely related genera. As a result, the name Megalleucosma Antoine, 1989 is here synonymised with Anelaphinis and a lectotype is designated for the type species, Cetonia dominula Harold, 1879. The genus Atrichelaphinis is redefined and a new subgenus, A. (Eugeaphinis), is proposed for Elaphinis simillima Ancey, 1883, Elaphinis vermiculata Fairmaire, 1894, Niphetophora rhodesiana Péringuey, 1907, Atrichelaphinis deplanata Moser, 1907 (with Anelaphinis kwangensis Burgeon, 1931 as junior synonym) and Anelaphinis sternalis Moser, 1914. Additionally, three new species and one new subspecies are recognised and described in this new subgenus: A. (Eugeaphinis) bomboesbergicasp. n. from South Africa; A. (Eugeaphinis) bjornstadisp. n. from Tanzania; A. (Eugeaphinis) garnierisp. n. from south–east Africa (Tanzania, Zimbabwe); and A. (Eugeaphinis) deplanata minettiissp. n. from central Africa (Malawi, Mozambique, Congo-Kinshasa, Congo-Brazzaville, South Africa, Rwanda, Zambia, Zimbabwe). The genus Atrichelaphinis is compared to its closest relatives and two separate keys are proposed, one for Atrichelaphinis and one for the sub-Saharan genera exhibiting completely or partially fused parameres.
Africa, Cetonia dominula , Anelaphinis , Atrichelaphinis , Heterelaphinis , Megalleucosma , Niphetophora , synonymy, new taxa
A number of new taxa closely associated with the genus Anelaphinis Kolbe, 1892 have recently been reported through intensified work in a number of Afrotropical countries. An attempt to integrate these into existing generic groups has led to a fresh analysis of the type specimens of the species previously included in this genus. This has revealed a state of relative confusion and great uncertainty about the allocation of previously described species to a number of closely related genera that have proliferated during the past century. This, combined with the realization that both genera Anelaphinis and Atrichelaphinis Kraatz, 1898 have effectively not been subject to any substantial revision since their original description (cf.
The two genera Atrichelaphinis and Anelaphinis exhibit simplesiomorphic similarities between each other and with a number of other closely related genera (
The description of morphological characters follows the terminology used in
Collection abbreviations used within the text are as follows:
BMNH The Natural History Museum, London, UK
CCEC Center for the Curation and Study of Collections, Lyon, France
IRSN Belgian Royal Institute of Natural Sciences, Bruxelles, Belgium
ISAM Iziko South African Museum, Cape Town, South Africa
MNHN National Museum of Natural History, Paris, France
MNHU Natural Sciences Museum of the Humboldt University, Berlin, Germany
MRAC Royal Museum for Central Africa, Tervuren, Belgium
NMKE National Museum of Kenya, Nairobi, Kenya
PCAB Private Collection Anders Bjørnstad, Skien, Norway
PCDC Private Collection Didier Camiade, Sallespisse, France
PCJT Private Collection Julien Touroult, Soyaux, France
PCPA Private Collection Philippe Antoine, Roubaix, France
PCRM Private Collection Robert Minetti, La Ciotat, France
PCRP Private Collection R Perissinotto & L Clennell, Port Elizabeth, South Africa
PCSR Private Collection Sébastien Rojkoff, Lyon, France
PCTB Private collection Thierry Bouyer, Chênée, Belgium
PCTG Private Collection Thierry Garnier, Montpellier, France
TMSA Ditsong National Museum of Natural History (formerly Transvaal Museum), Pretoria, South Africa
After clarifying an erroneous identification of Cetonia dominula Harold, 1879, contained in
A study of the type series of A. dominula reveals the presence of all characters used to define Megalleucosma Antoine, 1989 (type species: Eucosma breviceps Kolbe, 1892), especially in the structure of the aedeagus and the sclerite of the internal sac (Figures
It is not known what prompted
Among the other species usually placed within Anelaphinis are: Elaphinis simillima Ancey, 1883; E. vermiculata Fairmaire, 1894; Niphetophora rhodesiana Péringuey, 1907; Atrichelaphinis deplanata Moser, 1907; A. sternalis Moser, 1914; and A. kwangensis Burgeon, 1931. All have in common an aedeagus with parameres completely fused and without sclerite in the internal sac. Eventually, this character has been regarded as key to the diagnosis of the genus Anelaphinis (e.g.
Anelaphinis Kolbe, 1892
Alleucosma Schenkling, 1921, partim.
Megalleucosma Antoine, 1989: 248, 265, syn. n.
Type species: Cetonia dominula Harold, 1879
Anelaphinis allardi (Ruter, 1978)
Eucosma allardi Ruter, 1978
Megalleucosma allardi (Ruter) Antoine, 1989: 269
Anelaphinis bourgoini (Burgeon, 1932)
Alleucosma bourgoini Burgeon, 1932
Megalleucosma bourgoini (Burgeon) Antoine, 1989: 271
Anelaphinis breviceps (Kolbe, 1892)
Eucosma breviceps Kolbe, 1892: 253
Alleucosma breviceps (Kolbe, 1892)
Anelaphinis vaillanti De Lisle, 1947: 38
Megalleucosma breviceps (Kolbe) Antoine, 1989: 271
Anelaphinis dominula (Harold, 1879)
Cetonia dominula Harold, 1879
Anelaphinis maritima (Moser, 1914)
Eucosma maritima Moser, 1914
Alleucosma maritima Moser, 1914
Alleucosma maritimi Schein, 1956: 194
Anelaphinis pauliani (Antoine, 1989)
Megalleucosma pauliani Antoine, 1989: 268
Anelaphinis similis (Antoine, 1989)
Megalleucosma similis Antoine, 1989: 270
This genus was erected by
Following this,
The consequences of the confusion created with the genus Atrichelaphinis and with the species previously included in Anelaphinis are that currently their taxonomic position remains unclear (with the exception of A. dominula) and badly in need of revision. Only two options appear to be viable at the moment: 1) including them in an existing genus; or 2) erecting a new taxonomic entity especially for them. Upon completing an extensive analysis of many specimens for each species, the following key diagnostic characters are here selected for the new taxonomic order proposed in the section below.
Diagnostic characters:
ventral surface, with extensive scale-type hair cover;
clypeus, ratio of length/width;
anterior clypeal margin, with inflexions and/or ridges;
antennal club, longer in male (sexual dimorphism);
pronotal shape, of octogonal type;
pronotal structure, surface more or less bulbous/tectiform, tuberculate or without any modification at middle of the frontal margin, posterior margin more or less emarginate in front of the scutellum;
mesosternal apophysis, transverse;
elytra, tricostate;
protibiae, bi- or tridentate with variable denticle positions;
meso- and metatibiae, exhibiting external carina;
terminal spurs of metatibiae showing sexual dimorphism;
parameres, completely fused and showing apical expansions, apex more or less curved downwards, usually with small median incision/sinuosity;
female genitalia.
As suggested by
Firstly,
Secondly, Antoine also separated the nominal subgenus from Heterelaphinis through the shape of the aedeagal parameres, with apical median protrusion incised or not, the protibias bi- or tridentate and the antennal club, either equal in the two sexes of Atrichelaphinis s. s. or longer in the male of Heterelaphinis. However, observations undertaken during this study show that the antennal club is always longer in males than in females, in both subgenera, with maximum difference observed in Heterelaphinis. To appreciate correctly this character, it is necessary to compare specimens of the same size. This observation is also valid for the four species previously included in Anelaphinis mentioned above.
The ex-Anelaphinis species exhibit the main characteristics of the genus Atrichelaphinis s. l., which should be enough not to erect a new genus. There are, however, two features that do not allow a similar, full placement of some species within this genus. These are a transverse clypeus with sexual dismorphism, protibiae bidentate with wide separation between the teeth, in association with completely fused parameres without apical modification and just curved downwards, rather than reployed. To include species exhibiting such characters, we consider as necessary to erect a new subgenus, A. (Eugeaphinis) subgen. n. The implication of this is that the genus Atrichelaphinis and the two recognized subgenera must be redefined. Thus, the classification of
Atrichelaphinis (Atrichelaphinis) Kraatz, 1898
Elaphinis Péringuey, 1907.
Type species: Cetonia tigrina Olivier, 1789
Atrichelaphinis (Atrichelaphinis) tigrina (Olivier, 1789)
Cetonia tigrina Olivier, 1789
Cetoninus (Cetonia) tigrina (Olivier) MacLeay, 1838
Elaphinis tigrina (Olivier)
Cetonia furvata Fabricius, 1798
Atrichelaphinis furvata (Fabricius)
Euryomia furvata (Fabricius)
Atrichelaphinis (Atrichelaphinis) nigropunctulata (Péringuey, 1896)
Cetonia nigropunctulata Péringuey, 1896
Elaphinis nigropunctulata (Péringuey)
Elaphinis nigropunctata (Péringuey)
Atrichelaphinis (Heterelaphinis) Antoine, 2002
Type species: Elaphinis quadripunctata Lansberge, 1882
Atrichelaphinis (Heterelaphinis) quadripunctata (Lansberge, 1882)
Elaphinis quadripunctata Lansberge, 1882
Atrichelaphinis quadripunctata (Lansberge)
Cetonia quadripunctata (Lansberge)
Atrichelaphinis (Heterelaphinis) sexualis (Schein, 1956)
Leptothyrea sexualis Schein, 1956
Atrichelaphinis (Heterelaphinis) nigra Antoine, 2002
Atrichelaphinis (Eugeaphinis) subgen. n.
Pseudanelaphinis Antoine (in litteris)
Type species: Atrichelaphinis deplanata Moser, 1907
Atrichelaphinis (Eugeaphinis) deplanata deplanata (Moser, 1907)
Atrichelaphinis deplanata Moser, 1907
Atrichelaphinis deplanate (Moser)
Anelaphinis deplanata Moser
Anelaphinis kwangensis Burgeon, 1931
Atrichelaphinis kwangensis (Burgeon)
Atrichelaphinis (Eugeaphinis) deplanata minettii subsp. n.
Atrichelaphinis (Eugeaphinis) rhodesiana (Péringuey, 1907)
Niphetophora rhodesiana Péringuey, 1907
Anelaphinis rhodesiana (Péringuey)
Atrichelaphinis (Eugeaphinis) bomboesbergica sp. n.
Atrichelaphinis (Eugeaphinis) garnieri sp. n.
Atrichelaphinis (Eugeaphinis) simillima (Ancey, 1883)
Elaphinis simillima Ancey, 1883
Anelaphinis simillima (Ancey)
Atrichelaphinis simillima Müller, 1939
Atrichelaphinis (Eugeaphinis) sternalis (Moser, 1914)
Anelaphinis sternalis Moser, 1914
Atrichelaphinis (Eugeaphinis) vermiculata (Fairmaire, 1894)
Elaphinis vermiculata Fairmaire, 1894
Anelaphinis vermiculata (Fairmaire)
Atrichelaphinis vermiculata (Fairmaire)
Atrichelaphinis (Eugeaphinis) bjornstadi sp. n.
Clypeus rectangular; antennal club longer in male than in female; pronotum sub-octogonal, anterior border convex with or without projection, posterior border largely convex, more or less bisinuate, posterior angles not acute; scutellum longer than wide, apex from more or less acute to slightly dull; elytra tricostate; mesosternal apophysis tansverse; male abdomen concave with a median depression; protibia bi- or tridentate, meso- and metatibias with transverse carina on external side, metatibia widened and thickened at apex; aedeagus with parameres fused, except at apex, internal sac without sclerite; female genitalia (Figure
Atrichelaphinis Kraatz, 1898;
Elaphinis Péringuey, 1907;
Cetonia tigrina Olivier, 1789.
Clypeus transverse (almost as long as wide) without sexual dimorphism, bilobed; median part of pronotal posterior border either straight or slightly concave in front of scutellum; protibia tridentate, with two distal denticles very close to each other, proximal tooth sometimes very reduced or as slight sinuosity; metatibial apical spurs not enlarged in female; aedeagus with parameres fused and apically reployed downwards, except sometimes with small incision in downturning apical part (or sinuation when such incision is absent). Two species are currently included in the nominal subgenus.
Cetonia tigrina Olivier, 1789: 91;
Cetoninus (Cetonia) tigrina (Olivier) MacLeay, 1838: 46.
Elaphinis tigrina (Olivier) Blanchard, 1850: 12;
Atrichelaphinis tigrina (Olivier) Moser, 1907: 321;
Atrichelaphinis furvata (Fabricius) Marais & Holm, 1992: 11.
Cetonia furvata Fabricius, 1798: 130;
Elaphinis furvata (Fabricius) Burmeister, 1842: 597;
Euryomia furvata (Fabricius) Redtenbacher, 1868: 81;
Holotype unknown.
(n > 100). Size: length ♂, 8.6–15.2 mm; ♀, 9.6–15.0 mm; width ♂, 5.0–8.8 mm; ♀, 5.3–9.1 mm.
Body. Dorsum orange-brown, matt with black marks well defined and more or less developed, especially on head and pronotum; often with white tomentose spots on pronotum, scutellum and exceptionally on elytra; setae short on vertex, pronotum (mainly on lateral side) and elytral base, extremely short and barely visible on clypeus, elytra and pronotal disc.
Head. Clypeus almost as long as wide, bilobed and upturned on anterior margin, punctures deep, with setigerous punctures becoming confluent laterally and in front.
Pronotum. Angles round, postero-lateral ones sometimes vanishing; posterior margin straight to concave in front of scutellum; with reborded lateral margins.
Scutellum. Variably marked with black markings and white tomentose spots; setae barely noticeable, mainly at margins.
Elytra. Disc exhibiting three pairs of single to double geminate striae, with first two usually complete, third more or less complete; odd costae convex; sutural apex from blunt to protruding.
Pygidium. Black with some light to dark brown areas; lunulate setigerous punctures, sometimes forming a complete ring on the surface but near apex forming more or less horizontal and confluent lines; apex marginated.
Underside. Shiny, black with more or less developed brown areas and white tomentose spots on metasternum and abdomen; setae long, scattered and shorter on mesepimera and abdominal sternites; mesosternal apophysis transverse, sometimes very large, compressed between mesocoxae, usually flat, sometimes showing relief, covered with setae, but not protruding in front of them; metasternum laterally strongly sculptured with wrinkles, median part glabrous and smooth, with longitudinal mediam line; abdomen more densely sculptured laterally with horseshoe punctures; concave in males, sometimes with slight groove, convex in females.
Legs. Long setae particularly dense on femora and tibiae; metatibiae and metafemora thickened, without tomentum; metatibial spur thin and pointed in male, slightly enlarged and blunt in female.
Aedeagus. Parameres subparallel in their apical half, then enlarged; lateral apical angle showing more or less developed hook-like protrusion; downturned part of apex showing incision at middle.
One female from the MNHU (Coll. L.W. Schaufuss, labeled "Cap b. Sp.") exhibits protibiae bidentate, with teeth widely separated. Other female specimens have been observed with the same feature, but no males. This seems to be exceptional and possibly due to wearing during the fossorial action required to lay eggs underground.The species is mainly distributed in the eastern part of South Africa, reaching the Western Cape Province along the southern coastline. There are also occasional reports from Zimbabwe and southern Mozambique (
Cetonia nigropunctulata Péringuey, 1896: 163;
Elaphinis nigropunctulata (Péringuey) Péringuey, 1907: 448;
Elaphinis nigropunctata (Péringuey) Distant, 1897: 576;
Atrichelaphinis nigropunctulata (Péringuey) Moser, 1907: 321;
Holotype in ISAM.
(n > 30). Size: length ♂, 12.8–15.2 mm; ♀, 13.1–14.8 mm; width ♂, 7.7–8.8 mm; ♀, 7.7–8.8 mm.
Body. Orange with black markings on pronotum, scutellum and elytra, sometimes very reduced; occasionally showing some isolated small spots of white tomentum on pronotum, pygidium and venter; pilosity occasional and restricted to head.
Head. With vertex and lateral part of the frons black, clypeus slightly transverse, bilobed at apex, with anterior margin reborded and lobes slightly upturned. Sculpture deep, simple, becoming confluent in front, laterally and on frons; antennae darker.
Pronotum. With angles rounded, lateral margins almost entirely reborded except near posterior angles, lateral angles always marked, posterior part of lateral margin concave; posterior margin concave in front of scutellum, then laterally very convex; sculpture usually weak on disc, generally denser and deeper laterally.
Scutellum. Acute, grooved laterally; punctuation limited to anterior angles.
Elytra. Sculpture very scattered, disc with two pairs of geminate striae, usually consisting of virtually complete single lines, sometimes merged with horseshoe sculpture, lateral sculpture present or not, series of deep and large points along lateral margin always present; sutural apex from blunt to slightly protruding.
Pygidium. Sculpture usually of small points or lines, sometimes of wrinkles and/or horseshoe setigerous punctures; posterior margin slightly reborded; occasionally covered with short setae, particularly around margins.
Underside. Black except metepisternum, lateral parts of metacoxae, metasternum and sternites; mesepimeron black or orange; mesosternal apophysis orange with black sides, transverse, compressed between mesocoxae and not protruding; moderately covered with setae, except on abdominal sternites; metasternum with wrinkles and long pilosity laterally, grooved in the middle and poorly sculpted to smooth; abdomen poorly sculpted with setigerous horseshoe punctures, setae short; concave to grooved in males, convex in female.
Legs. Metafemora and metatibia enlarged apically, spurs not dilated in either sex; moderately covered with setae, particularly around base; metatibial spur thin and pointed in male, thin and less acute to sligthly blunt in female.
Aedeagus. Parameres almost twice as long as wide; basal half converging in front, apical divergent; lateral apical angles showing fairly developed hook.
The distribution of this species is restricted to the mountainous northeast part of South Africa. Some specimens could be confused superficially with some forms of A. tigrina, however they can be separated through analysis of the dorsal sculpture, shape of the pygidium and aedeagus. The species is most frequently found feeding on Protea spp. flowers.
Elaphinis quadripunctata Lansberge, 1882
Clypeus longer than wide with the apex slightly sinuous; anterior pronotal border tuberculate or tectiform, but minimally so in female; protibiae bidentate, with teeth normally separated; metatibial apical spurs strongly enlarged (spatuliform) in female; parameres of aedeagus fused with apex modified (in dorsal view), with protrusion in the middle deeply incised or not, apex laterally modified or not. Three species are currently included in this subgenus.
Elaphinis quadripunctata Lansberge, 1882: 24;
Atrichelaphinis quadripunctata (Lansberge) Marais & Holm, 1989: 30;
Cetonia quadripunctata (Lansberge) Antoine, 2002: 185.
(n = 7). Size: length LT ♂, 11 mm; width 5.5 mm.
Head. Dark brown with blackish areas, strongly sculpted, converging points forming deep striae; clypeus longer than wide, lateral and anterior margins strongly reborded, anterior slightly upturned and very slightly bilobed, lateral margins almost carinate in the basal part, then curved downwards, depressed in the middle and reborded in the apical part as the anterior margin, clypeal disc convex; frons with large striated protuberance between eyes, vertex with few smooth jointed areas between striae in apical part, posterior part only punctate; antennae brown with long clubs (as long as the flagellum in male).
Pronotum. Transverse, dark brown with transverse points of sculpture, disc poorly punctate, sculpture becoming more dense and confluent to striae in front and laterally; anterior margin slightly wider than head, medially slightly tuberculate; lateral margins reborded with very smooth lateral angles in posterior third; posterior margin bisinuate (concave in front of scutellum), posterior angles rounded.
Scutellum. Dark brown, longer than wide, apex acute, smooth, only a few setigerous points on lateral angles (scale pilosity); laterally grooved.
Elytra. Orange with four black markings, one on side of scutellum, one at middle split on each costa (discal and humeral), one on apical quarter near the suture and last on spiny apex; costae convex, smooth with only few points, discal costa incomplete, humeral costa concave to suture with concavity reaching elytral disc; sculpture of small longitudinal lines (near scutellum) and of horseshoe points in anterior half, becoming confluent posteriorly and leading to two formations: 1) laterally (i.e. between humeral costa and lateral margin) transverse lines becoming longer and denser toward apex; 2) longitunal lines between sutural and humeral costae becoming more numerous and strigillate towards apex; few minute and very short setae near apex.
Pygidium. Transverse, chestnut brown; sculpture horseshoe-like to annulate points drawing large irregular circles towards apex, some transverse striae along apex; few minute and very short setae.
Underside. From dark brown to chestnut brown, sculpture setigerous with long whitish pilosity, not dense except on femora and laterally on sternites 2–5; sculpture sparce, crescent on metasternum, denser to confluent laterally, disc poorly sculpted; abdomen with horseshoe sculpture, median part almost smooth, denser laterally near the margin; posterior coxae reborded laterally, setigerous sculpture made of transversal to backward-curved striae; mesosternal apophysis glabrous, transverse with minute points, strongly compressed between mesocoxae but not protruding; male abdomen concave with visible groove on sternites 2–5.
Legs. From dark brown to chestnut brown, with whitish pilosity; protibiae bidentate, meso- and metatibiae with median carina; profemora strigillate, mesofemora with crescent punctures or small striae, long setigerous stria along internal margin; metafemora slightly dilated with crescent punctures or small striae; all tarsal segments longer than first, metatarsi spiny, claws normal.
Aedeagus. Parameres fused and short, with two carinated lateral spines at apex, apical centre with short protrusion.
Only the MNHN type specimens are known. No recent material was found in the collections examined. Unfortunately,
Leptothyrea sexualis Schein, 1956: 196;
Holotype in NMKE: "Somaliland, Wardere, V.19 (THE Jackson)".
(n = unknown). After
Head. Clypeus longer than wide; lateral and anterior margins reborded and upturned, anterior margin flat and bilobed; deeply punctured; antennal club slightly longer than basal antennomeres, antennae orange/red.
Pronotum. Black or red, with white stripe along the lateral margin and two deep and round white maculae at base in male, red and without white maculae in female; almost as long as wide, posterior margin almost straight in front of scutellum, only slightly concave; posterior angles very blunt; lateral margin parallel in distal part, then strongly convergent in front.
Scutellum. Longer than wide, with lateral margins slightly concave, apex not acute.
Elytra. Black, with white macula at umbone (reaching the suture); 4–6 irregular stripes of broken white maculae and two white longitudinal stripes on disc, parallel to suture, made of irregular and interrupted spots in male; female without white maculae or only reduced marks in place of male stripes; white macula at sutural apex most often present; lateral margins subparallel, narrowing slightly towards apex; lateral costa forked and raised near the shoulder, reaching the humeral callus; sutural costa raised; third costa between sutural and lateral equally raised, joining the lateral costa near apical callus; suture and costae smooth, intervals exhibiting two thin geminate striae usually dissipating near lateral declivity.
Pygidium. Orange/red, covered by annular and ovoid sculpture; with two elongate and interrupted white maculae (separated in 4 parts) in male, absent in female.
Underside. Black, with last and penultimate segments orange/red in female; white maculae on epimeres, lateral parts of sternum and laterally on abdominal sternites 2–5 in male; female immaculate; fore coxae and sides of sternum with whitish pilosity; metasternal apophysis constricted between metacoxae, anterior part flat in shape of hammer; metasternum smooth at middle towards median sulcus, sides striated; abdominal segments widely punctated, with thin setae on sides; male without mid abdominal depression.
Legs. Protibiae widened towards apex, second tooth rounded in male, acute in female; metatibial spurs uneven and acute (longer one slightly curved) in male, enlarged with blunt apex in female; tarsi slender, metatarsus as long as as metatibia in male, shorter in female, first tarsal segment not spiny in either sexes.
Aedeagus. Apex of parameres round with two very small and short median protrusions, without space between them.
Described from Somaliland with no specification on number of type specimens. The description is based on the work of
Atrichelaphinis (Heterelaphinis) sexualis: A parameres. Atrichelaphinis (Heterelaphinis) quadripunctata: B apex of the parameres of the the lectotype. Atrichelaphinis (Heterelaphinis) nigra: C parameres D male pronotum E mesosternal lobe F left elytron (a male; b–d female). Scale bar = 1 mm (From
Holotype male in MNHN: "Somalie, Berbera Check, ex. Coll. Argod 1931". Two female paratypes in MNHN with the same label.
(n = 3). Size: length 8.8–10.3 mm; width 5.2–6.0 mm.
Body. Appearance stocky, black to dark-brown, from dull to slightly shiny, with white tomentose spots; lateral and irregular band on pronotal margin in male, narrower in female, occasionally reduced to line on lateral angle; three main spots on lateral margins of elytra in male, reduced and fragmented in female.
Head. Longer than wide, rectangular, with slightly sinuate anterior margin, slightly upturned and markedly thickened; disc convex; sculpture of large and deep punctures forming laterally some striae; lateral margin almost carinate at base, curving downwards and depressed at middle and reborded in apical part, as anterior margin; vertex and frons without protuberance, with same sculpture as clypeus; antennae dark-brown with club as long as flagellum in male, shorter in female.
Pronotum. Slightly transverse, larger at posterior angles; sculpture of transverse punctures with circular distribution centered at middle of posterior margin, middle unsculpted longitudinal line on disc, posterior margin in front of scutellum also unsculped; anterior margin slightly wider than head, slightly tectiform, lateral margins reborded with very smooth lateral angles at middle; posterior margin convex, straight to convex in front of scutellum.
Scutellum. Black to dark-brown, longer than wide, apex acute, smooth, with few punctures only on lateral angles and along lateral parts of basal third; grooved laterally.
Elytra. Dull, except costae and callus which are slightly shiny; strongly sculpted with two different punctures, fine on costae and horseshoe with central point (semi-annular) on remaining surface; sculpture of first two interstriae becoming confluent in apical half; costae strongly elevated, discal one almost complete to apical callus and strongly developed; apex angular but not produced; lateral margin reborded on basal half.
Pygidium. Transverse with horseshoe setigerous sculpture, setae thin and separate; medial line strongly convex, wide and smooth area just before apex reborded and depressed, depression exhibiting striae; two small depressions near anterior angles and one spot of white tomentum on each side.
Underside. With scattered lunulate setigerous sculpture, setae longer on sternum than on abdomen; wide crescent punctures on metasternum, disc poorly sculpted (few fine punctures), denser to confluent laterally; abdomen with horseshoe sculpture regularly distributed; posterior coxae reborded laterally, latero-posterior angles well marked, setigerous sculpture of transversal to backward-curved striae; mesosternal apophysis transverse with few setigerous punctures, compressed between the mesocoxae and not protruding; male abdomen concave with visible groove on the sternites 3–5; two small lateral spots on sternite 6 in male, absent in female.
Legs. Exhibiting whitish double setae, one long and simple, second scale-type; protibiae bidentate, meso- and metatibiae with carina in apical third; profemora strigillate, mesofemora with crescent punctures to small striae, long setigerous stria along internal margin; metafemora slightly dilated, with crescent punctures to small striae; first tarsal segment shorter than others, metatarsi not spiny; claws normal.
Aedeagus. Simple, with sides converging in front; apex rounded and slightly protruding at center, very short longitudinal incision just in front of protrusion.
This species is only known from the type series (male holotype and two female paratypes) and is apparently restricted to Somalia. Females exhibit a convex abdomen and enlarged to spatuliform metatibial spurs, while male spurs are slender and acute.
Atrichelaphinis deplanata Moser, 1907
Clypeus transverse, more or less upturned (this represents a very strong sexual dimorphic character in some species), with anterior angles at the level of the antennal insertion; anterior pronotal border from slightly tectiform (minimally in male) to tuberculate; protibiae bidentate, with denticles normally separated; metatibial apical spurs enlarged or not in female; parameres of aedeagus fused, with apex sometimes sinuate or projecting laterally into hook-like expansion, but without frontal protrusion at middle and never reployed on ventral side, only curved downwards at apex.
The type of Atrichelaphinis deplanata was labelled by
Atrichelaphinis deplanata Moser, 1907: 316–317;
Atrichelaphinis deplanata (Moser, 1908) Touroult & Le Gall, 2001: 34.
Atrichelaphinis deplanate (Moser) Antoine, 1988: 48.
Atrichelaphinis kwangensis (Burgeon) Marais & Holm, 1992: 11.
Anelaphinis deplanata (Moser) Antoine, 1991: 2;
Anelaphinis kwangensis Burgeon, 1931: 221–222;
A. deplanata, holotype in MNHU: "Dahomey"; A. kwangensis, holotype in MRAC: "Musée du Congo, Kwango V-1927, (D? Zoballo), Don R. Mayné".
(n > 30). Size: length ♂, 9.6–13.3 mm; ♀, 9.8–12.4 mm; width ♂, 5.6–6.9 mm; ♀, 6.2–7.0 mm.
Body. Dorsally velutinous, background colour from light-yellow to light-brown, with many black/dark brown markings and small white maculae; scale pilosity mainly on ventral suface, more extensive in male than in female, particularly dense on antero-lateral borders of pronotum, on mesepimerons and legs.
Head. Clypeus transverse, almost bilobed in front, anterior margin reborded, anterior angles rounded, lateral angle visible from above, large and dense simple punctures on disc, laterally wrinkled; small white maculae scattered throughout dorsal surface, scale pilosity laterally behind eyes; antennae concolor, with clubs slightly longer in male than in female.
Pronotum. Exhibiting strong development of black markings, reducing the background colour to margins in some specimens; octagonal, anterior margin from straight to slightly tectiform, disc bulbous in front and without punctures; lateral margin almost completely reborded, with posterior half straight from subparallel to convergent, lateral angles rounded but visible, posterior angles rounded; posterior margin straight to weakly convex laterally, medial part strongly emarginate in front of scutellum.
Scutellum. With apex from weakly rounded to acute, lateral margins from straight to weakly concave and with lateral grooves.
Elytra. Usually showing transverse area lighter than base and apical parts, which exhibit more black marks; tricostate, with the second costa raised only in basal half; three pairs of geminate striae, sculpture of horseshoe-like punctures diverging at basal part of each stria, becoming confluent and geminate on upper half; apico-sutural angle acute, longer in male than in female.
Pygidium. Light brown with black markings, with horseshoe punctures and dense scale pilosity; exhibiting many depressed areas.
Underside. Brown and black with white maculae; scale pilosity dense on lateral parts of sternum which are striated; abdominal pilosity thinner and reduced to lateral sides where punctuation consists of few horseshoe setigerous puncture; middle of sternum and abdomen without punctuation, only longitudinal line visible on metasternum, abdomen weakly concave in male, convex in female; mesosternal apophysis transverse, compressed and not protruding between mesocoxae, metasternal declivity with scale pilosity.
Legs. Light brown, with scale pilosity, metafemora widened, metatibia short, thickened at apex, tarsi unmodified and normal; latero-posterior angle of metacoxae rounded; metatibial spurs thin and acute in male, slightly thickened and acute in female.
Aedeagus. Parameres narrowing gently towards apex, more abruptly close to apex; apex truncate and curved downwards, apical curved part from bilobed to incised (in frontal view).
Through courtesy of the MNHU and the MRAC, an opportunity was provided to study both types of A. deplanata (Figure
Holotype male, Zambia: Central Province, Mfwanta, S13°07'247", E30°19'345", 1429 m, R. Minetti leg., XI-2010 (MNHN ). Paratypes, Angola: Huila Prov., 2 km S Negola, S14°08'53", E14°28'16", à vue S. Rojkoff rec., 9-XII-2012, 1♀ (PCSR). Congo-Brazzaville: Pool, Mabaya, Bruno Le Rü leg, III-1989, 1♂ (PCDC). Congo-Kinshasa: Katanga, Manika, Ch. Seydel leg, C 19101, X-1931, 1♀ (MRAC); Kafakumba, F.G. Overlaet, IV-1932, 1♂ (MRAC); Lualaba, Zilo, Dr. V. Allard leg., XI-1974, 1♂ (MNHN Coll. Ruter); Lulua, Kapanga, F.G. Overlaet, IX-1933, 1♀ (MRAC); Katanga, exploration du PNU, riv. Kapelo, Miss. Hasson & Bouyer, Projet ICCN-NA-SEA, PNU082A, 10/16-XI-2002, 1♂ (CPTB); Katanga, exploration du PNU, env. Lusinga, Miss. Hasson & Bouyer, Projet ICCN-NA-SEA, PNU063, 25-X/5-XI-2002, 1♀ (CPTB). Malawi: West, Dzelanyama Fst., 4200 ft, 25-II-1985, 1♂ (PCTG); Mzuzu, Nhorongoro, S11°29’ E33°59’, 1375 m, R.J. Murphy leg, 26-XII-1996, 1♀ (PCTG); same locality, 4500 ft, R.J. Murphy leg, 30-XII-1996, 1♂ (PCTG). Mozambique: Sierra de Choa, D. Camiade leg, XI/XII-2012, 1♀ (PCDC). Rwanda: Rinkwavu, J. Roggeman leg, VI-1970, 1♂ (MRAC); Rwinkwavu, Montfort leg, IV-1976, 2♂ (IRSN, Coll. Alexis I.G. 30 374); Mayaga, J. Roggeman leg, VI-1972, 3♀ (MRAC); Kigali, Dr. V. Allard leg, II-1971, 1♂, 1♀ (MNHN, Coll. Ruter); Nyarubuye (Kibinga), Dr. V. Allard leg, 5-XII-1972, 1♂? (abdomen absent), 2♀ (MNHN, Coll. Ruter). South Africa: Afriq. Austr., Linokana, Dr. E. Holubi, 1894 (170–357), 1♀ (MNHN); Transvaal, ex. Coll. Oberthür, 2♂ (MNHN); Pretoria N., Van Son G., II-1936, Transvaal Mus. don, 1♀ (MRAC). Zambia: same data as holotype, 1♀ (PCSR); Central Province, 50 km E Serenje, S. Rojkoff & K. Werner leg, 7/8-XII-2005, 1♂ (PCSR); SE Lusaka, S15°33'662", E28°30'646", 1281 m, in fruit-baited trap, J. Touroult leg, 22-XI-2006, 1 ♀ (PCJT). Zimbabwe: Rhodésie du Sud, Selukwe, A. Ellenberger 1915, 1♂ (MNHN), 1♀ (PCSR).
(n = 31). Size: length ♂, 9.1–11.5 mm; ♀, 10.2–11.5 mm; width ♂, 5.2–6.4 mm; ♀, 5.7–6.5 mm. This new subspecies differs from the nominal form by exhibiting the following characters: smaller size; black/brown markings more regularly disposed and reduced; background colour more reddish; anterior pronotal elevation more enhanced; lateral pronotal angles less round and hind part of lateral border slightly longer; antescutellar concavity weak; pilosity of sternum thinner, especially in male; parameres with lateral sides subparallel, apex with dull lateral angles, shape more squared.
This subspecies is named after the French collector Robert Minetti, who brought to the authors’ attention the holotype specimen from Zambia.
There was initially some reservation in erecting this new subspecies, despite the morphological differences with the other forms mentioned above. Only the study of a large series of A. (E.) deplanata from various localities made it possible to isolate the new subspecies, considering also its broad geographic distribution. It is here given subspecies status because some of the specimens from Rwanda and Kenya actually represent a transition between the two forms, exhibitng intermediate characteristics such as coloration, shapes of pronotum and aedeagus. However, no potential intermediate forms were available from Congo-Brazzaville, where both subspecies are known to occur, but in separate parts of the country. Despite the Rwanda-Kenya transition zone, the new subspecies has a separate geographical distribution area from the nominal subspecies, which is restricted to western and central Africa. The new subspecies is distributed from central to east Africa and throughout the eastern half of southern Africa.
Niphetophora rhodesiana Péringuey, 1907: 451;
Anelaphinis rhodesiana (Péringuey) Antoine & Holm, 1993: 102.
Holotype male: "S. Rhodesia, Umtali " (ISAM).
(n = 42). Size: length ♂, 10.2–12.6 mm; ♀, 10.4–12.2 mm; width ♂, 5.7–6.9 mm; ♀, 57–7.0 mm.
Body. Light brown mottle with dark marks from green to brown, dark color at times covering virtually entire surface; matt to shiny, white spots of tomentum scattered throughout; light pilisoty distributed on vertex, along lateral margins of pronotum, on mesepimeron, on elytra (mainly on sides and apex) and pygidium.
Head. Clypeus slightly transverse, anterior margin strongly upturned in male, reborded and slightly bilobed in female; disc convex; punctures scattered and superficial, striolated laterally and in front.
Pronotum. Transverse, lateral angles strongly rounded almost imperceptible to slightly discernible; lateral margin completely reborded; posterior margin concave in front of scutellum, laterally convex; anterior margin bluntly tuberculate at middle; punctuation sparse on disc, becoming denser and stronger laterally and in front.
Scutellum. With short setae and occasional round puctures at base; apex acute.
Elytra. With two pairs of striae between sutural costae; discolateral costae with lunulate punctures more or less complete and confluent, horseshoe sculpture also on lateral margins; apicosutural angle acute and more or less developed.
Pygidium. Parabolic with upturned posterior margin.
Underside. Shiny, generally with spots of white tomentum on abdomen and metasternum, sometimes also on metafemora; mesosternal apophysis transverse, compressed between the mesocoxae, anterior margin slightly convex; median part of metasternum and abdomen without pilosity and less sculpted.
Legs. Protibiae tri- to unidentate; meso and metatibae with tranverse carina under middle of external side slightly enlarged; metalegs more robust in female; second metatarsomere longer than third and fourth; with setae longer than in any other area; metatibial spurs thin and acute in male, slightly enlarged and blunt in female.
Aedeagus. Parameres (Figure
A large number of specimens from Zimbabwe and South Africa was analysed for this sudy (in IRSN, MNHN, MNHU, PCRP, PCSR). The South African distribution of the species is restricted to the eastern, wetter part of the country (
Holotype male: South Africa EC, Hofmeyr, 28-XI-2011, R. Perissinotto & L. Clennell leg (ISAM). Paratypes: 1♂, same data as holotype (ISAM, PCRP); 1♂, same data as holotype, but 10-XII-2011 (PCRP); 8♂ 3♀, same data as holotype, but 24-XII-2011 (TMSA, PCSR); 2♂ 1♀, same data as holotype, but 26-XII-2011 (PCRP); 8♀, same data as holotype, but 18-XII-2010 (PCRP, PCSR).
(n = 25). Size: length ♂, 9.4–11.7 mm; ♀, 10.1–12.8 mm; width ♂, 5.8–6.7 mm; ♀, 6.2 to 7.8 mm.
Body. Dorsal surface slightly shiny, ground colour from ochraceous to light-brown, with many black/dark brown markings and small white maculae; scale-like setae present and particularly well developed on pronotum, more extensive in male than in female.
Head. Anterior margin of clypeus sharply upturned, particularly in male, sligthly bilobed apically, anterior angles weakly rounded, lateral declivity visible from above; large crescent to horseshoe punctures, particularly dense on frons and vertex; scale-type setae particularly long and dense from frons to vertex; antennae with pedicel and flagellum reddish-brown, but clubs dark brown to black, club notably longer in male than in female.
Pronotum. With black markings not covering more than half of total area and particularly developed on anterior part of disc, on both sides of medial line; anterior margin tectiform; disc moderately tuberculate in front; with scale-like setae and round punctures diffuse but widespread thoughout surface, setae more dense and longer on lateral margins; lateral margins and angles smoothly rounded with ante-scutellar arch relatively straight.
Scutellum. With apex from weakly rounded to acute; lateral margins from straight to weakly concave, with shallow and narrow lateral grooves; prominent oblong medial black mark extending from base to middle of disc; exhibiting few fine punctures on apical half but no scale-like setae.
Elytra. Weakly tricostate, with costae barely visible in apical part; sutural costa bulging out towards middle of elytral length; striae partly geminate and with coarse horseshoe sculpture; black marking most developed around humeral and apical calluses and in mid area of lateral half; apical sutural end virtually straight in male but curving outwards in female.
Pygidium. Brown to reddish at centre, becoming dark brown to black towards lateral and lower margins; fine sculture and dense cover of scale-like setae throughout surface; exhibiting 2–3 pairs of depressed areas close to lateral margins.
Underside. Dark brown to black with white scattered maculae, particularly on metasternum and lateral margins of abdominal sternites; densely covered with long white setae, replaced in mid area of metasternum and abdominal sternites by few scattered scale-like setae; coarse and scattered horseshoe sculpture throughout, except on central areas of sternum and abdominal sternites; abdominal sternites weakly concave at middle in male, slightly convex in female; mesosternal apophysis ochraceous, small and rounded, with no projections extending between mesocoxae.
Legs. Tibia and femora light brown, with dark brown to black tips and joints; scattered white maculae present on both dorsal and ventral sides; tarsi dark brown to black; protibia unidentate but broadening remarkably towards apex, forming spade-like structure; numerous long setae throughout surface and scale-like setae at joints; metatibial spurs thin and acute in male, slightly enlarged and blunt in female.
Aedeagus. Parameres virtually straight from base to apical convergence, forming then a perfectly round apex, with slight indent at centre (dorsal view); apical margin curving downwards, but no ventral folding or projections visible in lateral view.
The species is named after the Bamboesberg mountain range of the Eastern Cape Province of South Africa, where it was discovered on its south-western slopes.
This new species represents the southernmost extension of the genus distribution range in the Afrotropical Region. Atrichelaphinis bamboesbergica appears to be restricted to a small area of the eastern Karoo semiarid region, where its larval stages develop exclusively in the dung middens of the antbear, Orycteropus afer (Pallas, 1766). Adults have a relatively short life span (2–3 weeks) and appear to be unable to feed, as none has yet been observed either on fruits, flowers or sap flows.
Holotype male: Tanzania, Mtandi Masasi reg., 19-III-2008 (IRSN). Paratypes: Tanzania, 2♂ 2♀, same data as HT (PCTG, PCSR, PCRP); 1♂ 1♀, same locality, but I-2006 (PCTG); 2♀, Morogoro reg., UIuguru Mts, M. Coache leg, IV-2006 (PCSR); Zimbabwe, 1♂, Rhod., Christon Bank, Dr. V. Allard don, 25-XI-1974 (MNHN).
(n = 10). Size: length ♂, 10.7–13.6 mm; ♀, 10.7–12.9 mm; width ♂, 6.1–7.4 mm; ♀, 6.2–7.4 mm.
Body. Light brown mottle with dark marks from green to brown, dark color at times covering virtually entire surface; matt to shiny, white spots of tomentum scattered throughout; light pilisoty distributed on vertex, along lateral margins of pronotum, on mesepimeron, on elytra (mainly lateral margins and apex) and pygidium.
Head. Clypeus slightly transverse, anterior margin strongly upturned in male, reborded and slightly bilobed and upturned in female; disc convex; sculpture scattered and superficial becoming striolated laterally and in front, few setae on frons and vertex.
Pronotum. Transverse, lateral angles strongly rounded and from almost imperceptible to slightly discernible; lateral margin completely reborded; posterior margin concave in front of scutellum, laterally convex; anterior margin tuberculate at middle; punctuation sparse on disc, becoming denser and stronger laterally and in front; pilosity present on lateral and frontal margins.
Scutellum. With very thin and short pilosity, occasional round puctures at base; apex acute.
Elytra. With two pairs of striae between sutural costae; discolateral costae with horseshoe sculpture more or less complete and confluent, horseshoe sculpture also on lateral margins; apicosutural angle acute and moderately developed.
Pygidium. Parabolic, with upturned posterior margin.
Underside. Shiny, generally with spots of white tomentum on abdomen and metasternum, sometimes also on metafemora; mesosternal apophysis transverse, compressed between the mesocoxae and with anterior margin slightly convex; median part of metaseternum and abdomen without pilosity and poorly sculpted.
Legs. Protibiae from bi- to tridentate (third tooth sometimes drastically reduced); meso and metatibae with tranverse carina under middle of external side; metafemora and metatibiae strongly enlarged in both sexes; second meta-tarsomere longer than third and fourth; male metatibial spurs large and acute, especially upper one; protarsi (excluding claws) longer than protibiae (from joint to apex of apical tooth); metatarsi robust, especially in female; metatibial spurs slightly enlarged and blunt in male, strongly enlarged to spatuliform and blunt in female.
Aedeagus. Less than twice as long as wide; width at base larger or equal to width at apex; lateral sides of parameres parallel to convergent, with apical margins showing sinuosity and/or hook-like projections.
The species is dedicated to the renowned French collector Thierry Garnier, who continues to contribute greatly to the knowledge of African entomofauna and brought to our attention several specimens of the type series.
A. (E.) garnieri is very similar to A. (E.) rhodesiana, from which it can be separated by the shape of the aedeagus and some external differences. Its body is slightly larger and the dorsal black marking is also usually darker and more developed than in A. (E.) rhodesiana (except where forms of A. (E.) rhodesiana do not exhibit the typical colour pattern). The pilosity of A. (E.) garnieri is thinner and longer than that of A. (E.) rhodesiana, especially on the underside but more difficult to appreciate on the upperside due to wear. The pronotal tubercule is also more pronounced and larger in A. (E.) garnieri than in A. (E.) rhodesiana. The male metatibial spurs are larger and blunt in A. (E.) garnieri, especially the upper one, while in the female they are are spatuliform. Finally, apart form exhibiting sinuosity and/or hook-like projections, the mean ratio of length to width of the aedeagus of A. (E.) garnieri is 1.59 versus the 1.88 of A. (E.) rhodesiana. The two species appear to be cryptic and are sympatric in Zimbabwe, which represents the northernmost geographic limit of A. (E.) rhodesiana and the southernmost for A. (E.) garnieri. The two females from the Uluguru Mountains show less enlarged metatibial spurs and very sligth differences in the formation of subcoxite IX. However, it is likely that these constitute simple population variations.
Elaphinis simillima Ancey, 1883: 94–95;
Anelaphinis simillima (Ancey) Schenkling, 1921: 306;
Atrichelaphinis simillima Müller, 1939: 299.
Holotype not traced; described from "Abyssinie" (collected by Raffray, with no date but probably ca 1881).
(n > 350). Size: length ♂, 9.4–13.4 mm; ♀, 10.3–14.8 mm; width ♂, 5.5–7.8 mm ♀, 6.1–8.7 mm.
Body. Ligth brown, velutinous to shiny with dark markings never covering whole surface, always lighter areas present; mesepimeron with setigerous sculpture in both sexes.
Head. With metallic sheen; vertex velutinous and hairy, sometimes reaching clypeal disc; clypeus transverse, reborded and slightly bilobed in front, sometimes weakly upturned; sculpture dense and deep, simple on disc and more or less confluent in front and latetally; vertex with smooth area and tomentum, large and smooth longitudinal carina extending from vertex to clypeal disc which is convex.
Pronotum. Exhibiting metallic sheen and setae on lateral margins; with lateral angles usually broadly rounded, sometimes almost undistinguished; lateral margin completely reborded, posterior margin weakly concave in front of scutellum, laterally convex towards posterior angles; diffuse tomentose lines along lateral margins, sometimes very reduced, two radial lines on each side of midline usually made of three groups of spots more or less developed; sculpture marked, not dense on disc, more or less confluent laterally; dark green marks sometimes very reduced.
Scutellum. Longitudinal with apex from acute to blunt, without sculpture except near angles, without tomentum; with lateral grooves and sides almost straight or weakly curved inwards.
Elytra. With weak posthumeral emargination, reborded laterally; dark green marks sometimes very reduced or absent, but never covering whole surface; disc without tomentum; sculpture variable in size and intensity, usually vertical series of horseshoe punctures, sometimes confluent; short setae on lateral declivity and apex; apex acute but not protruding backwards.
Pygidium. With short setae occasionally throughout surface.
Underside. Shiny, with metallic sheen; with dense pilosity; mesosternal apophysis transverse, finely punctate and glabrous, not or slightly protruding in front of mesocoxae, not clearly oriented in lateral view; metasternum and abdomen sculpted laterally (horseshoe to striolate punctures) and showing white tomentum usually more developed in male; abdomen concave in male, flat or slightly convex in female.
Legs. Meso- and metatibiae with carina on external side just under middle; metatarsomeres shorter and more robust in female; protibiae enlarged, metatibiae slightly broader and hind spurs enlarged in female; metatibial spurs thin and acute in male, slightly enlarged and less acute but not blunt in female.
Aedeagus. Parameres about twice as long as wide, side from parallel to slightly convergent in front, apex truncate with lateral angles rounded, sometimes weakly bulbous laterally, emarginated at middle of downturning apical part.
All specimens examined originated from Ethiopia. Some are labeled "Shoa-Somali" but without precise locality and were collected during the expedition of V. Erlanger. They were probably also collected within the current borders of Ethiopia. The type of
Anelaphinis sternalis Moser, 1914: 606–607;
Holotype male: "Abessinien" (MNHU).
(n = 24). Size: length ♂, 12.1–13.7 mm; ♀, 12.9–13.5 mm; width ♂, 7.1–8.2 mm; ♀, 7.5–7.8 mm.
Body. Gound color from brown orange to brown red, with green marks more or less developed, at times covering whole dorsal surface with exception of few areas of ground color; velutinous, with metallic reflections as in A. (E.) simillima; tomentum and pilosity well developed and with almost same distribution as in A. (E.) simillima; mesepimeron mainly glabrous and without sculpture in male (sometimes with tomentum), with setigerous sculpture in female.
Head. Vertex velvety sometimes reaching clypeal disc; clypeus transverse, reborded and slightly bilobed in front; sculpture dense and strong, simple on disc and more or less confluent in front and laterally; vertex with smooth area and tomentum; large and very smooth vertical carina extending from vertex to clypeal disc, which is convex.
Pronotum. Transverse; lateral margins with very rounded lateral angles and regularly curved from posterior to anterior angles, reborded except in front of posterior angles; posterior margin strongly concave in front of scutellum, then bisinuate on each side; sculpture very light, sometimes undiscernible, scattered on disc but slightly denser near the anterior angles, punctuation stronger in female; tomentose line along outer margins and two radial lines of three spots each at side of midline, sometimes extra spots between these and outer ones.
Scutellum. Longitudinal, acute to blunt, usually smooth but with few punctures in some specimens; grooved laterally, sides almost straight.
Elytra. With weak posthumeral emargination, reborded laterally; disc without tomentum and with sculpture consisting of simple to crescent small punctures forming simple striae and interstriae; dense horseshoe sculpture laterally, near humeral callus and apically; sutural apex acute, slightly protruding backwards in male but not in female.
Pygidium. With large tomentose spots and bands.
Underside. Shiny, with large tomentose areas on prosternum, procoxae, mesepimeron, metepimeron, metepisternum, sides of metasternum and abdomen; mesosternal apophysis finely punctate, not transverse, almost as wide as long, protruding in front of mesocoxae and orientated downwards; abdomen slightly concave in male and convex in female.
Legs. Shiny; meso- and metatibiae with carina on external side just below middle; female with protibiae and metatibial spurs enlarged, metatibiae stronger, metatarsomeres shorter and more robust than in male; metatibial spurs very thin and acute in male, very slightly enlarged and less acute but not blunt in female.
Aedeagus. Length of parameres less than twice their width, sides converging in front, apex rounded, not truncate and not bulbous laterally, incised in the mid downturning part of apex.
This species is currently only known from Ethiopia. It is very close to A. (E.) simillima from which it can be separated through the sculpture of the dorsal side, the shape of the mesosternal apophysis and, to a lesser extent, the aedeagus. The A. (Eugeaphinis) species from Ethiopia are sometimes difficult to identify. For example, the Alexis Collection (IRSN) holds specimens from Lake Tana that exhibit a color pattern typical of A. (E.) sternalis; however upon close scrutiny they were found by the authors to resemble most closely A. (E.) simillima. However, the general body shape, the laterally bulbous apex of the parameres, the slightly more upturned anterior margin of the clypeus and the very weakly protruding mesosternal apophysis in front of the mesocaxae without downturning could cast some doubt over this identification. All the other characters are similar to those found in A. (E.) simillima. It is also possible that these specimens could represent either a new species, subspecies or just an hybrid between the two species. Another possibility is that of marked intraspecific variation. It may be necessary to study extensive series of specimens from more localities in order to resolve this issue conclusively.
Elaphinis vermiculata Fairmaire, 1894: 653–654;
Anelaphinis vermiculata (Fairmaire) Antoine, 2002: 186.
Atrichelaphinis vermiculata (Fairmaire) Kraatz, 1898: 220;
Holotype not traced.
(n = 11). Size: length ♂, 12.1–13 mm; ♀, 10.7–13.9 mm; width ♂, 6.7–7.6 mm ♀, 6.1–7.9 mm.
Body. Velutinous, from light brown with dark marks to dark green with dark brown areas, white small irregular spots scattered throughout, sometimes becoming confluent on lateral declivity of elytra, pronotum and pygidium; light pilosity usually present on vertex and lateral margins of pronotum, mesepimeron, elytral apex and pygidium; mesepimeron with sculpture and pilosity limited to anterior half, posterior half smooth.
Head. With median vertical smooth carina extending from vertex to clypeal disc; clypeus clearly transverse, anterior margin reborded in both sexes but not strongly raised, slightly incised in the middle, more strongly punctate laterally and in front, where punctuation becomes confluent; disc exhibiting smooth areas.
Pronotum. Octagonal; not tuberculate on anterior margin; lateral angles well marked but rounded; lateral margins completely reborded, with posterior half parallel; posterior angles rounded; sculpture of setigerous crescents, usually sparce and poorly pronounced on disc, but stronger in front and on lateral marginss; posterior margin from straight to slightly concave in front of scutellum, lateral edges convex.
Scutellum. Not uniformly sculpted, laterally grooved, and with white tomentum.
Elytra. With lateral margins almost straight and parallel, posthumeral emargination weak; sculpture of strong and well marked horseshoe punctures sometimes confluent, especially in apical half; space between vertical lines of punctuation of same width, appearing not geminate; sutural apex blunt.
Pygidium. With white, small spots becoming confluent; light pilosity throughout.
Underside. Shiny, with small white spots on postero-lateral angles of sternites and metasternum, sometimes on the mesepimeron and metepimeron, some apical spots also on femora; pilosity long and thin, extending throughout surface except middle of metasternum and abdomen; mesosternal apophysis transverse, anterior margin straight, strongly compressed between mesocoxae and not protruding in front in lateral view; abdomen concave in male, convex in female, last sternite less sculpted at middle in male.
Legs. Metafemora sometimes with white spot of tomentum on underside close to joint; protibiae enlarged in female, meso- and metatibiae with transverse carina just after middle; metatibial spurs slender and more acute in male, larger and blunt in female.
Aedeagus. Parameres converging regularly at apex, without lateral expansions or modifications; apex with marked incision at middle of downturning margin.
Most of the specimens analysed in this study originated from Erythrea (PCDC, PCSR, MNHN, MNHU). Although the type was not traced, all specimens match
Holotype male: Tanzania, Mbulu, Mamamisara, 2000 m, J. Kielland leg, (Bjørnstad 34728), 6-III-1981 (IRSN). Paratypes: Tanzania, 1♂, same data as HT (Bjørnstad 34727) (PCSR); 4♂ 1♀, Babati D., Mt. Kwaraha, 1850 m, J. Kielland leg, 30-IV-1987 (Bjørnstad 35080, PCTG; Bjørnstad 35077-35079 and 35081, PCSR, PCRP and IRSNB); 1♀, Ngorongoro Crater, 2200 m, J. Kielland leg, 14-II-1980, (Bjørnstad 35052, PCSR); 1♀ same data as above but 2300 m (Bjørnstad 41980, PCAB).
(n = 9). Size: length ♂, 13.6–14.6 mm; ♀, 12–15 mm; width ♂, 7.9–8.5 mm; ♀, 7.1–8.7 mm.
Body. Velutinous, brown with green to dark green marks, with small white spots scattered throughout, sometimes becoming confluent on lateral declivity of elytra, pronotum and pygidium; light pilosity distributed on vertex, lateral margins of pronotum, apical part of elytra and pygidium; mesepimeron with sculpture and pilosity on whole surface.
Head. Clypeus slightly transverse, almost as long as wide, anterior margin reborded and slightly incised at middle; disc convex, regularly punctated on entire surface, except few small smooth areas, punctures denser and confluent laterally and in front.
Pronotum. Not tuberculate in front, with round and slightly detectable lateral angles; posterior half of lateral margins not parallel but convergent in front; posterior angles blunt; posterior margin strongly concave in front of scutellum, with lateral edges almost straight; sculpture of setigerous crescent punctures, almost absent on disc, denser in front and laterally.
Scutellum. Unsculpted, laterally grooved, with white tomentum.
Elytra. With lateral margins almost straight and parallel, posthumeral emargination weak; sculpture of thin and incomplete horseshoe punctures more developed laterally and at apex, sometimes confluent resulting in broken lines; lines of punctuation geminate; sutural apex acute.
Pygidium. With small white spots scattered throughout, becoming confluent.
Underside. Shiny, with white confluent tomentum laterally on anterior margin of sternites and on lateral sides of metasternum; pilosity long and thin; mesosternal apophysis transverse, anterior border slightly convex, strongly compressed between mesocoxae and not protruding in front in lateral view; abdomen concave in male, convex in female; last sternite less sculpted at middle in male.
Legs. Metafemora sometimes with white spots of tomentum on underside close to joint; meso- and metatibiae with transverse carina just after middle; metatibial spurs thinner and more acute in male, larger and blunt in female.
Aedeagus. Parameres forming slight concavity at middle of lateral margins; without projections at apex, but with marked incision at middle of downturning frontal margin.
This species is named after the Norwegian entomologist Anders Bjørnstad, who provided the type series for study.
This species is most closely related to A. (E.) vermiculata, from which it can be separated mainly by the shape of the clypeus. It has also a distinct pronotum, with lateral margins strongly diverging in a posterior direction and the lateral angles obliterated, which also allow easy separation from A. (E.) vermiculata. Its elytra exhibit visible but relatively shallow sculpture. The species has so far only been recorded from northern Tanzania.
Protibiae tridentate, with two apical teeth close to each other | 2 | |
– | Protibiae bi- or tridentate, with two apical teeth normally separated | 3 |
2 | Pygidium with setae and small round sculpture; first two elytral striae consisting of double grooves (Figure |
A. (A.) tigrina (Olivier, 1789) |
– | Pygidium with scattered crescent sculpture, asetose; first two elytral striae consisting of single grooves (Figure |
A. (A.) nigropunctulata (Péringuey, 1896) |
3 | Dorsum black, sometimes with red parts or with yellowish elytra; clypeus longer than wide; aedeagus with protrusion at middle of apex in dorsal view | 4 |
Dorsum never black, usually ground color light brown or green; clypeus transverse; aedeagus simple at apex, occasionally with lateral projections | 6 | |
4 | Elytra yellowish; parameres medially and laterally protruding (Figure |
A. (H.) quadripunctata (Lansberge, 1882) |
– | Elytra black to dark brown; parameres protruding only medially | 5 |
5 | Dorsum entirely black without red areas, mesosternal apophysis large; medial protrusion of parameres incised but without meatus (Figure |
A. (H.) nigra Antoine, 2002 |
– | Dorsum with red areas (pronotum, scutellum, pygidium and last sternites); medial protrusion of parameres more developed, deeply incised and with large meatus (Figure |
A. (H.) sexualis (Schein, 1956) |
6 | Pronotum tuberculate at middle of anterior margin | 7 |
– | Pronotum not tuberculate | 9 |
7 | Clypeus upturned in both sexes (less in female), apical half of scutellum punctate to striolate (Figure |
A. (E.) bambooesbergica sp. n. |
Clypeus upturned only in male, simply reborded in female, scutellum without sculpture on apical half | 8 | |
8 | Protarsi longer than protibiae; metafemora and metatibiae strongly enlarged (Figure |
A. (E.) garnieri sp. n. |
– | Protarsi shorter than protibiae; metafemora and metatibiae slightly enlarged (Figure |
A. (E.) rhodesiana (Péringuey, 1907) |
9 | Mesosternal apophysis prominent between mesocoxae and projecting downwards (in lateral view), lateral margins of pronotum incompletely reborded near posterior angles (Figure |
A. (E.) sternalis (Moser, 1914) |
– | Mesosternal apophysis not prominent between mesocoxae | 10 |
10 | Apicosutural angle of elytra acute and projecting backward; species of small size (9–12 mm) | 11 |
– | Apicosutural angle of elytra not projecting backward; species of larger size (12 to 15 mm) | 12 |
11 | Pronotum predominantly dark-brown, with light colour and white tomentum restricted to margins; with posterior border clearly concave in front of scutellum; elytra with two large lateral light brown areas adjacent to metacoxae, basal and apical parts dark; parameres converging towards apex, then more abruptly near apex, apex truncate in front (Figures |
A. (E.) deplanata deplanata (Moser, 1907) |
– | Pronotum light in colour, dark markings reduced but white tomentose spots more widespread; posterior margin weakly concave in front of scutellum; elytra light brown with dark markings regularly distributed; parameres almost parallel towards apex, truncate in front forming blunt angles (Figure |
A. (E.) deplanata minettii subsp. n. |
12 | Elytral sculpture well developed, showing series of regularly-spaced horseshoe punctures (Figure |
A. (E.) vermiculata (Lansberge, 1882) |
– | Elytral sculpture faint and incomplete, with intervals between punctures irregular | 13 |
13 | Posterolateral angles of metacoxae from subacute to blunt; medium size species usually with metallic sheen, elytra light brown with few, small dark marks; tomentum mainly restricted laterally on pronotum and elytra (Figure |
A. (E.) simillima (Ancey, 1883) |
– | Posterolateral angles of metacoxae widely rounded; larger species without metallic sheen on dorsum; ground colour brown with large green patches and white tomentum scattered on entire surface (Figure |
A. (E.) bjornstadi sp. n. |
1 | Parameres completely fused, except for occasional presence of small sinuosity or incision on downturning apical margin (frontal view); parameres with or without projections | 2 |
– | Parameres partially fused, with apex incised or modified (dorsal view) | 5 |
2 | Internal sac of aedeagus with sclerites (Figure |
Heteralleucosma Antoine, 1989 |
– | Internal sac without sclerites | 3 |
3 | Protibia tridentate, with two apical teeth close to each other (Figures |
Atrichelaphinis (Atrichelaphinis) Kraatz, 1898 |
– | Protibia bi- or tridentate, with two apical teeth widely separated; mesosternal apophysis transverse and flat | 4 |
4 | Aedeagus with protrusion at middle of apex (dorsal view) (Figure |
Atrichelaphinis (Heterelaphinis) Antoine, 2002 |
– | Aedeagus simple at apex, but often exhibiting lateral projections (Figures |
Atrichelaphinis (Eugeaphinis) subgen. n. |
5 | Internal sac of aedeagus without sclerites | 6 |
– | Internal sac with sclerites | 11 |
6 | Parameres flat and composed of two weakly sclerotized lateral lobes, with median azygous sclerotized lamina (Figure |
Niphetophora Kraatz, 1883 |
– | Parameres not as above | 7 |
7 | Parameres flat | 8 |
– | Parameres visibly convex in lateral view | 9 |
8 | Parameres with small incised protrusion at middle of apex (Figure |
Atrichelaphinis (Heterelaphinis) Antoine, 2002 |
– | Parameres with apical incision exhibiting two lateral, slightly sclerotized triangular parts; anterior border of clypeus separated from disc by deep groove (Figure |
Paranelaphinis Antoine, 1988 |
9 | Apex of parameres with expansion visible in lateral view (Figure |
Molynoptera Kraatz, 1897 |
– | Apex of parameres not expanded | 10 |
10 | Apical end of parameres with sharp but thin hook visible in lateral view and protruding on ventral side (Figure |
Pseudalleucosma Antoine, 1989 |
– | Apical end of parameres without modifications visible in lateral view, round with setae on ventral side (Figure |
Molynopteroides Antoine, 1989 |
11 | Internal aedeagal sac with three sclerites; parameres flat, incised at middle of apex and slightly sclerified laterally at apex (Figure |
Phaneresthes Kraatz, 1894 |
– | Internal aedeagal sac with one or two sclerites | 12 |
12 | Sclerites composed of two bands; parameres almost flat, slightly thickened and curved in apical third from lateral view, with apex rounded and exhibiting small median incision (Figure |
Paralleucosma Antoine, 1989 |
– | Only one sclerite present | 13 |
13 | Sclerite consisting of thin, ovoid, longitudinal and erect band; parameres usually with cavity on upper side just before apex, apex more or less modified at extremity, setae on ventral side virtually sclerified (Figure |
Anelaphinis Kolbe, 1892 |
– | Sclerite not as above | 14 |
14 | Parameres subparallel, sharply narrowing before apex, sclerite with complex shape (Figure |
Alleucosma (Alleucosma) Schenkling, 1921 |
– | Parameres triangular, regularly narrowing from base to upturned apex; sclerite small, oval or flat but not with complex shape (Figure |
Alleucosma (Eoalleucosma) Antoine, 1989 |
Schematic outlines of parameres (a frontal; b lateral; c chitinous appendage). A Heteralleucosma insignis Antoine, 1989 B Niphetophora hildebrandti hildebrandti (Harold, 1878) C Paranelaphinis signata Antoine, 1988 D Molynoptera multiguttata Kraatz, 1897 E Pseudalleucosma machatschkei (Ruter, 1960) F Molynopteroides guttiventris (Moser, 1914) G Phaneresthes flavovariegata Kraatz, 1894 H Paralleucosma glycyphanoides glycyphanoides (Moser, 1908) IAlleucosma (Alleucosma) viridula (Kraatz, 1880) JAlleucosma (Eoalleucosma) duvivieri (van der Poll, 1890) (Figure C after
We are grateful to the following curators and collectors for kindly making available material and observations towards this study: Philippe Antoine (PCPA); Anders Bjørnstad (PCAB); Didier Camiade (PCDC); Joel Clary and Harold Labrique (CCEC); Marc De Meyer (MRAC); Alain Drumont (IRSN); Fabien Dupuis (St Chamond, France); Gilles Flutsch (Gometz-le-Chatel, France); Johannes Fritsch (MNHU); Philippe Moretto (Toulon, France); Thierry Garnier (PCTG); Denis Keith (PCDK); Jean-Philippe Legrand (Dammarie-sur-Loin, France); Petr Malec (Sumpek, Czech Republic); Antoine Mantilleri and Olivier Montreuil (MNHN); Daniel Moore (Dulles, VA, USA), Robert Minetti (PCRM); José Pénancier (Meudon, France); Julien Touroult (PCJT); Simon van Noort and Aisha Mayekiso (ISAM). We also thank Michael and Lesley Lord of Hofmeyr (South Africa) for assisting with field observations and logistic support. Lynette Clennell and Nelson Miranda are thanked for taking photos of various specimens. We are very grateful to Fabien Dupuis for his assistance with the description of female genitalia.