The Smicronychini of southern Africa (Coleoptera, Curculionidae): Review of the tribe and description of 12 new species

This study reviews the species of the tribe Smicronychini Seidlitz, 1891 found in southern Africa. In total, 18 species are recognized in this region, 12 of them being described as new (Sharpia madibai sp. nov., Afrosmicronyx cycnii sp. nov., A. louwi sp. nov., A. marshalli sp. nov., A. nebulosipennis sp. nov., Smicronyx pseudocoecus sp. nov., S. australis sp. nov., S. drakensbergensis sp. nov., S. gracilipes sp. nov., S. paucisquamis sp. nov., S. san sp. nov., S. similis sp. nov.). The following genera and species are newly reported from this area: Afrosmicronyx Hustache, 1935; Sharpia Tournier, 1873; Smicronyx pauperculus Wollaston, 1864; S. albosquamosus Wollaston, 1854 and S. namibicus Haran, 2018. New host plant taxa and plant associations for the tribe are reported among the families Gentianaceae Juss., Orobanchaceae Vent. and Convolvulaceae Juss. A key to species and images of the habitus of adults and male genitalia are provided.


Introduction
The weevil fauna of southern African is still unevenly known (Proches 2003). While large-bodied groups have received attention from early and more recent authors (i.e., Marshall 1946;Louw 1995), the smaller-bodied groups (Borovec & Skuhrovec 2018), in particular among the subfamily Curculioninae Latreille, 1802, have scarcely been studied (but see Caldara et al. 2009). As a result, the few taxonomic revisions of genera conducted in the smaller-bodied clades usually lead to the discovery of a large number of new species (Caldara 1989(Caldara , 1996. Following the revisions of several genera among the Afrotropical Smicronychini Seidlitz, 1 891 (Haran & Perrin 2017;Haran 2018) the diversity of this tribe from southern Africa was explored.
Very little was known about the Smicronychini from the southern African region (here considered below the 20 th parallel South). An initial species, Smicronyx fallax (Gyllenhal,1836) was described from "Caffraria", an area currently including broadly the Eastern Cape Province of the Republic of South Africa. More recently, a second species distributed at several places within tropical Africa (S. zonatus Haran, 2018) was described based on specimens from the Western Cape Province of the Republic of South Africa. Lastly, in recent years, attempts were made to introduce S. lutulentus Dietz, 1894 from North America to South Africa for biological control of the Asteraceous weed Parthenium hysterophorus Linnaeus, 1753 (Strathie et al. 2011).
Knowledge on the diversity of Smicronychini in southern Africa is needed to ensure that native and introduced species can be effectively distinguished, especially when the establishment of introduced species is tracked based on fi eld sampling. Due to their associations with parasitic and hemi-parasitic plants of crops in the Afrotropical region (genera Orobanche Linnaeus, 1753, Cuscuta Linnaeus, 1753, Striga Loureiro, 1790, Buchnera Linnaeus, 1753) the genera of this tribe are of potential interest for biocontrol. The aim of this study was therefore to revise the Smicronychini of southern Africa. About 400 specimens obtained from Natural History Museums and newly collected ones were examined using a standard taxonomic approach based on morphology and molecular diagnostic tools. A key, a description or a diagnosis of each species is provided with images of the habitus of adults and penis of males.

Preparation and photographs
The abdomens of adult specimens were extracted and digested in KOH to obtain clean preparations of genital structures, as these are commonly used to distinguish between species in Smicronychini (Zumpt 1936;Péricart 1957;Haran & Perrin 2017;Haran 2018). The illustration of male structures was restricted to the body of the penis, as it shows substantial variation between species of Smicronyx. The endophallic structures of male genitalia and external genitalia of the female (spermatheca and sternum VIII) are not described or illustrated, as examination of these structures did not reveal any signifi cant or phylogenetically informative differences between most of the species from the area examined. The habitus and male genitalia were photographed using a Keyence® VHX5000 imaging system. Extended depth-of-fi eld images were obtained using the software of the imaging station. Measurements were taken with an optical micrometer. Body length refers to the distance from the apical margin of the head (excluding the rostrum) to the apex of the elytra. Rostrum length refers to the distance between the apical margin of the eyes and the apex of mandibles. The ratio of width to length (w/l) was measured at the widest point of the prothorax, the elytra and the penis. The length of the elytra was measured between the anterior part of the scutellum and the apex of the elytra. The length of the penis was measured between the base of the penis body (apodemes excluded) and the apex. The terminology used follows Lyal (2019). Scales are important features for distinguishing between Smicronyx species, but they are fragile and may be partly lacking on some individuals. The diagnoses and descriptions in this study refer to fresh specimens with an entire vestiture. The identity of the species described before recent revisions was established using the original description (S. fallax and S. lutulentus), or examination of type material and reference collections (S. albosquamosus Wollaston, 1854 andS. pauperculus Wollaston, 1864). Neotype designation follows ICZN (1999) article 75. Data on each label of holotypes are reported between single quotation marks, each line of the label is separated by a slash, additional information to help label interpretation is given in square brackets.

Molecular analysis
A specimen per species was sequenced for the standard barcode fragment when fresh material could be obtained. DNA was extracted from entire specimens using a DNeasy Blood & Tissue kit (Qiagen, Hilden, Germany). PCR amplifi cations were carried out using the standard primers for barcoding (mitochondrial cytochrome c oxidase subunit I, COI) for invertebrates: LCO1490: 5'-GGTCAACAAATCATAAAGATATTGG-3' and HCO2198: 5'-TAAACTTCAGGGTGACCAAAAAATCA-3' (Folmer et al. 1994). PCR reactions were carried out on a Mastercycler® Nexus (Eppendorf, Hamburg, Germany) in a volume of 10 μl of PCR mix containing 5 μl of Multiplex Master Mix (Qiagen, Hilden, Germany), 0.8 μl of primers (Forward and Reverse at 2 μM) and 2 μl of DNA. The PCR conditions were as follows: initial DNA denaturation at 94°C for 15 minutes, followed by ten cycles of 30 s each at 94°C, 1 min at 60-50°C (touchdown of -1°C per cycle) and 30 s at 70°C, followed by 30 cycles of 30 s at 94°C, 1 min at 55°C, and 1 min at 72°C with a fi nal extension of 20 min at 72°C. The PCR products were sequenced by Eurofi ns Genomics (http://www.eurofi nsgenomics.eu/). All voucher specimens sequenced were mounted, dried and deposited at the Centre de Biologie pour la Gestion des Populations, Montpellier, France (CBGP, CIRAD collection https://doi.org/10.15454/D6XAKL). Barcode sequences were aligned using CodonCode Aligner ver. 3.7.1. (CodonCode Corporation,Centerville,MA,USA), and checked to identify the presence of pseudogenes using standard detection methods (Haran et al. 2015). Sequences were subjected to a BLAST (implemented in the NCBI platform https://www.ncbi.nlm.nih.gov/) to compare them to the existing sequences of species of Smicronyx Haran 2018). Uncorrected p-distance values of pairwise genetic distances between species were computed with Mega 7 (Kumar et al. 2016). HEAD. Rostrum as long as head + prothorax and regularly downcurved in lateral view, thicker than protibiae in dorsal view, densely punctuate and striate (♂), smooth with small punctures in apical half (♀), basal ⅔ (♂) or basal half (♀) of upper side bearing dense cover of pale brown scales, upper side of scrobes forming a carina slightly expanding basally near eyes; transverse furrow at base of rostrum narrow, shiny, bare of scales, with two spots of erect white scales on side near eyes; head capsule short, densely covered with scales; antennae inserted at apical ⅓ (♂) or near middle of length (♀) with segment 1 longer than 2 + 3, 4 isodiametric, segments 5-7 wider than long.  ABDOMEN. Underside covered with greyish scattered scales, 3 × as long as wide, with a metallic-copper shine; ventrite 1 shorter than 2.

Class
LEGS. Femora clavate, armed with one ventral tooth; tibiae straight, bearing a moderate thickening ventrally, sometimes almost forming a tooth on protibiae, facing femoral tooth when legs are folded; claws equal in length.
GENITALIA. Body of penis short (ratio w/l: 0.42), as long as apodemes, sides straight, regularly converging from base to apex, apex rounded, curvature in lateral view regular (Fig. 5B).

Sexual dimorphism
Females are distinguished from males by their rostrum, which is slightly longer, and more downcurved in lateral view.

Life history
All the specimens of this species were collected on Cycnium adonense E. Mey. Ex. Benth., 1835 (Orobanchaceae) a hemiparasitic plant parasite of grass. Adults were collected in November in the Republic of South Africa.

Distribution
Republic of South Africa (Mpumalanga Province).

Etymology
This species is dedicated to the late Dr. Schalk Louw, for his contribution to the knowledge of southern African weevils.

Description
Male BODY LENGTH. 2.5 mm.
COLOUR. Body integument black; vestiture consisting of rounded recumbent scales, as long as wide or slightly elliptical, brown and greyish on elytra.
HEAD. Rostrum as long as head + prothorax in lateral view, moderately and regularly downcurved, slightly wider than protibiae in dorsal view, punctate and striate, smooth at apex, covered with scales in basal ⅔, less densely in apical ⅓, upper side of scrobes forming a carina with a small tooth basally near eyes, transverse furrow at base of rostrum shiny, bare of scales; antennae black, inserted before apical ⅓, segment 1 slightly longer than 2 + 3, 4 isodiametric, 5-7 wider than long.
PROTHORAX. Wider than long (ratio w/l: 1.14), widest slightly before middle of length, greatly narrowed at apex, sides rounded; integument shiny, densely granulate; scales brownish and greyish, longer than wide, generally concealing granules, forming two paler bands laterally to the median line.
ABDOMEN. Underside covered with rounded brownish-grey scales, not concealing the integument.
LEGS. Dark brownish, femora clavate, armed with one ventral tooth; tibiae bearing a moderate thickening ventrally, facing femoral tooth when legs are folded; claws equal in length.

Life history
Unknown.

Etymology
Afrosmicronyx marshalli sp. nov. is dedicated to Sir Guy A. K. Marshall for his outstanding contribution to the knowledge on African weevils and particularly on the southern African fauna. COLOUR. Body integument black; vestiture consisting of rounded recumbent scales, as long as wide or elliptical, greyish on elytra.

REPUBLIC OF SOUTH
HEAD. Rostrum equal (♂) or slightly longer than head capsule plus prothorax (♀) in lateral view, moderately downcurved, in dorsal view covered with scales only in basal ⅓, shiny and slightly punctate in apical ⅔; antennae inserted slightly after middle of length; transverse furrow at base of rostrum shiny, with two spots of whitish semi-erect scales on sides near eyes; head capsule glabrous; eyes fl at, subcontiguous ventrally, distance between eyes equal to the width of 1-2 ommatidia; antennal scape slightly bisinuate at base, moderately clavate at apex, segment 1 of funicle elongate, as long as 2 + 3, 3-7 wider than long.
ELYTRA. Sides convex, widest near middle (ratio w/l: 0.7); humeri raised; striae narrow, interstriae fl at, wide and shiny, striae ⅓ width of interstriae, covered with two or three series of aligned scales, apex of interstria 5 with slight declivital callosity, followed by a depression; scutellum small but visible.
ABDOMEN. Underside covered with elliptical greyish scales, not concealing the integument.
LEGS. Integument black, femora clavate, armed with one ventral tooth; tibiae bearing a thickening (♂) or a tooth (♀) ventrally, facing femoral tooth when legs are folded; claws equal in length.

Life history
Unknown.

Etymology
Afrosmicronyx nebulosipennis sp. nov. is named in reference to the ill-defi ned shades of its elytral vestiture.
COLOUR. Body integument black; vestiture consisting of rounded recumbent scales, elliptical, twice as long as wide, forming grey, brown and/or white, ill-defi ned transverse bands on elytra.
HEAD. Rostrum distinctly longer than head capsule + prothorax in lateral view, downcurved at base, almost straight in the rest of the length, covered by scales on the upper face only in basal ½ (♂) or in basal ⅓ (♀); integument punctuate; antennae inserted at apical ⅓; transverse furrow at base of rostrum covered with whitish scales, with two spots of semi-erect whitish or brownish scales on sides near eyes; head capsule glabrous except near transverse furrow; eyes fl at, subcontiguous ventrally, distance between eyes equal or shorter than width of 1-2 ommatidia; antennal scape slightly bisinuate at base, moderately clavate at apex, segment 1 of funicle elongate, as long as 2 + 3, 3-5 isodiametric, 6-7 wider than long.
ELYTRA. Sides subparallel in basal half, widest near middle (ratio w/l: 0.74), convex in apical half; humeri raised; striae narrow, interstriae fl at, 4 × as wide as striae, integument shiny and rough, covered with two or three series of aligned scales, apex of interstria 5 with a declivital callosity, followed by a depression; base of interstria 3 with a spot of white scales; scutellum rounded, small but visible.
LEGS. Integument black, femora clavate, armed with a strong ventral tooth; tibiae bearing an acute tooth ventrally, before middle of length, facing femoral tooth when legs are folded; claws equal in length.
GENITALIA. Body of penis short (ratio w/l: 0.55), as long as apodemes, sides parallel, apex fl at with an acute cuticular expansion, curvature in lateral view moderate and regular (Fig. 5E).

Life history
Unknown.

Etymology
The species epithet refers to the remarkable long and thin legs encountered in this species. COLOUR. Body integument black, antennae dark brown; vestiture of elytra made of recumbent scales, elliptic, not overlapping, twice as long as wide, grey, aligned in 2-3 ill-defi ned series on each interstria; HEAD. Rostrum slightly longer than head capsule + prothorax in lateral view, moderately downcurved, in dorsal view covered with scarce scales in basal ⅓ and very short white setae in apical ⅔, punctate over entire length, smooth at apex; antennae inserted near middle of length; transverse furrow at base of rostrum shiny, with two spots of whitish erect scales on sides near eyes; head capsule glabrous, with slight cover of scales near transverse furrow; eyes fl at, subcontiguous ventrally, distance between eyes equal to the width of two ommatidia; antennal scape slightly bisinuate at base, moderately clavate at apex, segment 1 elongate, as long as 2 + 3, 4-5 isodiametric, 6-7 wider than long.

Holotype
PROTHORAX. Slightly wider than long (ratio w/l: 1.05), widest near middle, sides rounded, narrowed in apical ¼; integument smooth, shiny, with small punctures; scales brown, elongate, not concealing the integument, with grey scales forming a pale band along the median line and two on the sides of prothorax.
ABDOMEN. Underside mostly covered with non-overlapping greyish scales, twice as long as wide.
LEGS. Femora clavate, unarmed or bearing a very small ventral tooth in middle; tibiae straight, unarmed, claws equal in length. Males unknown.

Life history
Host plant unknown. The holotype specimen was collected on fl owers of an unidentifi ed Anacardiaceous shrub surrounded by a large grassland. This specimen was collected in December.

Distribution
Republic of South Africa (KwaZulu-Natal Province).  Table 1 Differential diagnosis Smicronyx similis sp. nov. is mostly similar to S. guineanus Voss, 1956, a species widely distributed in tropical Africa. The two species can be distinguished by their general appearance, which is more robust in S. similis sp. nov. The second segment of the funicle is isodiametric in S. similis sp. nov. while it is distinctly longer than wide in S. guineanus, the elytra are wider in S. similis sp. nov. than in S. guineanus (ratio w/l: 0.68 and 0.58 respectively), and the integument of the prothorax of S. similis sp. nov. is more coarsely punctate than in S. guineanus. The body of the penis in S. similis sp. nov. is also thicker and less narrowed in the middle than in S. guineanus. In southern Africa, S. similis sp. nov. is closest to S. gracilipes sp. nov., but the latter species has distinctly longer legs and rostrum and a fi nely punctate prothoracic integument (coarsely granulate in S. similis sp. nov.).

Etymology
The species name ʻsimilisʼ refers to the similar appearance of this species with S. guineanus, which is a very widespread and abundant species in tropical Africa. COLOUR. Body integument black, shiny, antennae and legs reddish; vestiture of elytra with scattered, recumbent, brownish and whitish, rounded or elliptic scales, forming two ill-defi ned series on each interstria.

Holotype
HEAD. Rostrum slightly longer than head capsule + prothorax, moderately downcurved in lateral view, slightly longer and more downcurved in ♀ than in ♂, covered with scarce whitish scales dorsally (♂) or scales only in basal ¼ (♀), shiny and slightly punctate at apex (♂) or at apical ⅔ (♀); antennae inserted slightly after middle of length (♀) or at apical ⅓ (♂); transverse furrow at base of rostrum shiny, with two spots of whitish semi-erect scales on sides near eyes; head capsule glabrous, with slight cover of scales near transverse furrow; eyes fl at, subcontiguous ventrally, distance between eyes equal to the width of 1-2 ommatidia; antennal scape slightly bisinuate at base, moderately clavate at apex, segment 1 of funicle elongate, slightly longer than 2 + 3, 4-7 wider than long.
ABDOMEN. Underside mostly covered with non-overlapping greyish scales, twice as long as wide.
LEGS. Femora clavate, armed with a small ventral tooth in middle; tibiae straight, unarmed; claws equal in length.

Life history
Unknown.

Distribution
Republic of South Africa (Northern Cape Province), Zimbabwe.  (Fig. 1H-I) and a more sclerifi ed body of the penis (Fig. 5G-H) than S. paucisquamis sp. nov..

Etymology
Named in reference to the similar appearance of this species and S. coecus (Reich, 1797).    COLOUR. Body integument entirely black, shiny, mostly bare of scales; vestiture of elytra consisting of white recumbent, elliptical, scales twice as long as wide, grouped in ill-defi ned transverse bands, a spot on humeri and a longitudinal strip at base of interstria 3; each interstria with series of recumbent, white setae.

Holotype
HEAD. Rostrum slightly longer than head + prothorax in lateral view, strongly downcurved in basal half, less downcurved in apical half (Fig. 3H), dorsal face punctate, covered with suberect whitish setae forming four distinct series in basal ⅔ and two spots near eyes, apical ⅓ glabrous; transverse furrow at base of rostrum shiny, bare of scales; head capsule glabrous; eyes fl at, contiguous on ventral side; scape of antennae straight, slightly and regularly widening toward apex, clavate at apical ⅓, segment 1 of funicle elongate, as long as 2 + 3, 4-7 × wider than long.
PROTHORAX. As long as wide (ratio w/l: 1), widest near middle of length, narrowed at apex, sides rounded; integument punctuate and granulous, punctures forming concentric circles at basal half; scales white, elongate, rare, mostly present along apical margin, and forming a small spot near scutellum.
ABDOMEN. Underside mostly covered with overlapping white scales, more condensed on metanepisterna.

Sexual dimorphism
Females are distinguished from males by their rostrum, which is more downcurved in apical half in lateral view, and by the antennal insertion which is near middle of length (near apical ⅓ in males).

Life history
Adults of S. pseudocoecus sp. nov. were collected on Cuscuta nitida E. Mey ex Choisy, 1842  and other unidentifi ed species of Cuscuta. Most Cuscuta sp. are red-listed endangered species in the Republic of South Africa. Adults were collected in January, March, September and November.

Etymology
Smicronyx paucisquamis sp. nov. is named in reference to the very limited number of scales found on its elytra compared to other species in the genus. COLOUR. Body integument entirely black, shiny, mostly bare of scales; vestiture of elytra consisting of white recumbent, elliptical, scales more than twice as long as wide, grouped into spots at base of interstria 3, basal ⅓ of interstriae 4-5 and middle of length of interstriae 3-5; each interstria with a series of short, recumbent, white setae.

REPUBLIC OF SOUTH
HEAD. Rostrum as long as head + prothorax in lateral view, downcurved near base, less downcurved in apical half (Fig. 4A); dorsal face punctate, covered with suberect whitish setae forming four distinct series in basal ⅔ and two spots near eyes, apical ⅓ glabrous; transverse furrow at base of rostrum shiny, bare of scales; head capsule glabrous; eyes fl at, sub-contiguous on ventral side, space between them wider than the diameter of two ommatidia; scape of antennae slightly bisinuate, regularly widening toward apex, clavate in apical ⅓, segment 1 of funicle elongate, as long as 2 + 3, 4-7 wider than long.
PROTHORAX. Isodiametric (ratio w/l: 1), widest near middle of length, narrowed at apex, sides rounded; integument with large punctures, diameter of punctures wider than distance between punctures; scales whitish, elongate, semi-erect, not concealing the integument, mostly present at base of median line near scutellum and on sides of prothorax.
ABDOMEN. Underside mostly covered with overlapping white scales, more condensed on metanepisterna.
LEGS. Covered with whitish, recumbent, elongate scales; femora clavate, armed with a very small ventral tooth usually concealed by scales; tibiae straight on external side, slightly bisinuate ventrally; claws equal in length.
GENITALIA. Body of penis elongate (ratio w/l: 0.44), as long as apodemes, sides sub-parallel, slightly expanding from base to apex, converging near apex, truncate at apex, curvature in lateral view moderate and regular, tapering near apex (Fig. 5H).

Sexual dimorphism
Females are distinguished from males by their rostrum, which is slightly longer, and covered with scales only in basal ⅓ (in basal ½ in males).

Life history
Adults were found in February on an unidentifi ed Cuscuta sp.

Differential diagnosis
Of the southern African Smicronychini, Smicronyx fallax is closely related to S. albosquamosus, which may be found in sympatry with this species. They can be distinguished by the shape of their prothorax (isodiametric in S. fallax, wider than long in S. albosquamosus) and their claws (equal in length in S. fallax, unequal in S. albosquamosus in most specimens). These species show an interspecifi c genetic distance of 12.3% for the barcode fragment.

Material examined
Neotype ( COLOUR. Body integument entirely black, shiny; vestiture of elytra consisting of whitish and brownish, recumbent, elliptical scales, twice as long as wide; whitish scales forming two ill-defi ned, oblique bands from humeri to apical ⅔ of elytral suture and a spot on declivital callosities. HEAD. Rostrum longer than head + prothorax in lateral view, almost straight (Fig. 4B), dorsal face punctate, covered with suberect whitish scales forming four distinct series in basal ⅔ and two spots near eyes, apical ⅓ bearing scattered white setae; transverse furrow at base of rostrum shiny, bare of scales; head capsule glabrous; eyes fl at, sub-contiguous on ventral side; scape of antennae slightly bisinuate, slightly and regularly widening toward apex, not clavate at apex, segment 1 of funicle elongate, longer than 2 + 3, 2 square, 3-7 wider than long.
PROTHORAX. Isodiametric (ratio w/l: 1), widest near middle of length, sides moderately rounded, not greatly narrowed at apex; integument punctuate and granulous, punctures forming weak concentric circles in basal ½; scales whitish and brownish, elongate, mostly condensed along the median line and on the sides.
ABDOMEN. Underside mostly covered with non-overlapping greyish scales, not condensed on metanepisterna.
LEGS. Covered with whitish, semi-erect, elongate scales; femora clavate, armed with a tiny ventral tooth, usually concealed with scales; tibiae straight on external side, slightly bisinuate ventrally; claws equal in length. Genitalia. Body of penis elongate (ratio w/l: 0.33), 1.3 × as long as apodemes, sides sub-parallel, broadly rounded at apex, curvature in lateral view moderate and regular, tapering at apex (Fig. 5I).

Sexual dimorphism
Females are distinguished from males by their rostrum, which is slightly longer, and covered with scales only in basal ¼ (in basal ⅓ in males).

Life history
All recently collected specimens of this species were found on dodders (Cuscuta spp.) including the introduced weed Cuscuta campestris Yunck, 1932 (Fig. 6A-B). Labels indicate that this species might be collected with light traps (UV light). Adults were collected from October to March.

Distribution
Smicronyx fallax is widely distributed in the subtropical and desert areas of southern Africa. It has been recorded from Namibia to Zimbabwe, and in the Gauteng, KwaZulu-Natal, Limpopo, Mpumalanga and Orange provinces of the Republic of South Africa.

Remarks
The type specimen of S. fallax was deposited by Gyllenhal in the collection of Ecklon & Zeyer (Gyllenhal 1836), currently lost. In the absence of further information about this type, and in order to stabilize the taxonomy in this genus, a neotype was designated based on the description of this species. Gyllenhal described S. fallax as a species very closely related to the Palaearctic taxon S. jungermanniae Reich, 1797. This description corresponded very well to the specimens used to re-describe this species, both regarding external and internal morphology. In addition, COI sequences obtained from these specimens (JHAR01630_0101-1694_0101 [GB accession: MT370316-17]) showed that they are closest to S. jungermanniae out of all the species of Smicronyx for which a barcode sequence is available (uncorrected p-distance of 7.0%). The neotype (here designated) was designated for a male specimen from a population from the western side of KwaZulu-Natal Province in the Republic of South Africa, which is close to the original type locality of S. fallax ("Caffraria", probably corresponding to the Eastern Cape Province). Three male and nine unsexed specimens stored in ethanol from this series were deposited in the institutions CBGP, SAMC and SANC.
Smicronyx australis sp. nov. urn:lsid:zoobank.org:act:11BB79AF-69BD-4596-B063-01C2E7782E04 Figs 1K, 4C, 5J; Table 1 Differential diagnosis Smicronyx australis sp. nov. is closest to S. pauperculus, a species widely distributed in the Mediterranean region and in Africa, including the Republic of South Africa. The two species can be distinguished by the colour of their integument (elytra black in S. australis sp. nov., at least partly red in S. pauperculus). The sequences of COI of S. australis sp. nov. (GB accession: MT370318) showed an interspecifi c p-distance of 3% with specimens of S. pauperculus from the Near East and Canary Islands (Genbank accessions: KU942318 and KU942313, respectively).

Etymology
Smicronyx australis sp. nov. is named in reference to the southern distribution of this species, compared to the wide distribution of its neighbour species S. pauperculus. COLOUR. Body integument entirely black, shiny, femora and tibiae reddish in middle; vestiture of elytra consisting of whitish and brownish recumbent elliptical scales, more than twice as long as wide; whitish scales forming ill-defi ned transverse bands and a spot on prothorax and elytra near scutellum.

Holotype
HEAD. Rostrum as long as head capsule + prothorax in lateral view, moderately downcurved (Fig. 4C); dorsal face weakly punctate, covered with suberect whitish scales forming 4 more-or-less distinct series in basal ⅔ (♂) or ⅓ (♀), more condensed near eyes, apical ⅓ bearing scattered white setae; transverse furrow at base of rostrum shiny, bare of scales; head capsule glabrous; eyes fl at, sub-contiguous on ventral side, separated by a distance as wide as the width of 1 ommatidium; scape of antennae very slightly bisinuate, regularly widening toward apex, clavate at apex, segment 1 of funicle elongate, slightly longer than 2 + 3, segments 2-7 transverse, segment 7 more than twice as wide as long.
ABDOMEN. Underside mostly covered with non-overlapping whitish scales, not condensed on metanepisterna.
LEGS. Covered with whitish semi-erect elongate scales; femora clavate, bearing a minute ventral tooth usually hidden by scales; tibiae straight on external side, slightly bisinuate ventrally at basal third; claws equal in length.
GENITALIA. Body of penis short (ratio w/l: 0.60), 0.6 × as long as apodemes, sides sub-parallel or slightly expanding apicad, apex truncate, curvature in lateral view moderate and regular (Fig. 5J); endophallus sclerifi ed, forming longitudinal striae at base of body in dorsal view.

Sexual dimorphism
Females are distinguished from males by their rostrum, which is slightly longer, and covered with scales only in basal ⅓ (in basal ⅔ in males).

Life history
This species was collected on stems and fl owers of Cuscuta nitida (Fig. 6D) and other Cuscuta spp. Adult specimens were also collected by light trap and by beating fynbos bushes. Adult collection times ranged from August to February.

Distribution
This species is distributed in the Gauteng and Western Cape provinces of the Republic of South Africa.

Remarks
Slight differences in the external and internal morphology and the genetic distances between these species enabled S. australis sp. nov. to be distinguished as a distinct species. The black specimens of S. pauperculus reported previously (Haran 2018) correspond to S. australis sp. nov. Wollaston, 1864 Figs 1L, 5K; Table 1 Smicronyx pauperculus Wollaston, 1864: 317.

Differential diagnosis
Of the southern African Smicronychini, S. pauperculus can be distinguished by the red colour of integuments and the absence of a wide transverse band on its elytra.
HEAD. Rostrum moderately and regularly downcurved in lateral view, longer than head capsule + prothorax.
LEGS. Femora moderately clavate, armed with a small but distinct ventral tooth, tibiae straight; claws equal in length.
GENITALIA. Body of penis moderately elongate (w/l ratio: 0.60), 0.6 × as long as apodemes, sides straight, subparallel, apex truncate, moderately and regularly curved in lateral view (Fig. 5K); endophallus slightly sclerifi ed, not forming any longitudinal striae at base of body in dorsal view.

Life history
This species is known to feed on fl owers of Cuscuta campestris (Anderson 1974; developing on plants of open and disturbed agricultural areas. It is often recorded by light trapping. In sub-Saharan Africa, adults were collected almost all year round (February-May, July-September, November-December).

Distribution
Smicronyx pauperculus is widely distributed in the Mediterranean region (Caldara 2013;. In sub-Saharan Africa, it has been recorded in West (Mali) and East Africa (Kenya, Tanzania) (Haran 2018) and is newly reported here from southern Africa (Republic of South Africa [Gauteng and Limpopo and Mpumalanga provinces] and Namibia).

Remarks
The identity of this species was established based on the examination of the lectotype as reported by Haran (2018). In southern Africa, this species is closest to its sister species S. australis sp. nov.; see differential diagnosis and remark sections on that species for diagnostic traits.
Smicronyx san sp. nov. urn:lsid:zoobank.org:act:F652DB37-9106-483C-A89F-9F2375C52910 Figs 2A-C, 4D, 5L, 6E; Table 1 Differential diagnosis In the Republic of South Africa, S. san sp. nov. is closest to S. drakensbergensis sp. nov.; it can be distinguished from this species by the coating on the elytra (white and brown scales, grey in S. drakensbergensis sp. nov.), the ratio of its prothorax (isodiametric, slightly longer than wide in S. drakensbergensis sp. nov.), and its shorter penis body .

Etymology
This species is dedicated to the San people, hunter gatherers whose fi rst traces date back 44 000 years in southern Africa. They left remarkable marks of their presence along the coast in Western Cape Province, where S. san sp. nov. forms dense populations due to the local abundance of its host plants. COLOUR. Body integument entirely black, shiny; vestiture of elytra consisting of recumbent, elliptical, twice as long as wide, overlapping scales, pale brown with transverse shades of white scales, densely covering but not concealing the integument.
HEAD. Rostrum as long as head + prothorax in lateral view, moderately and regularly downcurved (Fig. 4D), in dorsal view punctate, covered with suberect brown scales in basal ⅓-½, glabrous in apical half; transverse furrow at base of rostrum shiny, bare of scales; head capsule glabrous; eyes fl at, close to each other on ventral side, separated by a distance equal to width of 2-3 ommatidia; scape of antennae straight, slightly and regularly widening apicad, clavate in apical ⅓, segment 1 of funicle elongate, as long as 2 + 3, 4-7 wider than long.
PROTHORAX. Isodiametric (ratio w/l: 1), widest near middle of length, narrowed at apex, sides rounded; integument greatly punctuate and granulous, shiny; scales pale brown, overlapping, usually concentrated on sides, with a spot of white scales near humeri; disc usually bare of scales or with narrow scales not concealing integument, forming a broad longitudinal strip where integument remains visible; median line sometimes bearing a narrow band of brown scales, generally visible at base near scutellum.
ABDOMEN. Underside mostly covered with overlapping pale brown scales, more condensed on metasternum and metanepisterna.
LEGS. Covered with pale brown semi-erect elongate scales; femora clavate, bearing a tiny ventral tooth usually concealed by scales; tibiae straight on external side, slightly bisinuate ventrally at basal third; claws equal in length.
GENITALIA. Body of penis very short (ratio w/l: 1), 0.8 × as long as apodemes, sides sub-parallel or slightly expanding from base to apex in basal ⅔, converging in apical ⅓, acuminate at apex, curvature in lateral view regular (Fig. 5L).

Sexual dimorphism
Females are distinguished from males by their rostrum, which is slightly longer, and less densely covered with scales in basal ½.

Distribution
Republic of South Africa (Western Cape Province).

Remarks
Smicronyx san sp. nov. belongs morphologically to the S. reichi (Gyllenhal, 1835) species group (Dieckmann 1990), which is associated with Gentianaceae, and shows a much shorter body of the penis than other Smicronyx species (Pericart 1957). S. san sp. nov. is a polymorphic species, specimens obtained from Orphium frustescens (Fig. 2B) are larger and have a less dense coating than specimens developing on Chironia (Fig. 2A). Specimens obtained from Sebaea have a more elongate body (Fig. 2C) than those on Chironia. However, the sequences of the gene COI revealed no genetic divergence between populations from each host plant.

Etymology
This species is named in reference to the mountain range where the type series was collected. Dietz, 1894 Figs 2F, 4F, 5O; Table 1 Smicronyx lutulentus Dietz, 1894: 170.

Differential diagnosis
Smicronyx lutulentus is morphologically similar to S. fallax and S. albosquamosus. It can be distinguished from these species by its elytra pattern, bearing broad and rounded scales, and its short penis body. COLOUR. Body integument black, rostrum black or dark brown, antennae and tarsi dark brown; vestiture of elytra generally consisting of rounded, grey-brown scales, isodiametric or slightly longer than wide, with a few scattered white scales; white scales concentrated on humeri, on sides and at base of median line near scutellum on prothorax.

Material examined
HEAD. Rostrum in lateral view downcurved at base and almost straight over the rest of length; slightly longer than head capsule + prothorax.

Life history
Smicronyx lutulentus develops in the seeds of the capsules of Parthenium hysterophorus L. (Asteraceae; Dhileepan et al. 1996).

Differential diagnosis
In southern Africa, S. albosquamosus is closest to S. fallax. See differential diagnosis section under this species for distinguishing traits.

Life history
Smicronyx albosquamosus is found across its distribution range on pods and fl owers of Cuscuta spp. Adults were collected in southern Africa from November to April.

Distribution
This species is widely distributed across Africa and the Mediterranean region. In the Republic of South Africa, this species is only found in subtropical areas (Gauteng, KwaZulu-Natal, Limpopo and Mpumalanga provinces)

Remark
The identity of this species was established by comparisons of specimens with reference material from the Mediterranean region reported by . Haran, 2018 Figs 2H, 5P; Table 1 Smicronyx namibicus Haran, 2018: 282.

Differential diagnosis
Of the African Smicronychini, Smicronyx namibicus can distinguished by the scales on elytra forming a contrasting black square on basal half of interstriae 1-4, surrounded with white scales (Haran 2018). COLOUR. Body integument black, antennae, femora and tibiae dark reddish; vestiture of elytra generally consisting of recumbent, rounded or slightly longer than wide scales, forming a dark-brown square in basal half and between interstriae 5 of each elytron; rest of scales on elytra greyish to brown, paler near dark-brown square; vestiture of prothorax consisting of brownish scales, with 5 longitudinal whitish stripes.

Material examined
HEAD. Rostrum slightly longer than head capsule + prothorax in lateral view, moderately downcurved, curvature stronger near base and near antennal insertion.
PROTHORAX. Slightly wider than long (w/l ratio: 1.06), widest before middle of length, sides greatly rounded in basal ⅔.

Life history
Unknown. The adults were collected in December in the Republic of South Africa and in November in Tanzania.

Remarks
Smicronyx namibicus was wrongly described from Namibia due to a misinterpretation of the handwritten label of the holotype. The locality of this specimen is 'Zanzib' (Island of Zanzibar, Tanzania) and not 'Namib' (Namib Desert) as reported in Haran (2018). I thank Helene Perrin (MNHN) for highlighting this misreading.

Discussion
This study enabled the recognition of 18 species of Smicronychini in southern Africa, of which 12 are described as new. The genera Sharpia and Afrosmicronyx are reported for the fi rst time from this area. Southern African Smicronychini contain some typical tropical African elements (Genus Afrosmicronyx; species allied to Smicronyx guineanus) in its northern and central parts (northeastern South Africa, Zimbabwe). It also includes some widely distributed elements across the whole of Africa and the Mediterranean area (S. albosquamosus, S. pauperculus). Several species endemic to southern Africa are morphologically and molecularly closely related to West Palearctic species: S. jungermanniae and S. fallax; S. pauperculus and S. australis sp. nov.; S. coecus and S. pseudocoecus sp. nov. These pairs of species suggest a somewhat recent contact zone and then an allopatric speciation between these regions, as has been shown in several groups of organisms, including in plants and weevils (Hernández-Vera et al. 2013;Pirie et al. 2016).
The plant associations reported in this study are consistent with the known host families of Smicronychini in the old world: Gentianaceae, Convolvulaceae and Orobanchaceae (Caldara et al. 2014;Dieckmann 1990). However, the following genera are newly reported: Cycnium (Orobanchaceae), Chironia, Orphium and Sebaea (Gentianaceae). Several species of Smicronyx were found on the introduced dodder Cuscuta campestris and can be considered as candidate species for its biocontrol. Apart from this species, most Cuscuta of southern Africa are rare and red-listed species that do not threaten crops.