Revision of the enigmatic South African Cryptolaryngini (Coleoptera, Curculionidae), with description of a new genus and twenty-two new species

The weevil genus Cryptolarynx Van Schalkwyk, 1966 is endemic to the Northern and Western Cape provinces of South Africa. The two previously known species of the genus, C. vitis (Marshall, 1957) and C. estriatus (Marshall, 1957), have an aberrant globular body and head shape, which has made it diffi cult to place the genus into the classifi cation systems of the Curculionoidea. This paper presents the description of 21 new species of Cryptolarynx from South Africa (C. subglaber Haran sp. nov., C. squamulatus Haran sp. nov., C. muellerae Haran sp. nov., C. hirtulus Haran sp. nov., C. robustus Haran sp. nov., C. namaquanus Haran sp. nov., C. carinatus Haran sp. nov., C. variabilis Haran sp. nov., C. pyrophilus Haran sp. nov., C. pilipes Haran sp. nov., C. armatus Haran sp. nov., C. falciformis Haran sp. nov., C. oberprieleri Haran sp. nov., C. spinicornis Haran sp. nov., C. cederbergensis Haran sp. nov., C. homaroides Haran sp. nov., C. marshalli Haran sp. nov., C. endroedyi Haran sp. nov., C. oberlanderi Haran sp. nov., C. san Haran sp. nov., and C. luteipennis Haran sp. nov.) and of one new genus and species, Hadrocryptolarynx major Haran gen. et sp. nov., also from South Africa. A redescription of the European Journal of Taxonomy 877: 1–89 (2023) 2 genus Cryptolarynx is provided to incorporate the characters of the new species. The plant genus Oxalis (Oxalidaceae) is recorded as larval host for several species of Cryptolarynx and for Hadrocryptolarynx Haran gen. nov., as their larvae develop in the subterranean bulbs of members of the genus, and the egg, larva and pupa of C. variabilis are described. The characters of the Cryptolarynx larva confi rm that Cryptolaryngini are an early-diverging group of Curculionidae, with a placement among taxa currently classifi ed in the subfamily Brachycerinae sensu lato, and although their exact taxonomic position remains unresolved, some larval characters, and also pupal ones, suggest a close relationship between Cryptolaryngini and Stenopelmus Schoenherr. Potential use of species of Cryptolarynx in the biological control of weedy South African species of Oxalis is discussed.


Introduction
In 1957, G.A.K. Marshall published the description of Cryptopharynx, a new weevil genus from the Northern and Western Cape provinces of the Republic of South Africa, and recognised two new species in it, Cryptopharynx vitis and C. estriatus.  regarded the genus as aberrant in shape and characters and could not place it in any of the known weevil subfamilies because of its globular body shape, its head almost entirely retracted into the pronotum and its very short, muzzle-like rostrum undifferentiated from the head. He consequently erected a new subfamily of Curculionidae Latreille, 1802 for it, named Cryptopharynginae.  further recorded the adults of C. vitis feeding on grapevine leaves in Stellenbosch. A few years later, Van Schalkwyk (1966) changed the name of the genus and subfamily to Cryptolarynx and Cryptolarynginae, respectively, as the name Cryptopharynx is preoccupied in Protozoa and the subfamily name therefore needed to be emended as well (Art. 39 of the ICZN).
Palaearctic Coleoptera. In contrast, Kuschel (1995), Oberprieler et al. (2007) and Oberprieler (2014) placed Cryptolaryngini Van Schalkwyk, 1966 as a tribe of Brachycerinae Billberg, 1820, the latter treated as a subfamily of Curculionidae. Morrone (1997) and Bouchard et al. (2011) adopted a different classifi cation system again, placing Cryptolarynginae as a subfamily of Brachyceridae, but Alonso-Zarazaga et al. (2017), in an update of the Catalogue of Palaearctic Curculionoidea, accepted the Kuschel-Oberprieler classifi cation, which is the one in current usage, in particular as recent molecular phylogenetic analyses (e.g., Mugu et al. 2018;Shin et al. 2018;Song et al. 2020) have confi rmed the phylogenetic position of Brachycerinae as a basal lineage of Curculionidae vis-à-vis their sister group, the family Brentidae Billberg, 1820. Oberprieler et al. (2007) reported the existence of another 16 undescribed species of Cryptolarynx and adults having been found feeding on leaves of Moraea Mill. (Iridaceae) and provided a fi rst colour photograph of a live specimen, and Oberprieler (2014) presented a comprehensive account of the morphological characters of Cryptolaryngini and a summary of the taxonomy, diversity and biology of the tribe.
More than a century after the collection of the fi rst specimens , Cryptolarynx remained a mystery in many ways. Recent fi eld sampling conducted in South Africa greatly increased the number of species and provided new insights into the life history of the genus. Based on this material, the objectives of this study are to i) describe new taxa, ii) redescribe the genus Cryptolarynx to account for the characters of the newly described species, iii) provide a description of the larval and pupal stages of the genus and iv) assess the position of the tribe Cryptolaryngini in the Curculionoidea in the light of all new evidence. In all, one new genus and 22 new species are described. The immature stages and life history of Cryptolarynx are described for the fi rst time. Photographs of the habitus of the adult weevils and of the distinguishing characters of their male genitalia are presented. Mitochondrial DNA barcode and nuclear-gene sequences are provided for most species. The potential use of Cryptolarynx in the biological control of alien invasive species of Oxalis L. is briefl y discussed.

Sampling
Field collecting was conducted in the Republic of South Africa from early 2018 to late 2019 at the localities of the two previously known species of Cryptolarynx as well as at other localities in the Western Cape and Northern Cape provinces, for a total of about 40 sites. Opportunistic sampling for Cryptolaryngini was also undertaken in the Eastern Cape and KwaZulu-Natal provinces. Specimens were obtained by beating and sweeping vegetation with an insect net and by visually searching for them on the ground. Plants found near specimens were sampled and plant tissues (fl owers, fruits, stems, bulbs, root system) were dissected to search for immature stages. Plants were identifi ed using fi eld guides, and identifi cations were subsequently confi rmed by plant taxonomists. The specifi c epithets of Oxalis host plants follow Salter (1944) and Manning & Goldblatt (2008). Weevil larvae were reared in vials with a sample of plant tissue or with soil collected on site. All specimens were collected directly into 96% ethanol and stored at room temperature. Dry pinned specimens in four collections (see depositories below) were borrowed and studied.

Preparation and taxonomic treatment
The abdomens of males were detached from the body and macerated in a 10% potassium hydroxide (KOH) solution to obtain clean preparations of genital structures. The habitus of adults and genitalia were photographed with a Keyence ® VHX-5000 imaging system. Measurements were made with an ocular micrometer. Body length (L) of adults was measured as the distance from the pronotal apex to the elytral apex in dorsal view. Widths (W) were measured in dorsal view at the widest parts of the pronotum, the elytra and the body of the penis. Lengths were measured in dorsal view, along the median line, of the pronotum from apex to base, of the elytra from the anterior margin of the scutellar shield to the elytral apex, of the penis from the base to the apex of the body of the penis (excluding the temones) (Fig. 8D,J). The width of the eyes was measured in facial (anterior) view. The terminology for the external and internal sclerites and structures of adults used mainly follows Lyal (2021). Deviations include the following: 'forehead' refers to the area between the eyes (Fig. 8A) and 'epifrons' to the dorsal surface of the rostrum between the anterior margins of the eyes posteriorly and the level of the antennal insertions anteriorly (Fig. 8A). The South African Cryptolaryngini exhibit signifi cant intraspecifi c variation in body size and shape, in elytral colour patterns and in ratios of the dimensions of the antennal segments. Morphological characters that may distinguish species of Cryptolaryngini are mainly found on the ventrites and tibiae, in head ratios and in genital structures of the males. Females of most species exhibit few stable morphological interspecifi c differences, and females are therefore not considered in the species descriptions below.
About 500 specimens were examined for this study. Identifi cations of the previously described species were made by comparison with type specimens and study of the original descriptions. Label data of name-bearing types are reported verbatim, with consecutive lines separated by a slash and data from a single label enclosed by double quotation marks. Additional information and interpretations appear between square brackets. The format of the label data of other specimens is standardised.
Valuable additional bio-ecological information was obtained from the Coleoptera fi eld collection books of the Ditsong National Museum of Natural History (the former Transvaal Museum) in Pretoria (TMSA). The handwritten entries in these books are numbered with the prefi x 'E-Y', but they reference not only Sebastian Endrödy-Younga but all beetle collectors associated with the Ditsong Museum from 1973 to 2018. For approximately the three decades after about 1975, and sometimes until even more recently, South African insect collections (here SAMC, SANC and TMSA) formatted geographic coordinates according to the notation of the then Times Atlas of the World. As an example, the coordinates for Cape Town were accordingly rendered as "33.55S 18.25E". In recent years, an increasing number of authors treated these decimal-looking coordinates as decimal degrees, an error that may lead to considerable misinterpretation. The coordinates for Cape Town in degrees-minutes notation rightly are 33°55′ S, 18°25′ E and correspond to 33.92° S, 18.42° E in decimal-degree notation. We retained historic notations in label data citations of name-bearing types, but in all other cases we standardised such coordinates to the degrees-minutes notation.
Following Article 50.1 and Recommendation 50A of the Code (ICZN 1999), JMH is responsible for the new names introduced in this paper. The lectotype designation is in accordance with Article 74.1 (ICZN 1999).
Larvae and pupae used for morphological descriptions were preserved in 80% ethanol. Habitus photos of the larvae were taken with a JVC KY-F75U camera associated with a Leica MZ16f stereo microscope. The pupa was photographed with a Keyence ® VHX-5000 imaging system. Larvae were dissected and slide-mounted for microscope study under high magnifi cation, basically following the technique of May (1993May ( , 1994. The larval head was extracted and the body was cut into two longitudinal halves and around the terminal abdominal segments (except for the smallest specimen). The dissected parts were cleared in 10% KOH, rinsed in distilled water and transferred to ethanol. After clearing, the mouthparts, head capsule and body cuticles were mounted in Euparal on permanent microscope slides. Illustrations of larval structures were made with the aid of a drawing tube attached to a Leitz Laborlux S compound microscope. The pupae were examined directly in ethanol under the stereo microscope. Terminology used for larval characters follows Marvaldi (1999), as also employed by Oberprieler et al. (2014).
Terminology for pupal characters follows May (1994). Setal numbers of bilateral structures are given for one side only.

Molecular analysis
For each species, when available, one or several specimens were sequenced for the standard DNA barcode region for invertebrates (mitochondrial cytochrome c oxidase subunit I: COI, Hebert et al. 2003) to test the interspecifi c boundaries identifi ed from morphological examination. Because genetic divergences in maternally inherited mitochondrial DNA can be high in fl ightless insects and overestimate divergence between populations, we also sequenced the nuclear gene Elongation Factor 1 (EF1α-F2, EF1) to cross-validate interspecifi c genetic divergences. DNA was extracted from whole specimens or a leg, using a DNeasy Blood & Tissue Kit (QIAGEN, Hilden, Germany). PCR amplifi cation was carried out using a mix of primers for amplifi cation of COI, and the primers described in Brady et al. (2006) were used for the amplifi cation of EF1 (Appendix 1). PCR reactions were carried out in a Mastercycler ® Nexus (Eppendorf, Hamburg, Germany) in a fi nal volume of 10 μL containing 5 μl of Multiplex PCR Master Mix (QIAGEN, Hilden, Germany), 2 μM of each primer and 2 μl of DNA template. The PCR conditions were as follows: initial DNA denaturation at 94°C for 15 minutes, followed by 35 cycles of 30 s at 94°C, 1 min at 52°C and 1 min at 72°C, with a fi nal extension of 15 min at 72°C. The PCR products were paired-end sequenced by Eurofi ns Genomics (http://www.eurofi nsgenomics.eu). All voucher specimens were mounted, dried and deposited at CBGP, Montpellier, France, in the CIRAD collection (https://doi.org/10.15454/D6XAKL) or in the institutions listed in each Material examined section. Nucleotide sequences were aligned and manually checked using CodonCode Aligner ver. 3.7.1. (CodonCode Corporation,Centerville,MA,USA) to verify the absence of pseudogenes. Uncorrected p-distance values of pairwise genetic distances between species were computed with Mega 7 (Kumar et al. 2016). Reconstruction of phylogenetic trees was carried out with COI and EF1 sequences (658 and 517 base pairs, respectively) using PhyML (Guindon & Gascuel 2003) with 1000 bootstrap replicates. The trees were rooted with three species of two tribes of Brachycerinae sensu Oberprieler (2014), Brotheus crenelatus Marshall, 1907 (JHAR01250_0101), Synthocus hopei Boheman, 1842 (JHAR02578_0101) (both Brachycerini) and Ocladius baccicollis Boheman, 1838 (JHAR02406_0101) (Erirhinini), as well as with a species of the brentid genus Episus Billberg (JHAR02469_0101) (Microcerinae). These are all South African species. All sequences were deposited in GenBank (Appendix 2).

Abbreviations of depositories
The specimens on which this study was based are lodged in the following institutions and specimen collections. COLOUR AND VESTITURE. Body integument black; antennae, tibiae and tarsi generally reddish. Dorsal vestiture (pronotum + elytra) consisting of short, recumbent clothing scales, these isodiametric to 2 × as long as wide, not aligned on interstriae, more or less concealing integument, colour ranging through black, dark brown to pale brown and grey to white, orange to yellow in one species; darker scales usually concentrated medially on pronotum and from there in broad stripe on elytral interstriae 1-3; paler scales concentrated at sides of dorsum or pale areas reduced to strips laterally on pronotum and along elytral interstriae 4; elytra with a pair of pale spots on interstriae 2-3 at apical ⅔ of length, spots sometimes hardly discernible or confl uent to form a pale transverse band that may further merge with pale areas laterally; scales arising from strial punctures suberect in some species and then up to 4 × as long as wide.
HEAD. Head capsule globose, in repose deeply retracted into prothorax, leaving only vertex and often eyes visible in dorsal view. Eyes subcircular or slightly oval, usually only slightly convex, situated dorsally or sublaterally, surrounded by a ring of mostly pale recumbent scales. Forehead fl at, width ranging from less than to twice width of eye; fovea absent. Distance between eye and scrobe smaller than width of eye. Rostrum very short and broad, not differentiated from head; mandibles abutting anterior part of procoxae when head in repose. Epifrons short, width ¾ × to subequal to width of forehead, medially deeply longitudinally depressed, epifrontal scales recumbent or suberect and orientated towards mouthparts. Frons largely indistinguishable but set off from epistome by slight carina, with 1 or more pairs of long setae laterally. Epistome crescentic, anterior margin medially notched, sometimes medially with single elongate seta about half as long as frontal setae. Mandibles beak-like, paucisetose (4-8 setae on each), with 2 setae long and erect and the others shorter; without scales. Maxillae with galea and lacinia separate, both bearing apical setae. Antennae inserted subdorsally at approximately midlength of rostrum; scapes slender, as long as or longer than width of epifrons between antennal insertions, regularly curved, clavate at apex and bearing erect setae, in repose folding into narrow scrobes extending onto underside of rostrum; funicles 7-segmented, longer than scape, segment 1 longer than wide, longer than or as long as 2, 1-4 fl attened dorsoventrally, 2 and 4 or 2-4 angular or toothed ventrally, 5-7 globular or moderately elongate; funicles entirely hidden between head and pronotum when head in repose; clubs 4-segmented, fusiform, acuminate, shorter than funicle.
THORAX. Pronotum convex, at least moderately transverse, widest at base or near midlength, sides arcuate; integument fi nely and densely punctate, dull between punctures; anterior margin bisinuate, posterior margin more or less bisinuate, fi tting closely to elytral bases up to level of elytral humeri. Prothorax anteriorly on each side produced into a large sharp-rimmed ventrolateral lamina extending from lower level of eye down to anterior edge of procoxa, concealing anterior prothoracic margin beneath it, rim of lamina asetose but anterior margin fringed with row of dense plumose scales, longer below eyes but shorter ventrally along prosternum. Prosternum broad, very short, depressed below anterior edge of procoxae, declivous, abutting rostrum when head in repose; procoxal cavities medially confl uent, hypomeral lobes behind them short, suture of median junction faintly discernible. Mesoventrite deeply depressed, almost vertically declivous, intermesocoxal process subtuberculate; mesepimera narrowly triangular, fully separating mesanepisterna from elytral margin. Metaventrite narrower than width of metatarsus; metanepisterna fully fused to metaventrite, metanepisternal suture completely obliterated.
ELYTRA. Globular to broadly ovate, sides convex, widest near or anterior of midlength; jointly rounded at apex; elytral base broadly concave, not marginate; integument fl at, dull or shiny, 10-striate but striae generally indiscernible on outer surface, mixed with regular punctures and covered by scales.
ABDOMEN. Ventrites more or less concave medially, median impressions surrounded with cuticular ridges or not, surface more or less densely clothed with pale, recumbent scales, intermixed with suberect setae; ventrite 1 medially about twice as long as laterally, as long as or longer than each of ventrites 2-4, intercoxal process usually ogival in shape, with apex pointed or rounded, medially slightly concave to convex; ventrite 5 fl at or medially slightly concave, apically devoid of scales.
MALE TERMINALIA. Body of penis elongate or moderately elongate (W:L ratio 0.25-0.6), 0.5-2.0 × as long as temones, acuminate at apex, moderately curved in profi le; tectum narrow but distinct; endophallus with symmetrical copulatory sclerite, sometimes divided into two symmetrical structures, and with rows of cuticular denticles more or less visible between copulatory sclerite and base of penis body. Parameroid lobes of dorsal plate of tegmen separate, divided by median notch, not fused; apical setae always present but variable in length, number and arrangement. Spiculum gastrale asymmetrical; divergence of basal arms V-or U-shaped, right arm often angulate medially or bearing a tooth externally.

Sexual dimorphism
The sexes are distinguishable by their body shapes, males being smaller and globular and females larger and broadly ovate, and by the structure of the ventrite 1, in males centrally depressed but covered by plumose scales and in females fl at.

Distribution
South Africa.

Redescription (♂)
MEASUREMENTS. Body length 1.6-3.6 mm. COLOUR AND VESTITURE. Body integument black, scapes and tarsi reddish; vestiture of dorsum (pronotum + elytra) consisting of short, recumbent, subtriangular clothing scales, isodiametric or slightly longer than wide, partly concealing integument, overlapping or subcontiguous, and longer, suberect clothing scales at least 2 × as long as wide in strial punctures; colour pattern highly variable, scale colours brown and  Van Schalkwyk, 1966 (dorsal view). A. C. vitis   . K. C. pyrophilus Haran sp. nov. L. C. pilipes Haran sp. nov. Not to scale. grey; grey scales usually condensed on pronotum at base and apex of median line, on two sublateral longitudinal stripes and on elytra on striae 4-5, usually forming a transverse band at apical ⅔, sometimes divided into a pair of spots spanning interstriae 1-3.
HEAD. Forehead narrower than width of an eye, surface slightly concave. Eyes fl at, in dorsal view only slightly exceeding outline of head, surrounded by a ring of short pale scales directed towards centre of eye; distance between eye and scrobe larger than width of antennal club. Epifrons narrow, distance between antennal insertions 0.5 × length of scape, scales at least 3 × as long as wide, suberect, overlapping. Frons with a single pair of long lateral setae. Epistome with single median seta. Antennae variable, with funicle segments 1 elongate, at least 2 × as long as wide; 2-4 at least slightly longer than wide, compressed, on inside slightly angled; 5-7 isodiametric or wider than long, globular or slightly compressed.
PRONOTUM. Strongly transverse, nearly 2 × as wide as long (W:L ratio 1. 6-1.8), almost semicircular in dorsal view, widest at base, sides parallel or slightly arcuate in basal ⅓, straight or slightly curved and converging in apical ⅔; anteriorly 2 × narrower than posteriorly.
LEGS. Tibiae with black apical mucro; protibiae with outer margin straight, inner margin with series of small black teeth in apical half; metatibiae bisinuate, distal side of mucro perpendicular to external margin of metatibiae. Tarsi short, segment 2 much wider than long (W:L ratio 1.7-2.0).

Sexual dimorphism
Males and females can be distinguished by their body shape (smaller and globular in male, larger and broadly ovate in female), the structure of the fi rst ventrite (with a central cuticular depression concealed by plumose scales in male, fl at in female) and the width of the forehead (at most slightly wider in male, 2 × as wide as epifrons between antennal insertions in female).

Life history
Large numbers of specimens of Cryptolarynx vitis were collected in monospecifi c stands of Oxalis glabra Thunb. at various sites, but it was not reared from the bulbs of this plant species and its exact association with this species of Oxalis therefore needs verifi cation. Oxalis glabra is a weedy species that forms dense covers in vineyards in winter. High population densities of C. vitis in these stands can lead to damage to vine leaves when adults emerge and undertake maturation feeding . In this study, adults of this species were mostly found between August and November but once, in a population  at Wellington, in January and February. The seasonal occurrence reported by Van den Berg (1968), if applicable to the same species, indicates an activity of adults from late September to early December, with a peak in mid-November.

Distribution
The species occurs in lowland valleys of the Cape Town area, from Cape Town to Malmesbury and Wellington in the north and to Gansbaai in the south-east, from sea level to 400 m in elevation (Fig. 13). The specimens from Ceres reported by  could not be examined for this study and may represent a different species.

Remarks
A redescription of this species is provided to accommodate the variability in characters encountered in the wider geographical context than known to  and Van den Berg (1968). Together with C. hirtulus sp. nov., C. robustus sp. nov., C. squamulatus sp. nov. and C. subglaber sp. nov., C. vitis forms a species group characterised mainly by a narrow forehead, slightly erect elytral scales and a narrow apex of the parameroid lobes. Preliminary reconstruction of the phylogenetic relationships between the species of Cryptolarynx also suggests that these species form a cluster distinct from the other species of the genus (Fig. 12). It should be noted that the weakly supported preliminary phylogenetic relationships in this species group were not fully resolved with EF1 (Fig. 12). In C. vitis, intraspecifi c distances ranged up to 10.0% for COI and 2.5% for EF1 between specimens from different localities. This species may comprise several genetic lineages in the process of speciation, but stable morphological differences could not be identifi ed.
The description of C. vitis was based on 24 males and 24 females , of which 7 males and 10 females are preserved in the NHMUK and 2 males and 2 females in the SAMC. A female syntype in the NHMUK, bearing a type label in Marshall's hand, is here designated as the lectotype of C. vitis and was labelled accordingly (see above), and all other syntypes examined were labelled as paralectotypes.

Differential diagnosis
Cryptolarynx subglaber sp. nov. is closely related to C. vitis, see the Differential diagnosis and Remarks sections under that species for diagnostic characters and genetic distances.

Etymology
This species is named in reference to the glabrous appearance of the integument, seemingly unique among known species of Cryptolarynx. The specifi c epithet is an adjective in the masculine form.

Material examined
Holotype REPUBLIC OF SOUTH AFRICA • ♂; "REPUBLIC OF SOUTH AFRICA. Western Cape province, [Somerset West,]  COLOUR AND VESTITURE. Body integument black, antennae and tarsi reddish; vestiture of dorsum (pronotum + elytra) consisting of short, recumbent, rounded or subtriangular clothing scales, isodiametric or slightly longer than wide, not contiguous, and longer, suberect scales, at least 2 × as long as wide, brown or white, in each puncture of striae; scale colours brown and pale grey; grey scales usually condensed on two lateral stripes on pronotum and on transverse band on elytra at apical ⅔ of length, sometimes divided into a pair of large spots on interstriae 1-3.
HEAD. Forehead at most slightly wider than epifrons near antennal insertions, narrower than width of an eye, scales not concealing integument. Eyes fl at, in dorsal view only slightly exceeding outline of head, surrounded by a ring of short pale scales directed towards centre of eye; distance between eye and scrobe larger than width of antennal club. Epifrons narrow, distance between antennal insertions 0.5 × length of scape, scales at least 3 × as long as wide, erect, overlapping. Frons with a single pair of long lateral setae. Epistome with single median seta. Antennae variable, funicles with segment 1 elongate, at least 2 × as long as wide; 2-4 at least slightly longer than wide, compressed, slightly angular on inside; 5-7 isodiametric or wider than long, globular or slightly compressed.
PRONOTUM. Strongly transverse, nearly 2 × as wide as long (W:L ratio 1.7-2), almost semicircular in dorsal view, widest at base, sides parallel or slightly arcuate in basal ⅓, straight or slightly emarginate and converging in apical ⅔; apex 2 × narrower than base.
LEGS. Tibiae with black apical mucro; protibiae with outer margin straight, inner margin with series of small black teeth in apical half; metatibiae bisinuate, distal margin of mucro perpendicular to external margin of metatibia. Tarsi short, segment 2 much wider than long (W:L ratio 1.7-2.0).

Sexual dimorphism
The sexes can be distinguished by their body shape (globular in males (W:L ratio 0.7), broadly ovate in females (W:L ratio 0.6)) and by the cuticular elevation on ventrite 1 of males forming two ridges laterally (ventrite only slightly concave in females).

Life history
Specimens of Cryptolarynx subglaber sp. nov. were collected in mixed patches of species of Oxalis (O. lanata L. f., O. glabra and O. purpurea L.), and the exact host plant of the species is yet unknown. Adults were collected in August and October.

Distribution
The species occurs on western slopes of the Hottentots Holland Mountains, in Stellenbosch and the Helderberg Nature Reserve (elevation 130-170 m; Fig. 13).

Differential diagnosis
Cryptolarynx squamulatus sp. nov. belongs to the C. vitis group and in this group is most closely related to C. hirtulus sp. nov. These two species can be distinguished by their body shape, being globular in C. squamulatus and elongate in C. hirtulus, and their interspecifi c genetic distance was found to be 18.6% for COI and of 3.7% for EF1.

Etymology
This species is named in reference to the dense cover of suberect scales (squamae) on its elytra. The specifi c epithet is an adjective in the masculine form. COLOUR AND VESTITURE. Colour as in females (single known male teneral). Body integument black, antennae and tarsi reddish. Dorsal vestiture (pronotum + elytra) consisting of short, recumbent, subtriangular clothing scales, pearly grey, slightly longer than wide, subcontiguous on interstriae, and longer, suberect scales, at least 4 × as long as wide, brown or white, in each strial puncture; pale scales usually concentrated at base of interstriae 4 and corresponding area at pronotal base as well as forming an ill-defi ned transverse band at apical ⅔ of elytra. HEAD. Forehead slightly wider than epifrons near antennal insertions, narrower than width of an eye, scales not concealing integument. Eyes fl at, in dorsal view only slightly exceeding outline of head, surrounded by a ring of short pale scales directed towards centre of eye; distance between eye and scrobe larger than width of antennal club. Epifrons narrow, distance between antennal insertions 0.5 × length of scape, scales at least 3 × as long as wide, suberect, overlapping. Frons with single pair of long lateral setae. Epistome with single median seta. Antennal funicles with segment 1 elongate, 2 × as long as wide; 2-4 slightly longer than wide, compressed, slightly angular on inside; 5-7 globular, slightly compressed.    . K. C. pyrophilus Haran sp. nov. L. C. pilipes Haran sp. nov. Not to scale. PRONOTUM. Strongly transverse (W:L ratio 1.6), almost semicircular in dorsal view, widest at base, sides arcuate in basal ⅓, almost straight and converging in apical ⅔; apex 1.5 × narrower than base.
LEGS. Tibiae with black apical mucro; protibiae with outer margin straight; metatibiae with inner margin bisinuate, distal margin of mucro perpendicular to external margin of metatibia. Tarsi short, segment 2 much wider than long (W:L ratio 1.7).
TERMINALIA. Body of penis elongate (W:L ratio 0.3), twice as long as temones, sides straight, diverging from base to apical ¾, converging in apical quarter; curvature in profi le weak and regular, dorsoventrally narrowed before apex. Copulatory sclerite weakly sclerotised (or not discerned in single examined specimen). Parameroid lobes separate, divided by deep median notch, each lobe narrowed before apex, spatulate, bearing long marginal setae and shorter setae discally, all setae orientated centrifugally. Spiculum gastrale with basal arms regularly curved, right arm with lateral tooth.

Sexual dimorphism
The sexes can be distinguished by their body shape (more globular in male, broadly ovate in female).

Life history
Specimens of Cryptolarynx squamulatus sp. nov. were collected in a patch of Oxalis obtusa Jacq., but the exact host association of the species was not verifi ed. Freshly emerged adults were found in October.

Distribution
The species occurs in the West Coast National Park, Postberg Section (elevation 170 m; Fig. 13).

Differential diagnosis
Cryptolarynx muellerae sp. nov. has the general appearance of the species in the C. vitis group, but the male can be distinguished from these by its wider forehead, the absence of a mucro on the metatibiae and its non-constricted parameroid lobes. Among other species of the genus, it is the only species with long metatibial setae that has a body length of less than 2.5 mm.

Paratypes
REPUBLIC OF SOUTH AFRICA -Western Cape • 1 ♂; same collection data as for holotype; CBGP • 1 ♀; same collection data as for holotype; TMSA • 1 ♀; same collection data as for holotype; SANC.
HEAD. Forehead wider than epifrons near antennal insertions, as wide as width of an eye, scales not concealing integument. Eyes fl at, in dorsal view only slightly exceeding outline of head, surrounded by a ring of short pale scales, those on forehead directed towards occiput; distance between eye and scrobe as large as width of antennal club. Epifrons narrow, distance between antennal insertions 0.5 × length of scape, scales at least 3 × as long as wide, suberect, overlapping. Frons with single pair of long lateral setae. Epistome without single median seta. Antennal funicles with segments 1-2 elongate, 2 × as long as wide; 3-4 slightly longer than wide, compressed, slightly angular on inside; 5-6 globular; 7 transverse. PRONOTUM. Strongly transverse (W:L ratio 1.4), almost semicircular in dorsal view, widest near midlength, sides arcuate; width of apex 0.7 × width of base.
LEGS. Tibiae with black apical mucro; protibiae with both outer and inner margins straight; metatibiae with inner fringe of long setae, the setae longer than segment 5 of metatarsus. Tarsi with segment 2 isodiametric or slightly wider than long.
TERMINALIA. Body of penis moderately elongate (W:L ratio 0.4), 0.7 × as long as temones, sides straight, subparallel, converging abruptly near apex; curvature in profi le weak and regular, dorsoventrally narrowed before apex. Copulatory sclerite weakly sclerotised or not discerned in examined specimens. Parameroid lobes separate, divided by a deep median notch, each lobe parallel-sided, rounded at apex; distal margin setose, two setae long, exceeding length of others. Spiculum gastrale with basal arms regularly curved, right arm laterally angulate.

Sexual dimorphism
The sexes can be distinguished by their body shape (more globular in male, more broadly ovate in female) and the width of the forehead (slightly wider than epifrons in male, nearly 2 × wider in female).

Life history
Unknown. All known specimens of this new species were collected by sweeping vegetation.

Distribution
The species is only known from the type locality, near Vanrhynsdorp in the Western Cape Province (Fig. 13

Differential diagnosis
Cryptolarynx hirtulus sp. nov. belongs to the C. vitis species group, see the Remarks section under that species for details. In this group it is most closely related to C. squamulatus sp. nov., see the Differential diagnosis section under that species for diagnostic characters and genetic distances.

Etymology
The species name is derived from the Latin adjective 'hirtus', meaning 'hairy', and refers to the suberect scales in the strial punctures, which give the species a slightly hairy appearance. The specifi c epithet is an adjective in the masculine form. COLOUR AND VESTITURE. Body integument black, antennae, tarsi and sometimes tibiae reddish. Dorsal vestiture (pronotum + elytra) consisting of short, recumbent, subtriangular clothing scales, 1.2-2 × as long as wide, subcontiguous on interstriae, and longer, suberect scales, at least 4 × as long as wide, in each strial puncture, visible in lateral view; scales creamy-white, brown and black; pale scales usually concentrated in two longitudinal stripes on pronotum and on elytral interstriae 4 and forming a pair of pale spots surrounded by black scales on interstriae 1-4 at apical ⅔ of elytra.

Material
HEAD. Forehead as wide as epifrons near antennal insertions, narrower than width of an eye, scales not entirely concealing integument. Eyes moderately convex, in dorsal view only slightly exceeding outline of head, surrounded by a ring of short pale scales directed towards centre of eye; distance between eye and scrobe larger than width of antennal club. Epifrons narrow, distance between antennal insertions 0.5 × length of scape, scales at least 3 × as long as wide, suberect, overlapping. Frons with single pair of long lateral setae. Epistome without single median seta. Antennal funicles with segments 1-2 elongate, 2.5 × as long as wide; 3-4 slightly longer than wide, compressed, slightly angular on inside; 5-6 globular; 7 transverse. PRONOTUM. Strongly transverse (W:L ratio 1.4), widest near midlength, sides arcuate; apex and base subequal in width.
LEGS. Tibiae with black apical mucro; protibiae with outer margin straight, inner margin bisinuate; metatibiae with inner setal fringe, the setae shorter than segment 5 of metatarsus. Tarsi with segment 2 isodiametric or slightly wider than long.
ABDOMEN. Ventrite 1 fl at or slightly concave, bearing plumose scales in impression; other surfaces with overlapping creamy-white scales, partly concealing integument.
TERMINALIA. Body of penis elongate (W:L ratio 0.25), 1.5-2.0 × as long as temones, sides moderately convex, widest near midlength; curvature in profi le weak and regular, dorsoventrally slightly narrowed before apex. Copulatory sclerite weakly sclerotised or not discerned in examined specimens. Parameroid lobes separate, divided by deep median notch, each lobe narrowed anteapically and rounded at apex, bearing long setae marginally and discally, all setae orientated centrifugally. Spiculum gastrale with basal arms regularly curved.

Sexual dimorphism
The sexes can be distinguished by the width of the forehead (as wide as epifrons in male, distinctly wider in female), and females also lack the metatibial mucro and plumose scales on ventrite 1.

Life history
Adults of C. hirtulus sp. nov. were found at several localities in homogenous stands of Oxalis obtusa. Some were extracted from galleries in the soil, emerging from the bulbs of O. obtusa (JHAR02444, Fig. 6E-F). Adults were collected between July and September, with freshly emerged specimens encountered in July and August (JHAR02356; JHAR02444).

Distribution
Cryptolarynx hirtulus sp. nov. occurs at various localities in the southern and northern parts of the Cederberg mountain range and in the area of Malmesbury (Fig. 13). Specimens were collected from 120 to 1544 m above sea level.

Remarks
The amplifi cation of COI sequences from several populations of this species failed repeatedly, probably due to mismatches in the primer sequences. At intraspecifi c level, genetic distances between distant populations were found to be up to 1.3% in EF1 (JHAR02356; JHAR02561), suggesting that several lineages may exist in the species as described here. It should be noted that the preliminary phylogenetic relationships were not fully resolved for EF1 between this species and C. robustus sp. nov. (Fig. 12).

Differential diagnosis
Cryptolarynx robustus sp. nov. belongs to the C. vitis species group, see Remarks section under that species for details. In this group it is distinguishable from C. hirtulus sp. nov. by the width of its forehead being greater than the width of an eye (narrower in C. hirtulus). Uncorrected p-distances between C. robustus (JHAR02560) and C. hirtulus (JHAR02561) were found to be 17.4% for COI and 1.7% for EF1 (Supp. fi le 1).

Etymology
The species name is derived from the Latin adjective 'robustus' and refers to the stocky appearance of the species. The specifi c epithet is an adjective in the masculine form.

Paratypes
REPUBLIC OF SOUTH AFRICA -Western Cape • 1 ♀; same collection data as for holotype; SAMC • 1 ♀ (preserved in ethanol); same collection data as for holotype; CBGP.
HEAD. Forehead slightly wider than epifrons near antennal insertions, wider than width of an eye, scales suberect, not entirely concealing integument. Eyes fl at, in dorsal view only slightly exceeding outline of head, surrounded by a ring of short pale scales directed towards centre of eye; distance between eye and scrobe larger than width of antennal club. Epifrons narrow, distance between antennal insertions 0.5 × length of scape, scales at least 3 × as long as wide, suberect, overlapping. Frons with single pair of long lateral setae. Epistome with two median setae arising from same puncture. Antennal funicles with segments 1-2 elongate, subequal, about 3 × as long as wide; 3-4 slightly longer than wide, compressed, slightly angular on inside; 5-7 globular, isodiametric.
LEGS. Protibiae with outer margin straight, inner margin slightly bisinuate; metatibiae with black apical mucro and inner setal fringe, the setae shorter than segment 5 of metatarsus. Tarsi with segment 2 slightly wider than long.
TERMINALIA. Body of penis elongate (W:L ratio 0.3), 1.5-1.7 × as long as temones, sides moderately convex, widest near midlength; curvature in profi le weak and regular, dorsoventrally slightly narrowed before apex. Copulatory sclerite weakly sclerotised or not discerned in examined specimen. Parameroid lobes separate, divided by deep median notch, each lobe narrow, narrowed anteapically and rounded at apex, bearing long setae marginally and discally, all setae oriented centrifugally. Spiculum gastrale with basal arms regularly curved.

Sexual dimorphism
Males can be distinguished from females by the width of their forehead (as wide as epifrons in male, distinctly wider in female), and females also lack a mucro on the metatibiae and plumose scales on ventrite 1.

Life history
Adults of C. robustus sp. nov. were collected in September, at the bases of plants of Oxalis cf. purpurea in patches of Renosterveld.    . K. C. pyrophilus Haran sp. nov. L. C. pilipes Haran sp. nov. Not to scale.

Distribution
The species was only found at the type locality in the Western Cape province (Fig. 13).

Remarks
Cryptolarynx robustus sp. nov. and C. hirtulus sp. nov. have a similar general appearance and can be found in sympatry at the same localities.

Differential diagnosis
Cryptolarynx namaquanus sp. nov. can be distinguished from other species of Cryptolarynx by its wide epifrons (subequal to length of scapes), its narrow forehead (narrower than width of eyes) and its proximally cylindrical metatibiae. Its broad parameroid lobes of the male genitalia, bearing only very short setae, and the shape of the spiculum are also unique among the species of the genus. Cryptolarynx namaquanus is most similar to C. variabilis sp. nov. and C. carinatus sp. nov., but in the male it can be easily distinguished from these by its proximally cylindrical metatibiae (bearing an inner carina in C. variabilis and C. carinatus) and the conformations of its male genitalia. Uncorrected p-distances between C. namaquanus and these two species were found to span 12.8-15.1% for COI and 3.7-4.2% for EF1 with C. variabilis and 9.3-10.4% for COI and 3.4% for EF1 with C. carinatus (Supp. fi le 1).

Etymology
The species name namaquanus refers to the area where this species was found, the Namaqualand region of the Northern Cape province and part of the traditional home of the Nama people (Namaqua). The specifi c epithet is an adjective in the masculine form.

Description (♂)
MEASUREMENTS. Body length 2-3 mm. COLOUR AND VESTITURE. Body integument black, antennae and tarsi reddish. Dorsal vestiture (pronotum + elytra) consisting of overlapping, recumbent, parallel-sided clothing scales, 2 × as long as wide, truncate at apex; colour of scales varying from evenly greyish or creamy-white to brown or black; pale scales concentrated in two longitudinal stripes on pronotum and on elytral interstriae 4 as well as forming a pair of whitish spots surrounded by black scales at apical ⅔ of elytra; scales on interstriae recumbent, in lateral view not distinct from rest of vestiture.
HEAD. Forehead as wide as epifrons near antennal insertions, slightly narrower than width of an eye, scales recumbent. Eyes convex, in dorsal view exceeding outline of head, surrounded by a ring of short pale scales, on forehead directed towards occiput; distance between eye and scrobe as large as or slightly smaller than width of antennal club. Epifrons wide, distance between antennal insertions as large as length of scape, scales at most 2 × as long as wide, recumbent, subcontiguous. Frons with pairs of erect lateral setae. Epistome without median seta. Antennal funicles with segments 1-2 elongate, subequal, about 3 × as long as wide; 3-4 as long as wide, compressed, slightly angular on inside; 5-7 globular, isodiametric; 7 sometimes wider than long.
TERMINALIA. Body of penis moderately elongate (W:L ratio 0.5), 2 × as short as temones, sides subparallel in basal half, converging in distal half; curvature in profi le weak and regular, dorsoventrally slightly thickened at apex. Copulatory sclerite weakly sclerotised or not discerned in examined specimens. Parameroid lobes separate, divided by very deep median notch, each lobe broad, rounded at apex, bearing a series of very short setae. Spiculum gastrale with basal arms short and feebly curved.

Sexual dimorphism
Males can be distinguished from females by the shape of ventrites 1 and 5 (concave in male, fl at or slightly convex in female).

Life history
Adults of C. namaquanus sp. nov. were found in monospecifi c stands of Oxalis obtusa, in the month of August.

Distribution
All specimens collected were found at the type locality, the only one thus far known for the species (Fig. 13

Differential diagnosis
Cryptolarynx carinatus sp. nov. is most similar to C. variabilis sp. nov. but distinguishable from it by the inner carina on the metatibiae, whose proximal edge forms a smaller angle with the tibial axis than in C. variabilis (Fig. 8E). The copulatory sclerite of the penis is divided into two blocks in C. carinatus, whereas it forms an entire sclerite in C. variabilis. Genetic distances between these two species were found to range between 10.5% and 17.4% for COI and 3.1% and 3.6% for EF1. They were also not found in sympatry, C. carinatus apparently restricted to the eastern slopes of Table Mountain and C. variabilis only found at several localities on the western slopes of the Hottentots Holland Mountains.

Etymology
The species name carinatus refers to the distinct proximal inner carina of the metatibiae in this species. The specifi c epithet is an adjective in the masculine form. COLOUR AND VESTITURE. Body integument black, scapes and tarsi reddish. Dorsal vestiture (pronotum + elytra) consisting of overlapping, recumbent, parallel-sided clothing scales, 2-3 × as long as wide, truncate or rounded at apex; colour of scales varying from greyish to pale brown to black; pale scales concentrated in two longitudinal stripes on pronotum and at base of elytral interstriae 4, white scales forming a pair of spots surrounded by black scales at apical ⅔ of elytra; scales of interstriae recumbent, in lateral view not distinct from rest of vestiture.

Material
HEAD. Forehead as wide as epifrons near antennal insertions, as wide as width of an eye, scales recumbent. Eyes convex, in dorsal view slightly exceeding outline of head, surrounded by a ring of short pale scales, on forehead directed towards occiput; distance between eye and scrobe slightly smaller than width of antennal club. Epifrons with distance between antennal insertions 0.67 × length of scape, scales at least 2 × as long as wide, recumbent, subcontiguous. Frons with pairs of erect lateral setae. Epsitome without median setae. Antennal funicles with segment 1 elongate; 2 half as long as 1, at most 2 × as long as wide; 3-4 globular, isodiametric, compressed, slightly angular on inside; 5-7 globular, isodiametric; 7 sometimes wider than long.
TERMINALIA. Body of penis moderately elongate (W:L ratio 0.4), as long as temones, sides subparallel, converging near apex; curvature in profi le weak and regular, dorsoventrally narrowed before apex. Copulatory sclerite serrate, divided into two sections, a long part near body of penis and a second, shorter part near apices of temones. Parameroid lobes separate, divided by moderate median notch, each lobe broad, bearing a series of long setae directed apicad. Spiculum gastrale with basal arms long and regularly curved.

Sexual dimorphism
Female unknown.

Life history
All specimens of C. carinatus sp. nov. were collected at the base of Oxalis purpurea L. plants, in the month of June.

Distribution
The species in only known to occur in the Rondebosch Common and at Constantia in Cape Town (Fig. 13

Differential diagnosis
Cryptolarynx variabilis sp. nov. is most similar to C. carinatus sp. nov., see the Differential diagnosis and Remarks sections under that species for diagnostic characters and genetic distances.

Etymology
The species name variabilis refers to the substantial morphological variation encountered in this species in terms of size, body ratios and elytral pattern. The specifi c epithet is an adjective in the masculine form.

Material examined
Holotype REPUBLIC OF SOUTH AFRICA • ♂; "REPUBLIC OF SOUTH AFRICA. Western Cape Province, Stellenbosch. 20.vii.2018 COLOUR AND VESTITURE. Body integument black, scapes and tarsi reddish. Dorsal vestiture (pronotum + elytra) consisting of overlapping, recumbent, parallel-sided clothing scales, 2-3 × as long as wide, truncate or rounded at apex; colour of scales varying from greyish to pale brown to black; pale scales concentrated in two longitudinal stripes on pronotum and on elytral interstriae 4, white scales forming a pair of spots surrounded by black scales at apical ⅔ of elytra; scales of striae recumbent, in lateral view not distinct from rest of vestiture.
HEAD. Forehead as wide as epifrons near antennal insertions, as wide as width of an eye, scales recumbent. Eyes convex, in dorsal view slightly exceeding outline of head, surrounded by a ring of short pale scales, on forehead directed towards occiput; distance between eye and scrobe slightly smaller than width of antennal club. Epifrons with distance between antennal insertions 0.75 × length of scape, scales at least 2 × as long as wide, recumbent, not contiguous. Frons with pairs of erect setae laterally. Epistome without median seta. Antennal funicles with segment 1 elongate; 2 half as long as 1, at most 2 × as long as wide; 3-4 globular, isodiametric, compressed, slightly angular on inside; 5-7 globular, isodiametric; 7 sometimes wider than long.
LEGS. Protibiae with outer margin straight, inner margin almost straight; metatibiae with apical mucro and inner setal fringe, setae shorter than segment 5 of metatarsus; metatibiae proximally with inner carina set off at an angle of ca 20° to outer margin. Tarsi with segment 2 wider than long.
TERMINALIA. Body of penis elongate (W:L ratio 0.4), as long as temones, sides subparallel, converging close to apex; curvature in profi le weak and regular, dorsoventrally narrowed before apex. Copulatory sclerite serrate. Parameroid lobes separate, divided by modest median notch, each lobe broad, bearing a series of long setae directed apicad. Spiculum gastrale with basal arms long, right arm angled in midlength.

Sexual dimorphism
The sexes can be distinguished by the shape of ventrite 1 (in males concave with long setae divided from their bases, in females fl at or slightly convex with short setae not or only slightly divided at their apices).

Life history
The larvae of C. variabilis sp. nov. develop in the bulbs of various species of Oxalis (O. pes-caprae L., O. purpurea and O. lanata); see Life history and Morphology of immature stages subsections of the Results section for details. Adults of the species were collected between July and October. The heat tolerance of this species was assessed in a comparative study of weevils associated with fi re-prone ecosystems (Javal et al. 2022). Adults of C. variabilis were found to survive temperatures above 50°C, which is rare for arthropods, especially in small insects. This tolerance was hypothesized to be an adaptation for the survival of the larvae and teneral adults when enclosed in the bulbs during summer, a few centimetres below the surface of the ground.

Distribution
Based on the samples examined, C. variabilis sp. nov. seems to be restricted to the western slope of the Hottentots Holland Mountains range, from the Franschhoek Pass to Somerset West (Fig. 13).

Differential diagnosis
Cryptolarynx estriatus can be distinguished from all other species of the genus by its suberect scales on epifrons, the two short carinae on ventrite 1 of the male and the unique long seta on the parameroid lobes.

Material examined
Holotype REPUBLIC OF SOUTH AFRICA • ♂; "Type" "S Africa. COLOUR AND VESTITURE. Body integument black, scapes and tarsi reddish; vestiture of dorsum (pronotum + elytra) consisting of short, appressed, narrowly elliptical clothing scales partly concealing integument, slightly overlapping or subcontiguous, and longer, suberect clothing scales at least 2 × as long as wide on strial punctures; colour pattern consisting of broad median stripe of pearly brown scales on pronotum and elytra, fl anked on either side by narrower stripe of white scales, laterally with paler brown scales intermixed with sparser white scales, without distinct white spots on elytral declivity.
HEAD. Forehead narrower than width of an eye, surface fl at. Eyes fl at, in dorsal view only slightly exceeding outline of head, surrounded by a ring of short pale scales directed towards centre of eye except mesal ones directed posteriad; distance between eye and scrobe as wide as antennal club. Epifrons narrow, distance between antennal insertions 0.5 × length of scape, scales narrow, 3 × as long as wide, suberect. Frons with double pair of long lateral setae. Epistome with single median seta ca half as long as frontal setae. Antennae with funicle segment 1 and 2 elongate, more than 2 × as long as wide; 3-4 slightly longer than wide, slightly compressed, slightly angulate on inside, 2 and 4 on inside slightly angled; 5-7 isodiametric, subglobular.
PRONOTUM. Strongly transverse, nearly 1.5 × as wide as long (W:L ratio 1.4-1.5), subcircular in dorsal view, widest just behind middle of length, sides strongly arcuate in basal half, straight and converging in apical half; anteriorly 2 × narrower than posteriorly.
LEGS. Tibiae with black apical mucro; protibiae with outer margin straight, inner margin with series of small black teeth in apical half; metatibiae with outer and inner margins faintly sinuate, mucro subperpendicular to external margin. Tarsi short, segment 2 wider than long (W:L ratio 1.4).
ABDOMEN. Ventrites with creamy-white to pale brown plumose scales almost concealing integument, medially intermixed with long suberect setae; ventrite 1 concave medially, fl anked on either side by a short but high, fl at, black peg midway between anterior margin at metacoxae and posterior margin.
TERMINALIA. Body of penis elongate (W:L ratio 0.45), 0.9 × as long as temones, sides subparallel, strongly rounded inwards distally, apex subacuminate; curvature in profi le weak and regular, dorsoventrally strongly fl attened before apex. Copulatory sclerite indistinct, consisting of short, very thin, outwardly curved sclerotised rods at apices of longer and thicker crescentic fi elds of very fi ne asperities. Parameroid lobes separate, divided by deep median notch, each with anteapical constriction, bearing long marginal setae and shorter setae discally, all setae orientated centrifugally. Spiculum gastrale with basal arms regularly curved.

Sexual dimorphism
The sexes can be distinguished foremost by the conspicuous pair of fl at black pegs on ventrite 1 in the male (absent in the female).

Life history
Specimens of C. estriatus were found on the ground or feeding on vegetation, but no precise data of its host plants are known. A series of adults was found feeding on leaves of Moraea by S. Neser near Nieuwoudtville in 1995, and they also fed on this plant in captivity (Oberprieler et al. 2007: fi g. 17), but, in view of the wide use of Oxalis as larval host in Cryptolarynx, the larvae of C. estriatus probably also develop in bulbs of an Oxalis species rather than on Moraea. Adults were collected in July, September and November.

Distribution
Most specimens were collected in the Nieuwoudtville area, on the escarpment (Fig. 13).

Remarks
Marshall described this species based on a single female, but examination of the genitalia of ts specimen showed that it is actually a male. No fresh material of this species could be obtained.

Differential diagnosis
Cryptolarynx pyrophilus sp. nov. can be distinguished from other species of the genus by its very wide forehead (2 × as wide as width of an eye) and by the suberect scales on the epifrons. It is most similar to C. cederbergensis sp. nov., the two species showing interspecifi c genetic distances ranging from 20.9% to 23.3% for COI and 5.2 % for EF1 among the specimens available for study.

Etymology
The species was named in reference to the migration of specimens towards recently burnt areas observed in this species, formed from combining the Greek nouns 'pyr' ('fi re') and 'philia' ('affection'). The specifi c epithet is an adjective in the masculine form.  COLOUR AND VESTITURE. Body integument black, antennae and tarsi reddish. Dorsal vestiture (pronotum + elytra) consisting of overlapping, recumbent, parallel-sided clothing scales, 2-3 × as long as wide, truncate or rounded at apex; colour of scales varying from creamy-white to pale brown to dark brown; white scales concentrated in two longitudinal stripes laterally on pronotum and on elytral interstriae 4, creating a broad, dark medial stripe on pronotum and basal ⅔ of elytra; white scales forming a pair of spots surrounded by black scales on interstriae 3 at apical ⅔ of elytral length; scales of striae recumbent, in lateral view not distinct from rest of vestiture.
HEAD. Forehead very wide, 1.3 × as wide as epifrons near antennal insertions, 2 × as wide as width of an eye, scales recumbent. Eyes convex, in dorsal view slightly exceeding outline of head, surrounded by a ring of short pale scales, on forehead directed towards occiput; distance between eye and scrobe as large as width of antennal club. Epifrons with distance between antennal insertions 0.75 × length of scape, scales at least 2 × as long as wide, suberect, contiguous. Frons with a single pair of erect lateral setae. Epistome with single elongate median seta. Antennal funicles with segment 1 elongate; 2 shorter, at most 1.5 × as long as wide; 3-4 globular, isodiametric, compressed, slightly angular on inside; 5-6 globular, isodiametric; 7 wider than long.
LEGS. Tibiae with apical mucro; protibiae with outer margin straight, inner margin bisinuate; metatibiae with inner setal fringe, the setae shorter than segment 5 of metatarsus. Tarsi with segment 2 wider than long.

Sexual dimorphism
The sexes can be distinguished by the shape of the elytra (wider than long in male, longer than wide, more broadly ovate in female), and females also have a wider forehead and smaller metatibial mucrones.

Life history
Large numbers of adults of C. pyrophilus sp. nov. were collected in and close to a recently burnt area (seven months prior to sampling), at the bases of emerging Oxalis pes-caprae L. plants; see the subsection Behaviour below for details. The heat tolerance of this species was assessed in a comparative study of weevils associated with fi re-prone ecosystems, but the adults showed a lower tolerance to heat than those of C. variabilis sp. nov. (Javal et al. 2022). Adults were collected in September and October.

Distribution
The species occurs in the inland valleys of Montagu and the Hex River (Fig. 13).

Differential diagnosis
Cryptolarynx pilipes sp. nov. can be distinguished from other species of the genus by the combination of the inner fringe of elongate setae on the metatibiae and the suberect scales on the epifrons.

Etymology
The species name pilipes is derived from the Latin nouns 'pilus' ('hair') and 'pes' ('foot') and refers to the fringe of long setae on the metatibiae of the species. The specifi c epithet is a noun in apposition. COLOUR AND VESTITURE. Body integument black, antennae and tarsi reddish. Dorsal vestiture (pronotum + elytra) consisting of overlapping, recumbent, parallel-sided clothing scales, at least 2 × as long as wide, truncate or rounded at apex; colour of scales varying from white to pale brown to dark brown, white scales concentrated laterally on pronotum and at base of elytral interstriae 4 and forming a pair of subcontiguous white spots surrounded by dark scales at apical ⅔ of interstriae 1-4; scales of striae recumbent, in lateral view not distinct from rest of vestiture.

Material
HEAD. Forehead wide, 1.5 × wider than epifrons near antennal insertions, 1.25 × as wide as width of an eye, scales suberect. Eyes convex, in dorsal view slightly exceeding outline of head, surrounded by a ring of short pale scales, on the forehead directed towards occiput; distance between eye and scrobe as large as width of antennal club. Epifrons with distance between antennal insertions 0.5 × length of scape, scales at least 2 × as long as wide, suberect, contiguous. Frons with single pair of erect lateral setae. Epistome without median seta. Antennae with funicle segment 1 elongate; 2 slightly shorter, at most 2 × as long as wide; 3-5 longer than wide, compressed; 6-7 globular, isodiametric.
LEGS. Protibiae with both outer and inner margins straight; pro-and mesotibiae with small apical mucro, metatibiae amucronate; metatibiae with inner fringe of setae as long as segment 5 of metatarsus. Tarsi with segment 2 wider than long.
TERMINALIA. Body of penis elongate (W:L ratio 0.3), slightly shorter than temones, sides subparallel, converging strongly near apex; curvature in profi le weak and regular, dorsoventrally narrowed at apex. Copulatory sclerite weakly sclerotised or not discerned in examined specimens. Parameroid lobes separate, divided by deep median notch, each lobe broad, rounded at apex, bearing a series of long setae directed apicad, setae as long as depth of median notch. Spiculum gastrale with basal arms short, regularly and moderately curved, laterally slightly angulate.

Sexual dimorphism
Female unknown.

Distribution
The species was found only at the type locality, Clanwilliam in the Western Cape province (Fig. 13

Differential diagnosis
Cryptolarynx armatus sp. nov. is most similar to C. falciformis sp. nov. and C. oberprieleri sp. nov., in possessing apically thickened protibiae in the male. In addition to their distinct copulatory sclerites of the endophallus, these species can also be distinguished by the setae on the ventrites (simple in C. armatus, bifi d at least apically in C. falciformis and C. oberprieleri). Genetically, C. armatus is closest to C. spinicornis sp. nov., the distance between their EF1 sequences found to be 1.6%.

Etymology
The species name armatus refers to the apical cuticular expansion of the protibiae of the males, forming together with the mucro two strong teeth. The specifi c epithet is an adjective in the masculine form. COLOUR AND VESTITURE. Body integument black, scapes, tibiae and tarsi reddish in fully sclerotised specimens. Dorsal vestiture (pronotum + elytra) consisting of overlapping, recumbent, parallel-sided clothing scales, 2-2.5 × as long as wide, truncate at apex; colour of scales varying from pale brown to dark brown; pale scales concentrated in two longitudinal bands laterally on pronotum as well as broadly on elytral interstriae 4, creating a broad, dark medial stripe on pronotum and basal ⅔ of elytra; pale scales forming a pair of spots surrounded by black scales on interstriae 3 at apical ⅔ of elytral length; scales of striae recumbent, in lateral view not distinct from rest of vestiture.

Holotype
HEAD. Forehead wide, slightly wider than epifrons near antennal insertions, almost 2 × as wide as width of an eye, scales recumbent. Eyes convex, in dorsal view slightly exceeding outline of head, surrounded by a ring of short pale scales, on forehead directed towards occiput; distance between eye and scrobe as large as width of antennal club. Epifrons with distance between antennal insertions as large as length of scape, scales in middle of epifrons at most 2 × as long as wide, recumbent, not contiguous. Frons with a pair of long erect lateral setae. Epistome without median seta. Antennal funicles with segment 1 elongate, 2 × as long as wide; 2 shorter, at most 1.5 × as long as wide; 3-5 isodiametric, compressed, 4 slightly angular on inside; 6-7 wider than long.
LEGS. Protibiae with outer margin straight and inner margin bisinuate, expanded proximally of apical mucro; meso-and metatibiae with small apical mucro, metatibiae with inner setal fringe, setae shorter than segment 5 of metatarsus. Tarsi with segment 2 isodiametric or wider than long.
TERMINALIA. Body of penis moderately elongate (W:L ratio 0.4), almost 2 × as short as temones, sides subparallel, converging in apical quarter; in profi le almost straight, dorsoventrally narrowed close to apex. Copulatory sclerite small, comma-shaped. Parameroid lobes separate, divided by modest median notch, each lobe broad, bearing a series of setae directed apicad, the longest setae located closer to middle. Spiculum gastrale with basal arms long, regularly curved.

Sexual dimorphism
The sexes can be distinguished by the shape of the elytra (wider than long in male, longer than wide, more broadly ovate in female) and by the inner expansion of the apex of the protibiae (present in male, absent in female).

Life history
Adult specimens of C. armatus were collected in August, in stands of Oxalis obtusa and O. cf. suteroides T.M. Salter.

Distribution
All specimens were collected at the type locality near Nieuwoudtville (Fig. 13).

Remarks
Mitochondrial barcode sequences could not be obtained for this species, probably due to a mismatch in primer sequences.

Differential diagnosis
Cryptolarynx falciformis sp. nov. is most similar to C. oberprieleri sp. nov.; see Differential diagnosis section under that species for details.

Etymology
The species name falciformis refers to the falcate (sickle-shaped) copulatory sclerite in the endophallus of this species. The specifi c epithet is an adjective in the masculine form. COLOUR AND VESTITURE. Body integument black, antennae and tarsi reddish. Dorsal vestiture (pronotum + elytra) consisting of overlapping, recumbent, parallel-sided clothing scales, 2-3 × as long as wide, rounded or truncate at apex; colour of scales greyish and pale brown; greyish scales concentrated laterally of elytral interstriae 4 and in two ill-defi ned pale spots at apical ⅔ of interstriae 3; scales of striae recumbent, in lateral view not distinct from rest of vestiture.

REPUBLIC OF SOUTH
HEAD. Forehead wide, slightly wider than epifrons near antennal insertions, almost 2 × as wide as width of an eye, scales recumbent. Eyes convex, in dorsal view slightly exceeding outline of head, surrounded by a ring of short pale scales, on forehead directed towards occiput; distance between eye and scrobe smaller than width of antennal club. Epifrons with distance between antennal insertions as large as length of scape, scales in middle of epifrons at most 2 × as long as wide, recumbent, not contiguous. Frons with a pair of long erect lateral setae. Epistome without median seta. Antennal funicles with segments 1-2 elongate, subequal; 3-6 isodiametric, 4 slightly angular ventrally; 7 wider than long.
LEGS. Protibiae with outer margin straight and inner margin bisinuate, expanded proximally of apical mucro; meso-and metatibiae with apical mucro; metatibiae with inner setal fringe, setae shorter than segment 5 of metatarsus and with small ventral carina near base. Tarsi with segment 2 wider than long.
TERMINALIA. Body of penis moderately elongate (W:L ratio 0.5), almost 1.5 × as short as temones, sides subparallel, converging in apical quarter; in profi le almost straight, dorsoventrally narrowed before apex. Copulatory sclerite shaped like a pair of sickles. Parameroid lobes separate, divided by modest median notch, each lobe narrowing apicad, bearing two long setae apically. Spiculum gastrale with basal arms long, regularly curved.

Sexual dimorphism
Female unknown.

Life history
No data on host plants are available. The single known specimen was collected in November.

Distribution
The species was only found at the type locality, Tulbagh in the Western Cape province (Fig. 13).

Differential diagnosis
Cryptolarynx oberprieleri sp. nov. can be distinguished from other species of the genus by the combination, in males, of apically expanded protibiae, deeply divided setae on ventrite 1 and arrowheadshaped copulatory sclerite. It differs from the most similar congener, C. falciformis sp. nov., by the structure of the apex of the parameroid lobes and the copulatory sclerite in the endophallus (Fig. 2O). Among the species for which fresh tissue was obtained, C. oberprieleri was found to be genetically closest to C. marshalli sp. nov., the two species showing uncorrected p-distances ranging from 19.8% to 22.1% for COI and from 3.4% to 3.6% for EF1, whereas intraspecifi c distances were up to 3.5% for COI (JHAR01246; JHAR02585) and 0.6% for EF1 (JHAR02340; JHAR02585) (Supp. fi le 1).

Etymology
Cryptolarynx oberprieleri sp. nov. is dedicated to Rolf G. Oberprieler for his substantial contribution to weevil taxonomy and classifi cation, not least regarding the South African fauna. The specifi c epithet is a noun in the genitive case. COLOUR AND VESTITURE. Body integument black, scapes and tarsi reddish in fully sclerotised specimens. Dorsal vestiture (pronotum + elytra) consisting of overlapping, recumbent, parallel-sided clothing scales, 2-3 × as long as wide, rounded or truncate at apex; colour of scales pale to dark brown; pale scales concentrated in two lateral stripes on pronotum and on elytra from interstriae 4 laterad, as well as forming a pair of white spots surrounded by darker scales at apical ⅔ of interstriae 2-3; scales of striae recumbent, in lateral view not distinct from rest of vestiture. HEAD. Forehead slightly wider than epifrons near antennal insertions, as wide as an eye, scales recumbent. Eyes convex, in dorsal view slightly exceeding outline of head, surrounded by a ring of short pale scales, on forehead directed towards occiput; distance between eye and scrobe smaller than width of antennal club. Epifrons with distance between antennal insertions slightly smaller than length of scape, scales in middle of epifrons at least 3 × as long as wide, recumbent, not contiguous. Frons with a pair of long erect lateral setae. Epistome without median seta. Antennal funicles with segments 1-2 moderately elongate, subequal in length, 1.5-1.8 × as long as wide; 3 isodiametric; 4 slightly angular ventrally; 5-7 globular, isodiametric.
LEGS. Protibiae with outer margin straight and inner margin slightly bisinuate, with small expansion proximally of apical mucro; meso-and metatibiae with small apical mucro, metatibiae with inner setal fringe, setae shorter than segment 5 of metatarsus, and with small inner carina near base. Tarsi with segment 2 wider than long.

Sexual dimorphism
The sexes can be distinguished by the shape of the elytra (wider than long in male, longer than wide and more broadly ovate in female), by the shape of ventrite 1 (medially concave in male, convex in female) and by the protibiae (with small expansion near mucro in male).

Life history
Specimens of C. oberprieleri sp. nov. were collected at several localities in monospecifi c stands of Oxalis glabra and, at one location, at the base of a plant of O. livida Jacq. Larvae and teneral adults were found inside bulbs of O. glabra. Adults were collected between late June and late October, found to be active during the day at the base of their host plant but at sunset retreating into small holes in the soil, which they formed under debris and under the leaves of Iridaceae.

Distribution
This species was found on the western slopes of the Hottentots Holland Mountain range and adjacent valleys, from Stellenbosch and Klipheuwel in the north to Kogel Bay beach in the south. A single specimen was discovered in the De Hoop Nature Reserve (Fig. 13).

Differential diagnosis
Cryptolarynx spinicornis sp. nov. differs from all other species of the genus by its enlarged funicle segments 2 and 4, bearing an inner tooth (Fig. 8H). A mitochondrial barcode fragment could not be obtained for it, probably due to mismatch in the primers sequences. Uncorrected p-distances of EF1 show that this species is closest to C. san sp. nov. but distant from it by 1.1% (Supp. fi le 1).

Etymology
The species name spinicornis is derived from the Latin nouns 'spina' ('spine') and 'cornu' ('horn', 'antenna') and refers to the teeth on funicle segments 2 and 4 in this species. The specifi c epithet is an adjective in the masculine form. COLOUR AND VESTITURE. Body integument black, antennae, tarsi and sometimes tibiae reddish. Dorsal vestiture (pronotum + elytra) consisting of overlapping, recumbent, parallel-sided clothing scales, 3 × as long as wide, truncate at apex; colour of scales pale to dark brown; pale scales generally concentrated in two lateral bands on pronotum and on elytral interstriae 4 as well as forming a pair of white spots surrounded by darker scales at apical ⅔ of interstriae 2-3; dark scales concentrated in a pair of spots at base of pronotum near scutellar shield; scales of striae recumbent, in lateral view not distinct from rest of vestiture. HEAD. Forehead slightly wider than epifrons near antennal insertions, slightly wider than width of an eye, scales recumbent. Eyes convex, in dorsal view slightly exceeding outline of head, surrounded by a ring of short pale scales, on forehead directed towards occiput; distance between eye and scrobe smaller than width of antennal club. Epifrons with distance between antennal insertions as large as length of scape, scales in middle of epifrons at least 3 × as long as wide, recumbent, not contiguous. Frons with 3 pairs of erect lateral setae. Epistome without median seta. Antennal funicles with segment 1 very elongate, 2 × as long as 2; 2 and 4 compressed, strongly angular, toothed on inside; 3 isodiametric or slightly longer than wide; 5-7 globular, isodiametric.
LEGS. All tibiae with small apical mucro; protibiae with outer margin straight, inner margin slightly bisinuate; metatibiae with inner setal fringe, setae shorter than segment 5 of metatarsus. Tarsi with segment 2 wider than long on forelegs and longer than wide on hindlegs.
TERMINALIA. Body of penis moderately elongate (W:L ratio 0.4), slightly shorter than temones, sides subparallel in basal ⅔, converging in apical third; curvature in profi le moderate and regular, more strongly downcurved near apex, dorsoventrally narrowed before apex. Copulatory sclerite shaped like a pair of sickles. Parameroid lobes separate, divided by deep median notch, each lobe slightly bilobate, with median sublobes bearing several long setae apically. Spiculum gastrale with basal arms long, right arm angulate at its midlength.

Sexual dimorphism
The sexes can be distinguished by the shape of the elytra (shorter in male than in female) and the forehead (narrower in male than in female).

Life history
Specimens of Cryptolarynx spinicornis sp. nov. were found at the bases of plants of Oxalis cf. luteola Jacq. and O. cf. odorata J.C. Manning & Goldblatt at sites where O. obtusa was also present.

Distribution
The species occurs in the Vanrhynsdorp area and up to 1500 m above sea level on the Great Escarpment near Sutherland (Fig. 13).

Remarks
Mitochondrial barcode sequences could not be obtained for this species, probably due to a mismatch in the primer sequences.

Differential diagnosis
Cryptolarynx cederbergensis sp. nov. is most similar to C. pyrophilus sp. nov. but can be distinguished from it by the width of its forehead, which is less than twice the width of an eye (equal to twice width in C. pyrophilus), and by the arrangement of setae at the apex of the parameroid lobes (Fig. 2Q). The two species were found to have interspecifi c uncorrected p-distances ranging from 20.9% to 23.3% for COI and 5.2% for EF1 (Supp. fi le 1).

Etymology
The species name cederbergensis refers to the origin of this species, the Cederberg mountains in the Western Cape province. The specifi c epithet is an adjective in the masculine form. COLOUR AND VESTITURE. Body integument black, basal half of scapes reddish in mature specimens. Dorsal vestiture (pronotum + elytra) consisting of overlapping, recumbent, parallel-sided clothing scales, 2 × as long as wide, mostly rounded at apex; colour of scales white or pale brown to dark brown; white scales usually concentrated in a pair of longitudinal lateral stripes on pronotum and elytral interstriae 4, creating a broad darker stripe medially on pronotum and basal ⅔ of elytra; white scales also concentrated in a pair of ill-defi ned pale spots surrounded by black scales at apical ⅔ of interstria 3; scales of striae recumbent, in lateral view not or only very slightly distinct from rest of vestiture.
HEAD. Forehead very wide, slightly wider than epifrons near antennal insertions, ca 2 × as wide as width of an eye, scales suberect. Eyes convex, in dorsal view slightly exceeding outline of head, surrounded by a ring of short pale scales, on forehead directed towards occiput; distance between eye and scrobe as large as width of antennal club. Epifrons with distance between antennal insertions 0.8 × length of scape, scales at least 3 × as long as wide, suberect, contiguous. Frons with a pair of long erect lateral setae. Epistome with one or two elongate median setae. Antennal funicles with segment 1 elongate; 2 shorter, at most 1.5 × as long as wide; 3 longer than wide; 4 globular, isodiametric, compressed, slightly angular on inside; 5-7 globular, isodiametric.
LEGS. Tibiae with apical mucro; protibiae with both outer and inner margins straight; metatibiae with inner setal fringe, the setae shorter than segment 5 of metatarsus. Tarsi with segment 2 slightly longer than wide.
TERMINALIA. Body of penis elongate (W:L ratio 0.3-0.4), as long as temones, sides subparallel, converging in apical third; curvature in profi le weak and regular, dorsoventrally narrowed before apex. Copulatory sclerite forming a reversed V. Parameroid lobes separate, divided by deep median notch, each lobe broad, bearing a series of long setae directed apicad. Spiculum gastrale with basal arms long, regularly curved, right arm angulate near its base.

Sexual dimorphism
The sexes can be distinguished by the width of the forehead (as wide as or narrower than width of an eye in male, wider in female).

Life history
Specimens of C. cederbergensis sp. nov. were collected in monospecifi c stands of Oxalis obtusa, in the month of August.

Distribution
The species was found only at the type locality (Fig. 13

Differential diagnosis
Cryptolarynx homaroides sp. nov. is most similar to C. marshalli sp. nov., but the body of the latter species is distinctly more elongate and lacks the distinct lobster-like copulatory sclerite of C. homaroides.

Etymology
The species name homaroides is derived from the genus name of the European Lobster (Homarus gammarus Linnaeus) and refers to the remarkably similar shape of copulatory sclerite in the endophallus of this species. The specifi c epithet is an adjective with immutable ending. COLOUR AND VESTITURE. Body integument black, scapes at their bases, tibiae and tarsi reddish. Dorsal vestiture (pronotum + elytra) consisting of overlapping, recumbent, parallel-sided or subtriangular clothing scales, 1.5-2 × as long as wide, truncate or rounded at apex; colour of scales pale to dark brown; pale scales generally concentrated in two lateral bands on pronotum and at base of elytral interstriae 4, creating irregular shades on elytra, and concentrated in a pair of pale spots surrounded by darker scales at apical ⅔ of interstriae 2-3; scales of striae recumbent, in lateral view not distinct from rest of vestiture.

REPUBLIC OF SOUTH
HEAD. Forehead as wide as epifrons near antennal insertions, scales recumbent. Eyes convex, in dorsal view slightly exceeding outline of head, surrounded by a ring of short pale scales, on forehead directed towards occiput; distance between eye and scrobe smaller than width of antennal club. Epifrons with distance between antennal insertions as large as length of scape, scales in middle of epifrons at most 2 × as long as wide, recumbent, not contiguous. Frons with 3 pairs of long erect lateral setae. Epistome without median seta. Antennal funicles with segment 1 moderately elongate, 1.5 × as long as wide; 2-4 longer than wide, compressed; 5-6 globular, isodiametric; 7 wider than long.
LEGS. All tibiae with small apical mucro; protibiae with outer margin straight, inner margin slightly bisinuate; metatibiae with inner setal fringe, the setae shorter than segment 5 of metatarsus. Tarsi with segment 2 wider than long.
TERMINALIA. Body of penis moderately elongate (W:L ratio 0.45), slightly shorter than temones, sides convex; in profi le straight, slightly downcurved and dorsoventrally slightly narrowed near apex. Copulatory sclerite in dorsal view shaped like a lobster. Parameroid lobes separate, divided by deep median notch; each lobe bearing two brushes of erect setae. Spiculum gastrale with basal arms long, longer than ventral strut, right arm regularly curved.

Sexual dimorphism
Female unknown.

Life history
Unknown. The single known specimen of C. homaroides sp. nov. was collected under stones.

Distribution
The species was only found at the type locality, the Springbok area in the Northern Cape province (Fig. 13).

Differential diagnosis
Cryptolarynx marshalli sp. nov. is closely related to C. oberprieleri sp. nov. but distinctly more elongate, and the apex of its parameroid lobes is also distinct (Fig. 2O-S). See Differential diagnosis section under species C. oberprieleri for the genetic distances between these species.

Etymology
This species is dedicated to the late weevil expert Sir Guy A.K. Marshall, who described the genus and its original two species and discussed its unique characters among known weevils. The specifi c epithet is a noun in the genitive case. Paratypes REPUBLIC OF SOUTH AFRICA -Western Cape • 1 ♂, 2 specs; same collection data as for holotype; CBGP.

Description (♂)
MEASUREMENTS. Body length 2.5-2.9 mm. COLOUR AND VESTITURE. Body integument black, antennae, tibiae and tarsi reddish. Dorsal vestiture (pronotum + elytra) consisting of overlapping, recumbent, parallel-sided clothing scales, 3 × as long as wide, truncate at apex; colour of scales mostly brown, white scales interspersed with pale brown scales concentrated in two longitudinal bands on pronotum, at base of elytral interstriae 4, and in a pair of pale spots surrounded by black scales at apical ⅔ of interstriae 3; scales of striae recumbent, in lateral view not or only very slightly distinct from rest of vestiture.
HEAD. Forehead wide, slightly wider than epifrons near antennal insertions, scales suberect. Eyes convex, in dorsal view slightly exceeding outline of head, surrounded by a ring of short scales, on forehead directed towards occiput; distance between eye and scrobe smaller than width of antennal club. Epifrons with distance between antennal insertions slightly smaller than length of scape, scales at least 2 × as long as wide, recumbent, mostly non-contiguous. Frons with 3 pairs of erect lateral setae. Epistome without median seta. Antennal funicles with segment 1 moderately elongate, 1.5 longer than wide; 2 subequal in length to 1; 2 and 4 compressed, slightly angular on inside; 5-7 globular, isodiametric or wider than long.
ABDOMEN. Ventrite with creamy-white plumose scales not fully concealing integument, scales on ventrites 2-5 medially intermixed with long suberect setae, apically bifi d or not; ventrite 1 slightly concave medially, impression covered with long setae deeply divided from their bases; ventrite 5 with scales concentrated laterally and on basal third.
TERMINALIA. Body of penis moderately elongate (W:L ratio 0.45), slightly shorter than temones, sides convex; curvature in profi le weak, stronger in basal half, not dorsoventrally narrowed at apex. Copulatory sclerite weakly sclerotised or not discerned in examined specimens. Parameroid lobes separate, divided by modest median notch, each lobe broad, bearing a series of setae directed apicad, median setae longer. Spiculum gastrale with basal arms long, right arm slightly angulate at its midlength.

Sexual dimorphism
The sexes can be distinguished by the elytra (shorter in male) and by ventrite 1 in the female lacking the long, deeply divided setae.

Life history
All specimens of this species were collected in July at the base of Oxalis imbricata Eckl. & Zeyh. plants.

Differential diagnosis
Cryptolarynx endroedyi sp. nov. differs from all other known species of the genus by its distinctly elongate body and by the following two features (in the male): the presence of depressions on the pronotum and elytra and the temones of the penis being almost twice as long as the penis body.

Etymology
We dedicate this species to the late Sebastian Endrödy-Younga, coleopterist at the Ditsong National Museum of Natural History (formerly the Transvaal Museum) from 1973 to 1999. The extent of his fi eld collecting in South Africa and Namibia exceeds imagination with respect to the large numbers of species and of specimens he collected and the signifi cant number of localities he surveyed. He also collected many of the specimens of Cryptolarynx and Hadrocryptolarynx gen. nov. reported in this study. The specifi c epithet is a noun in the genitive case. Paratypes REPUBLIC OF SOUTH AFRICA -Western Cape • 2 ♀♀; same collection data as for holotype; TMSA • 1 ♀; same collection data as for holotype; CBGP.

Description (♂)
MEASUREMENTS. Body length 3.6 mm. COLOUR AND VESTITURE. Body integument black, antennae, tibiae and tarsi reddish. Dorsal vestiture (pronotum + elytra) consisting of overlapping, recumbent, parallel-sided clothing scales, 3-4 × as long as wide, mostly truncate at apex; colour of scales mostly brown; whitish scales interspersed with pale brown scales concentrated laterally on pronotum and on elytra laterally from interstriae 4, white scales forming a pair of pale spots surrounded by dark scales at apical ⅔ of elytral interstriae 2-3; scales of striae recumbent, in lateral view not distinct from rest of vestiture. HEAD. Forehead slightly wider than epifrons near antennal insertions, scales suberect. Eyes convex, in dorsal view distinctly exceeding outline of head, surrounded by a ring of short pale scales, on forehead directed towards occiput; distance between eye and scrobe slightly smaller than width of antennal club. Epifrons with distance between antennal insertions 0.33 × length of scape, scales at least 2 × as long as wide, recumbent, subcontiguous. Frons with 3 pairs of long erect lateral setae. Epistome without median seta. Antennal funicles with segments 1-2 elongate, subequal in length; 3-4 longer than wide; 5-6 globular; 7 wider than long.
PRONOTUM. Transverse (W:L ratio 1.35), widest near midlength, sides arcuate; apex and base subequal in width; with a depression at midlength on either side of dark median longitudinal stripe.
ABDOMEN. Ventrites with creamy-white plumose scales not concealing integument; scales on ventrites 2-5 medially intermixed with long suberect setae, bifi d or not at the apex; ventrite 1 slightly concave medially, impression covered with long setae deeply divided from their bases; ventrite 5 with scales concentrated in basal quarter.
TERMINALIA. Body of penis moderately elongate (W:L ratio 0.5), 2 × as short as temones, sides convex; in profi le straight, downcurved near apex, dorsoventrally narrowed at apex. Copulatory sclerite weakly sclerotised or not discerned in examined specimen. Parameroid lobes separate, divided by modest median notch, each lobe broad, bearing a series of setae directed apicad. Spiculum gastrale with basal arms long, regularly curved.

Sexual dimorphism
The sexes can be distinguished by the shape of the elytra (shorter in the male) and by ventrite 1 lacking the long and deeply divided setae in the female.

Life history
Specimens of Cryptolarynx endroedyi sp. nov. were collected in a fl owering meadow. No data about any host plant association of the species are available.

Differential diagnosis
Cryptolarynx oberlanderi sp. nov. is very similar to C. san sp. nov. and C. luteipennis sp. nov. but distinguishable from these by its narrower forehead and non-overlapping scales on the epifrons, and the apex of its parameroid lobes is also distinct among these species. Cryptolarynx oberlanderi was found to be genetically distinct from C. luteipennis and C. san by 8.1% and 15.1% for COI and by 3.7% and 4.0% for EF1, respectively.

Etymology
This species is dedicated to our colleague Kenneth Oberlander, specialist of the taxonomy of the genus Oxalis. Thanks to his substantial help with the location and identifi cation of species of this genus, numerous new species of Cryptolarynx were discovered, including this one. The specifi c epithet is a noun in the genitive case. COLOUR AND VESTITURE. Body integument black, scapes and tarsi reddish. Dorsal vestiture (pronotum + elytra) consisting of overlapping, recumbent, parallel-sided or subtriangular clothing scales, isodiametric to 3 × as long as wide, truncate at apex; colour of scales brown; dark brown scales forming a longitudinal stripe medially on pronotum and elytral interstriae 1-3, pale brown scales concentrated laterally on pronotum and on elytra laterally of interstria 4; white scales surrounded by black scales forming a pair of pale spots on elytral interstriae 2-3 at apical ¾ of elytral length; scales of striae recumbent, in lateral view not distinct from the of vestiture.
HEAD. Forehead wide, slightly wider than epifrons near antennal insertions, scales recumbent, not concealing integument. Eyes convex, in dorsal view slightly exceeding outline of head, surrounded by a ring of short scales, on forehead directed towards occiput; distance between eye and scrobe smaller than width of antennal club. Epifrons with distance between antennal insertions as large as length of scape, scales in middle of epifrons at most 2 × as long as wide, recumbent, non-contiguous. Frons with 3 pairs of long erect lateral setae. Epistome without median seta. Antennal funicles with segment 1 moderately elongate, 1.5 × as long as; 2 slightly shorter, 2 and 4 compressed, slightly angular on inside; 5-7 globular, isodiametric.
ABDOMEN. Ventrite with creamy-white plumose scales not concealing integument, scales on ventrites 2-5 medially intermixed with long suberect setae, apically bifi d or not; ventrite 1 slightly concave medially, impression covered with long setae deeply divided from their bases; ventrite 5 almost completely devoid of scales except for a series of small scales along suture, medially with a smooth area without setae or punctures.
TERMINALIA. Body of penis moderately elongate (W:L ratio 0.6), 2 × as short as temones, sides slightly convex, converging in apical quarter; in profi le straight, weakly downcurved and dorsoventrally narrowed at apex. Copulatory sclerite weakly sclerotised or not discerned in examined specimens. Parameroid lobes separate, divided by modest median notch, each bearing a series of setae directed apicad, longer medially, longest setae longer than depth of median notch. Spiculum gastrale with basal arms long, right arm slightly angulate near midlength.

Sexual dimorphism
The sexes can be distinguished by the elytra (longer in the female) and by abdominal ventrite 1 lacking long, deeply divided setae in the female.

Life history
The known specimens of C. oberlanderi sp. nov. were all found in July, at the base of plants of Oxalis depressa Eckl. & Zeyh.

Distribution
The species is only known from the Hex River valley and the Worcester area (Fig. 13).

Differential diagnosis
This species is most similar to C. luteipennis sp. nov. but distinguishable by the setae on ventrite 2 of the male (bifi d from the base in C. san sp. nov.; entire or only bifi d near apex in C. luteipennis). The shape of the body of the penis and the conformation of the apical setae on the parameroid lobes are also different between these species (Fig. 2V). Uncorrected genetic distances between them were found to be 16.3% for COI and between 3.5% and 4.1% for EF1.

Etymology
This species is dedicated to the San people, the fi rst inhabitants of southern Africa. This hunter-gatherer people left important traces of their activities (debris of bones, shells) on the coasts of the Western Cape province. The specifi c epithet is a noun in apposition. COLOUR AND VESTITURE. Body integument black, antennae, tibiae and tarsi reddish. Dorsal vestiture (pronotum + elytra) consisting of overlapping, recumbent, parallel-sided clothing scales, 2-3 × as long as wide, truncate at apex; colour of scales black, pale brown or white; black scales forming a medial longitudinal stripe over pronotum and on basal half of elytral interstriae 1-3; pale brown and white scales concentrated laterally on pronotum and on elytra laterally from interstriae 4; white scales forming a pair of pale spots on elytral interstriae 2-3 at apical ¾ of elytral length, these spots sometimes contiguous, confl uent and merged with pale areas laterally; scales of striae recumbent, in lateral view not distinct from rest of vestiture.

Holotype
HEAD. Forehead wide, distinctly wider than epifrons near antennal insertions, scales suberect, almost concealing integument. Eyes convex, in dorsal view slightly exceeding outline of head, surrounded by a ring of short pale scales, on forehead directed towards occiput; distance between eye and scrobe smaller than width of antennal club. Epifrons with distance between antennal insertions 0.33 × length of scape, scales in middle of epifrons at least 2 × as long as wide, recumbent, non-contiguous. Frons with 3 pairs of long erect lateral setae. Epistome without median seta. Antennal funicles with segment 1 elongate, 2 × as long as wide; 2 slightly shorter; 2 and 4 compressed, slightly angular on inside; 5-7 globular, isodiametric.
TERMINALIA. Body of penis elongate (W:L ratio 0.35), as long as temones, sides subparallel, slightly divergent from base to apex, strongly converging at apex; curvature in profi le weak and regular, dorsoventrally narrowed at apex. Copulatory sclerite weakly sclerotised or not discerned in examined specimens. Parameroid lobes separate, divided by modest median notch, each lobe bearing a series of setae directed apicad, longer medially, longest setae shorter than depth of median notch. Spiculum gastrale with basal arms moderately and regularly curved, each arm bearing a wide cuticular expansion.

Sexual dimorphism
The sexes can be distinguished by the elytra (longer in females) and by the centre of ventrite 1 (lacking long deeply divided setae in females).

Life history
Adults of C. san sp. nov. were collected in the vicinity of stands of various species of Oxalis (O. obtusa, O. luteola and O. hirta L.), but the exact host plant could not be confi rmed. Specimens were collected in July and September. The scales on the elytra can be white or pale brown, and their colour corresponds well with that of the soil where particular specimens were found. Compared with the other species of the genus, the adults of C. san can move very fast on the ground.

Distribution
The species occurs at various locations of the West Coast, in Bokpunt and near the West Coast National Park (Fig. 13).

Differential diagnosis
This species is most similar to C. san sp. nov.; see Differential diagnosis section under that species for diagnostic characters and genetic distances.

Etymology
The species name luteipennis refers to the orange or yellowish shades on elytra on many specimens of this species. These colours are seemingly unique to this species and provide an effi cient camoufl age of adults on the pinkish-orange sand on which they were found near Graafwater. The specifi c epithet is an adjective in the masculine form. COLOUR AND VESTITURE. Body integument black, antennae, tibiae and tarsi reddish. Dorsal vestiture (pronotum + elytra) consisting of overlapping, recumbent, parallel-sided clothing scales, 1.5-2 × as long as wide, truncate at apex; colour of scales black, dark brown to pale brown and yellow or orange; darkest scales forming medial longitudinal stripe over pronotum and on basal ⅔ of elytral interstriae 1-3; paler scales concentrated laterally on pronotum and on elytra from interstriae 4 laterad; white scales forming a pair of pale spots on elytral interstriae 2-3 at apical ¾ of elytral length; scales of striae recumbent, in lateral view not distinct from rest of vestiture.

REPUBLIC OF SOUTH
HEAD. Forehead wide, slightly wider than epifrons near antennal insertions, scales suberect, almost concealing integument. Eyes strongly convex, in dorsal view distinctly exceeding outline of head, surrounded by a ring of short scales, on forehead directed towards occiput; distance between eye and scrobe smaller than width of antennal club. Epifrons with distance between antennal insertions 0.33 × length of scape, scales at least 2 × as long as wide in middle, recumbent, non-contiguous. Frons with 3 pairs of erect lateral setae. Epistome without median seta. Antennal funicles with segment 1 elongate, globular, isodiametric; 7 wider than long.
ABDOMEN. Ventrites with creamy-white plumose scales not concealing integument, scales on ventrites 2-5 intermixed mostly medially with long suberect setae, bifi d from midlength or at least at apex; ventrite 1 slightly concave medially, devoid of scales, impression covered with long setae, deeply divided from their bases; ventrite 5 devoid of scales, bearing only erect setae.
TERMINALIA. Body of penis moderately elongate (W:L ratio 0.5), 0.66 × length of temones, sides convex; curvature in profi le weak, more strongly downcurved near apex, not dorsoventrally narrowed at apex. Copulatory sclerite weakly sclerotised or not discerned in examined specimens. Parameroid lobes separate, divided by modest median notch, each lobe bearing a series of setae directed apicad and converging. Spiculum gastrale with basal arms moderately curved, bearing a tooth near midlength.

Sexual dimorphism
The sexes can be distinguished by the shape of ventrite 1 (convex with deeply divided setae in male, concave with undivided setae in female).

Life history
Adults of C. luteipennis sp. nov. were collected in the vicinity of stands of various species of Oxalis (including O. obtusa), but the exact host plant of the species has not been identifi ed. All specimens were collected in July and August.

Undescribed species of Cryptolarynx Van Schalkwyk, 1966 left in abeyance
Among the specimens of Cryptolarynx recently collected or located in museums, six additional putative undescribed species were found, as based on morphological differences and/or genetic distance. These species are not described here as they were only represented by female specimens or because longer series are needed to identify reliable morphological features to distinguish them from similar species. To facilitate future research on this genus, the label data of these specimens, their depositories are reported here and links to their DNA sequences data in Appendix 2. Genus Hadrocryptolarynx Haran gen. nov. urn:lsid:zoobank.org:act:1983869F-4CDD-43AD-A2B0-CAEC1CED873B

Type species
Hadrocryptolarynx major Haran gen. et sp. nov., by present designation.

Differential diagnosis
Among the specimens of, or similar to, Cryptolarynx available for study, a series was found that differ from all species of Cryptolarynx in having a larger body size (length ca 6.0 mm; only ca 4.5 mm in the largest Cryptolarynx), a more elongate shape in the male (more or less globular in Cryptolarynx), elongate metatarsi with segment 2 at least 1.5 × as long as wide (shorter in Cryptolarynx) and apically fused and glabrous parameroid lobes of the aedeagus (divided and setiferous in Cryptolarynx). As already suggested by Oberprieler (2014), these differences are deemed signifi cant to assign these specimens to a different genus.

Etymology
This genus is named in reference to the remarkably larger size and bulk of its type species compared to that of species of Cryptolarynx. The name is a noun in the nominative singular case and its gender is masculine.

Description (♂)
MEASUREMENTS. Medium-sized, body length 2.0-6.0 mm. Body shape elongate in dorsal view, elytra and pronotum subequal in width. Pronotum widest at or slightly anteriorly of midlength; base narrower than elytra at humeri. Elytra widest near midlength. Body in lateral view (Fig. 3X) slightly hunched, highest just behind elytral base to middle of elytral length; head almost hypognathous.
COLOUR AND VESTITURE. Body integument black; antennae, tibiae and tarsi generally reddish. Dorsal vestiture (pronotum + elytra) consisting of short, recumbent clothing scales 2-5 × as long as wide, not aligned on interstriae, more or less concealing integument, colour ranging through black, dark brown to pale brown and grey to white; darker scales usually concentrated medially on pronotum and from there in broad median stripe along elytral interstriae 1-3; paler scales concentrated on a spot near middle of elytral interstriae 1-4. HEAD. Head capsule globose, in repose deeply retracted into prothorax, leaving only vertex and eyes visible in dorsal view. Eyes subcircular or slightly oval, convex, situated sublaterally, surrounded by a ring of pale recumbent scales. Forehead fl at, slightly narrower than width of an eye; median fovea absent. Rostrum very short and broad, not differentiated from head, dorsal surface perpendicular to longitudinal axis of prothorax when head in repose. Epifrons fl at, as wide as forehead, epifrontal scales suberect and orientated laterad, towards scrobes. Mandibles beak-like, densely setose (ca 20 setae each), with a pair of longer erect setae arising medio-laterally. Maxillae with galea and lacinia separate, both bearing apical setae (Fig. 8Q). Antennae inserted subdorsally at approximately middle of rostrum; scapes slender, as long as width of epifrons between antennal insertions, regularly and moderately curved, apically clavate and bearing erect setae, in repose folding into narrow scrobes extending onto underside of rostrum; funicles 7-segmented, longer than scapes, segment 1 elongate, funicles entirely hidden between head and cuticular anteroventral expansion of prothorax when head in repose; clubs 4-segmented, fusiform, acuminate, shorter than funicles.
THORAX. Pronotum moderately transverse; integument densely punctate, narrow spaces between punctures dull; anterior margin bisinuate, posterior margin arcuate, fi tting closely to elytral bases up to level of humeri. Prothorax anteriorly on each side produced into a large sharp-rimmed ventrolateral lamina extending from lower level of eye down to anterior edge of procoxa, concealing anterior prothoracic margin beneath it, rim of lamina asetose but anterior margin fringed with row of dense plumose scales, longer below eyes but shorter ventrally along prosternum. Prosternum broad, short, depressed below anterior edge of procoxae, declivous, abutting rostrum when head in repose; procoxal cavities medially confl uent, hypomeral lobes behind them short, suture of median junction obscure. Mesoventrite deeply depressed, almost vertically declivous, intermesocoxal process subtuberculate; mesepimera broadly separating mesanepisterna from elytral margin. Metaventrite between metacoxae as wide as metatarsus; metanepisterna fully fused to metaventrite, metanepisternal suture completely obliterated.

Paratypes
REPUBLIC OF SOUTH AFRICA -Western Cape • 1 ♂, 1 ♀; same collection data as for holotype; CBGP • 1 ♂, 1 ♀; same collection data as for holotype; NHMUK • 1 ♂; same collection data as for holotype; MNHN • 4 specs (preserved in ethanol); same collection data as for holotype; CBGP COLOUR AND VESTITURE. Body integument black, antennae, tibiae and tarsi reddish. Dorsal vestiture (pronotum + elytra) consisting of overlapping, recumbent, parallel-sided or bilaterally convex clothing scales, each 2-5 × as long as wide, truncate at apex, not aligned on interstriae, concealing integument. Colour of scales black, dark brown to pale brown, grey or white; darker scales usually concentrated medially over pronotum and on elytral interstriae 1-3; paler scales concentrated laterally and on pronotum and elytra, usually forming a transverse spot near middle of interstriae 1-3. Scales arising from strial punctures not different from those of rest of vestiture.
HEAD. Forehead slightly wider than epifrons near antennal insertions, scales recumbent. Eyes convex, in dorsal view distinctly exceeding outline of head, surrounded by a ring of short scales, on forehead directed towards occiput; distance between eye and scrobe less than width of an eye. Epifrons fl at, as wide as forehead, epifrontal scales suberect and orientated laterad, towards scrobes, overlapping. Frons with 3 pairs of long erect lateral setae. Epistome without median seta. Antennae with funicle segment 1 longer than wide, as long as 2 and 3 combined, 2-4 subcylindrical, slightly angular on inside, 5-7 globular or moderately elongate.
TERMINALIA. Body of penis elongate (W:L ratio 0.3), as long as temones, acuminate at apex, in profi le downcurved in apical third. Tectum narrow but distinct; endophallus with copulatory sclerite divided into two symmetrical, elongate structures. Parameroid lobes of dorsal plate of tegmen fused and jointly evenly rounded at apex, devoid of seta. Spiculum gastrale asymmetrical; divergence of basal arms V-shaped.

Sexual dimorphism
The sexes can be distinguished by the shape of the elytra (elongate and widest near midlength in male, shorter and widest at base in female) and by ventrites 1-2 (concave in male, convex in female), and half of females examined bear a series of long, suberect setae on interstriae 1-3-5-7. HABITUS. Body very robust, curved, widest at thorax and anterior abdominal segments; setae pallid and inconspicuous, the longer setae slender, fi ne; cuticle with minute asperities; posterior third of head capsule in repose partially retracted into pronotum.
STEMMATA. Present as two pigmented dark spots, anterior stemma larger than posterior one.
ANTENNAE. Situated on dorso-anterior margin of head; membranous basal segment bearing sensorium and eight smaller sensilla externally; sensorium broadly conical, about isodiametric or slightly longer than wide, circular in apical view.
MAXILLAE. Each stipe with 1 seta at base, 2 palpiferal setae and 1 minute seta plus sensillum near base of mala; maxillary malae with 4 vms, median 2 smaller, and with 6 elongated dms in a row; maxillary palps 2-segmented, 1 seta on basal segment, none on distal segment.
THORAX. Spiracles placed on pronothorax, oval, annular, without airtubes. Pronotum pigmented, not divided by median line, with 9 setae. Meso-and metathorax with 1 prs and 4 pds. Pedal area with 4 clearly distinct setae (t, u, v and w); the longest (v and w) placed on a lobe slightly distinct in the pedal area; some minute additional setae present (e.g., v´).
ABDOMEN. Spiracles of AI-VIII placed laterally, of similar size as thoracic spiracle, subcircular, without airtubes. Segments AI-VII with 3 dorsal folds, prodorsum and postdorsum both similarly prominent. AVIII with prodorsum and postdorsum distinct. AIX with dorsum somewhat expanded over AX (in mature larva). AX with anus subterminal; lateral anal lobes and dorsal anal lobe larger than ventral anal lobe.

Remarks
The smaller larva examined (Fig. 9B) may correspond to the second instar (body length 2.40 mm; head width 0.39 mm). It agrees with the last-instar larva in the characters described above (Fig. 10A-I), including the spiracles. Differences between early-and later-instar larvae involve relative dimension of structures, with the head (in relation to body), the antennae and stemmata being relatively larger in earlier instars, and the pigmentation and level of sclerotisation of body areas tend to increase in successive instars.

Molecular analyses
DNA-grade specimens were obtained for 18 of the 24 known species of South African Cryptolaryngini. Fragments of the COI and EF1 genes were successfully amplifi ed for, respectively, 16 and 17 species (Table 1). Preliminary PhyML analyses conducted on separate genes showed a consistent arrangement, with the C. vitis group forming a cluster nested with C. cederbergensis sp. nov., C. pyrophilus sp. nov. and C. sp. 2 (bt values: 91% and 84% for COI and EF1, respectively; Fig. 12). Relationships between other species or species groups were less stable between gene trees. Uncorrected p-distances between closely related species ranged from 10% to 20% for COI (Supp. fi le 1). At intraspecifi c level, distances ranged up to 10% for this DNA fragment between populations of C. vitis. In the C. vitis group, several closely related species showed incomplete phylogenetic sorting in their current concept in regard to the COI gene (C. vitis / C. subglaber; C. hirtulus / C. robustus). Although less densely sampled, relationships between these species were better resolved with EF1.

General life cycle
Weevils of Cryptolarynx and Hadrocryptolarynx gen. nov. are active in the austral winter, with adults encountered between the end of June and the middle of October and a peak of abundance of many species occurring in July and August. However, sporadic records of some species of Cryptolarynx also exist for February (C. vitis). In contrast, adults of Perieges in the Palaearctic Region are active during spring and autumn (April and September). Detailed observations of the life cycle are only available for C. variabilis sp. nov. (a population in Stellenbosch; JHAR01185). The fi rst adults there emerge in late June to early July and start their maturation feeding on aerial parts of surrounding vegetation. They remain mainly on the soil surface at the base of their host plants, becoming active and moving on the ground in open areas only during the hottest hours of the day. Mating was observed in early July, and gravid females with mature eggs were found at the end of July. Oviposition was observed in late September and occurs directly onto the immature and non-sclerifi ed bulbs of Oxalis pes-caprae and O. purpurea when the vegetative parts of the plants may begin to decline. Females reach the bulbs by digging vertical holes in the ground with their mandibles. In clayish soils, holes are distinct on the surface in the vicinity of stems of species of Oxalis. Larvae were observed in Oxalis bulbs from early December to March. Oviposition and early development of the larvae occur in such a way that the tunics of the bulbs are allowed to sclerify before weevil damage starts. As a result, attacked bulbs are intact from the outside and the larvae are sealed inside. Larvae entirely consume the fl eshy parts of the bulb, leaving no trace thereof. They do not produce any frass (Fig. 6K). The size of the last larval instar (and then of the resulting adult) depends on the size of the bulb, as also observed for other weevils that develop in limited substrates (de Medeiros et al. 2014). Variation in the size of Oxalis bulbs therefore explains the variability in the sizes of adult males and females of C. variabilis. After reaching the last instar, larvae undertake a period of inactivity during the summer. Pupation takes place inside the bulb around March (Fig. 6L). Mature larvae and freshly emerged adults of C. variabilis (Fig. 6K,M) are often found in the bulbs at this time, but pupae are rare, indicating a very brief pupal stage. Teneral adults of C. variabilis remain in the bulbs until emergence. Emergence takes place via a circular hole made in the basal or lateral parts of the bulbs (Fig. 6I). Adults of C. luteipennis sp. nov. newly emerged from bulbs were collected in late July and those of C. hirtulus sp. nov., C. squamulatus sp. nov. and C. armatus in August, suggesting that different species of Cryptolarynx may have chronologically specifi c and partly different life cycles.

Host plants
Cryptolarynx and Hadrocryptolarynx gen. nov. are closely associated with species of Oxalis. The weevils track the appearance and growth of their host plants by emerging at the base of the aerial parts early in the growing season in autumn and then being active during and after the fl owering season in winter and spring, and they spend the heat and drought of summer as immature stages in the protective environment of the subterranean bulbs of their host plants (Fig. 6I, K-L). Host plant species were established for C. variabilis sp. nov. and C. oberprieleri sp. nov., of which larvae, pupae and teneral adults were found in the bulbs of O. pes-caprae, O. purpurea and O. glabra. For the other species, species of Oxalis growing in close vicinity were only recorded and specifi c host associations could not be confi rmed. The fact that different species of Oxalis often occur in local sympatry makes it diffi cult to link particular species of Cryptolarynx with a specifi c host plant, especially given that the weevils may still be active when the aerial parts of their Oxalis host have already withered away. Adults of Cryptolarynx and Hadrocryptolarynx appear to be polyphagous, as they have been found feeding on leaves of several plant species in the surrounding vegetation, such as Vitis vinifera (C. vitis;, Moraea sp. (C. estriatus; Oberprieler et al. 2007), Poaceae (C. variabilis, C. vitis; pers. obs. JMH) and Arctotheca calendula (C. pilipes sp. nov.). They seem, however, to avoid consumption of the foliage of their own Oxalis larval host and would eat its leaves only in non-choice lab conditions or when all surrounding vegetation is senescent. The level of damage to Oxalis bulbs is variable. In C. variabilis, for example, between 5% and 25% of bulbs were attacked in the few sites investigated. The rearing of specimens from bulbs collected in the fi eld did not reveal any natural enemies such as parasitoids, but remains of adults were found in spider webs under stones. Bulbs, including those containing weevil larvae, are also intensively eaten by rodents during summer.

Distribution and habitat
Cryptolarynx and Hadrocryptolarynx gen. nov. are so far only known to occur in the Mediterraneanclimate and semi-arid regions of the Greater Cape Floristic Region in the Western and Northern Cape provinces of South Africa, with Cryptolarynx recorded from Namaqualand in the north to the Swellendam area in the south and inland to the Great Escarpment (Sutherland area) and Hadrocryptolarynx from Namaqualand to Vanrhynsdorp and Worcester (Fig. 13).
The habitats of Cryptolarynx and Hadrocryptolarynx gen. nov. correspond with those of their Oxalis hosts. Cryptolarynx species occur in Fynbos, Nama Karoo, Renosterveld and Succulent Karoo vegetation types and on various soil types, from clayish to sandy and from very fi ne to coarse sandstones and on fl at habitats from sea level to 1500 m on the Great Escarpment, but they seem to be absent from steep slopes and mountain tops. Hadrocryptolarynx major gen. et sp. nov. was recorded mainly in the Succulent Karoo biome, more rarely in the Fynbos biome, and on more or less coarse sandy soils from the seashore to about 60 km inland, always on plains. The species of both genera occur mainly in open habitats but sometimes also under dense forest canopy.

Behaviour
The adults of Cryptolarynx and Hadrocryptolarynx gen. nov. are diurnal, spending the day on the soil surface in the vicinity of their larval host plants (Fig. 7A, E). On open fi elds they start moving during the warmer hours of the day, occasionally climbing onto vegetation in large numbers for a few hours on hot afternoons. The adults of some species can run very fast on hot sands, possibly to avoid predation or heat (C. san sp. nov.; Fig. 7L). Around sunset they disappear from the soil surface to hide in little holes they form under debris, stones or leaves that effectively conceal the ground, such as those of Haemanthus L. (Amaryllidaceae) or Lachenalia J.Jacq. (Hyacinthaceae) (Hadrocryptolarynx major gen. et sp. nov., Fig. 7O). The broad, short and fl exible head with its fl at eyes and stout mandibles seems to be adapted for digging holes and tunnels into the soil. The weevils are well camoufl aged on the ground, their colour and vestiture largely matching the colour and texture of the soil substrate (Fig. 6A-B). When disturbed or threatened, they feign death and retract their head and appendages (Figs 6B, 7A, 7E), as many weevils do. When the danger recedes, they start moving again but with the antennae remaining hidden in the space between head and prosternum (Fig. 8C) and only the clubs protruding below the mouthparts. Seven months after a natural fi re near Montagu, a large migration of adults of C. pyrophilus sp. nov. (JHAR01528) took place from unburnt vegetation to the burnt areas, where plants of some species of Oxalis (mainly O. pes-caprae) were emerging. Oxalis plants often emerge and fl ower prolifi cally in the fi rst or second year after a fi re, leading to vigorous growth of the plants, which the weevils appear to be able to detect from a distance. Species of Cryptolarynx show a moderate to high tolerance to heat in the adult stage, when compared to other weevils (Javal et al. 2023). As larvae and teneral adults are enclosed in bulbs and cannot move, this tolerance has been proposed to be an adaptation to survive during the summer, when bulbs can be very close to the surface of the ground.

Species diversity
This study highlights the large diversity of Cryptolaryngini that exists in the southwestern parts of South Africa. Only two species were known since  original description of Cryptolarynx. Oberprieler et al. (2007), from an assessment of particularly the material collected by S. Endrödy-Younga and colleagues of the former Transvaal Museum in Pretoria during the 1970s and 1980s, estimated the number of species in the genus as 18, including an aberrant species recognised as representing a different genus (Oberprieler 2014) and here described as Hadrocryptolarynx major gen. et sp. nov. The discovery of the larval hosts of the South African Cryptolaryngini and focussed surveys of stands of Oxalis by the fi rst author of this paper (JMH) during 2018 and 2019 resulted in the recognition of 22 species as here newly described, which brings the number of described species of Cryptolaryngini in South Africa to 24, and another six are left undescribed for the moment. However, the level of endemism in this group and the rate of discoveries of new species over the two years of sampling by JMH suggest that the diversity of this tribe in southern Africa is far from being fully known. Oxalis comprises about 200 species in southern Africa, and even if Cryptolaryngini are restricted to the Greater Cape Floristic Region, as appears to be the case, their diversity may be considerably larger and their distribution extend northwards into southern Namibia and eastwards along the southern coast of the Western Cape province.
Cryptolarynx is remarkably homogeneous in habitus, but several stable morphological features of the males (mainly the genitalia and ventrite 1) provide reliable diagnostic characters to distinguish the species. Interspecifi c genetic distances based on mitochondrial (COI) and nuclear (EF-1) gene fragments were found to be consistent and corroborate the species distinctions derived from morphological characters. For two pairs of species, incongruences were found between morphological and nucleargene species delimitation, suggesting that these species as currently recognised may comprise multiple cryptic genetic lineages. Interspecifi c distances found between closely related species for COI (10-20%) are high for insects (Hebert et al. 2003) including beetles (Woodcock et al. 2013). At intraspecifi c level, distances ranged up to 10% for this DNA fragment in the current concepts of the species, suggesting that the commonly applied "barcode-gap" to defi ne species genetically is inapplicable in this group. It is likely that the fl ightless condition of Cryptolaryngini enhances the intraspecifi c and probably also the interspecifi c genetic divergences by geographical isolation and drift of populations and species. It should be noted that Cryptolarynx species in the Cape Town area occur mainly in fl at Renosterveld patches that are isolated from each other by unsuitable Fynbos patches. In addition, the frequent cycles of natural fi res in this area may strengthen the barrier effect of Fynbos patches for such small and fl ightless insects.
At most localities, several species of Cryptolarynx were found together in stands of multiple Oxalis species. This suggests that some species divergence may have occurred in alignment with larval host species. However, at least one species (C. variabilis sp. nov.) was found to be able to complete its larval development on two co-occurring species of Oxalis, O. purpurea and O. pes-caprae, and host records suggest that an Oxalis species can serve as larval host for several species of Cryptolarynx, depending on geographical location. Thus, it is unclear to what extent host plant divergence may have driven speciation in these weevils, but the system represents an interesting model to explore the processes of diversifi cation of phytophagous insects in the Greater Cape Floristic Region. Due to their local diversity, abundance and limited dispersal, the South African Cryptolaryngini also represent potential candidates to estimate the conservation value of Renosterveld patches from an arthropod perspective.

Systematic position of Cryptolaryngini
Due to their pedotectal male genitalia, the Cryptolaryngini phylogenetically fall outside the "higher" Curculionidae (Oberprieler 2014;Shin et al. 2018), i.e., the CEGH and CCCMS clades (Gunter et al. 2016). The "lower" Curculionidae comprise a clade formed by the subfamilies Dryophthorinae Schönherr, 1825 and Platypodinae Shuckard, 1840 and a range of taxa of uncertain relationships to each other (Shin et al. 2018), which retain the plesiomorphic condition of the pedotectal aedeagus and lack the synapomorphies that defi nes the clade of "higher" Curculionidae. Because of the widely varying and confl icting classifi cations of these taxa, Oberprieler (2004) combined the tribes with pedotectal aedeagus into a single subfamily Brachycerinae (sensu lato), though acknowledging that this likely constitutes a paraphyletic grade (Oberprieler et al. 2007). Oberprieler (2014) discussed the chequered classifi cation history of these taxa, in particular of the Erirhinini, in greater detail and recognised seven tribes in Brachycerinae: Brachycerini Billberg, 1820, Cryptolaryngini, Erirhinini Schoenherr, 1825, Himasthlophallini Zherikhin, 1991, Tanysphyrini Gistel, 1856, Myrtonymini Kuschel, 1990and Raymondionymini Reitter, 1913, with Bagoini Thomson, 1859 possibly constituting an eighth. Molecular phylogenetic analyses of weevils undertaken since (Gillett et al. 2014(Gillett et al. , 2018Gunter et al. 2016;Mugu et al. 2018;Shin et al. 2018;Baird et al. 2021) affi rmed the non-monophyly of Brachycerinae sensu lato, but have not resolved this conundrum mainly because of insuffi cient taxon sampling, even though several key genera, such as Brachycerus Olivier, 1789, Ocladius Schoenherr, 1825, Echinocnemus Schoenherr, 1843 and Bagous Germar, 1817 have been included in most of them. As a result, and because Cryptolaryngini have never been included in any large-scale molecular phylogenetic analysis, the phylogenetic position and classifi catory status of this tribe remain indeterminate. Preliminary results of a study in progress, analyzing the phylogeny of this group using nuclear ribosomal markers and mitochondrial COI, corroborate the placement of Cryptolarynx among the earliest-divergent curculionids and suggest a close relationship with Stenopelmus Schoenherr, 1835 (Marvaldi et al. 2022) itself of uncertain position (Erirhininae, Stenopelmini sensu Alonso-Zarazaga & Lyal (1999); Tanysphyrini sensu Oberprieler 2014).
Resolution of this issue is beyond the scope of the present study, but its assessment of a much larger number of species and especially of the larval and pupal characters of Cryptolarynx enables it to make an additional contribution to the debate.
Cryptolarynx was initially considered as unique and unrelated to other weevil groups due to its globular body shape and the head and short rostrum being retractable into a large ventral recession in the prothorax ) marked by prominent lateral fl anges abutting onto the procoxae. Such a short, broad ventral recession also exists in Perieges as well as in some other genera of Brachycerinae s. lat. with a relatively short rostrum, such as Brotheus Stephens, 1829, Byrsops Germar, 1829 and Synthocus Schoenherr, 1842 of Brachycerini, and comparable recessions of the prosternum also occur in numerous brachycerine genera with a longer rostrum, varying from broad and shallow with distinct lateral carinae, as in Raymondionymini, to narrower and deeper and forming a rostral canal that may reach posteriad to between the mesocoxae, as in the Ocladius group of Erirhinini (Oberprieler 2014). The prosternal recession of Cryptolarynx is unusual in that the row of dense setae that brush over the eyes when the head in retracted is not located on the rim of the lateral fl anges but much deeper inside, so that the fl anges do not represent the anterior prothoracic margin, whereas in other brachycerines with a ventral prothoracic recession (including Perieges) these setae are located on the rim or just inside and usually on a pronounced ocular lobe of the prothoracic margin. A similar condition to that of Cryptolarynx occurs in Bagoinae, however, although the lateral fl anges in this taxon do not directly abut onto the procoxae.
With the larger number of South African Cryptolaryngini now known, some differences between Cryptolarynx and Perieges as reported by Oberprieler (2014) no longer hold: the number of setae on the mandibles ranges from 4 to 8 in Cryptolarynx and is ca 20 in Hadrocryptolarynx gen. nov. (4 placed, maxillary mala with 6 elongated and slightly curved dms labial palps 1-segmented (also in some species of Bagous and some Tanysphyrini); the proximal pair of postmental setae of labium (pms1) more widely apart than pms2,3 (also in Ocladius). Their pupae also show similarities in general appearance and structure, i.e., the posterior processes being very small and pointed with a sclerotised apex and the abdominal segments lacking setae on the ventrolateral areas.
Regarding life history traits, this study found that Cryptolarynx larvae develop endophytically in the bulbs of species of Oxalis. This lifestyle resembles that of Brachycerini, whose larvae develop in the bulbs or tubers of geophytes or stems of aloes (e.g., Louw 1990;May 1993;Howden 1995;Germann 2003;Oberprieler 2014). The life history of Brachycerus differs from that of Cryptolarynx in that the mature larvae leave the bulb and pupate in an earthen cell in the soil (Louw 1990;Germann 2003), but those of Synthocus (and perhaps other genera of Byrsopina) seem to pupate inside the bulbs as adults of S. ornithoglossi Marshall, 1956 have been found in the bulbs of Ornithoglossum (Marshall 1956). The larval hosts of Brachycerini also differ from those of Cryptolarynx in being exclusively monocotyledonous plants (Liliopsida) (Oberprieler 2014), phylogenetically unrelated to the eudicotyledonous Oxalidaceae, and the subterranean bulbs of Oxalis are only superfi cially similar storage organs and not homologous with monocot bulbs.

Potential biological control of invasive Oxalis
Several South African species of Oxalis have been introduced overseas and become invasive weeds. Oxalis pes-caprae is listed among the ten worst invasive species in Europe (Vilà et al. 2010) and has an ecological and economic impact on invaded ecosystems (reviewed in McLaughlan et al. 2014). Control of weedy species of Oxalis is diffi cult and largely ineffective (Marshall 1987), and only one potential biological control agent has so far been identifi ed, a moth whose larvae feed on the aerial parts of the plants (Kluge & Claassens 1990). We here record the fi rst phytophagous insects feeding on the bulbs of species of Oxalis including O. pes-caprae, weevils of the genera Cryptolarynx and Hadrocryptolarynx gen. nov. As the feeding of their larvae results in the destruction of the bulbs of the plants, they have the potential to control the vegetative proliferation, which is the main reproductive means of species of Oxalis in invaded areas. Further, available biological data indicate that the larvae of Cryptolarynx and Hadrocryptolarynx are specialised feeders of Oxalis and have no other natural host plants, which makes these weevils potentially suitable candidates for biological control of weedy Oxalis species outside of their native ranges. The adult weevils, however, have the potential to feed on various other plants, including grapevine (Vitis vinifera) and are consequently regarded as quarantine pests in some countries, e.g., Israel.