Coniophloeus, a new genus of Embrithini from South Africa, with description of four new species (Coleoptera: Curculionidae: Entiminae)

A new genus, Coniophloeus gen. n., is described for fi ve species of South African weevils known from the Western and Eastern Cape and assigned to the tribe Embrithini Marshall, 1942: Coniophloeus squalidus (Boheman, 1842) comb. n. (type species, transferred from the genus Trachyphloeus Germar, 1817 and tribe Trachyphloeini Lacordaire, 1863), C. alternans sp. n., C. oberprieleri sp. n., C. robustus sp. n. and C. obrieni sp. n. All fi ve species are illustrated and keyed. ZooBank Article LSID: F6DEACAD-1127-4A2D-8E9C-6923069EF96F


INTRODUCTION
The tribe Embrithini Marshall, 1942, as redefi ned by Borovec & Oberprieler (2013), currently comprises 67 genera. Almost all known species of Embrithini are arboricolous or fl oricolous, active during the day on plants, shrubs or trees. There are some exceptions only in several South African embrithine genera, whose species live among litter under plants, such as some undescribed species of Cycliscus Schoenherr, 1842 and Glyptosomus Schoen herr, 1847 as well as Ellimenistes humeralis Marshall, 1947 and three similar undescribed species. The only described genus in which all the species are terricolous is Afrophloeus Borovec & Oberprieler, 2013, which comprises three South African species, one of them introduced to Australia. However, there are several other terricolous species among as yet unidentifi ed material, which are also assignable to Embrithini but can not be accommodated in any described genus, for example those described from the Afrotropical region in the Palaearctic tribe Trachyphloeini (Borovec & Skuhrovec, 2017). One such species is Trachyphloeus squalidus Boheman, 1842, which is here removed from Trachyphloeus Germar, 1817 and Trachyphloeini Lacordaire, 1863. The description of a new genus and discussion of its taxonomic position are the aims of the present paper.
The change in lifestyle from fl oricolous or arboricolous to living in soil or litter have to include also some morphological changes to the body, mainly shortening of extremities, which can be also more robust, diminution in clay; MNBG -Museum für Naturkunde der Humboldt Universität, Berlin, Germany, Manfred Uhlig; NHRS -Naturhistoriska Riksmuseet, Stockholm, Sweden, Johannes Bergsten; SANC -National Collection of Insects, Pretoria, South Africa, Riaan Stals; SDEI -Senckenberg Deutsches Entomologisches Institut, Müncheberg, Germany, Lutz Behne; TMSA -Ditsong National Museum of Natural History (formerly Transvaal Museum), Pretoria, South Africa, Ruth Müller.

TAXONOMY
Genus Coniophloeus gen. n.  ZooBank taxon LSID: 8D5D314D-3A3B-43AB-BCC7-CACD2D62778D Solakryl BMX (epoxy resin soluble in toluene; Medika, Prague) and male genitalia were mounted dry on the same card as the respective specimen, with tegmen and sternite IX embedded in Solakryl BMX. Habitus images were taken with a Canon EOS 5D Mark II camera fi tted with a Canon MP-E65 1-5 × macro lens. Images were stacked using the program Zerene Stacker and edited in Adobe Photoshop CC 2015.
The terminology used to describe the rostrum and the genitalia follows Oberprieler et al. (2014).
Exact label data of type material are cited; with different labels indicated by a slash (/). Author's remarks and comments are given in square brackets.

Type species. Trachyphloeus squalidus Boheman in
Description . Body length 3.2-5.6 mm. Body (Figs 1-5) dark brownish to blackish, in some species tarsi or only spines at apex of tibiae, claws and antennal club brownish. Dorsal and ventral part of body densely covered with oval to rounded, small, appressed scales, 6-8 scales across width of one interstria, scales longitudinally fi nely striate, dense, hiding integument or partly isolated. Scapes, funicles, femora, tibiae and tarsi densely covered by identical appressed scales, only clubs shortly and densely setose. Elytra with each interstria having 1-2 dense rows of subspatulate, conspicuous, appressed to semi-erect setae; pronotum, head with rostrum, antennae except club and legs with similar but more slender and shorter, densely irregularly scattered setae, semi-appressed to semi-erect.
Rostrum (Figs 6,7,12,13,18,19,22,23,31, 32) 1.1-1.8 × wider than long, apical half rounded; in lateral view distinctly convex. Epifrons at base as wide as or narrower than distance between eyes, tapering anteriorly with concave and somewhat swollen edges, longitudinally deepened, posteriorly separated from head by transverse shallow to deep sulcus, often covered by vestiture. Epifrons beneath scales longitudinally distinctly and regularly impressed, in transverse profi le V-shaped, its posterior corners in dorso-lateral view distinctly prominent, creating blunt notches; base of epifrons arched or straight, sharply, tooth-like constricted to form transverse and sometimes a very deep furrow separating it from head. Frons short, squamous, bearing 3-4 pairs of stout setae. Epistome moderately large, V-shaped, posteriorly sharply and narrowly carinate, not reaching anterior border of scrobes, not squamous. Antennal scrobes in dorsal view scarcely or partly visible, if visible then reniform; in lateral view short, weakly curved, directed just towards eyes, separated from anterior margin of eye by slender or wide squamous stripe. Vertex fl at and wide, beneath scales with numerous, sometimes ill-defi ned longitudinal striae radiating from middle of base of epifrons. Head in lateral view fl at, distinctly obliquely declivous to transverse furrow separating head from epifrons. Eyes small, circular in outline, convex, weakly prominent from outline of head, in lateral view placed at middle of head height. Mandibles small, not squamous, trisetose but most of specimens without setae, which were possibly abraded; in two specimens with setal remnants. Submentum with two long setae.
Antennae robust, with scape (Figs 6,12,18,22,31) moderately to very robust, wide, slightly exceeding anterior margin of pronotum when at rest, longer than funicle, in some species strongly S-shaped at base, enlarged apically, widest at apex or at anterior third, at apex wider than club and equally as wide as protibiae at midlength; funicle 7-segmented, funicular segments wide and short, segments 1 and 2 conical, the others transverse; club small, in some species barely wider than last funicular segments.
Etymology. The genus name is derived from the Greek nouns kónis (dust), referring to the dusty appearance of the body, and the Greek word phloio (bark) as used in the compound name of the morphologically superfi cially similar genus Trachyphloeus, the name under which most of the specimens considered here were originally deposited in collections. The name is masculine in gender.

Biology.
Unknown, there are no data where collected except for O. robustus sp. n., which was sifted from forest litter.
Distribution. The genus is known only from South Africa (the Western and Eastern Cape provinces).
Remarks. This genus includes fi ve species, four of them newly described below. The species of Coniophloeus resemble large species of Trachyphloeus or other Trachyphloeini, but are easily distinguishable mainly by the wide, squamous metatibial corbels and from Trachyphloeus by the distinct transverse sulcus between head and rostrum (rostrum continuous with head in Trachyphloeus), connate tarsal claws (free claws), tegmen with parameres (lacking parameres) and sternite VIII with apodeme terminating inside plate with ill-defi ned posterior border (apodeme creating distinct basal margins of plate). The fi rst three characters (presence of corbel, sulcus between head and rostrum and tarsal claws connate in basal third), combined with the trisetose mandibles and dorso-laterally placed scrobes, indicate that the newly described genus belongs to the tribe Embrithini.
Within the Embrithini Coniophloeus gen. n. and Afrophloeus are the only Embrithini with short and robust antennae and very wide and short scapes, short and robust tibiae and laterally enlarged protibiae with apices armed with moderately long spines. Coniophloeus gen. n. can be distinguished from Afrophloeus by the following characters: Afrophloeus: Apex of protibiae rounded with single laterally prominent spine. Elytra with small subhumeral tubercles visible in dorso-lateral view. Eyes fl at. Metaventral process 1.5 × as wide as transverse diameter of metacoxa. Suture between ventrite 3, 4 and 5 straight. Tegmen lacking parameres. 2.3-3.3 mm.
Coniophloeus gen. n.: Head and rostrum in profi le clearly separated by transverse furrow. Sulcus between head and rostrum in dorsal view with ill-defi ned margins. Eyes without supraocular tubercles. Protibiae laterally enlarged, apically armed with conspicuous spines, united with inner, middle and outer groups, outer spines connected. Ventrite 1 twice as long as ventrite 2; ventrite 2 as long as ventrite 3 and 4 combined. 3.2-5.6 mm.
Coniophloeus differs from Heisonyx, Lalagetes and Phaylomerinthus by the following characters: epifrons posteriorly separated from head by a wide transverse sulcus with ill-defi ned borders (clearly defi ned narrow sulcus in Heisonyx, Lalagetes and Phaylomerinthus); frons densely squamous (glabrous); scapes short and robust (long and slender); funicular segments 3-7 short and transverse (long and slender); clubs barely wider and hardly separated from last funicular segments (distinctly wider and well separated from last funicular segments); pronotum short and wide, at least 1.7 × wider than long (narrower, at most 1.5 × wider than long); apex of protibiae widened laterally, apically with distinct spines (straight laterally, apically with fringe of setae or short spines) and metatibiae with broad, distinct corbels (corbels slender, sometimes barely visible). Redescription. Body length 3.66-4.19 mm. Body (Fig.  5) dark brownish to blackish, only clubs and tarsi paler, brownish; incrustate with indistinct appressed vestiture of oval scales except clubs and tarsi, 5-6 scales across width of one elytral intestria; setae on elytra semi-erect, conspicuous, long, moderately slender, subspatulate, in regular row on interstriae, slightly longer than half width of one interstria, intervals between setae slightly larger than setal length; setae on pronotum similar but densely irregularly scattered, semi-appressed and slightly broader; on head and rostrum similar but semi-appressed and distinctly shorter; femora, tibiae, scapes and funicles with very slender, long, semi-erect setae.
Rostrum (Figs 31, 32) 1.11-1.14 × wider than long, in basal half with sides concave, then widely rounded around antennal insertions, at apical third equally wide as at base. Epifrons narrow, 0.6 × as wide as rostrum in middle, with sides slightly concave, at base distinctly narrower than space between eyes, weakly longitudinally depressed; posteriorly V-shaped, separated from head by deep transverse furrow; beneath scales weakly shiny, fi nely sparsely granulate. Vertex irregularly and fi nely longitudinally striate, striae radiating from middle of base of epifrons. Scrobes dorsally reniform, distinctly visible in anterior half; laterally slender and short, narrow, weakly curved, directed to middle of eye, separated from it by wide squamous stripe. Eyes small, weakly convex, dorsally scarcely prominent from outline of head; laterally circular, placed in middle of head. Head wide, widest at base, distinctly tapering anteriorly with slightly convex sides, interocular space very wide.
Remarks. This species is distinguishable from all others except C. alternans sp. n. by its long semi-erect elytral setae, dorsally clearly visible scrobes, long rostrum, basally straight scapes and glabrous apical surface of the metatibiae, and from C. alternans sp. n. by elytral setae on all intervals. Description. Body length 3.25-3.78, holotype 3.25 mm. Body ( Fig. 1) dark brownish to blackish, only clubs somewhat paler, brownish; incrustate with indistinct appressed vestiture. Setae on elytra semi-erect, conspicuous, long, moderately slender, subspatulate, in regular row only on odd interstriae, each seta slightly longer than half width of one interstria, intervals between setae somewhat larger than setal length; setae on pronotum similar but semi-appressed and slightly wider, densely irregularly scattered; on head and rostrum similar, but semi-appressed and distinctly shorter; on femora, tibiae, scapes and funicles very slender, long, semi-erect.
Rostrum (Figs 6, 7) 1.05-1.08 × wider than long, basal half with concave sides, then widely rounded around antennal insertions, in apical third equally wide as at base to 1.05 × wider. Epifrons narrow, 0.6 × wider than rostrum in the middle, with sides slightly concave, at base distinctly narrower than space between eyes, weakly longitudinally depressed; posteriorly U-shaped, separated from head by deep transverse furrow, beneath scales weakly shiny, indistinctly sparsely granulated; vertex irregularly and fi nely longitudinally striate, striae radiating from middle of base of epifrons. Scrobes dorsally reniform, clearly visible on anterior half; laterally slender and short, weakly curved, directed to ventral border of eye, separated from it by wide squamous stripe. Eyes small, weakly convex, dorsally scarcely prominent from outline of head; laterally circular, placed in middle of head. Head wide, widest at base, distinctly tapering anteriorly with sides slightly convex, vertex very wide.
Etymology. The Latin name refers to the raised elytral setae, forming rows only on alternate odd interstriae.
Remarks. The holotype lacks the right funicle with club, left mesotarsus and the whole right hind-leg. This species differs from all others except C. squalidus by the long semi-erect elytral setae, the dorsally clearly visible scrobes, a longer rostrum, the basally straight scapes and the glabrous apical surface of the metatibiae, and from C. squalidus by having semi-erect elytral setae only on the odd interstriae. Description. Body length 4.19 mm. Body (Fig. 2) dark brownish, only clubs slightly paler brownish, except clubs and tarsi with appressed small rounded scales, leaving distinct interspaces, 6-8 scales across width of one interstria; on elytra semi-appressed, in dense regular row on each interstria, subspatulate, widest before tip, fi nely longitudinally striate, about as long as half the width of one interstria, intervals between setae slightly larger than setal length; on pronotum similar but wider, densely irregularly scattered; on head and rostrum similar but distinctly shorter; on scapes, femora and tibiae very dense, semi-appressed, as long as setae on head with rostrum; on funicles more slender.

Coniophloeus oberprieleri
Rostrum (Figs 12, 13) wide and short, 1.79 × wider than long, widest at base, here 1.09 × as wide as at apex, basal half regularly tapering anteriorly, apical half regularly rounded. Epifrons moderately narrow, at base slightly narrower than space between eyes, with sides concave, in middle 0.6 × wider than the base of the rostrum; beneath scales moderately shiny and roughly irregularly punctate, with posterior border almost straight, separated from head by deep transverse furrow; head longitudinally striate, striae radiating from middle of base of epifrons. Scrobes not visible dorsally; laterally short and wide, curved, directed towards eyes and separated from them by narrow squamous stripe. Eyes small, convex, dorsally clearly visible, prominent from outline of head; laterally circular, placed at middle of head. Head short and wide, weakly tapering anteriorly, laterally distinctly obliquely declivous to base of rostrum. Vertex wide.
Etymology. This newly described species is dedicated to my friend and colleague Rolf Oberprieler (CSIRO, Australia), collector of the holotype and outstanding expert on South African weevils.
Remarks. C. oberprieleri sp. n. is easily distinguishable from all other species of the genus mainly by having the outer protibial margin enlarged to form a laterally prominent rounded projection, the pronotum 2.2 × wider than long, as wide as the elytra, widest at the base and with the posterior corners pointed, elongate posteriorly. Description. Body length 4.63 mm. Body (Fig. 3) dark brownish; except for the tarsi covered with small, irregularly angular appressed scales leaving slender interspaces, 7-8 scales across width of one elytral interstria; on elytra semi-appressed, subspatulate, widest before tip, fi nely longitudinally striate, slightly shorter than half width of one interstria, in 1-2 irregular dense row(s), intervals between setae slightly longer than setal length; on pronotum similar but slightly wider, very densely irregularly scattered; on head and rostrum similar, but shorter; on scapes, femora and tibiae slender, dense, subspatulate, semi-appressed.
Rostrum (Figs 18, 19) 1.35 × wider than long, widest at base, regularly tapering anteriorly with sides straight, at base 1.08 × as wide as at apex. Epifrons widest at base, here as wide as space between eyes, regularly tapering anteriorly with sides slightly concave, in middle 0.7 × wider than the width of the middle of the rostrum; beneath scales roughly punctate, longitudinally deepened with narrow median longitudinal stria, with base weakly arched, separated from head by deep transverse furrow; head shiny, distinctly longitudinally striate, striae radiating from middle of base of epifrons. Scrobes not visible dorsally; laterally curved, moderately short, distinctly enlarged posteriorly, posterior part as wide as diameter of eye, directed towards eyes and separated from them by narrow squamous stripe. Eyes small, fl at, dorsally scarcely visible; laterally placed at middle of head. Head short and wide, tapering anteriorly; laterally declivous to base of epifrons; interocular space wide.
Outside and inside of the apex of protibiae (Fig. 20) enlarged, armed with 4 connected yellowish brown spines on outer lobe and 9 isolated spines of different sizes on anterior border, plus brownish curved short mucro. Apical surface of metatibiae densely squamous. Tarsi with segment 2 1.3 × wider than long; segment 3 1.4 × wider than long and 1.4 × wider than segment 2; onychium short, 1.2 × longer than segment 3; claws shortly fused at base, moderately divergent, brownish. Penis (Fig. 21) ventrally long and slender, with base and apex similarly wide with weakly concave sides, apically rounded with short, subtrapezoidal tip; laterally wide with ventral surface straight and dorsal surface curved, regularly tapering apically with tip pointed.
Female genitalia unknown.
Etymology. The new species is named in honour of the eminent entomologist Charles W. O'Brien (Phoenix, Arizona, USA).
Remark. The holotype lacks both funicles and clubs, the complete left anterior and posterior legs and tarsus of the right middle leg. Its locality was explained to me by Riaan Stals and Elisabeth Globbelaar (SANC), as: "The Hex River Valley in the current Cape Winelands District Municipality, Western Cape Province; in 1885 located in the colony of the Cape of Good Hope. The only settlement in the Hex River Valley is the small town of De Doorns [33°28´39˝S, 19°40´01˝E], which already existed in 1885." Coniophloeus obrieni sp. n. is easily distinguishable from all other species of the genus mainly by the pronotum being widest in the middle, the laterally fl at elytra with straight sides and the eyes barely visible dorsally. Description. Body length 3.78-5.56 mm, holotype 4.63 mm. The whole body (Fig. 4) dark brownish to blackish, only clubs slightly paler, brownish, except for clubs and tarsi covered with small, irregularly angular appressed scales, leaving very slender interspaces, 7-8 scales across width of one interstria; setae on elytra semi-appressed, fi nely longitudinally striate, widest at apex, slightly shorter than half width of one interstria, in one regular row on each interstria, intervals between setae about twice the setal length, on disc slender, subspatulate, in posterior declivity twice as wide as those on disc, spatulate; on pronotum, head and rostrum similar, but densely irregularly scattered, on pronotum as long as on elytra, on head and rostrum shorter; on scapes, femora and tibiae similar, on funicles appressed, more slender.
Rostrum (Figs 22, 23) very wide and short, 1.53-1.67 × wider than long, weakly regularly tapering anteriorly. Epifrons wide, at base almost as wide as space between eyes, in basal half distinctly tapering anteriorly, in apical half subparallel-sided, with sides distinctly concave; beneath scales moderately shiny, irregularly roughly punctate, longitudinally shallowly deepened with posterior border arched with short concavity at middle, separated from head by deep transverse furrow. Head irregularly longitudinally roughly striate, at middle of anterior part with small and deep triangular incision. Scrobes dorsally almost not visible; laterally short and wide, subtriangular, distinctly curved, posteriorly about as wide as diameter of eye, directed towards eye, separated from them by wide squamous stripe. Eyes small, weakly convex and weakly prominent from outline of head; laterally circular, in middle of head. Head wide and short, weakly tapering anteriorly, interocular space very wide.
Antennae extremely robust; basal third of scapes ( Fig.  22) conspicuously S-shaped, apical two thirds weakly curved, widest at apical third and here twice as wide as clubs and about as wide as middle of protibiae; funicle with segments 1 and 2 obconical, the others transverse; segment 1 1.6-1.7 × longer than wide and 1.1 × longer than segment 2, which is 1.4-1.5 × longer than wide, segments 3-4 1.5 × wider than long, segments 5-7 1.7-1.8 × wider than long; clubs short, 1.3 × longer than wide, about as wide as last funicular segments. Pronotum (Fig. 4) 1.64-1.83 × wider than long, basal half the widest, here subparallel-sided with slightly rounded sides, apical half conspicuously constricted; disc regularly convex; base weakly arched; in lateral view weakly convex. Elytra (Fig. 4) long-oval, 1.20-1.31 × longer than wide, with sides weakly rounded, laterally convex; at posterior declivity with small, sometimes indistinct tubercles, in some specimens bigger on odd intervals than on inner ones.
Wide outer and inner sides of apices of protibiae (Fig.  24) armed with 15-20 yellowish short spines, on outer and middle part aggregated in two groups, on inner part isolated, with short brownish mucro at inner corner. Apical sur-face of metatibiae densely squamous. Tarsi with segment 2 1.7-1.8 × wider than long; segment 3 1.5 × wider than long and 1.2-1.3 × wider than segment 2; onychium moderately long, 1.5 × longer than segment 3; claws shortly fused at base, moderately divergent, brownish. Penis (Fig. 25) ventrally short, widest at base, tapering apically with slightly concave sides, apex rounded with very short, obtuse tip; laterally wide, ventral surface straight, dorsal surface of anterior third distinctly enlarged, tip distinctly extended.
Female genitalia. Spermatheca (Fig. 28) with cornu almost straight; corpus long; ramus about 1.5 × longer than wide, apically rounded; nodulus twice a long as ramus, straight. Gonocoxites (Fig. 30) subtriangular, evenly tapering apically, with short apical styli. Sternite VIII (Fig. 29) with plate 1.4 × wider than long; apodeme terminating on apical half. Remarks. This is the largest species in this genus, similar only to C. obrieni sp. n. in its epifrons being basally as wide as the space between the eyes Distinguishing characters between these two species are cited in the key to species.