A revision of the Palaearctic Pimeliini (Coleoptera: Tenebrionidae): a comparative analysis and systematic position of Eastern European and Asian taxa with dorso-lateral eyes

. A taxonomic review of tenebrionid platyopoid genera of the subfamily Pimeliinae from Eastern Europe, Central Asia, Afghanistan, Iran and Pakistan is given. This group of taxa was known before 1994 as the tribe Platyopini Motschulsky, 1849, which is now interpreted as a junior synonym of Pimeliini Latreille, 1802. The group is different from other Pimeliini in having dorso-lateral eyes, located above the level of the genae, and it includes the following ultrapsammophilic genera at least from Central and Southern Asia: Apatopsis Earophanta 1903. These genera are distributed in almost all large deserts of Palaearctic Asia: Karakum, Kyzylkum, Muyunkum, Taklamakan, Gobi, Registan, Dasht-e-Kawir, Dasht-e-Lut, as well as in other arid and semi-arid sandy landscapes from European Russia to the south of Eastern Siberia. The group of platyopoid genera is polyphyletic. We propose at least two monophyletic branches: the Habrobates genus group (the ﬁ rst four genera mentioned above), which represents the subtribe Bogatchev, 1957 (holotype). 2022. A revision of the Palaearctic Pimeliini (Coleoptera: Tenebrionidae): a comparative analysis and systematic position of Eastern European and Asian taxa with dorso-lateral eyes. European Journal of Taxonomy 809: 1–71. https://doi.org/10.5852/ejt.2022.809.1719 long, very Spiculum common with strongly sub-angularly curved apical piece. revision of the Pimeliini (Coleoptera: Tenebrionidae)


Introduction
Pimeliini is a large tenebrionid tribe in the subfamily Pimeliinae Latreille, 1802 with 64 genera distributed mainly in the Palaearctic and transition biogeographic zones of the Afrotropic and the Indo-Malayan regions (Gebien 1937;Iwan et al. 2020). The majority of genera is distributed in deserts of Central Asia, Iran, Pakistan, Afghanistan, the Near East and North Africa. Many authors hypothesised that the ecological diversifi cation of the tribe took place since the Pleistocene (Kryzhanovsky 1965) or no later than the Pliocene (Medvedev 2005;Abdurakhmanov & Nabozhenko 2016;Abdurakhmanov et al. 2016a;Abdurakhmanov et al. 2016b) in Central Asia. Kwieton (1981) proposed an Iranian origin of Pimeliini.
The classifi cation of genera within the tribe is discussed by Semenov (1893), Skopin (1962), Pierre (1964), Koch (1969), Kwieton (1981), Kaszab (1982) and Doyen (1993). Semenov (1893) erected the subtribe Platyopina (written originally as "Platyopidae") on the basis of dorsally located eyes, unlike the laterally located eyes in other pimeliine genera. Skopin (1962) was the fi rst to question the need for this tribe and treated it as a junior synonym of Pimeliini. He studied larval structures (Skopin 1959(Skopin , 1962 and concluded that larvae of Pimeliini and Platyopini have no fundamental differences. Pierre (1964) erected the tribe Leucolaephini for six Saharo-Arabian genera mainly on the basis of structures of antennae, profemora and the sub-cylindrical prothorax, but Koch (1969) challenged this decision and suggested that Leucolaephini does not differ from Platyopini and cannot be retained, while considering that even representatives of the tribes Pimeliini and Platyopini do not always clearly differ. Later, Kwieton (1981) synonymized the tribe Leucolaephini with Platyopini without any comments. Doyen (1994) studied adult structures of Pimeliinae and placed Platyopini as a junior synonym of Pimeliini as a result of cladistic analysis. Despite this, some authors continued to interpret Pimeliini and Platyopini as separate tribes (Ren & Yu 1999;Ren & Dong 2001;Lillig & Pavlíček 2003;Medvedev 2005;L. Soldati 2009).
Despite such abundance of works on Central Asian Pimeliini, the morphological basis to build a balanced classifi cation of the tribe is very poor. The current classifi cation of the genera is based mainly on the external morphology of adults, especially on the structure of eyes, legs, antennae, pronotal and elytral structures. Internal structures (genitalia) are briefl y presented in the work of Doyen (1994), who fi gured the ovipositor of Lasiostola and the female genital ducts of Ocnera and Sternoplax. Some fi gures of male genitalia and ovipositor for species of the genera Platyope and Ocnera are presented in the works of Chinese authors mentioned above. Leo (2016) photographed the ovipositor of the species Spectrocnera anguliceps Kwieton, 1981 from the Arabian Peninsula. Svetlana Chigray (2019) fi gured male and female genitalia of Podhomala Solier, 1836. The majority of Central Asian genera have not been studied since the original descriptions.
The aim of this work is to compare the Asian genera of Pimeliini with dorso-lateral eyes, which were previously united in the tribe Platyopini. Kaszab (1982) listed 15 genera of this group, three of which (Leucolaephus Lucas, 1859, Pseudoplatyope Pierre, 1964, Pseudostorthocnemis Gridelli, 1953 are distributed in North Africa, including the Afrotropical regions (Pierre 1964), two occur in the Near East, including the Arabian Peninsula (Paraplatyope Löbl, Bouchard, Merkl & Bousquet, 2020, Astorthocnemis Lillig & Pavlíček, 2003 and one genus Storthocnemis Karsch, 1881 is Saharo-Arabian. Species of all the genera mentioned above have lateral eyes located on the level of the genae and have ovipositors that are typical for the subtribe Pimeliina. The mentioned genera were erroneously interpreted as members or relatives of the former tribe Platyopini. bpar = baculi of the paraproct bptg = baculi of proctiger cI, cII, cIII, cIV = lobes of the coxite dbpar = dorsal baculi of the paraproct par = paraproct ptg = proctiger sv = median sclerotization of vulva

Diagnosis
Body entirely covered with dense scales. Pronotum and elytra without large tubercles or strong sculpture, or only with small sparse tubercles. Eyes rounded or weakly oval, dorsoventrally placed above genal level. Legs thin, slender, long, not fossorial, with long sparse setae. Ovipositor very short and weakly sclerotized, transverse or of subequal length and width; paraproct very short, often with reduced or very short baculi; proctiger strongly widened, with very wide weakly sclerotized baculi; lobes 4 of coxite membranous, rounded, with long, very dense hair brush. Spiculum ventrale very short, strongly widened and without common stem. Aedeagus with strongly sub-angularly curved apical piece.

Notes
Semenov described two species in this genus but did not designate a type species. Later, he placed Apatopsis conradti Semenov, 1891 as a junior synonym of A. grombczewskii Semenov, 1891(Semenov 1903aSemenov-Tjan-Shansky 1907) and clearly indicated that the genus Apatopsis includes only the one latter species. This indication (Semenov 1903a) corresponds to the subsequent designation of the type species of the genus according to article 69.1.1 of ICZN (1999). Gebien (1937) also listed Apatopsis as monotypical genus. We do not know why these two species of Apatopsis were given as valid in both editions of the Palaearctic Catalogue (Löbl et al. 2008;Iwan et al. 2020) and we here resurrect the synonymy after the examination of the lectotypes of these conspecifi c taxa: A. grombczewskii Semenov, 1891 = Apatopsis conradti Semenov, 1891, syn. resurr.
Elytra translucent (see in transmitted light), with wide, oval depressions on lateral sides, surface of depressions densely covered with very long yellowish setae (Fig. 1A). Surface of elytra with longitudinal rows of very small sparse granules; each granule with long, erect, yellowish setae (Fig. 8D). Scutellar

Female genitalia
Spiculum ventrale (Fig. 14I) very short and thickened, with short common stem, rods connected by membranous plate at apex; apophyses of sternite VIII long, acutely angulate at apex. Ovipositor ( Fig. 14J-L) weakly sclerotized, very short, transverse. Paraproct short, with narrow baculi; coxite lobes not merged; lobe IV transformed into weakly sclerotized and very densely pubescent pair plate, rounded at apex; lobe III forms additional elongate apical process, similar to lobe IV; lobes I-II with shorter setae; lobe I with oblique, narrow baculi, widened at apices. Proctiger wide, with straight apical margin and densely pubescent middle, baculi of proctiger strongly widened along all length. Fig. 6. Species of the genus Platyope Fischer von Waldheim, 1820, habitus, dorsal view. A. P. proctoleuca proctoleuca Fischer von Waldheim, 1820, ♂ (ZIN). B. P. ordossica Semenov-Tjan-Shansky, 1907, ♂ (ZIN). (Fig. 23A) Vagina sacciform, narrower before oviduct and after spermatheca, apical part of vagina elastically curved. Spermatheca consists of six tubes: two tubes fl ow into vagina independently, other four tubes combined into bundle with common base; two tubes (one independent, one in bundle) bifurcated. Accessory gland with four constrictions, one (as narrow tube) at base and three others in middle.
Trochanters with very short dense setae. Meso-and metafemora curved outward along elytral lateral vertical side. Pro-and metatibiae weakly curved outward, protibiae (Figs 26A, 27E) widened at apex, outer margin of protibia projecting into short rounded process at apex with one (H. primaeveris) or several acutely angulate and closely located (H. vernus) short spines; outer margin armed with very short tubercles with long and thin spines and additionally with long and sparse setaesetae (Fig. 27E). Protibial terminal spurs strongly elongated, extending to base of protarsomere 5, mesotibial terminal spurs extending to base of mesotarsomere 3, metatibial terminal spurs extending to midlength of metatarsomere 1. Outer and inner protibial terminal spurs subequal in length, inner meso-and metatibial terminal spurs longer than outer one; spurs fl attened, outer spur moderately narrowed to apex, acutely angulate at apex. Protarsi laterally not fl attened, covered with short strong setae. Meso-and metatarsi fl attened from sides, covered with long fi ne setae dorsally and shorter and stronger setae ventrally. Tarsal claws elongated, thin, weakly curved outward.

Male genitalia
Male genitalia identical to those in Apatopsis.

Female genital ducts
Destroyed during preparation.

Notes
This species was originally described from a single specimen. The name 'primaeveris' is incorrectly written as "primaeverris" in both editions of the catalogue of Palaearctic Coleoptera (Löbl et al. 2008;Iwan et al. 2020). Semenov Tjan-Shansky used the original epithet 'primae-veris'.

Diagnosis
Body slender, elongate-oval, brown ( Trochanters with brush of dense long setae. Meso-and metafemora curved outward along lateral vertical side of elytra. All tibiae weakly curved outward. Protibiae weakly widened at apex, outer margin of protibiae with small, spinose granules and long, erected setae along all length and projecting apical process (Figs 26B, 27C-D). Protibial terminal spurs more or less widened and fl attened, extending to base of protarsomere 4, mesotibial terminal spurs extending to apex of mesotarsomere 1, metatibial terminal spurs extending to midlength of metatarsomere 1. Outer and inner protibial terminal spurs subequal in length, inner meso-and metatibial terminal spurs longer than outer ones. Protarsi not fl attened from lateral sides, completely covered with spinose setae ventrally and simple setae dorsally. Meso-and metatarsi fl attened from lateral sides, covered with long, fi ne setae dorsally and shorter, stronger setae ventrally. Tarsal claws long, thin, curved outward.
Male genitalia Inner sternite VIII (Fig. 16I) weakly sclerotized along lateral margins and in middle, densely setose, especially at outer margin; anterior margin with short median emargination; pair gland present.
Ovipositor ( Fig. 16K-M) very short and weakly sclerotized. Paraproct very short, with reduced baculi (with only weakly sclerotized inner apices); lobe I of coxite with very short baculi (in form of short sclerotization of inner apices); lobe II weakly sclerotized, transverse; lobe III conical, slightly stronger sclerotized; lobe IV not paired and not sclerotized, membranous, rounded at apex, densely pubescent by long setae. Proctiger very wide, widely emarginate at apex, sparsely pubescent on lateral margins and with very wide trianglular baculi. (Fig. 23B) Vagina strongly widened, sacciform, narrower before oviduct and after spermatheca, apical part of vagina elastically, sharply curved. Spermatheca short, single-tube. Accessory gland widened at apical ⅔, with thick basal canal and one short constriction between gland and vagina.

Notes
Only one character, the protruding and raised prosternal process, distinguishes Kawiria from Habrobates. This character cannot be used as the main character for diagnostics of genera within Pimeliini, because it is often used for diagnosis of different species within one genus, for instance, in Diesia Fischer von Waldheim, 1820, Sternoplax, Lasiostola. A complex of other characters shows that the type species of both Habrobates and Kawiria are congeneric: body completely covered by fl attened scales without central ridge, head with erect sparse setae, pronotum with microgranules for scales and slightly small granules for short setae, elytra with 10 longitudinal rows of small granules (each granule with short seta) and dense granules along apical margin, width of intercoxal process of abdominal ventrite 1 less than width of one metacoxa, outer margin of protibiae with thin short spines, sparse long setae and projecting process at apex. Characters such as body colour and density of setation of meso-and metatibiae are of species level. Consequently, the following synonymy is proposed: Habrobates Semenov, 1903= Kawiria Schuster, 1935. So, the following new combination is established: Habrobates gabrieli (Schuster, 1935) comb. nov. (from Kawiria).
It is interesting that Schuster did not compare Kawiria with Habrobates in the original description (but he compared it with Mantichorula and Homopsis), although he noted that his new genus is very similar to Habrochiton. In addition, he interpreted scales on the body as microwrinkles, which give a silky shine (Schuster 1935).
In general, early authors gave unjustifi ably high importance to the prosternal process and used this species-level character for descriptions of genera. The same situation with the prosternal process is observed in two species of Dietomorpha and discussed below in the case of the synonymy of the genera Platyope and Homopsis.

Bionomics
Detailed bionomics, behaviour and adaptations of Habrobates vernalis were described by Medvedev (1965b) and Kaplin (2019). Brief information about bionomics of H. agnesae and H. gabrieli comb. nov. was given by Schuster (1935Schuster ( , 1938. Species of this genus inhabit non-fi xed barchans sand dunes, but often climb bushes and trees (usually Haploxylon) for thermoregulation. Schuster noted that Kawiria was described from a salt desert, and Habrobates is known from sand deserts (Schuster 1938). However, two collectors, Dr Alfons Gabriel and his wife Agnes, informed him via the privy councillor Professor Meinhard v. Pfaundler, that they collected these beetles (Kawiria) on saxaul barchan sand dunes near Halvan village (South Khorasan Province), without getting off their camels. So, Habrobates gabrieli comb. nov. has very similar behaviour and bionomics to other species of Habrobates, and it is not associated with salt marshes. Schuster, 1938 Material examined

Diagnosis
Body robust ( Trochanters with elongate, sparse setae. Meso-and metafemora curved outward. Protibiae with projecting process at apex of outer margin, armed with tuft of spines (Fig. 26B). Lateral margins of protibiae with long, fi ne setae and short spines. Protibial terminal spurs elongated, extending to base of protarsomere 4; mesotibial terminal spurs extending to apex of mesotarsomere 2; metatibial terminal spurs extending to midlength of metatarsomere 1. Outer and inner protibial terminal spurs subequal in length, inner mesoand metatibial terminal spurs longer than outer ones, spurs fl attened from sides. All tarsi fl attened from sides, covered with long setae dorsally and short setae ventrally. Tarsal claws elongated, thin, weakly curved outward.
Rods of spiculum gastrale (Fig. 17G) widely spaced, thickened, arcuately connected at apex, with wide membrane along this connection and near derivatives of inner sternite IX; these derivatives elongated, ladle-shaped, evenly sclerotized in middle and slightly more sclerotized terminally; apical margin of derivatives covered with short dense setae.
Tegmen of aedeagus thickened, obtuse ( Fig. 17A-C). Basal piece of tegmen much shorter and wider than apical piece, with tubercle-shaped apex ( Fig. 17A-B). Apical piece strongly curved, narrowly rounded at apex, covered with short setae in apical half; ventral apophyses not expressed, dorsal apophyses moderately long, triangular. Median lobe (Fig. 17D, F) widened in basal ⅔ and narrowed in apical third, with weakly separated apex, strongly curved; basal part with additional four sclerotized armatures; baculi not merged at apex.
Ovipositor ( Fig. 17G-L) short, weakly sclerotized. Coxite with four distinct not merged lobes. Baculi of coxite lobe I short, shorter than paraproct baculi; lobe II small, sclerotized; lobe III larger, conical, sclerotized; lobe IV membranous (only at apex slightly sclerotized), conical, with dense pubescence of long setae. Paraproct V-shaped on each side ventrally, baculi sharply widened in base and narrow in apical third. Vulva with narrow sclerotization in the middle of ventral side. Apical margin of proctiger with deep V-shape emargination, baculi of proctiger strongly widened. (Fig. 23C) Vagina moderately widened, sacciform, narrower before oviduct and after spermatheca, apical part of vagina not elastically curved. Spermatheca very short, single-tube. Accessory gland long, with two tubelike constrictions, short in base and long in middle. Basal part between two constrictions not sacciform.

Distribution
Iran and Pakistan (Balochistan region in both countries), Afghanistan. Medvedev (2005) listed D. pardalis for Registan Desert (Afghanistan) and adjacent areas of Pakistan, but this record was omitted from the Palaearctic Catalogue (Iwan et al. 2020

Notes on paratypes (Maxwell Barclay, personal communication)
Ernest Vredenburg (the donor of the specimens, Superintendant of the Geological Survey of India) and Captain Frank C. Webb Ware (the collector of the paratypes, at Quetta, 'Political Assistant' at Chagai and who was working on the Sistan-Quetta caravan route) went to Quetta together. It is likely that the beetles were collected at Quetta in 1899, given the Indian Museum registration in 1914, and eventually given the BMNH registration in 1973.
Pronotum transverse (1.7 × as wide as long), widest before middle, where it 1.2 × as wide as head, with two dark, broad fi elds and pale spot in middle; lateral margins at base together with contiguous part of prothoracic hypomera with glabrous and smooth black spot. Ratio of width of pronotum at anterior margin to its maximum width and to width at base: 8 : 8.5 : 8. Anterior margin and base of pronotum widely weakly emarginated at middle. Lateral margins rounded, emarginated at base. Anterolateral angles not expressed, posterolateral angles obtuse, widely rounded. Disc of pronotum transversely convex, with two triangular depressions at base. Apical half and middle of pronotum covered with projecting spine-like spinose tubercles (mainly on sides of pronotal disc). Prosternal process narrow, elongated (3 × as long as wide), not raised between procoxae, slightly protruding beyond procoxae.
Elytra weakly elongate (1.2 × as wide as long), 3.4 × as long and 1.8 × as wide as pronotum, 2.1 × as wide as head. Each elytron with fi ve brown, longitudinal stripes on creamy background. Elytra without humeral ribs, with four longitudinal rows of spine-like tubercles: one row on each lateral vertical side, one humeral row and four discal rows. Apex of elytra with dense tubercles. Scutellar shield not concealed by base of elytra; surface around scutellar shield with triangular depression. Metepisterna, meso-and metaventrites with fi ne sparse spinose granules. Transverse length of one metacoxa subequal to intercoxal process of abdominal ventrite 1.

Comparative diagnosis
This new species differs from D. pardalis by the more elongated body, the slightly protruding prosternal process (strongly protruding beyond procoxae in D. pardalis), the striped pattern (spotted in D. pardalis), the longitudinal rows of spine-like setae on elytra (the tubercles located only in spots in D. pardalis), the absence of elytral humeral ribs and not concealed scutellar shield.

Diagnosis
Body ( Fig. 4A-B) from pale brown to dark brown, robust, elytra rounded (semispherical), sometimes slightly fl attened dorsally, completely covered with recumbent, white or yellowish and brown setae, with longitudinally striped pattern on elytra. Body length 7-18 mm. Eyes small, circular, convex in dorsal view; surface behind eyes concealed by anterior margin of pronotum.
Pronotum almost square (width subequal to length) (Fig. 3A), with lateral margins straight or weakly emarginated basally. Anterolateral angles strongly projecting, acutely angulated (but rounded at apex), visibly raised above level of eyes. Disc of pronotum moderately convex, with rounded, deep depression in middle and two triangular, deep depressions at base, surface around depressions densely covered with very large semispherical tubercles with seta ( Fig. 11A-B), with setae between them. Prosternum long, 1.8 × as short as longitudinal diameter of one procoxa (Fig. 4B), without depression along anterior margin. Prosternal process narrower between middle level of coxae and strongly widened to apex (1.5 × as long as wide), not raised between procoxae and not protruding beyond procoxae.
Elytra semispherical (2 or more × as wide as pronotum), humeral rib weakly expressed, not strongly elevated, with row of sparse tubercles. Elytral pubescence has longitudinally striped pattern (from 3 in P. trinkleri to 7 in P. dilatata) of recumbent, yellowish setae (Fig. 4A) as a result of presence of denser or sparser setae (Fig. 11C-D). Przewalskia trinkleri has glabrous anterior half and striped apical and lateral parts of elytra. Each elytron with three (P. trinkleri) to 4-6 longitudinal rows of very small granules with long seta in denser pubescent (light) striae. Scutellar shield not hidden under base of pronotum, widened to apex; surface around scutellar shield without depression. Transverse length of one metacoxa subequal to intercoxal process of abdominal ventrite 1.
At least pro-and mesotrochanters with very long setae. Meso-and metafemora curved along elytral lateral vertical side. Protibiae widened from proximal to distal part, weakly curved outward, with strong, short spines (on outer margin) and very dense, long setae (Figs 26E, 28A-B); spines denser at apex. Mesotibiae curved outward, with smaller and sparser spines and dense pubescence on inner margin. Metatibiae straight, with small, sparse spines and dense pubescence at least in apical part. Protibial terminal spurs usually lanceolate (at least outer spur), large, widened and fl attened, slightly curved inward, subequal in length, extending to base of protarsomere 3; meso-and metatibial terminal spurs different in length, narrowed from base to apex, mesotibial terminal spurs extending to base of mesotarsomere 2, the latter ones extending to apex of metatarsomere 1. Pro-and mesotarsi fl attened from lateral sides, covered with long fi ne setae dorsally and shorter, stronger setae ventrally; metatarsi also fl attened from lateral sides, with long setae dorsally and very short ones ventrally; setae form dense, fl attened brushes. Tarsal claws elongate, narrow, weakly curved outward.
Tegmen of aedeagus ( Fig. 18A-C) long, slender, with basal piece slightly shorter than apical one; apical piece bare, slightly curved, acutely angulate at apex ventral apophyses not expressed, dorsal apophyses long, merged in triangular plate; basal piece with narrowly rounded apex; median lobe with wide sclerotization in apical and basal parts and small membranous area in middle, base narrowly rounded, not bifurcate, apex acutely angulate (Fig. 18D-F).
Ovipositor (Fig. 18J-L) moderately sclerotized, elongate. Coxite with four distinct, not merged lobes. Baculi of coxite lobe I well expressed, but much shorter than paraproct baculi; lobe II short, transverse; lobe III elongate; lobe IV strongly sclerotized, transformed into curved spatulate structure. Paraproct elongate, with narrow, long baculi. Proctiger also elongate and with narrow baculi, anterior margin rounded. Vulva with pair of sclerotized 'spicula' in the middle on ventral side. Paraproct in base, coxite and proctiger with very dense and long pubescence. (Fig. 23D) Vagina elongated, with longitudinal wrinkles and separated anterior part. Spermatheca as short single tube. Accessory gland long, sacciform, with one thick constriction at base.

Diagnosis
Body black, wide, dorsoventrally fl attened, disc-shaped, without dense pubescence dorsally, only some parts of pronotum and head with setae, elytra dorsally and laterally with very short, fi ne, sparse spines ( Fig. 4C-D) and sometimes with small area of setation along apical part of suture. Body length 10-20 mm. Eyes large, circular, moderately convex dorsally.
Pronotum strongly transverse (more than 2 × as wide as long). Anterolateral angles weakly projecting. Base of pronotum strongly widely emarginated, and posterolateral angles deeply protruding and impressed into elytral base (Figs 4C, 11E). Disc of pronotum with one medial and two lateral longitudinal stripes of recumbent, moderately dense, short setae and coarse, semispherical tubercles on lateral sides ( Fig. 11E-F); base widely, sharply depressed and fl attened. Prosternum before procoxae near 2 × shorter than longitudinal diameter of one procoxa. Prosternal process very large and broad, raised between procoxae, strongly protruding beyond procoxae, extending to mesoventrite (Fig. 4D).
Protrochanters and inner side of profemora covered with very long, dense setae. Femora robust, weakly curved outward. All tibiae weakly curved outward. Protibiae strongly gradually widened to apex, triangular, fl attened, without projecting process at apex of outer margin, with dense, short, strong spines, but with longer and fi ner spines and dense, long setae along lateral margins (more recumbent on inner side) (Figs 26D, 27F-G); strong spines denser at apex of outer margin. Meso-and metatibiae with similar structures, but not dorsoventrally fl attened and widened, often additionally covered with cream coloured scales. Protibial terminal spurs large and widened, not lanceolate ( Fig. 27F-G), subequal in length, extending to base of protarsomere 5. Meso-and metatibial terminal spurs different in length, inner tibial terminal spurs longer than outer ones, extending to protarsomere 3 and half length of metatarsomere 1 subsequently. Protarsi with long, recumbent spines dorsally and ventrally and short setae laterally; meso-and metatarsi fl attened from sides, with very dense, long setae on dorsal side.
Ovipositor (Fig. 19J-L) is very similar to that in Przewalskia, but differs in the absence of sclerotization on vulva, apical lobes turned to dorsal side and widely weakly rounded proctiger. (Fig. 23E) Vagina elongated, tube-shaped, not widened, with short primary bursa copulatrix at apex. Spermatheca absent. Accessory gland of spermatheca very short, tube-shaped, not modifi ed, without constrictions.

Distribution
Mongolia, China (Inner Mongolia, Ningxia, Gansu, Shanxi provinces). Schuster (1940) believed that all three species of Mantichorula should be regarded as valid, and that M. grandis distinctly differs from M. semenowi and M. mongolica because of their short antennae, which do not extend far beyond the base of the pronotum. Medvedev (1990) interpreted the genus Mantichorula as monotypic. We studied the type series of M. grandis and discovered that this species and some specimens standing as M. semenowi in the ZIN collection do indeed have shorter antennomeres. Studies of the type specimens of M. semenowi and M. mongolica, as well as a detailed examination of male and female genitalia, are necessary to revise this genus.

Diagnosis
Body black, elongate-oval (Figs 4E-F, 5G-I, 6), weakly fl attened dorsally, densely covered with recumbent grey, yellowish, cream or white setae; elytra often with striped or mixed spotted and striped pattern partly or completely. Sometimes (P. ordossica) body dorsally glabrous (Fig. 6B). Body completely or partly granulated, each granule with spine or strong seta. Body length 9-15 mm. Eyes circular, very weakly convex in dorsal view; surface behind eyes partly concealed by anterior margin of pronotum.
Protibia wide and fl attened, triangular, without projecting process at apex of outer margin, with strong teeth on outer margin (Figs 26H, 28D-E). Maximal length of teeth much shorter than maximal width of protibia. Teeth spinose or not, often spines abraded. Meso-and often metatibiae curved outward, with conical, coarse granules bearing spines and dense or sparse, long setae. Protibial terminal spurs moderately elongated, extending to protarsomere 4, subequal; longest mesotibial terminal spurs extending to or slightly not extending to apex of mesotarsomere 1; longest metatibial terminal spur not extending to apex of metatarsomere 1. Tarsal claws comparatively (compared to other genera) short, weakly curved outward. Spiculum gastrale (Figs 20F-G, 21G-H) arcuately connected at apex, slightly curved in lateral view; derivatives of inner sternite IX large, weakly oval, unevenly sclerotized, with membranous C-shaped area along outer margins; apical margins of these derivatives pubescent with long sparse setae.   Tegmen of aedeagus long and slender (Figs 20A-C, 21A-C), basal piece more or less longer than apical piece. Apical piece bare, weakly curved, fusiform, acutely angulate at apex; ventral apophyses presented, short; dorsal apophyses long, triangular, not merged. Basal piece slightly wider than apical piece. Median lobe with baculi not distinct, bifurcate or narrowly rounded apex and sub-acutely angulate base (Figs 20D-E, 21D-F).

Female genitalia
Spiculum ventrale (Figs 20I, 21J) long, with long common stem. Inner sternite VIII can be with long or short acutely angulate apophyses. Ovipositor long, moderately sclerotized (Figs 20J-L, 21K-M). Coxite with not merged lobes. Baculi of lobe I transverse, narrow, strongly sclerotized only near middle of ovipositor; pair of lobe I forms cruciform sclerotization area ventrally together with median sclerotization of vulva; lobes I and II membranous; lobe III widely sclerotized; apical lobes fossorial, strongly sclerotized, fl attened, subacutely angulate or widely rounded at apex. Coxite laterally and on inner side of apical lobe densely  covered with very long setae. Apical margin of proctiger narrowly or densely rounded. Paraproct basally, proctiger and coxite densely pubescent with long setae.

Female genital ducts (Fig. 23F-G)
Vagina strongly widened, sacciform, sometimes with poorly expressed bursa copulatrix, sometimes in apical third elastically turned back. Spermatheca short, single-tube (Fig. 23F) or with thick basal tube with tuft of small tubes at apex (P. grumi) (Fig. 23G). Accessory gland of spermatheca elongate, shortly constricted at base and with narrow basal duct, sacciform after this duct.

Notes
The genus Homopsis was described by Semenov (1893) based on the structure of the prosternal process, which is slightly raised and slightly protruding beyond the posterior margin of procoxae, in contrast with Platyope, which has the vertically rounded prosternal process, not raised in ventral aspect and not protruding beyond procoxae. Semenov (1893) placed Homopsis and Mantichorula in one couplet, as genera having protruding and raised prosternal process, in contrast to other genera of 'Platyopidae'. In our opinion, the prosternal process in the latter genera is strongly different from each other. Mantichorula has a process that is strongly protruding, extending to mesoventrite and horizontally (in lateral view) fl attened, while Homopsis has only a slightly protruding, angularly rounded prosternal process. Homopsis is very similar to Platyope and differs only by some species-rank characters, including the prosternal process, the structure of the pronotum, the elytra, the male and female genitalia (especially the ovipositors are similar). On the other hand, a single species of Homopsis has peculiarities in the structure of the female genital ducts: vagina elastically V-curved and spermatheca multi-tubed at the apex. However, the main congeneric character of Platyope and Homopsis is externally open procoxal cavities (postcoxal bridge is discontinuous in the middle). As a result, the following synonymy is proposed: Platyope Fischer von Waldheim, 1820 = Homopsis Semenov, 1893 syn. nov. Consequently, the following combination is proposed: Platyope grumi Semenov, 1893 comb. nov (Iwan et al. 2020), but Medvedev (1990) recorded P. proctoleuca proctoleuca also for Russian Altai and the Irtysh valley in Kazakhstan.   Semenov, 1903 (ZIN). C. Dietomorpha pardalis Kühnelt, 1957 (ZIN). D. Przewalskia dilatata (Reitter, 1887)  Platyope ordossica Semenov-Tjan-Shansky, 1907  Platyope grumi (Semenov, 1893)   Genus Earophanta Semenov, 1903Figs 7, 13E, F, 22, 23H, 24E, 25B, 26G, 28C Earophanta Semenov, 1903b: 172 (replacement name for the elimination of the homonymy). Earophila Semenov, 1903a: 9 (type species: Platyope serrata Semenov, 1893, by original designation; as subgenus of Platyope; homonym, nec Earophila Gumppenberg, 1887). Earophilina Strand, 1917: 99 (unnecessary replacement name for the elimination of the homonymy).

Diagnosis
Body black, robust, wide, oval, completely or partially covered with dense white, grey or yellowish recumbent setae, elytra with moderate or large tubercles and serrate lateral margins (Fig. 7A-D). Body length 7.5-17 mm. Eyes circular, convex in dorsal view.
Pro-and mesotrochanters usually with long, dense pubescence. Pro-and mesofemora weakly curved outward, protibiae straight, meso-and metatibiae curved along elytral lateral vertical side. Outer margin of protibiae with 5-7 very large and long sparse teeth and sometimes with several small teeth, widened to apex, sub-triangular, without projecting process at apex of outer margin (Figs 26G, 28C). Protibiae of Earophanta beludzhistana have four teeth in basal part and fl attened undulate lamina (merged teeth) in distal part (Fig. 7F). Length of largest teeth is at least one third of width of protibia at apex. Tibial terminal spurs as in Platyope. Protibiae often curved outward (Fig. 7A, C-F), but sometimes straight (E. planidorsis) (Fig. 7B). Mesotibiae always curved outward, metatibiae straight. Tarsi as in Platyope.

Female genitalia
Spiculum ventrale (Fig. 22I) with long and sharply widened at apex common stem, thin rods and often with weak sclerotized area between them, apophyses of sternite VIII comparatively short, acutely angulate at apex. Ovipositor (Fig. 22J-L) long and moderately sclerotized. Coxite lobe I with baculi, strongly sclerotized distally; lobes II and III merged into one moderately sclerotized plate; apical lobe fossorial, transformed to strongly sclerotized and rounded apically scoop-like plates. Paraproct with long and narrow ventral baculi and very short additional dorsal baculi. Vulva without sclerotized areas. Only coxite with sparse pubescence of moderately long and short setae; paraproct with very short setae on margins near coxite. Proctiger with long, narrow baculi and rounded apical margin.
Female genital ducts (Fig. 23H) Very similar to those in Platyope, with single-tube long spermatheca and long, constricted basally accessory gland.

Notes
Earophanta loudoni was described from a single specimen and was synonymised by Medvedev (2005). The specimens from Turkmenistan, which were standing in the collection of ZIN as E. loudoni, differ from the specimens from Afghanistan (determined by Z. Kaszab as E. pilosissima) by much more pubescent elytra and visibly smaller smooth tubercles on the elytra. A larger series of specimens is necessary to establish the status of these populations.  Skopin, 1960 (Skopin 1960(Skopin , 1968.

Note
Skopin and curators marked two specimens of E. pubescens from HNHM as a holotype and a paratype, respectively. However, Skopin (1960) did not include these specimens in the type series. Bogatchev, 1957 Fig. 7E-F

Note
This species was originally described from a single specimen and known only from the type locality.  6. Elytra smooth and glabrous, only with very small, sparse granules and one elevated line of slightly larger granules along granulated lateral margin. Pronotum with strongly arcuately emarginated base, posterolateral angles protruding far into impressed base of elytra. Prosternal process very broad, weakly narrowed between procoxae, slightly raised between procoxae and horizontally fl attened (lateral view), strongly protruding beyond posterior margin of procoxae to mesoventrite. Protibiae with dense, long, thin spines and setae along outer margin ....................Mantichorula Reitter, 1889 -Elytra with coarse granules or/and smooth tubercles completely or at least on sides and pubescence between them or at least apical striped pubescence (sometimes elytra without pubescence as in Platyope ordossica Semenov-Tjan-Shansky, 1907). Pronotum with straight or very weakly emarginated base, posterolateral angles not protruding. Prosternal process broad, but usually visibly narrowed between procoxae, not raised between procoxae, not protruding or very weakly protruding beyond posterior margin of procoxae, rounded down. Protibiae with teeth on outer margin .......... 7 7. Procoxal cavities externally closed. Protibiae with large teeth or teeth partly merged into thin entire lamella (as in Earophanta beludzhistana Bogatchev, 1957); length of largest teeth is at least one third of width of protibia at apex.

Status of the former tribe Platyopini
The family name "Platyopes" was originally established by Motschulsky (1849) to hold two Iberian genera Morica Dejean, 1834 and Akis Herbst, 1799(Herbst 1799 and fi ve unknown (Herbst 1799: 58) genera from "à Astrabad en Perse" (in Astrabad in Persia). Bouchard et al. (2005Bouchard et al. ( , 2011 listed Platyope as the type genus of "Platyopes" according to Article 29.1 of ICZN (1999), but Motschulsky did not include this genus in the mentioned family because no species of Platyope occur in Iran and adjacent countries.
The subtribe "Platyopidae" (original spelling) was secondarily erected on the base of dorso-lateral eyes, located above the level of the genae (Semenov 1893). Later, Semenov-Tjan-Shansky (1907) raised its rank to a tribe. Pierre (1961Pierre ( , 1964 erected from Pimeliini the additional tribe Leucolaephini (correct name Leucolaephusini according to Bouchard et al. (2011)) on the base of callus-like swellings on the inner side of male profemora and the structure of antennae with 3-segmented club and very small antennomere 11 with the basal part placed into the penultimate antennomere. Koch (1969) noted that both these characters occur in different genera of Pimeliini and Platyopini, therefore Leucolaephusini is a doubtful tribe. Later Leucolaephusini was synonymized with Platyopini (Kwieton 1981). It should be noted that Koch (1969), in an endeavor to analyse the relationships between Pimeliini, Platyopini and Leucolaephusini and to establish a status of these tribes, used the structure of the protibiae, which has an adaptive function and cannot be used as a basis for classifi cation.
Many subsequent authors did not consider other structures and were guided by this single character of the location of eyes for Platyopini. Only Skopin (1962) did not accept the tribe Platyopini after the analysis of structures of the larvae of Platyope and Earophanta. He concluded that it cannot be saved even as a subtribe because larvae are identical to those of Pimeliini. Doyen (1994) also proposed the Platyopini as a junior synonym of Pimeliini and concluded that these two groups only slightly differ in the structure of mouthparts and antennae, but he included only one species of Platyopini (Platyope) and three species of Pimeliini in his analysis. He also fi gured the structures of the ovipositor and the female genital ducts, but only for two species of Pimeliini. We analysed a comprehensive complex of external and internal structures and found that the former Platyopini (s. str., excluding Saharo-Arabian genera) is a polyphyletic group and includes two branches, which were previously united in one tribe on the basis of the adaptive character of dorso-lateral eyes, which appeared independently in their evolution.
The fi rst group (subtribe Habrobatina) united related ultrapsammophilic genera, which sharply differ from other Pimeliini by the structure of ovipositor and aedeagus. The second group (Platyope genus group) forms a separate subgroup within the tribe, differing in the dorso-lateral eyes, the large hemispherical tubercles on the pronotum and the absence of the uncus on the lacinia (Medvedev 1959), but it has no signifi cant differences in the structure of the male genitalia and the ovipositor. Thus, the tribe Platyopini with the type genus Platyope was correctly synonymized with Pimeliini by Skopin (1962) and Doyen (1994).
Since both groups inhabit sandy deserts with the dominance of Arenosols -soils with very initial features of pedogenesis and alteration of the mineral part (IUSS Working Group WRB 2015), they have acquired a number of similar adaptations. Below, we analyse adaptive structures, which originated independently in psammophilic Pimeliini, and characters suitable for a future phylogeny.

Structural transformations and adaptations to life in sandy deserts
Surface of integument Many desert darkling beetles have a waxy coating pigmentation or various structures on the surface of the integument, partially to protect the body from overheating (Pierre 1958;Cloudsley-Thompson & Chadwick 1964;Edney 1971;Medvedev & Nepesova 1985;Medvedev 1990;Abushama 1990). Pale colouration can lower the temperature of the corresponding parts of the body by several degrees (Edney 1971). Turner & Lombard (1990) argue that colouration only partially affects the beetle's body temperature. Black tenebrionids heat up much more than white-black ones, at least at direct absorption of visible light warms, and "a black beetle is also warmed more by refl ected visible radiation than the white beetle" (Turner & Lombard 1990: 311). In Pimeliini, the protective structures of the integument are usually pale, white, yellow, pale brown, pale grey. We can observe several directions for the formation of pale colouration among diurnal Pimeliini: white waxy coating (Trigonoscelis Dejean, 1834, Sternoplax, Pisterotarsa Motschulsky, 1860, Sternodes Fischer von Waldheim, 1837), setae, spines and coarse elytral and pronotal sculpture for the formation of a non-natural covering (clay crust) (Lasiostola Dejean, 1834, often Pimelia Fabricius, 1775), entire or partial cover of pale felt setae (Pterocoma Dejean, 1834, Platyesia Skopin, 1971, the Platyope group of genera, some Pimelia), scale cover (Leucolaephus, Paraplatyope, some Storthocnemis, Pimelia senegalensis Olivier, 1795, Habrobatina). A combination of bristles and scales is also not uncommon. In Habrobatina and platyopoid genera, the latter two types of integument are shown. Leaf-like scales with micro-grooves in the form of venation (Figs 8B, D, 9B, 10) or scales with medial elevated ridge (Figs 9D, F) cover the integument of species of Habrobatina. These scales originated from setae as it is well illustrated on fi g. 10A(b) with intermediate scale-like setae. The pubescence of tomentose recumbent white, pale grey, and yellow setae is characteristic for the Platyope genus group (Figs 4A, E, 5G, I, 6A), and it can either be entire or between tubercles (Earophanta) (Figs 7A-C). It should be noted that the covering of hairs and scales also provides the function of protective colouration. Many species hide in the shade of desert vegetation, and the pattern on the elytra is often associated with the pattern of this shade. Thus, the spotted pattern on the dorsal side of the body of Dietomorpha pardalis (Fig. 3A) is probably associated with dappled shadows, as in the ground beetle genus Graphipterus Latreille, 1802, which becomes completely invisible in the shade of plants with small rounded leaves (personal observations of M.V. Nabozhenko in Morocco). Structures on the elytra of many species of Earophanta apparently have a similar function of protective colouration, combining a dark smooth large round tubercles and a grey or yellow tomentose cover of hairs (Fig. 7E, C). Platyope leucogramma with a striped pattern (Fig. 4E) is hardly visible among the elongated shadows of cereals (personal observations of M.V. Nabozhenko in the Rostov Region of Russia). A similar camoufl age effect was noted for Zophosini from the Namib Desert (Matthews et al. 2010). Some species of Platyope have mixed camoufl age with spotted and striped patterns (Fig. 5G, I). Species of some genera (for example Mantichorula, some Platyope) have the bare body or dorsal surface (Figs 4C, 6B), covered with dark-brown scales (Habrobates gabrieli comb. nov.) ( Fig. 2A-E), and probably their adaptations to the high temperatures in deserts are associated with behaviour, diurnal activity, and phenology, as in other black-coloured desert darkling beetles (Matthews et al. 2010).

Some features of the structure of the head
The most characteristic feature for Habrobatina and platyopoid genera is the eyes, located dorsolateral, above the level of genae (Figs 2C, E, 3, 3E, G). Such location of eyes is associated with general compaction of the body and transformation of the head from prognathous to sub-hypognathous.
Mouthparts are tenebrionoid, the mentum is slightly enlarged but does not hide the cardo and stipes, unlike many other desert Pimeliinae (Erodiini, Zophosini, the majority of Tentyriini, etc.) with a large mentum (Medvedev 1959). Antennae with a weak 3-segmented club, moreover the antennomere 11 looks like deeply with the basal part placed into the penultimate antennomere (Fig. 24). Habrobatina and platyopoid genera, like all other Pimeliini (Skopin 1964(Skopin , 1971, have a distinct border between the apical sensorial area and the remaining antennomeres, which are bare. The difference from other Pimeliini is that the antennomere 11 is always very small, and its apex is hemispherical, not acutely angulate. A similar structure of the apical antennomere occurs among other Pimeliini, in the genus Pimelia Fabricius, 1775. Only Habrobates gabrieli comb. nov. has the intermediate structure of antennomere 11, which is slightly smaller than the penultimate one and has a large sensorial area with a sub-acutely angulate apex. In general, there are no clear differences in the antennal structures between different groups of Pimeliini, as it was noted by Koch (1969).
The structure of the thorax Doyen (1993) noted that at least Platyope (Fig. 25A) and Ocnera Fischer von Waldheim, 1822 of the tribe Pimeliini have externally open procoxal cavities, which is a secondary condition. However, species of Ocnera have procoxal cavities closed by thin postcoxal bridge, which can be broken near the prosternal process in collection specimens (we studied 10 specimens of two species of Ocnera and postcoxal bridge was broken and discontinuous in some specimens). Probably, he examined such a specimen. Doyen assoсiates the secondarily open procoxal cavities in some groups of Tenebrionidae with the close association of the pro-and pterothoraces, unlike the primary internally and externally open ones in Zolodininae (Doyen 1993;Matthews & Bouchard 2008) and Kuhitangiinae (Medvedev 1962;Nabozhenko & Sadeghi 2017) (Doyen erroneously wrote that one species of the tribe Kuhitangiini has internally closed procoxal cavities). However, he noted that the prothorax in Pimeliini is relatively free but internally closed procoxal cavities indicate the secondary reduction of the postcoxal bridge in Platyope. In fact, the prothorax is fi xed in relation to the pterothorax (although the pro-and mesothorax are joined by a thickened membrane rather than fused through various mechanisms as in Edrotini, Adesmiini, Erodiini, and Cryptochilini, see Doyen 1994), which is associated with the general compaction of the thoracic region. All at least platyopoid genera have an identical fusion of the proand pterothorax, but within Pimeliini only Platyope has the discontinuous postcoxal bridge (Fig. 25A). Therefore, the open procoxal cavities in Platyope can not be explained by the close association of the pro-and pterothorax. Probably, this is associated with the strengthening of the muscles of the fossorial protibiae and, accordingly, an increase in the area of the inner surface of the procoxae for attaching this musculature (and, accordingly, an increase of the size of the procoxae). Thus, the procoxae are increased due to the reduction of the postcoxal bridge. In contrast, an increase in the size of the procoxae in some Earophanta is achieved by a strong shortening of the prosternum before procoxae (Fig. 7D).
The pterothorax in all Pimeliini is characterized by mesocoxal cavities that are partially closed by the mesepimeron, by the large trochanters and the very wide distance between the coxae, which is associated with the increase of the volume of the subelytral cavity. Doyen (1994) noted that mesocoxal cavities that are externally closed by the mesepimeron, certainly represents the primitive condition, which occurs in the non-pimeliine lineages, while the majority of pimeliines has mesocoxal cavities closed by meso-and metaventrites.
The structure of protibiae Various structures of protibiae are closely assoсiated with adaptations to locomotion in sandy soils. All the genera of Pimeliini in this study inhabit sand deserts and have different adaptations to moving on sandy soil. Medvedev (1965b) analysed in detail the adaptations of legs of the Central Asian darkling beetles and indicated several directions of tibial and tarsal transformations in association with the structure of sandy soils. Two types of specialization (with some intermediate types) are observed in the tribe Pimeliini in general and in platyopoid genera and Habrobatina in particular: 1. Inhabitants of migratory sands. Sand dunes are characterised by a weak development of the root system of plants and an extremely low aggregation of textured grains of fi ne sand; therefore, protibiae in species living on such sands are adapted to sand sweeping, which is accompanied by an increase of the tibial speed (when digging) in some groups, such as Habrobates (Medvedev 1965b). The following directions of adaptation of non-fossorial protibiae can be observed among genera with dorso-lateral eyes: a) Protibiae narrow, with the sharp, projecting process at the apex of outer margin ( Fig. 26A-C). This process can by reinforced by longer merged spines at the protibial apex (Habrochiton) (Figs 26A, 27E) or with a series of shorter strong spines (Habrobates, Dietomorpha) (Figs 26C, 27C-D) or large, but not reinforced additionally (Apatopsis) (Figs 26F, 27A-B). The outer margin of protibiae bears sparse, long setae and sparse, spinose tubercles. Combinations can be from very short narrow tubercles and thin, long spines (Habrochiton) (Fig. 27E) to the similar tubercles, but stronger and shorter spines (Habrobates, Dietomorpha) (Figs 26C, 27C-D) and to large tubercles with short, strong spines (Apatopsis) (Fig. 27A). Spines can be abraded during the lifetime of the adults (Fig. 27B).
b) Protibiae strongly gradually widened to the apex, fl attened, with a weak, rounded process at the apex of the outer margin and with dense, thin, long spines and long setae along the outer margin (Mantichorula) (Figs 26D, 27F-G). Spines can be of two types: stronger and slightly shorter mainly dorsally (Fig. 27F) and fi ner and longer ventrally (Fig. 27G).
Przewalskia has the intermediate type of protibiae, with long and very dense setae, two types of dense spines, but protibiae are thin and not fl attened as in the version a) (Fig. 28A-B).
Thus, the adaptive transformation of protibiae in Habrobatina and platyopoid genera living on sand dunes is accompanied by an increase of the surface area of the protibiae due to the development of long setae and long thin spines on the outer margin, necessary for the displacement of dry moving sand. Spines can be reinforced for digging in deeper and denser sand layers.
Mantichorula and Sternodes Fischer von Waldheim, 1837, which have a more fl attened strongly widened body and a very similar structure of tibiae, share the same life form and therefore they probably have similar bionomics and move on the sand as described by Medvedev (1965b) for the latter genus.
2. Inhabitants of fi xed or wet (in early spring periods of rains) sands have widened and fl attened tibiae with various teeth or tubercles with strong spines, as well as the apex of the protibiae often sharply projecting outward. All these adaptations are necessary for digging in the sand with high particle adhesion, which is achieved in different ways (in order of increasing complexity of specialization): a) Protibiae curved and moderately widened to the apex, that is compensated by very large and sparsely spaced teeth on the outer margin (Figs 26G, 28C), which (teeth) sometimes merged into an entire undulate lamina (Fig. 7F); the outer margin is without a protruding process at the apex (Earophanta); b) Protibiae strongly widened to the apex and have dense, strong, but much shorter teeth on the outer margin than Earophanta; the outer margin is not projecting as the process at apex (Platyope) (Figs 26H, 27F-G).
Thus, the strengthening of the fossorial function of protibiae occurs due to the increase of their area at the apex of the outer margin or the development of various spines and teeth on the outer margin.
The structure of the abdomen Habrobatina and platyopoid genera, as well as other Pimeliini, have intersegmental membranes between abdominal ventrites 3-5, in contrast with other Pimeliinae. Doyen (1994) believes that these membranes in Pimeliini are secondary exposed, because they do not have defensive glands, as in all other Pimeliinae. The correlation between the presence of intersegmental membranes and defensive glands was noted by Doyen (1972) himself, therefore reasons for this "possibly secondarily evolved specialization" (Doyen, 1994: 470) are unclear.

Male genitalia
The aedeagus in genera with dorso-lateral eyes is typical for Pimeliini, inverted 180°, and the apical piece without a long ventral apophysis. Species of the subtribe Habrobatina differ from species of the other Pimeliini by their strongly sub-angularly curved apical piece of the tegmen (Figs 14C, 15C, 16C, 17C). Other characters of the aedeagus can help to differentiate genera and generic sub-groups. Apatopsis and Habrochiton have small elevated sclerotized ventral apophyses of the apical piece (Figs 14A, 15A). The majority of genera of Habrobatina have more or less long, triangular, not merged dorsal apophyses of the apical piece (Figs 14B, 15B, 17B), excluding Habrobates with Y-like, thin, merged apophyses (Fig. 16B). The apical piece consists of paired lobes, which are usually merged, therefore the merged apophyses can be interpreted as the advanced state. The apical piece of species in Dietomorpha are covered with short, fi ne spines apically ( Fig. 17A-B), while species of other Habrobatina and Pimeliini, in general, have a bare apical piece. The median lobe is more or less uniform in the subtribe, only Apatopsis and Habrochiton have a rolled baсuli at the apex (Figs 14E, 15E), which is not typical for other pimeliine genera, and Habrobates has rod-like baculi that are fused apically (Fig. 16D-E). Other genera we studied have rod-like baculi that are separated, this character state is probably a symplesiomorphy within Tenebrionidae, as it is also shown for the evolution of the median lobe in Heleini (Matthews 1993).
The aedeagus in platyopoid genera is characterized by a weakly curved and glabrous apical piece ( The inner sternite VIII is similar in all Pimeliini, having a V-shaped or roundly-emarginate anterior margin. The most important is the structure of the gland of the sternite. This gland is well expressed among darkling beetles of the tribe Blaptini (Medvedev 2001;I. Chigray et al. 2018I. Chigray et al. , 2019I. Chigray & Ivanov 2020;etc.) and some Helopini (Nabozhenko & Ando 2018;Nabozhenko & Grimm 2019;Nabozhenko & Purchart 2019;etc.), but it has not been imaged or discussed for other Tenebrionidae. We did not fi nd this gland in some samples because it often dissolves when boiled in alkali, but several species have a well-expressed gland. Habrobates, for example, has paired short gland of the sternite VIII with separate ducts weakly sclerozited at the apex (Fig. 16I). Platyope and Earophanta have long not paired ducts of the gland, whereas the gland in Platyope is shorter and sclerotized only in the apical part (Figs 20H, 21I), while Earophanta has a very long and completely strongly sclerotized gland (Fig. 22H). The paired duct of glands, as well as its extension and sclerotization can be interpreted as advanced states.
The structure of the spiculum gastrale is similar in all pimeliine genera and differs only at the species level (sclerotization and form of derivatives of inner sternite IX).

Ovipositor
The Pimeliini is distinctly divided into two groups by the structure of the ovipositor. The majority of Pimeliini (subtribe Pimeliina) has an ovipositor that is partly transformed into a fossorial structure (Doyen 1994;Ren & Ba 2009;Leo 2016;S. Chigray 2019): the paraproct and proctiger are long and armed with long distinct baculi, apical lobe of coxite strongly sclerotized and transformed into glabrous shovel-like derivatives of inner sternite IX (Figs 18J-L, 19J-L, 10J-L, 21K-M, 22J-L). Species of the subtribe Habrobatina have the ovipositor transformed to a structure for sweeping away sand: paraproct and proctiger short, with reduced baculi, apical lobe of coxite sub-conical, membranous, with a brush of dense, long setae (Figs 14J-L, 15J-L, 16K-M, 17J-L). The coxite lobe III is the most transformed structure in Habrobatina. The ovipositor of Apatopsis has four apices with hair brushes (apomorphic state): two of these are on pair coxite lobe IV and two additional ones are formed from a strongly projecting apical part of pair coxite lobe III on the ventral side of the ovipositor (Fig. 14J-K). The coxite lobe III is conical in other genera, without additional processes apically, very short in Habrochiton (Fig. 15L) and much longer and larger than lobes II and IV in Habrobates (Fig. 16K) and Dietomorpha (Fig. 17J).
The ovipositor of the Platyope genus group does not differ from that in other Pimeliina, but many species have very dense pubescence of the coxite, which combines fossorial and the sand brushing functions (Figs 18J-L, 19J-L, 20J-L, 21K-M). The characters of the ovipositor of the ultrapsammophilic group Habrobatina are regarded as the advanced state.

Female genital ducts
The most simple structure (plesiomorphic state) is presented in the genus Mantichorula, species of which do not have the spermatheca (instead, only a short primary bursa copulatrix), and accessory gland of spermatheca short, thin, not branched and not modifi ed (without constriction and the basal narrow part or duct) (Fig. 23E). The further ways of specialization are associated with an increase in the volume of the spermatheca due to the lengthening of the single duct as in Earophanta (Fig. 23H) or by increasing the number of short ducts fl owing directly into the vagina as in Apatopsis (Fig. 23A) like those in Edrotini (Doyen 1994;S. Chigray 2018), or branched at the apex of the thickened basal duct of the spermatheca as in Platyope grumi (Fig. 23G). The complication of the accessory gland is achieved by increasing its volume and the formation of constrictions, similar to those in some Tentyriini (S. S. Chigray & Abakumov 2019): one basal in Habrobates, Przewalskia, Platyope, Earophanta (Fig. 23B, D, F-H,), two in Dietomorpha (Fig. 23C) or four in Apatopsis (Fig. 23A).
linguistic review and valuable comments and two reviewers for helpful corrections and questions. We also cordially thank Maxwell Barclay and Dmitry Telnov (BMNH) for providing material, important information on paratypes of Dietomorpha gonzalesi S. Chigray & Nabozhenko sp. nov. and valuable corrections. The authors are very grateful to Győző Szél, Aranka Grabant (HNHM), Christoph Germann (NMB), Eduard Khachikov (ZMMSU) and V.Yu. Savitsky (ZMMU) for provided material and some photographs.