taxonID	type	description	language	source
2161879CFFCE8A18FF7A36726465DBB6.taxon	diagnosis	Characteristics. Adults. Body ant-like, distinctly constricted between head and pronotum and between pronotum and elytra, elongate, convex or flattened, from yellowish to black, sometimes bicolored, with nearly black, dark brown or dark reddish-brown head and pronotum and testaceous, reddish or yellowish elytra; body length 1.10 – 8.70 mm. Head capsule elongate or transverse, prognathous; occipital constriction distinctly (often much) narrower than maximum width of head, dividing head capsule into exposed anterior part and narrow, subcylindrical ' neck' region largely retracted into prothorax; eyes typically located closer to mandibular bases than to occipital constriction, except for Papusus, in which eyes are submediam or subposterior; in some species eyes strongly reduced or absent; clypeus demarcated from frons by transverse groove or at least rapidly declined and at middle not confluent with frons. Labrum transverse, with variously deep anteromedian emargination (except in some Clidicus). Mandibles diverse, typically subtriangular, with mesal teeth (often more than one) and setose prostheca (except for Leptomastax, which has strongly modified and specialized mandibles). Maxillae with variously enlarged 4 - segmented palps, which are highly diverse in shape, different in each tribe. Submentum not demarcated laterally by sutures; mentum subtrapezoidal; prementum prominent; labial palpomere II enlarged, typically broadest and longest, palpomere III much narrower than II, strongly elongate and pointed at apex. Hypostomal ridges present, incomplete or nearly reaching posterior tentorial pits, the latter present and distinct, elongate or circular and always located in front of transverse impression demarcating ventrally ' neck' region. Gular plate lacking gular sutures, weakly transversely reticulated or nearly smooth. Antennal insertions located anterodorsally, between or in front of eyes; scape enlarged, often strongly so, with deep apical emargination located typically lateroventrally or ventrally, so that antennae can bend between scape and pedicel; antennae very weakly and gradually thickened distally or not thickened. Pronotum about as long as broad or (more frequently) elongate, broadest distinctly in front of middle or, rarely, at middle, with anterior corners rounded, posterior corners not marked or variously distinct and obtuse-angled, lateral edges or carinae absent; pronotal base lacking pits and grooves, or with an impressed transverse row of several pits, or with faint paired pits only. Basisternal part of prosternum as long as coxal part or longer, laterally fused with prothoracic hypomera, in some genera rudiments of notosternal sutures visible as short notches at anterior sternal margin or anterolateral margins of procoxal cavities; procoxal cavities broadly open or posteriorly delimited by overlapping (but not fused) postcoxal hypomeral lobes and lateral lobes of coxal portion of prosternum; prosternal process absent or developed as weakly elevated and diffuse carina, often broad anteriorly and narrowing posteriorly, in intact specimens hidden between contiguous procoxae. Hypomeral ridges absent. Mesoscutellum in intact beetles entirely or nearly entirely covered by posterior pronotal margin, subtriangular, cordiform or subtrapezoidal; scutoscutellar suture absent. Mesoventrite anteriorly forming a compact ' collar', which in intact beetles is inserted into posterior opening of prothorax (collar reduced in some Mastigini), typically with median subtriangular expansion directed posterad (absent in some Mastigini and some Clidicini) and posteriorly demarcated by a transverse setose impression (reduced in some Mastigini). Mesoventral intercoxal process distinctly but moderately broadly separating mesocoxae, variously developed but always flat or very weakly convex. Posterior margins of mesocoxal cavities typically carinate (indistinctly or not carinate in Mastigini). Metanotum largely reduced; structures associated with wing articulation absent, but alacristae, though somewhat shortened, are present even in entirely wingless genera. Metaventrite broadening posteriorly, with metanepisterna largely exposed in ventral view in intact beetles, usually conspicuously broadened (except in Leptomastax and Mastigini), metepimera strongly broadened; metacoxae broadly separated, posterior margin of metaventrite between metacoxae typically concave (straight in some Mastigini). Metendosternite (= furcasternum) with short, but sometimes very broad, laminar stem or virtually lacking stem and then lateral furcal arms are broadly separated and inserted each close to mesal margin of metacoxa. Abdomen longer than metaventrite; six abdominal ventrites visible, ventrite I longer than any subsequent ventrite. Elytra entire, suboval, lacking basal impressions, basal foveae and subhumeral carinae; humeri typically lacking humeral calli; elytral disc often with longitudinal rows of variously distinct punctures, sometimes very fine and barely discernible, rarely entirely absent. Hind wings absent or (less frequently) present. Legs with metacoxae about as long as broad, clearly divided into subglobose basal and subconical distal part. Aedeagus elongate, median lobe symmetrical or asymmetrical; parameres present (in Mastigini one paramere may be reduced to various extent), lacking apical macrosetae; endophallus lacking sclerotized anchoring structures, except for dense tiny denticles on its inflatable membrane, flagellum short and straight or long and then forming several loops; diaphragm present and located on the opposite wall in relation to basal foramen; aedeagus, when asymmetrical, often twisted inside abdomen in such a way that terms ' dorsal' and ' ventral' are ambiguous and instead ' parameral' and ' abparameral' are used. Ductus ejaculatorius with ' sperm pump', which is a variously developed sclerotized elongate thickening of the ductus provided with external longitudinal muscle fibers that allow for changing its length, and consequently its inner volume, the pump often has one or both its ends provided with sclerotized funnel-like structures. Spermatheca sclerotized, globular or strongly elongate, with a small accessory gland, ductus spermathecae moderately long, not coiled. Characteristics. Larvae. Body campodeiform, subcylindrical or flattened, subparallel-sided to strongly narrowing posterad; legs long and slender. Pigmentation from creamy-white to orange, brown, up to nearly black, typically head, and tergal plates darker than remaining regions. Body moderately densely setose, setae unmodified or thickened, in some taxa additional tiny leaf-like setae present, in some taxa cuticle covered with dense asperities. Head prognathous, usually weakly tilted downward in living larvae, with or without an annuliform ' neck' and with one pair of stemmata; epicranial stem and frontal sutures distinct; nasale lacking teeth, with a pair of long teeth or with several small setiferous papillate protuberances. Antenna longer than head (from slightly to strikingly so) and very slender, not clubbed, long antennomeres I and II of similar diameter, antennomere III small and in some taxa vestigial, in some genera antennomere II subdivided into three parts, accessory appendage of antennomere II strongly elongate, spatulate or subconical. Mandibles falciform, moderately to very slender, pointed, each with one submedian mesal tooth or lacking teeth; maxillary mala undivided or stipital projection of maxilla with two lobes; maxillary palp long, with all palpomeres elongate. Thoracic tergites with distinct ecdysial lines at least on pronotum and mesonotum; sterna with paired (2 – 4) small setose sternal plates. Ten abdominal segments present, urogomphi absent or present and then each composed of one elongate segment fused with posterolateral margin of abdominal segment IX. Spiracles annular, ventrolateral or lateral, nine pairs: one on mesothorax and eight pairs on abdominal segments I – VIII.	en	Paweł Jałoszyński (2018): World genera of Mastigitae: review of morphological structures and new ecological data (Coleoptera: Staphylinidae: Scydmaeninae). Zootaxa 4453 (1): 1-119, DOI: 10.11646/zootaxa.4453.1.1
2161879CFFCE8A18FF7A36726465DBB6.taxon	discussion	Remarks. Although Mastigitae can be easily distinguished from all other Scydmaeninae on the basis of the general appearance of adults, diagnosis of this group is problematic because of high degree of morphological diversity. They share with Scydmaenini the basisternal part of prosternum fused laterally with prothoracic hypomera, and the broadly separated metacoxae, but various forms of the prosternal-hypomeral fusion can also be found in some Glandulariini, and the metacoxae are broadly separated in Cephenniitae. The great diversity of the forms of maxillary palps makes it difficult to use this structure to define a supertribe, but this character system can be used to distinguish Mastigitae from similar Scydmaenini. In the latter group the maxillary palpomere IV is always minute, dome-shaped and symmetrical, always narrower than the apex of palpomere III. In Mastigitae, the palps are typically strongly, usually conspicuously enlarged and the palpomere IV can be strongly asymmetrical, forming a compact oval together with the palpomere III, or flattened and subtriangular, or suboval and strongly elongate. The apical emargination of the enlarged scape in Mastigitae is typically located ventrally or ventrolaterally, whereas a similar emargination of not enlarged or indistinctly enlarged scape in Scydmaenini is typically dorsolateral (or absent). Mastigitae and Scydmaenini (one of the four currently recognized tribes of Scydmaenitae) were suggested to be sister groups (Jałoszyński 2012 a), and Scydmaenitae requires a profound reclassification as a paraphyletic group. An unexplored character system that may represent an autapomorphy of the tribes currently classified in Mastigitae is the tentorium with its dorsal arms fused with the vertex, so that at least in some taxa sites of fusion are marked on the surface as shallow impressions or dark spots visible through a lightly pigmented cuticle. This fusion may be unique for Mastigitae, but the tentorium remains poorly studied, especially in Scydmaenini and Glandulariini. Composition and distribution. Mastigitae currently comprise six tribes with nine extant and six extinct genera; they include 128 extant species and subspecies, and fifteen extinct species. Two other extinct species, explicitly placed in or informally associated with Mastigitae, are here treated as Scydmaeninae incertae sedis or Coleoptera incertae sedis. The distribution of Mastigitae is strikingly disjunct and demonstrates the relict status of this group. Extant species of Mastigitae occur in the northern and north-eastern parts of the Mediterranean Basin, Southeast Asia and north-eastern Australia, South Africa, western United States, Central and northern part of Southern America. The largest numbers of genera and species are known from the Mediterranean Basin.	en	Paweł Jałoszyński (2018): World genera of Mastigitae: review of morphological structures and new ecological data (Coleoptera: Staphylinidae: Scydmaeninae). Zootaxa 4453 (1): 1-119, DOI: 10.11646/zootaxa.4453.1.1
2161879CFFCD8A1EFF7A36D261A7D8C6.taxon	diagnosis	Diagnosis. Leptomastacini differ from all other Mastigitae in several unique apomorphies: maxillary palpomeres III ̄ IV forming a compact oval, with distal margin of palpomere III and basal margin of IV strongly oblique, and palpomere IV broader than long (e. g. Fig. 17); most antennal flagellomeres transverse (e. g. Fig. 14); body with numerous modified setae (thickened, lanceolate or leaf-like) (e. g. Figs 19, 21, 65, 67), especially submental setae always conspicuously thickened (e. g. Fig. 15); basisternal part of prosternum largely asetose (even in taxa with dense setae on other body parts) except for rows of modified setae along anterior and posterior margins and rarely some median setae (e. g., Figs 18, 66); metaventrite with three pairs of anterolateral foveae (e. g., Fig. 20); aedeagus with symmetrical median lobe and parameres, but in repose positioned asymmetrically inside abdomen (Fig. 20). Additionally, Leptomastacini have a set of synapomorphies that occur in other genera, but never all together: mandible with one mesal tooth or lacking teeth; hypostomal ridges reaching to about middle between anterior submental margin and posterior tentorial pits; anterior and posterior margins of mesoventral setose impression with subtriangular or subtrapezoidal median asetose projections, so that the impression is horizontally 8 - shaped (e. g., Fig. 24); mesoventral intercoxal process slender, weakly but distinctly narrowing posterad (e. g., Fig. 21); and a row of several (3 – 5) long and thick coxal bristles on each mesocoxa (e. g., Fig. 25). Characteristics. Adults. Body small, 1.10 ̄ 2.89 mm in length, slender and flattened, usually light brown or even pale yellowish, rarely moderately dark brown, covered with setae, of which all or some are modified, thickened, lanceolate or short and leaf-like. Head of various shape, about as long as broad or transverse; antennal insertions anterodorsal, broadly separated, and each broadly separated from side of head and from mandibular base; vertex transverse, distinctly demarcated from ' neck' region, with a pair of conspicuously large posterolateral setiferous punctures; frons confluent with vertex, forming a subtriangular ' platform' between antennal insertions with step-wise anterolateral margins; clypeus strongly transverse; eyes vestigial or absent, often reduced to one ommatidium at each side located laterally at the level of antennal insertions. Gular plate transversely reticulated; posterior tentorial pits small, slot-like, C-shaped or straight; hypostomal ridges arcuate, posteriorly extending to middle between anterior submental margin and posterior tentorial pits. Postgenae with variously distinct longitudinal ridges, each extending anterad ventrally from occipital constriction (in one genus ridges absent). Antennae moderately slender, shorter than body; scape weakly but distinctly enlarged, subequal to length of head, about as long as 3 subsequent antennomeres, with distinct, deep ventral or ventrolateral apical emargination; pedicel weakly elongate, subtriangular, 1 / 2 ̄ 1 / 3 as long as scape; antennomeres III ̄ X transverse or some about as long as broad, rarely antennomere X indistinctly elongate; antennomere XI elongate and slightly asymmetrical. Antennomeres III ̄ XI each with narrow subcylindrical basal stalk and sharp basal marginal ring; all antennomeres densely covered with modified setae of two forms, short and subcylindrical or lanceolate, and long and lanceolate, setae inserted on small papillate elevations, so the surface of antennomeres appears coarse. Mouthparts. Labrum strongly transverse and with a deep anteromedian subtriangular emargination, with only a few dorsal setae. Mandibles symmetrical, each with one mesal tooth shifted slightly dorsad, or lacking teeth, subtriangular and robust or extremely slender and falciform, if subtriangular than with setose prostheca, when falciform then lacking prostheca. Maxillae generalized, as in all Scydmaeninae, except for maxillary palps, which have palpomeres III and IV forming a compact oval, without any constriction between strongly oblique apex and base of palpomeres III and IV; palpomere IV broader than long, rounded at apex; all palpomeres densely covered with modified setae. Submentum transverse, with a pair of long and conspicuously thick setae near anterior margin; submentum subtrapezoidal, with anterior margin straight or nearly straight; prementum prominent, largely membranous, with its anteromedian surface devoid of setae; insertions of labial palps broadly separated, labial palpomere I small and weakly elongate, palpomere II strongly enlarged, long and broadening distad; palpomere III narrow and distinctly longer than half length of palpomere II (in one genus palpomeres II and III subequal in length), pointed at apex. Pronotum elongate (sometimes weakly so), broadest in anterior third or fourth, with anterior and posterior margins nearly straight or arcuate, anterior corners broadly rounded, posterior corners strongly obtuse-angled, sides rounded, sometimes nearly straight in posterior half; pronotal base lacking pits or groove. Prosternum laterally fused with hypomera, but sometimes with traces of notosternal sutures marked just at anterior sternal margin or near procoxal cavities, with basisternal part about as long as coxal part or distinctly longer, largely asetose, only with a row of setae along anterior and posterior margin, in one genus additionally with some median setae; coxal part demarcated anteriorly and laterally by complete arcuate ridge extending posteriorly to postcoxal hypomeral lobes; prosternal carina short, narrow or diffuse and weakly elevated, not separating procoxae; procoxal cavities broadly open; each hypomeron behind procoxal cavity forming a small subtriangular postcoxal hypomeral lobe directed mesad. Mesoscutellum in intact beetles not visible between elytral bases, subtriangular or subtrapezoidal; scutoscutellar suture absent. Metanotum largely membranous, but with only slightly shortened alacristae; hind wings absent. Mesoventrite with prepecti and anteromedian area forming a massive transverse scaly sculptured ring or ' collar' inserted into prothorax, with a shallow and strongly transverse medioventral impression just behind anterior ridge, ventral area behind collar strongly concave, forming transverse setose impression filled with modified, dense and leaf-like setae, anteromedian and posteromedian margins of setose impression each with asetose subtriangular or subtrapezoidal projection directed respectively posterad and anterad, so that setose impression is horizontally 8 - shaped. Mesoventral intercoxal process slender and elongate, weakly but distinctly narrowing posterad, weakly convex or nearly flat, posteriorly fused with metaventrite. Mesepimera partly exposed in ventral view, demarcated from mesanepisterna by a ridge or fused, and from metepimera by an indistinct suture. Metaventrite subquadrate or slightly transverse; mesocoxal cavities with their anterior and posterior margins carinate; posterior adcoxal margin of ventrite strongly concave and with short adcoxal carina at each side; metaventral intercoxal process very short and broad, with concave or nearly straight posterior margin and subtriangular posterolateral corners. Metanepisterna broad or narrow, exposed in ventral view; metepimera very broad. Metaventrite with three pairs of foveae: lateral meso-metaventral foveae between meso- and metaventrite laterad mesocoxal insertions; lateral mesocoxal foveae posterolaterad mesocoxal insertions; and postmesocoxal foveae posterad mesocoxal cavities. In one genus metaventrite with median longitudinal carina. Elytra elongate, oval, lacking humeral calli, basal impressions and basal foveae; in one genus each elytron with one basal or sub-basal setiferous puncture. Elytral disc with longitudinal rows of punctures, in some species weakly marked. Abdomen longer than metaventrite, with 6 sternites visible; sternite III (first visible) much longer than each of IV ̄ VIII; posterior margin of sternite VIII in males not emarginate; suture between sternite VII and VIII distinct. Aedeagus elongate, weakly sclerotized (translucent), with symmetrical median lobe and symmetrical parameres, their apices rapidly curved mesad; flagellum straight or nearly straight. Aedeagus in repose asymmetrically positioned inside abdomen. Spermatheca globular or strongly elongate. Characteristics. Larvae. Only larva of the Mediterranean Leptomastax hypogea Pirazzoli, 1855 was described (Vít & De Marzo 1989); description given at Leptomastax.	en	Paweł Jałoszyński (2018): World genera of Mastigitae: review of morphological structures and new ecological data (Coleoptera: Staphylinidae: Scydmaeninae). Zootaxa 4453 (1): 1-119, DOI: 10.11646/zootaxa.4453.1.1
2161879CFFCD8A1EFF7A36D261A7D8C6.taxon	description	Composition and distribution. Leptomastacini currently comprise 30 species and subspecies classified into three genera. All species occur in the Western Palaearctic region, predominantly in the northern and eastern part of the Mediterranean Basin, with a few representatives known from the southern Carpathians, western Caucasus and the southern coastal region of Caspian Sea (" Hyrcania ") (Figs 7 – 9).	en	Paweł Jałoszyński (2018): World genera of Mastigitae: review of morphological structures and new ecological data (Coleoptera: Staphylinidae: Scydmaeninae). Zootaxa 4453 (1): 1-119, DOI: 10.11646/zootaxa.4453.1.1
2161879CFFCD8A1EFF7A36D261A7D8C6.taxon	discussion	Remarks. Leptomastacini are adapted to subterranean life, which is reflected by their small and flattened bodies, light pigmentation and reduction of composite eyes, wings and associated structures of the wing base. They have many unique apomorphies and are one of the best defined tribes, strongly supported as monophyletic unit in phylogenetic analyses (Jałoszyński 2012 b, 2016 a; Jałoszyński et al. 2018). In addition to characters given in the diagnosis, Leptomastacini may also be unique among Mastigitae (and all Scydmaeninae) in having the tarsal claws with longitudinal costae (Figs 76 – 77), but further study is required to verify whether it is stable or variable within genera.	en	Paweł Jałoszyński (2018): World genera of Mastigitae: review of morphological structures and new ecological data (Coleoptera: Staphylinidae: Scydmaeninae). Zootaxa 4453 (1): 1-119, DOI: 10.11646/zootaxa.4453.1.1
2161879CFFCB8A00FF7A36506249DB0E.taxon	diagnosis	Diagnosis. Ablepton differs from other Leptomastacini in two unique apomorphies: anterolateral corners of head capsule (i. e. sides of clypeus at each mandibular base) forming rounded lobes projecting anterolaterad (Figs 10 – 11); postmesocoxal foveae directed mesad (Fig. 20); and in a combination of synapomorphies that occur in some other tribes and genera, but never all together: head, pronotum and elytra uniformly covered with setae (Fig. 1); head subquadrate (Fig. 10); ' neck' region broader than half width of head (Fig. 12); postgenae with distinct longitudinal ridges (Fig. 12; lr); each mandible with a median mesal tooth (Fig. 11); median projections of anterior and posterior margins of setose impression of mesoventrite not meeting (Fig. 24); mesoscutellum subtriangular, with pointed tip (Fig. 26); metaventrite with median longitudinal carina (Fig. 21). Characteristics. Adult. Body (Figs 1 – 2) small, 1.40 ̄ 2.10 mm in length, light to moderately dark brown, flattened, sparsely setose, setae mostly modified, either flattened and broadened but strongly elongate or very short and broad, leaf-like. Head capsule (Figs 10 – 12) divided into large and exposed anterior part and smaller, subcylindrical ' neck' region retracted into prothorax and demarcated by distinct occipital constriction; ' neck' region broader than half width of head. Anterior part of head flattened and about as long as broad or slightly longer, subquadrate in shape, with rounded sides of vertex, broadest behind middle but far from posterior margin. Composite eyes (Fig. 10; ce) slightly anterior to middle, on sides of head, vestigial, each composed of 2 ̄ 3 small ommatidia which are barely discernible under stereoscopic microscope. Vertex transverse, convex, with posterior margin nearly straight at middle and a pair of large lateroposterior setiferous punctures (Fig. 10; sp). Tempora long and rounded. Frons between antennal insertions subtriangular, anteriorly demarcated by deep and narrow frontoclypeal groove (Fig. 10; fcg), which is obliterated laterally. Clypeus transverse, with subtriangular and rounded lateral projections behind external margin of each mandibular base. Antennal insertions (Fig. 10; ai) laterodorsal, in submedian area of head. Gular plate (Fig. 12; gp) lacking sutures, with transverse reticulation; posterior tentorial pits (Fig. 12; ptp) C-shaped and located in front of transverse impression demarcating ' neck' region ventrally. Head densely but finely punctate and covered with sparse flat, broad and moderately long setae. Antennae (Figs 1 – 2, 12, 14) much shorter than body; scape (Fig. 12; sc) only 3 ̄ 4 times as long as broad; pedicel only slightly longer than antennomere III and distinctly broadening from relatively narrow base to apex; antennomeres III – X (Fig. 14) mostly transverse and thickening distad, each with short and narrow stalk and sharp basal ring, antennomeres covered with dense and evenly distributed short and only slightly flattened setae and much longer, strongly flattened and broadened sparse setae around distal margin of each antennomere except I, II and XI, all setae inserted on small papillae, so that surface of antennomeres appears coarse. Labrum (Figs 10 – 11; lbr) strongly transverse, with slightly rounded and anteriorly divergent lateral margins and concave anterior margin with deep median subtriangular emargination; dorsal surface with only several long and thick setae. Mandibles (Figs 10 – 12) symmetrical, each subtriangular but relatively slender, with broad base and long, curved and moderately sharp apical tooth; mesal tooth (Figs 10 – 11; mt) present, subtriangular and shifted dorsad; setose prostheca (Fig 11; pst) present, small. Maxilla (Figs 15 – 16) with large and nearly semicircular cardo (Fig. 15; cd); basistipes (Fig. 15; bst) subtriangular and elongate; mediostipes (Figs 15 – 16; mst) large and sharply demarcated from lacinia (Figs 15 – 16; lac) and galea (Figs 15 – 16; gal), which are both elongate and curved mesad and each bears a dense row of flattened distal setae; palpifer (Figs 15 – 16; ppf) broad and elongate; maxillary palp (Figs 15 – 17) composed of minute palpomere I (Figs 15 – 16; mxp 1), elongate and broadening distad palpomere II (Figs 15, 17; mxp 2), and palpomeres III (Fig. 17; mxp 3) and IV (Fig. 17; mxp 4) forming large compact oval in which palpomere III is pedunculate and with strongly oblique apex and palpomere IV is broader than long and broadly subconical with rounded but distinctly marked apex. Labium (Fig. 15) with broad submentum (Fig. 15; smn) posteriorly not demarcated from gular region and laterally indistinctly demarcated from postcardinal portions of hypostomae by weakly marked and incomplete ridges; mentum (Fig. 15; mn) subtrapezoidal with anterior margin slightly concave; prementum (Figs 15 – 16; pm) long, subtrapezoidal, broadest distally, without demarcated ligula, with broadly separated bases of labial palps; lateral lobes of hypopharynx (Fig. 15; lhl) moderately large; labial palp composed of three palpomeres: palpomere I (Fig. 15; lp 1) small, elongated, palpomere II (Fig. 15; lp 2) largest, strongly elongated and broadened distally, palpomere III (Fig. 15; lp 3) narrow, long and pointed. All mouthparts and clypeus covered with sparsely distributed porous fields (Fig. 13; pf); maxillary palpomeres II ̄ IV covered with elongate and only slightly broadened and flattened setae (Fig. 17). Prothorax (Figs 1 – 2, 18 – 19) flattened and elongate, broadest near anterior third. Pronotum with all margins rounded; anterior corners not marked, posterior corners distinct, obtuse-angled; pronotal base lacking pits, impressions, grooves or carinae. Prosternum (Figs 18 – 19) with basisternal part (Figs 18 – 19; bstr) about as long or only slightly longer than coxal part (Fig. 19; cxst), laterally completely fused with hypomera and lacking any noticeable traces of notosternal sutures. Coxal region demarcated anteriorly by rounded carina extending laterally up to apices of subtriangular postcoxal hypomeral lobes (Fig. 18; pchl) projecting mesad. Procoxal cavities broadly open; profurcal foveae (Fig. 10; pff) small but distinct. Prosternal intercoxal process (Fig. 19) subtriangular and posteriorly very narrow, diffuse, indistinctly demarcated laterally and very weakly elevated, so that procoxae are not separated. Ventral surface of prothorax largely asetose and glabrous; short, broad and strongly flattened setae present along anterior and posterior margins of basisternal area (Figs 18 – 19). Mesoventrite (Figs 20 – 24) subtrapezoidal, broadening posteriorly. Prepecti (Fig. 21; pre) long and together with anteromedian mesoventral area forming continuous ' collar', with a narrow and shallow transverse groove just behind its anterior ridge (Fig. 21; ar) and deeply bisinuate posterior margin with rounded and asetose subtriangular posteromedian expansion. Region just behind collar strongly and abruptly constricted laterally and impressed medially, forming a transverse setose impression (Figs 20 – 23; si) filled with dense, strongly flattened, leaf-like setae. Posterior margin of setose impression forming a large median subtriangular projection directed anterad, but not meeting with anterior projection of collar. Mesoventral intercoxal process (Fig. 21; msvp) long, narrow and weakly convex, fully separating mesocoxae, slightly narrowing posteriorly and fused with metaventrite. Mesanepisterna (Fig. 23; aest 2) large and subtriangular, demarcated from mesepimera by carinate ridge; mesepimera (Fig. 23; epm 2) indistinctly demarcated from metepimera and partly exposed in ventral view (Fig. 21). Mesonotum with subtriangular and pointed mesoscutellum (Fig. 26; scl 2) not visible between elytral bases in intact specimens, scutoscutellar suture absent. Metaventrite (Figs 20 – 22) short, subtrapezoidal, broadest at level of metacoxae, with lateral margins rounded; mesocoxal cavities with nearly continuous marginal carina encompassing nearly entire cavity except for its external portion; posterior margin of metaventrite deeply bisinuate laterally in admetacoxal region and with a broad metaventral intercoxal process (Fig. 21; mtvp) with nearly straight posterior margin and subtriangular lateral corners weakly projecting posteriorly; metaventrite with median longitudinal carina (Fig. 21; mc) and three pairs of foveae, whose openings are filled with leaf-like setae: lateral meso-metaventral foveae (Figs 20 – 23; lmf) laterad mesocoxal insertions; lateral mesocoxal foveae (Figs 20; 22; lmcf) posterolaterad mesocoxal insertions; and postmesocoxal foveae (Figs 20 – 24; pmcf) posterad mesocoxal cavities, in submedian region of ventrite. Admetacoxal margin of metaventrite with a short thickening demarcated by a groove, forming adcoxal carina (Fig. 21; acxc). Metanepisterna (Figs 20 – 21, 23) partly visible in ventral view, relatively narrow, distinctly broadening posterad; metepimera (Fig. 23; epm 3) about three times as broad as metanepisterna, with indistinctly demarcated inner and outer component, posteriorly extending far behind metacoxae. Metendosternite (metafurca) Y-shaped, with very short but distinct stem and divergent lateral furcal arms (Fig. 20; lmfa), additionally with a short anteromedian projection. Legs (Figs 1 – 2, 21, 25, 28) moderately long, robust. Pro- and mesocoxa short subconical, metacoxa with a nearly hemispherical basal part and subconical distal part. Mesocoxa (Fig. 25; cx 2) with impressed lateral adtrochanteral area, border between concave and convex surface with a row of 3 ̄ 5 long coxal bristles (Fig. 25; cxb). All trochanters short and subtriangular. Femora weakly clavate. Tibiae robust, all nearly straight. Tarsi short, subcylindrical, tarsomeres reducing in length but not in width from I to IV, tarsomere V strongly elongate, with curved and slender claws and empodium with a pair of modified empodial setae. Elytra (Figs 1 – 2, 26) oval, flattened, lacking humeral calli and basal impressions, with rounded apices; elytral disc with large and deep punctures arranged in nearly complete longitudinal rows. Hind wings absent. Abdomen (Fig. 20) with sternite III not fused with metaventrite, about as long as sternites IV and V together; sternite VIII in male with rounded posterior margin. Sternite III with distinct anteromedian and indistinct anterolateral (postmetacoxal) impressions filled with leaf-like setae. Aedeagus (illustrated in (Jałoszyński et al. (2015, 2018 )) symmetrical and lightly sclerotized, strongly elongate, parameres partly fused to lateral walls of median lobe, with apices strongly curved mesad; flagellum simple and broad, not coiled. Ejaculatory duct with strongly elongate and narrow sperm pump, its distal end with a sclerotized funnel-like collar. Spermatheca (not illustrated) globular and thick-walled, nearly spherical, with ' stalk' as long as globular part and as broad as 1 / 3 of globular part; ductus spermathecae and duct of accessory gland inserted into the stalk. Sexual dimorphism expressed in protrochanters, which in males are subquadrate and with a subtriangular ventral (posterior) projection (Fig. 27). Larva. Unknown.	en	Paweł Jałoszyński (2018): World genera of Mastigitae: review of morphological structures and new ecological data (Coleoptera: Staphylinidae: Scydmaeninae). Zootaxa 4453 (1): 1-119, DOI: 10.11646/zootaxa.4453.1.1
2161879CFFCB8A00FF7A36506249DB0E.taxon	description	Composition and distribution. Only three species are known (but see Remarks), distributed in Romania and Serbia (Fig. 7). Natural history. Some of the Serbian species of Ablepton were collected by pitfall traps with vinegar placed between roots of old beech trees on a limestone-rich ground near a stream, some others in vicinity of caves. Specimens of Ablepton treforti studied by the author were obtained by sifting leaf litter in beech forests in Romania. Nothing else is known about the natural history of this genus.	en	Paweł Jałoszyński (2018): World genera of Mastigitae: review of morphological structures and new ecological data (Coleoptera: Staphylinidae: Scydmaeninae). Zootaxa 4453 (1): 1-119, DOI: 10.11646/zootaxa.4453.1.1
2161879CFFCB8A00FF7A36506249DB0E.taxon	discussion	Remarks. Within Leptomastacini, Ablepton closely resembles Taurablepton. These genera can be distinguished on the basis of the following characters: the shape of head (subquadrate in Ablepton vs. subtriangular in Taurablepton); anterolateral corners of head capsule at each mandibular base (forming rounded lobes projecting anterolaterad in Ablepton vs. not projecting in Taurablepton); median projections of anterior and posterior margins of setose impression of mesoventrite (not meeting in Ablepton vs. nearly touching in Taurablepton); postmesocoxal foveae (directed mesad and very close to each other in Ablepton vs. directed anteromesad and broadly separated in Taurablepton); the mesoscutellum (subtriangular with pointed tip in Ablepton vs. subtrapezoidal with truncate tip in Taurablepton). Additionally, Ablepton has the median area of prosternum asetose, whereas in Taurablepton there are some median setae on the basisternal region. The three species placed in Ablepton differ in so minor characters that they may in fact represent variability within one species. Nonveiller & Pavičevič (1990, 1999) stated that the two Serbian species do not differ in the structure of the aedeagus from the type species of the genus, A. treforti. Also external diagnostic characters of their newly described two Serbian species seem rather elusive. A larger material from the entire known distributional range of Ablepton must be studied to address this issue.	en	Paweł Jałoszyński (2018): World genera of Mastigitae: review of morphological structures and new ecological data (Coleoptera: Staphylinidae: Scydmaeninae). Zootaxa 4453 (1): 1-119, DOI: 10.11646/zootaxa.4453.1.1
2161879CFFD58A0DFF7A368865BDD856.taxon	description	Pyladus Fairmaire, 1856 a: 179. Type species: Pyladus coquereli Fairmaire, 1856 a (monotypy). Synonymized with Leptomastax by Fairmaire (1856 b). Pylades Fairmaire, 1856 b: 526. Type species: Pylades coquereli Fairmaire, 1856 a (monotypy). Probably a misspelling of Pyladus. Leptomastoides Karaman, 1962: 170 (as subgenus of Leptomastax; nomen nudum, no type species designated; treated as synonym of Leptomastax by Castellini (1996 )).	en	Paweł Jałoszyński (2018): World genera of Mastigitae: review of morphological structures and new ecological data (Coleoptera: Staphylinidae: Scydmaeninae). Zootaxa 4453 (1): 1-119, DOI: 10.11646/zootaxa.4453.1.1
2161879CFFD58A0DFF7A368865BDD856.taxon	diagnosis	Diagnosis. Leptomastax differs from other Leptomastacini in the following unique apomorphies: very large and flat head which is often broader than prothorax and strongly transverse (Figs 29 – 31); mandibles extremely long and slender, falciform and lacking mesal teeth (Figs 29 – 31); body appears asetose and only under high magnification flattened and broadened setae can be seen on antennae, maxillary palps, prosternum, and in foveate structures of metaventrite and first visible abdominal sternite (Figs 33 – 34, 37, 42); metanepisterna conspicuously narrow, so that in ventral view entirely visible, and also extremely broad metepimera partly exposed in intact beetles (Figs 42 – 44). Additionally, ' neck' region is subequal to half width of head, a character known in other tribes of Scydmaeninae, but not in other Leptomastacini; and postgenae have indistinct longitudinal ridges (Fig. 30; lr) (among Mastigitae present also in Ablepton). Characteristic. Adult. Body (Figs 3 – 4) small, 1.24 ̄ 2.89 mm in length, light brown or yellowish, flattened, dorsally virtually asetose, setae mostly modified, either flattened and broadened but strongly elongate or very short and broad, leaf-like, distributed mostly on antennae and ventral side of thorax. Head capsule (Figs 29 – 32) divided into large and exposed anterior part and much smaller, rounded and flattened ' neck' region retracted into prothorax and demarcated by distinct occipital constriction; ' neck' region about as broad as half width of head. Anterior part of head strongly flattened and remarkably broad, strongly transverse and as broad as pronotum or broader, broadest at anterior margin. Composite eyes (Fig. 29; ce) dorsolateral, near anterior margin of head, from moderately large and composed of about a dozen ommatidia to very small and indistinct, in some cases only one small ommatidium present (barely discernible under stereoscopic microscope); eyes, even when large, flat, in dorsal view not projecting laterad. Vertex strongly transverse and weakly, evenly convex, with posterior margin straight or nearly straight at middle and a pair of long posterolateral setae (often only setiferous punctures remain; Fig. 29; sp). Tempora very long and rounded. Frons between antennal insertions forming a short and broad subtrapezoidal ' platform', entire frons anteriorly demarcated by a deep, narrow frontoclypeal groove (Fig. 29; fcg) obliterated laterally. Clypeus very short and broad, with rounded sides. Antennal insertions (Fig. 29; ai) located dorsally in submedian area of head, broadly separated. Gular plate (Fig. 30; gp) lacking sutures, transversely reticulated; posterior tentorial pits (Figs 30, 32; ptp) arcuate, in front of transverse impression demarcating ' neck' region ventrally; hypostomal ridges (Fig. 30; hr) arcuate, posteriorly reaching middle between anterior submental margin and posterior tentorial pits. Head impunctate or very finely punctate and dorsally asetose. Antennae (Figs 3 – 4, 29 – 30, 33) much shorter than body; scape (Figs 29 – 31; sc) 4 ̄ 5 times as long as broad, often with dorsal longitudinal ridge; pedicel (Figs 30 – 31; pd) distinctly longer than antennomere III and strongly broadening from narrow base to apex; antennomeres III – X mostly transverse and weakly thickening distally, each with narrow and short basal stalk and sharp basal ring; antennomere XI elongate and slightly asymmetrical. Antennomeres covered with dense and evenly distributed short and slightly flattened setae and much longer, strongly flattened and broadened sparse setae around distal margin of each antennomere except I, II and XI (Fig. 33); all setae inserted on small papillae, so that surface of antennomeres appears coarse. Mouthparts. Labrum (Fig. 29; lbr) strongly transverse, with slightly concave or straight and anteriorly strongly convergent lateral margins and anterior margin with a deep subtriangular emargination occupying nearly its entire width; dorsal surface with only several long and thick setae anteriorly. Mandibles (Figs 29 – 31, 35 – 36) symmetrical, each extremely long and slender, longer than head, falciform, without mesal teeth and without prostheca, mandible in cross-section nearly circular or oval, surface near base with dense oblique rugae and in distal part with longitudinal grooves (Figs 35 – 36). Maxilla (Fig. 30) with large but relatively short cardo (Fig. 30; cd); basistipes (Fig. 30; bst) subtriangular and elongate; mediostipes (Fig. 30; mst) large and sharply demarcated from lacinia and galea, which are both elongate and curved mesad and with dense row of distal flattened setae; palpifer (Fig. 30; ppf) broad and elongate; maxillary palp composed of minute palpomere I (Fig. 30; mxp 1), elongate and broadening distally palpomere II (Figs 30 – 31; mxp 2), and palpomeres III (Figs 30 – 31, 34; mxp 3) and IV (Figs 30 – 31, 34; mxp 4) forming large compact oval, in which palpomere III is pedunculate and with strongly oblique apex and palpomere IV is broader than long and broadly subconical with rounded but distinctly marked apex. All palpomeres densely covered with lanceolate setae, only IV mostly with short and subcylindrical setae with admixture of lanceolate ones (Fig. 34). Labium (Fig. 30) with broad submentum (Fig. 30; smn) posteriorly not demarcated from gular region and laterally indistinctly demarcated from postcardinal portions of hypostomae by incomplete and weakly marked ridges, bearing a pair of long and thickened setae near its anterior margin; mentum (Fig. 30; mn) subtrapezoidal and strongly transverse, with anterior margin slightly concave; prementum (Fig. 30; pm) long, subtrapezoidal, broadest distally, lacking demarcated ligula, with broadly separated bases of labial palps (Fig. 30; lp); lateral lobes of hypopharynx moderately large; labial palp (Fig. 30; lp) composed of three palpomeres: palpomere I small, elongated, palpomere II largest, strongly elongated and slightly broadening distad, palpomere III narrow, long and pointed. All mouthparts and clypeus covered with sparsely distributed porous fields. Prothorax (Figs 37 – 39) flattened and elongate, broadest near anterior third or fourth. Pronotum with all margins rounded or sides in posterior half nearly straight; anterior corners broadly rounded, posterior corners indistinct, obtuse-angled and often completely rounded; pronotal base lacking pits, impressions, grooves or carinae. Prosternum (Figs 37 – 39) with basisternal part (Fig. 37; bstr) twice or more as long as coxal part (Fig. 39; cxst), laterally completely fused with hypomera but lateroposteriorly with short remnants of notosternal sutures (Fig. 37; nss) extending to 1 / 4 or less of basisternal length. Coxal region demarcated anteriorly by carina extending laterally up to apices of subtriangular adcoxal hypomeral lobes projecting mesad. Procoxal cavities broadly open. Prosternal intercoxal process narrowly subtriangular and posteriorly carinate, sharply demarcated laterally and weakly elevated, so that procoxae are not separated. Ventral surface of prothorax largely asetose and glabrous; sparse leaf-like setae distributed only along anterior and posterior margins of basisternal area (Figs 37, 39). Mesoventrite (Figs 40, 42 – 44) subtrapezoidal, broadening posteriorly. Prepecti long and together with anteromedian mesoventral area form a massive continuous ' collar', which bears a transverse groove just behind its anterior ridge, its posterior margin deeply bisinuate with subtriangular or subtrapezoidal posteromedian projection, which meets a similar anteromedian projection of posterior margin of setose impression (Figs 40, 42 – 43; si), separating the latter into two lateral halves. Region just behind collar strongly and abruptly constricted, ventrally forming setose impression (Figs 40, 42 – 43; si) filled with dense, strongly flattened, leaf-like setae. Mesoventral intercoxal process (Figs 40, 42; msvp) long, narrow and weakly convex but distinctly carinate at middle, fully separating mesocoxae, weakly but distinctly narrowing posterad and fused with metaventrite. Mesanepisterna (Fig. 44; aest 2) large and subtriangular, fused with mesepimera (Fig. 44; epm 2), the latter partly exposed in ventral view (Figs 40, 42). Mesonotum with subtriangular and pointed mesoscutellum (Fig. 41; scl 2), which in intact specimens is visible only between basal elytral articulating lobes; scutoscutellar suture absent. Metanotum largely membranous, but with alacristae only slightly shortened; hind wings absent. Metaventrite (Figs 42 – 44) short, subquadrate, with lateral margins rounded; mesocoxal cavities with marginal carina encompassing each cavity nearly entirely, except narrow lateral area; posterior margin of metaventrite deeply bisinuate laterally (in front of each metacoxa) and with a broad metaventral intercoxal process (Fig. 42; mtvp) with a deeply concave posterior margin and subtriangular posterolateral corners; metaventrite with three pairs of foveae, whose openings are filled with leaf-like setae: lateral meso-metaventral foveae (Figs 42 – 44; lmf) laterad mesocoxal insertions; lateral mesocoxal foveae (Fig. 43; lmcf) posterolaterad mesocoxal insertions; and postmesocoxal foveae (Figs 42 – 44; pmcf) posterad mesocoxal cavities, in submedian region of ventrite. External admetacoxal part of posterior metaventral margin with additional marginal thickening demarcated by a groove, forming adcoxal carinae (Fig. 42; acxc) at each side. Metanepisterna (Figs 42 – 44; aest 3) entirely visible in ventral view, very narrow, narrowing both anterad and posterad; metepimera (Figs 42 – 44; epm 3) more than eight times as broad as metanepisterna, with indistinctly demarcated inner and outer component, posteriorly extending far behind metacoxae. Metendosternite (metafurca) Y-shaped, with short but distinct stem and divergent lateral furcal arms, additionally with short anteromedian projection. Legs (Figs 3 – 4, 40, 42 – 43) moderately long, slender. Pro- and mesocoxa short subconical, metacoxa with nearly hemispherical basal part and subconical distal part. Mesocoxa with impressed lateral adtrochanteral area, border between concave and convex surface with 3 ̄ 4 long coxal bristles (Fig. 40; cxb). All trochanters short and subtriangular. Femora weakly clavate. Tibiae robust, all nearly straight. Tarsi short but slender, nearly subcylindrical, tarsomeres reducing in length but not in width from I to IV, tarsomere V strongly elongate, with curved and slender claws and small empodium bearing a pair of lanceolate empodial setae. Elytra (Figs 3 – 4, 41) oval, flattened, lacking humeral calli and basal impressions, with rounded apices; elytral disc with distinct large and deep punctures arranged in nearly complete longitudinal rows. Base of each elytron with large setiferous punctures (Fig. 41; sp) bearing a long and thickened seta (often broken off in dry-mounted specimens). Abdomen (Figs 42 – 43) with sternite III not fused with metaventrite, about as long as sternites IV and V together; sternite VIII in male with rounded posterior margin. Sternite III with indistinct anterolateral (postmetacoxal) impressions filled with leaf-shaped setae. Aedeagus (illustrated in Castellini (1996) and Jałoszyński et al. (2015, 2018 )) symmetrical and lightly sclerotized, strongly elongate, parameres partly fused to lateral walls of median lobe, with apices strongly curved mesad; flagellum simple and broad, not looped. Ejaculatory duct with strongly elongate and narrow sperm pump lacking funnel-like collars. Spermatheca (Fig. 43) globular and thick-walled, nearly spherical, with small accessory gland. Sexual dimorphism typically not evident, females can be slightly larger than males. Only in two species (L. orousseti Castellini, 1996 and L. piniphila Franz, 1988) males differ from females in having slightly broader profemora and protibiae, the latter with an apical projection or tooth; rarely the tarsomere I of male is enlarged. Characteristics. Larva. Body (Fig. 45) campodeiform, subparallel and flattened, whitish with testaceous head, moderately densely covered with thickened setae. Head prognathous, conspicuously large, broader than thorax, rhomboidal, strongly transverse, broadest near middle; posteriorly head capsule with a short annuliform ' neck'; with one pair of dorsolateral stemmata; epicranial stem and frontal sutures distinct; nasale with two long and broad teeth separated by a deep U-shaped emargination. Mandibles falciform and strikingly long and slender, much longer than head or antennae, apices pointed, mesal margins smooth; stipital projection of maxilla with two lobes, subapical lobe elongate and densely setose, apical lobe broad and bearing two modified thick setae; maxillary palp long, with all palpomeres elongate, palpomeres II and III comparable in length; labial palpomere II longer than I. Antenna only slightly longer than head, weakly clubbed, antennomeres I and II each strongly elongate and of similar length, but antennomere II broader than I and slightly broadened distad, broadest near apex; antennomere III small, strongly elongate, shorter than accessory appendage of antennomere II, which is strongly elongate, slightly asymmetrical, subconical and slightly bent near middle. Thoracic tergites with demarcated tergal plates and distinct ecdysial lines at least on pronotum and mesonotum. Abdomen with ten segments, all except X transverse; segment X elongate; urogomphi present, each composed of one elongate segment fused with posterolateral margin of abdominal segment IX and bearing several long apical bristles. Legs long and slender, densely covered with relatively short, spiny setae. Spiracles annular, lateral, nine pairs: one on mesothorax and eight pairs on abdominal segments I – VIII. Composition and distribution. Leptomastax comprises 23 species (one with two subspecies) distributed (Fig. 8) mainly in the northern and eastern parts of the Mediterranean Basin, with single species occurring east to the Black Sea and south to the Caspian Sea; in Spain, France, Italy, Croatia, Montenegro, Bosnia & Herzegovina, Serbia, Macedonia, Albania, Bulgaria, Romania, Greece, Turkey, Israel, Jordan, Lebanon, Georgia (Abkhazia) and Iran. Majority of species occur in mountains or highlands and in proximity of the Mediterranean Sea, Black Sea and Caspian Sea coastlines.	en	Paweł Jałoszyński (2018): World genera of Mastigitae: review of morphological structures and new ecological data (Coleoptera: Staphylinidae: Scydmaeninae). Zootaxa 4453 (1): 1-119, DOI: 10.11646/zootaxa.4453.1.1
2161879CFFD58A0DFF7A368865BDD856.taxon	description	Natural history. Leptomastax seems to be adapted to life in deep layers of soil, and its characters are most derived among Leptomastacini. Species of this genus can be collected by sifting the upper organic layers of soil (i. e. leaf litter), but also by the ' soil washing' technique, from mineral layer of soil. They can be found in deciduous forests in moist places (e. g., Figs 52, 55 – 56), but also in relatively dry habitats, as shrublands, in leaf litter accumulated under bushes (e. g., Figs 53 – 54). The extremely long mandibles clearly suggested predatory habits of Leptomastax, although no published records exist supporting this suspicion. I had an opportunity to make observations of three living adults of L. rousi, collected in Romania from the mineral soil layer. Beetles survived a week under laboratory conditions. During that time, they were not observed actively hunting for quickly-moving springtails, mites and other potential prey provided. Most of the time beetles vividly walked around the plaster of Paris arena, penetrating all spaces between particles of soil, rotten wood and mosses, inserting their flat heads and long mandibles into small crevices and pits in the surface of plaster of Paris, and easily climbing smooth plastic walls of the Petri dish. As a vented Petri dish was used, there was a tiny space between the vertical walls of the bottom dish and its lid to allow for air exchange. This space was large enough for beetles to insert the head between the lid and the upper surface of the wall of the bottom dish. This made it possible to make observations from above, through transparent plastic, of what beetles were doing with their heads inserted into this narrow space, providing an analog for similarly narrow spaces between soil particles. The flat head with very long mandibles functioned as a capturing device to detect any objects and drag them out, as long as it was possible to pierce such an object at least with one mandible. The ' probing' of this space (as well as those between other objects on the arena) was performed by beetles for most of the time, and one beetle would walk for hours on the wall of the arena just below the lid, inserting its head into the narrow space, spreading and closing its mandibles for several seconds, then retracing the head, moving one or two millimeters further on, inserting the head and again spreading and closing the mandibles. This behavior was repeated hundreds of times. When small dead isotomid springtails were placed in this space, they were sooner or later detected by one of the beetles on the arena, quickly captured in closing mandibles and pulled out. A successful beetle was observed manipulating the prey with its mandibles and front legs to move the springtail close to the maxillae, and the prey was nearly completely eaten within a few minutes. The space under the lid was even more frequently searched for food than spaces between soil, wood and moss particles on the arena. If wounded, slowlymoving, and often relatively large mesostigmatan mites and isotomid springtails were placed in such narrow crevices or under twigs of moss, they were found, pulled out and eaten in a similar way. Such a large prey (Figs 46 – 51), still moving (although not very vigorously), was observed being carried in the mandibles for 15 – 30 min, the beetle's mandibles were moving, but only slightly spreading and then strongly closing again, pulling the pray close to maxillae and inflicting more damage to the cuticle, which presumably facilitated penetration of prey's tissues by digestive juices. Living springtails and mesostigmatan mites of various sizes, when touched by beetles, were able to quickly escape and were not seen being captured even once during the one-week observation period. Although the above observations are only preliminary, they are valuable as representing the first record of feeding-related behavior of any member of Leptomastacini. It seems that Leptomastax uses a complicated threedimensional system of narrow spaces between soil particles to search for food, and its searching technique relies on many repeated insertions of its flat head into small crevices, where potential prey may hide, spreading and closing its enormously long, slender and curved mandibles, and, if a springtail or other invertebrate has been found, it is then momentarily pierced and pulled out of its shelter. In natural conditions, even a large, strong and fast-moving prey may probably be captured, if there is no space to jump or run. The very flat head and the very long and strongly curved mandibles form a perfect long-reach tool to probe narrow spaces, the slender and sharp mandibular apices pierce through prey's cuticle quickly, and closing, overlapping curved mandibles prevent a wounded prey from freeing itself from this firm grip. Moreover, the pair of long setae on the vertex of Leptomastax seems to function as tactile sensors, informing the beetle how deep it can insert its head into a crevice, as inserting the anterior body too deep could impede the function of front legs and slow down the process of retracting mandibles with a large and resisting prey. The unusually broad separation between mandibular bases, yet increased by the grotesquely broadened head, has a functional explanation. If the mandibles were similarly long (to allow for a long reach) but with narrowly separated bases, it would be impossible for the beetle to eat its prey while still held with mandibular apices. But when broadly separated and strongly curved mandibles are being closed, with their tips sunken into the prey's body, a springtail or mite is quickly pierced through and moved close to maxillae and the mouth opening, which increases damage to its cuticle and at the same time brings it into the range of digestive juices exuded from the beetle's mouth. As the larva of Leptomastax has a very similar shape of mandibles and head as adults, it seems possible that it also feeds in a similar way.	en	Paweł Jałoszyński (2018): World genera of Mastigitae: review of morphological structures and new ecological data (Coleoptera: Staphylinidae: Scydmaeninae). Zootaxa 4453 (1): 1-119, DOI: 10.11646/zootaxa.4453.1.1
2161879CFFD58A0DFF7A368865BDD856.taxon	discussion	Remarks. As reviewed and clarified by Castellini (1996), the grammatical gender of the name Leptomastax was problematic for several authors, and the name of the type species of the genus, L. hypogea, was several times emended to become masculine or neuter in grammatical gender, often gaining an additional " a " after " g ". Even Pirazzoli, who described this species as L. hypogeum, labeled a box with his specimens " hypogaeus mihi " (Castellini 1996). In fact, the Greek stem " mastax " (mouth) is feminine. In Leptomastax this stem is combined with the prefix derived from the Greek leptos (thin), and clearly refers to the slender mandibles. Morphological structures of Leptomastax were described and illustrated by Castellini (1996), but this author used a rather unorthodox Italian terminology, not always congruent with standard morphological terms used by other coleopterists. Moreover, some structures were either omitted or misinterpreted. The head in dorsal view was shown with a pair of dorsal tentorial pits (" fosetta tentoriale ") (Castellini 1996, fig. 2), whereas in fact there are no such pits. Because of the typically light pigmentation, the sites of fusion of dorsal tentorial arms with the head capsule can be seen through the cuticle as a pair of dark dots (similarly darkened insertions of dorsal tentorial arms are illustrated in a transparent mount of Taurablepton in Fig. 69; idta), which in dry-mounted specimens and in transparent mounts may be easily confused with tentorial pits. SEM examination demonstrates clearly that the vertex has not even traces of tentorial pits (Fig. 29). The hypostomal ridges were misinterpreted as sutures (Castellini 1996, fig. 3); the posterior tentorial pits were illustrated as circular (Castellini 1996, fig. 3), whereas in fact in Leptomastax the pits are developed as narrow, long and typically arcuate ' slots'. The stipes was illustrated as a structure composed of basistipes and palpifer completely and indistinguishably fused together (Castellini 1996, figs. 3, 9), whereas in all Scydmaeninae, including Leptomastax, the subtriangular basistipes is sharply demarcated from surrounding structures by sutures well-visible not only under SEM, but also in transparent mounts under light microscope. Details of meso- and metaventrite were simplified (Castellini 1996, fig. 13), and the conspicuous metaventral foveae filled with strongly modified, leaf-like setae, were not illustrated at all.	en	Paweł Jałoszyński (2018): World genera of Mastigitae: review of morphological structures and new ecological data (Coleoptera: Staphylinidae: Scydmaeninae). Zootaxa 4453 (1): 1-119, DOI: 10.11646/zootaxa.4453.1.1
2161879CFFD88A36FF7A35A064DFDB2B.taxon	diagnosis	Diagnosis. Taurablepton differs from other Leptomastacini in one unique apomorphy: mesoscutellum subtrapezoidal with truncated tip (Fig. 70); and a combination of synapomorphies shared with other tribes and genera, but not all together: head, pronotum and elytra uniformly covered with fine setae (Fig. 5); head subtriangular (Figs 57 – 58); ' neck' region broader than half width of head (Fig. 57); each mandible with a median mesal tooth; anterolateral corners of head capsule (i. e. sides of clypeus at each mandibular base) lacking projecting lobes; postgenae lacking longitudinal ridges; median projections of anterior and posterior margins of setose impression of mesoventrite nearly touching at middle (Fig. 67); postmesocoxal foveae directed anteromesad (Fig. 74); metaventrite lacking median longitudinal carina (Fig. 66). Characteristics. Adult. Body (Figs 5 – 6) small, 1.10 – 1.50 mm, light brown or yellowish, flattened, finely setose, most setae unmodified. Head capsule (Figs 57 – 58, 69; 72) divided into large and exposed anterior part and smaller, subcylindrical ' neck' region retracted into prothorax and demarcated by distinct occipital constriction; ' neck' region broader than half width of head. Anterior part of head flattened and subtrapezoidal in shape, narrower than pronotum and broadest near posterior margin. Eyes rudimentary or absent. Vertex strongly transverse and weakly, nearly evenly convex except for posteromedian subtriangular impression, posterior margin of vertex weakly concave, with a pair of small setiferous punctures (Figs 58 – 59; sp). Tempora very long and rounded. Frons between antennal insertions subtrapezoidal and forming ' platform' sharply demarcated anterolaterally; frontoclypeal groove marked only at middle and obliterated laterally. Clypeus short and broad, with sides rounded and not projecting anteriorly. Antennal insertions (Fig. 58; ai) located dorsally in submedian area of head. Gular plate (Fig. 57; gp) without sutures, transversely reticulate; posterior tentorial pits (Figs 57, 72; ptp) arcuate and located in front of transverse impression demarcating ' neck' region ventrally. Head finely and densely punctate and dorsally covered with short, sparse and slightly suberect setae. Antennae (Figs 5 – 6) much shorter than body; scape (Figs 57, 69, 72; sc) 3 ̄ 4 times as long as broad; pedicel (Figs 69, 72; pd) distinctly longer than antennomere III and strongly broadening from narrow base to apex; flagellomeres (Fig. 62) mostly transverse and weakly thickening distad; antennomeres covered with two kinds of setae: dense and evenly distributed short and only slightly flattened setae and much longer, strongly flattened and broadened sparse setae around distal margin; all setae inserted on small papillae, so that surface of antennomeres appears coarse. Basal rings of antennomeres III – XI (Fig. 63) finely serrate. Labrum strongly transverse, with slightly rounded and anteriorly divergent lateral margins and concave anterior margin with deep median subtriangular emargination; dorsal surface with only several long and thick setae. Mandibles symmetrical, each subtriangular but relatively slender, with broad base and long, curved and moderately sharp apical tooth; mesal tooth present, subtriangular and shifted dorsad in relation to the coronal plane of mandible; setose prostheca present, small. Maxilla (Figs 57, 72) with large and arcuate cardo (Figs 57, 72; cd); basistipes (Figs 57, 72; bst) subtriangular and elongate; mediostipes (Figs 57, 72; mst) large and sharply demarcated from lacinia (Figs 57, 72; lac) and galea (Figs 57, 72; gal) which are both elongate and curved mesally and each with dense row of distal flattened setae; palpifer (Figs 57, 72; ppf) broad and elongate; maxillary palp composed of minute and elongate palpomere I (Figs 57, 72; mxp 1), strongly elongate and broadening distally palpomere II (Figs 57, 72; mxp 2), and palpomeres III (Figs 57, 72; mxp 3) and IV (Figs 57, 72; mxp 4) forming large compact oval, in which palpomere III is pedunculate with strongly oblique apex and palpomere IV is shorter than broad and broadly subconical with rounded but distinctly marked apex. Palpomeres II – IV densely covered with lanceolate setae forming irregular longitudinal rows (Fig. 61). Labium (Figs 57, 72) with broad submentum (Figs 57, 72; smn) posteriorly not demarcated from gular region and laterally indistinctly demarcated from postcardinal portions of hypostomae by diffuse ridges; mentum (Figs 57, 72; mn) subtrapezoidal with anterior margin slightly concave; prementum (Figs 57, 72; pmn) long, subtrapezoidal, broadest near apex, without demarcated ligula, with broadly separated bases of labial palps; lateral lobes of hypopharynx moderately large; labial palp (Figs 57, 72; lp) composed of three palpomeres: palpomere I small, elongated, palpomere II largest, strongly elongated and broadening distally, palpomere III narrow, long and pointed. All mouthparts and clypeus covered with sparsely distributed porous fields. Prothorax (Figs 64 – 66, 71, 73) flattened and weakly elongate or nearly as broad as long, broadest near anterior forth. Pronotum (Fig. 64) with all margins rounded; anterior corners not marked, posterior corners also indistinct, obtuse-angled; pronotal base without pits, impressions, grooves or carinae. Prosternum with basisternal part (Figs 66, 73; bst) about as long or only slightly longer than coxal part (Fig. 73; cxst), laterally completely fused with hypomera, but in one species barely discernible traces of notosternal sutures (Fig. 66; nss) can be seen at anterior sternal margin. Coxal region demarcated anteriorly by carina extending laterally to middle of procoxal cavities. Procoxal cavities broadly open. Prosternal intercoxal process subtriangular and posteriorly narrow, sharply demarcated laterally and very weakly elevated, in intact specimens hidden between procoxae. Ventral surface of prothorax largely asetose and glabrous; short leaf-like setae present along midline and along anterior and posterior margins of basisternal area (Fig. 65). Mesoventrite (Figs 66, 71, 74) subtrapezoidal, broadening posteriorly. Prepecti (Fig. 74; pre) long and fused together with anteromedian mesoventral area to form a ' collar' with anteromedian transverse groove and posterior margin with rounded subtriangular posteromedian projection. Region just behind collar strongly and abruptly constricted laterally and impressed medially, forming setose impression (Figs 66, 71, 74; si), with its posterior margin forming a subtrapezoidal or subrectangular median projection nearly meeting anteromedian projection, so that setose impression is transversely 8 - shaped (Fig. 67). Mesoventral intercoxal process Figs 66, 68, 74; msvp) long, moderately narrow and nearly flat, fully separating mesocoxae, distinctly narrowing posteriorly where it is variously distinctly demarcated from metaventrite (border may be indistinct under SEM, as in Fig. 68, but distinctly visible in transparent mounts, Fig. 74). Mesepimera (Figs 66, 74; epm 2) partly visible in ventral view. Mesonotum (Fig. 70) with subtrapezoidal mesoscutellum, its apex truncate, scutellum not visible between elytral bases in intact specimens, scutoscutellar suture absent. Metaventrite (Figs 66, 71, 74) short, subquadrate or transversely subrectangular, with lateral margins rounded; mesocoxal cavities with nearly continuous marginal carina interrupted only laterally; posterior margin of metaventrite deeply bisinuate laterally and with a broad metaventral intercoxal process (Figs 66, 74; mtvp) with subtriangular posterior emargination; metaventrite with three pairs of foveae, whose openings are filled with leaflike setae: lateral meso-metaventral foveae (Fig. 74; lmf) laterad mesocoxal insertions; lateral mesocoxal foveae (Fig. 74; lmcf) posterolaterad mesocoxal insertions; and postmesocoxal foveae (Fig. 74; pmcf) posterad mesocoxal cavities, in submedian region of ventrite. Admetacoxal region of posterior metaventral margin with additional marginal thickening at each side, forming adcoxal carina (Figs 66, 74; acxc). Metanepisterna (Figs 66, 71, 74; aest 3) partly visible in ventral view, distinctly broadening posteriorly; metepimera broad, not exposed in intact specimens. Metendosternite (metafurca; Figs 71, 74) Y-shaped, with short but distinct stem and divergent lateral furcal arms (Figs 71, 74; lmfa), additionally with a short anteromedian projection. Legs (Figs 5 – 6, 66, 68, 71, 74 – 77) moderately long, robust. Pro- and mesocoxa suboval, metacoxa with nearly hemispherical basal part and subconical distal part. Mesocoxa with impressed lateral adtrochanteral area, border between concave and convex surface with 3 ̄ 4 long coxal bristles (Fig. 68; cxb). Pro- and mesotrochanters short and subtriangular, metatrochanters distinctly elongate. Femora weakly clavate. Tibiae robust, all nearly straight. Tarsi (Figs 75 – 77) short, nearly subcylindrical, tarsomeres reducing in length but not in width from I to IV, tarsomere V strongly elongate, with robust claws covered with longitudinal costae (Figs 76 – 77); empodium with a pair of lanceolate empodial setae (Figs 76 – 77, emps). Elytra (Figs 5 – 6) oval, flattened, lacking humeral calli and basal impressions, with rounded apices; elytral disc with distinct small, moderately deep or shallow and dense punctures arranged in indistinct rows, in some species barely discernible. Hind wings absent. Abdomen (Figs 66, 71, 74) with sternite III not fused with metaventrite, about as long as sternites IV and V together or slightly longer. Sternite III with indistinct and shallow anterolateral (postmetacoxal) impressions filled with short leaf-like setae. Aedeagus (illustrated in Jałoszyński et al. (2018 )) symmetrical and lightly sclerotized, strongly elongate, parameres partly fused to lateral walls of median lobe, with apices strongly curved mesad; flagellum simple and broad, not coiled. Ejaculatory duct with relatively short and narrow sperm pump, its both ends with broad sclerotized funnel-like structures. Spermatheca (Fig. 71; sp) thin-walled, strongly elongate, recurved. Larva. Unknown.	en	Paweł Jałoszyński (2018): World genera of Mastigitae: review of morphological structures and new ecological data (Coleoptera: Staphylinidae: Scydmaeninae). Zootaxa 4453 (1): 1-119, DOI: 10.11646/zootaxa.4453.1.1
2161879CFFD88A36FF7A35A064DFDB2B.taxon	description	Composition and distribution. Only three nominal species are known, inhabiting the south-central and southeastern part of Turkey (Fig. 9). Natural history. Species of Taurablepton were collected from leaf litter accumulated along a stream in a pine forest (Franz 1988), in leaf litter in a deep gorge (Besuchet 1969) and in soil under a partially rotten oak (Besuchet 1969). Nothing else is known about life of this enigmatic genus.	en	Paweł Jałoszyński (2018): World genera of Mastigitae: review of morphological structures and new ecological data (Coleoptera: Staphylinidae: Scydmaeninae). Zootaxa 4453 (1): 1-119, DOI: 10.11646/zootaxa.4453.1.1
2161879CFFD88A36FF7A35A064DFDB2B.taxon	discussion	Remarks. Morphological structures of Taurablepton and Ablepton are very similar, but these genera can be easily distinguished at first sight on the basis of a clearly different shape of the head (subtriangular in Taurablepton and subquadrate with rounded tempora in Ablepton). Other differences include: the sides of clypeus (in Taurablepton not forming rounded lobes projecting anterolaterad vs. with such lobes in Ablepton); the postgenae (in Taurablepton lacking longitudinal ridges vs. with distinct ridges in Ablepton); the median projections of anterior and posterior margins of setose impression of mesoventrite (nearly touching each other in Taurablepton vs. broadly separated in Ablepton); the postmesocoxal foveae (directed anteromesad in Taurablepton vs. mesad in Ablepton); the mesoscutellum (subtrapezoidal with truncated tip in Taurablepton vs. subtriangular and pointed in Ablepton); the median longitudinal carina of metaventrite (absent in Taurablepton vs. present in Ablepton). Possible differences are also the shape of spermatheca (strongly elongate in Taurablepton vs. globular in Ablepton) and basal rings of antennomeres III – XI (serrate in Taurablepton vs. smooth in Ablepton), but these characters were not studied in all species of Taurablepton and may be variable with this genus. Taurablepton shows a tendency toward reducing the rows of punctures on elytra; in some species they are relatively distinct, although less regular than those in Ablepton treforti and most species of Leptomastax, but in some the rows are indistinct, irregular, incomplete or even barely discernible. The holotype of Taurablepton subterraneum Franz, 1988, the type species of the genus, was not found in the Franz Collection neither by myself nor by the NHMW curators (H. Schillhammer, pers. comm.), but the shape of head described by Franz (1988) agrees with that in the two nominal and several undescribed species from the same area examined during the present study. Many males and females, possibly representing several species, are preserved in the collection of Heinrich Meybohm, all collected in Turkey. They include also specimens collected in the type localities of the nominal species. However, males that differ considerably in body length and proportions of body parts have nearly identical aedeagi; differences are rather subtle, and on the basis of external characters it is difficult to assign specimens to ' morphospecies', because shapes and measurements seem to change gradually. It is either one or two highly variable species, or a complex of populations that has only recently started differentiating. A larger sample of specimens and molecular methods used along with morphological analysis should be used to clarify this problem.	en	Paweł Jałoszyński (2018): World genera of Mastigitae: review of morphological structures and new ecological data (Coleoptera: Staphylinidae: Scydmaeninae). Zootaxa 4453 (1): 1-119, DOI: 10.11646/zootaxa.4453.1.1
2161879CFFE38A34FF7A36466567DFE8.taxon	diagnosis	Diagnosis. Clidicini differ from all other Mastigitae in two unique apomorphies: some setae on frons and vertex conspicuously long and erect among much shorter and denser basic vestiture (Fig. 84, but in some species indistinct); one or two conspicuously long and erect ventral setae present in basal half of each profemur (reduced in some species); and a combination of synapomorphies that can be found in other tribes and genera, but not occurring all together: only scape enlarged and as long as head or (more frequently) longer (e. g., Fig. 84); head capsule about as long as broad or slightly transverse (e. g., Fig. 90); antennal insertions broadly separated (e. g., Fig. 90); occipital constriction about half as broad as width of head or narrower (e. g., Fig. 90); eyes small, much shorter than tempora, anterior (e. g., Fig. 90); hypostomal ridges nearly reaching posterior tentorial pits; maxillary palpomere IV subtriangular (e. g., Fig. 93); mandibles with at least three (usually more) subapical teeth of various lengths, some located above, and some beneath the coronal plane of mandible (Fig. 92); pronotum with posterior collar demarcated by a transverse row of pits (e. g., Figs 78 – 79); mesoventral intercoxal process narrow, carinate, with subparallel lateral margins (Fig. 97); metacoxal margin of metaventrite with an angulate adcoxal expansion at each side (Fig. 97); metaventrite lacking foveae; elytral rows of punctures deep, distinct and regular (Figs 78 – 83); abdominal sternite VIII in males not emarginate; aedeagus heavily sclerotized and in repose positioned symmetrically inside abdomen; larva with subtrapezoidal head with short annular ' neck', each larval mandible with a preapical tooth, larval antennomere II undivided, larval antennomere III much longer than broad, sensory appendage spatulate, urogomphi present. Characteristics. Adults. Body (Figs 78 – 83, 86 – 88) small to large, in extant species 2.99 ̄ 8.5 mm in length, in nominal extinct species 2.23 – 7.33 mm, typically from light to dark, often reddish brown, in one species red with dark brown head; strongly convex, dorsally densely setose, setae typically conspicuously long and erect, unmodified except for variously developed longer and more erect bristles distributed on antennae, maxillary palps, head capsule and legs.	en	Paweł Jałoszyński (2018): World genera of Mastigitae: review of morphological structures and new ecological data (Coleoptera: Staphylinidae: Scydmaeninae). Zootaxa 4453 (1): 1-119, DOI: 10.11646/zootaxa.4453.1.1
2161879CFFE38A34FF7A36466567DFE8.taxon	description	Head capsule (Figs 84 – 85, 90 – 91) divided into large and exposed anterior part and much smaller, subcylindrical ' neck' region retracted into prothorax and demarcated by distinct occipital constriction; ' neck' region about as broad as half width of head. Anterior part of head strongly flattened, typically subequal in width with prothorax, weakly transverse or (rarely) about as long as broad, broadest behind middle or (rarely) at middle. Composite eyes dorsolateral, near anterior margin of head, small but composed of numerous small ommatidia, weakly convex. Vertex strongly transverse and posteriorly impressed at middle, convex at sides, with posterior margin concave; in one problematic extinct genus with a pair of long posterolateral setae. Tempora long and rounded. Frons between antennal insertions broadly subtrapezoidal, not forming a demarcated ' platform', anteriorly demarcated by a deep and complete frontoclypeal groove (Fig. 90; fcg). Clypeus very short and broad, with rounded or straight sides. Antennal insertions (Fig. 90; ai) located anterodorsally, broadly separated. Gular plate (Fig. 91; gp) lacking sutures, nearly smooth; posterior tentorial pits (Fig. 91; ptp) arcuate, in front of transverse impression demarcating ' neck' region ventrally; hypostomal ridges (Fig. 91; hr) nearly straight or arcuate, posteriorly nearly reaching posterior tentorial pits. Head with or without punctures, densely setose, often with conspicuously long setae (Fig. 84). Antennae (Figs 78 – 88, 95) shorter or about as long as body, slender; scape (Fig. 95; sc) 4 ̄ 6 times as long as broad, usually slightly broadening distad, with very deep lateroventral emargination; pedicel (Fig. 95; pd) typically shorter than, or less frequently about as long as antennomere III and broadening from narrow base to apex; antennomeres III – X elongate (usually strongly, less frequently weakly) and weakly thickening distad, basal stalks not exposed, basal rings absent or indistinct; antennomere XI elongate and slightly asymmetrical. Antennomeres covered with variously dense, long setae; surface of antennomeres in most specimens smooth. Mouthparts. Labrum (Fig. 92; lbr) strongly transverse, variable in shape, with lateral margins parallel or (more frequently) divergent anterad, nearly straight or rounded, and with anterior margin straight, concave, emarginated or notched at middle, or with a row of variously shaped anterior teeth or denticles, with a transverse dorsoanterior row of long setae. Mandibles (Figs 90 – 92) symmetrical, subtriangular and robust, with at least three (usually more) subapical teeth of various lengths, some located above, and some beneath the coronal plane of mandible, setose prostheca present and long. Maxilla (Fig. 94) with large but relatively short cardo (Fig. 94; cd); basistipes (Fig. 94; bst) subtriangular and elongate; mediostipes (Fig. 94; mst) large and sharply demarcated from lacinia (Fig. 94; lac) and galea (Fig. 94; gal), which are both elongate and curved mesally and each with a dense row of distal setae; palpifer (Fig. 94; ppf) broad and elongate; maxillary palp (Figs 84 – 85, 93 – 94) composed of minute palpomere I, elongate and broadening distad palpomere II, palpomere III (Fig. 93; mxp 3) subtriangular, broadening distad, with oblique distal margin, palpomere IV (Fig. 93; mxp 4) subtriangular, with pointed apex, its base typically narrower than apex of palpomere III or subequal in width; palpomeres III and IV often somewhat flattened. Maxillary palps can be much longer than head capsule (Fig. 84) or about as long as head (Fig. 85), typically in strongly elongate species with very slender antennae and legs also palps are long and slender (Fig. 82), and in those with relatively stout body and short antennae and legs (Fig. 80) the palps are proportionally shorter and with broader palpomeres III and IV. All palpomeres densely covered with variously long setae, palpomere II and / or III with some long and more erect setae among suberect and shorter basic vestiture (well-visible in Fig. 84, in some species indistinct). Labium (Figs 91 – 94) with broad submentum (Fig. 91; smn) posteriorly not demarcated from gular region, bearing a pair of long thin setae near its anterior margin or in subanterior region; mentum (Fig. 94; mn) subtrapezoidal and strongly transverse, with anterior margin straight or broadly rounded; prementum (Fig. 94; pm) long, subtrapezoidal, broadest distally, lacking demarcated ligula, with a pair of anterolateral setae, with broadly separated bases of labial palps; lateral hypopharyngeal lobes (Fig. 94; lhl) moderately large; labial palp (Fig. 91; lp) composed of three palpomeres: palpomere I (Fig. 94; lp 1) small, elongate, broadening distad, palpomere II (Fig. 94; lp 2) largest, strongly elongate and only indistinctly broadening distad, only in one problematic extinct taxon narrow at base and strongly broadened in distal half, palpomere III (Fig. 94; lp 3) narrow, long and pointed, in one problematic extinct taxon extremely slender, bristle-like. Porous fields on mouthparts and clypeus not found. Prothorax (Figs 78 – 85, 96) about as long as broad or elongate, strongly convex, broadest near anterior third. Pronotum with anterior and posterior margins arcuate or nearly straight, sides rounded in anterior half and usually sinuate in posterior half; anterior corners broadly rounded, posterior corners obtuse-angled; pronotal base with a short posterior ' collar' demarcated by a dorsal transverse row of several variously distinct pits, often connected by a groove. Prosternum (Fig. 96) with basisternal part (Fig. 96; bstr) subequal in length to coxal part, but proportions depend on the shape of pronotum, in species with short pronota the basisternal part tends to be indistinctly shorter than coxal part, in species with elongate pronota the basisternal part can be longer than coxal part. Prosternum laterally completely fused with hypomera. Coxal region demarcated anteriorly by carina extending laterally up to apices of very weakly developed adcoxal hypomeral lobes. Procoxal cavities broadly open. Prosternal intercoxal process indistinct, usually developed as a diffuse and weakly elevated ridge hidden between procoxae. Ventral surface of prothorax typically densely setose. Mesoventrite (Figs 84 – 85, 97) subtrapezoidal, broadening posteriorly. Prepecti (Fig. 97; pre) long and together with anteromedian mesoventral area forming a massive ' collar', which bears a transverse groove just behind its anterior ridge (Fig. 97; ar), its posterior margin bisinuate with short subtriangular or subtrapezoidal posteromedian projection, which is broadly separated from a similar anteromedian projection of posterior margin of setose impression (Fig. 97; si). Region just behind collar strongly and abruptly constricted, ventrally forming setose impression (Fig. 97; si) filled with dense, unmodified setae. Mesoventral intercoxal process (Fig. 97; msvp) long, narrow and weakly convex, nearly parallel-sided or indistinctly broadened near middle of mesocoxal cavities, fully separating mesocoxae, posteriorly fused with metaventrite or with variously distinct posterior tip. Mesanepisterna large and subtriangular; mesepimera (Fig. 97; epm 2) partly exposed in ventral view. Mesonotum with subtriangular, elongate mesoscutellum with rounded apex, in intact specimens only its very tip visible between elytral bases; scutoscutellar suture absent. Metanotum fully developed or somewhat reduced, with alacristae not or only slightly shortened; hind wings present or absent. Metaventrite (Fig. 97) short, subquadrate or subrectangular and slightly transverse, with lateral margins rounded; each mesocoxal cavity with posterior marginal carina, anterior margins not carinate; posterior margin of metaventrite deeply bisinuate laterally (in front of each metacoxa) and with a broad metaventral intercoxal process (Fig. 97; mtvp) with variously deeply concave posterior margin and subtriangular posterolateral corners; metaventrite lacking foveae. External admetacoxal part of posterior metaventral margin with additional marginal thickening demarcated by a groove, forming adcoxal carina (Fig. 97; acxc) at each side. Metanepisterna (Fig. 97; aest 3) broad and partly visible in ventral view, broadened posteriorly; metepimera broader than metanepisterna, with indistinctly demarcated inner and outer component, posteriorly extending far behind metacoxae. In one extinct species metaventrite with longitudinal median carina. Metendosternite (metafurca) Y-shaped, with short but distinct stem and divergent lateral furcal arms. Legs (Figs 78 – 88, 97) long and slender, often conspicuously so. Pro- and mesocoxa short subconical, metacoxa with nearly hemispherical basal part and subconical distal part. Mesocoxa lacking coxal bristles, but often with a comb of thin long setae. All trochanters short and subtriangular. Femora weakly clavate, profemur typically with at least one conspicuously long and erect ventral seta (rarely setae reduced). Tibiae robust, protibiae often curved in distal third or fourth. Tarsi long and slender, nearly subcylindrical, tarsomeres I – V subequal in length or reducing in length, tarsomere V strongly elongate, with curved and slender claws lacking elongate costae; empodial region was not studied. Elytra (Figs 78 – 83) oval, strongly convex, with variously distinct humeral calli, lacking basal impressions, with rounded or pointed apices; elytral disc with distinct large and deep punctures arranged in nearly complete and regular longitudinal rows. Elytra variously densely setose, setae typically long and erect. Abdomen with sternite III not fused with metaventrite, about as long as sternites IV and V together; sternite VIII in male with rounded posterior margin. Aedeagus (illustrated in Besuchet (1971), Jałoszyński (2009, 2018), O’Keefe & Monteith (2000), Orousset (2014), Zhou & Li (2015 )) strongly elongate, with symmetrical median lobe, parameres heavily sclerotized, symmetrical or one longer than the other; flagellum simple, not coiled, but often broadened and forming a sac-like structure. Ejaculatory duct with elongate and narrow sperm pump, typically with funnel-like structures at both ends, but in some species with one or both funnels absent. Aedeagus in repose positioned symmetrically inside abdomen, with basal orifice facing up. Spermatheca (illustrated e. g., by Orousset (2014 )) globular, variable in shape, in some species nearly spherical, in others elongate, thick-walled, with relatively large accessory gland. Characteristics. Larvae. Only a larva of the Australian Clidicus abbotensis O'Keefe, 2000 was described (O’Keefe & Monteith 2000); characteristics given at Clidicus.	en	Paweł Jałoszyński (2018): World genera of Mastigitae: review of morphological structures and new ecological data (Coleoptera: Staphylinidae: Scydmaeninae). Zootaxa 4453 (1): 1-119, DOI: 10.11646/zootaxa.4453.1.1
2161879CFFE38A34FF7A36466567DFE8.taxon	discussion	Composition and distribution. Clidicini, as currently defined, comprise 32 extant and extinct species classified in three genera. The extant species are distributed (Fig. 89) in the Oriental region and northern Australia; definitive fossils come from Cenomanian of Myanmar and Priabonian of central-eastern Europe. Extinct taxa that may belong to Clidicini are known from Upper Eocene Baltic amber, Lower Cretaceous Charentese French amber and Middle Miocene Chiapas amber. Remarks. Historically, Clidicini included Clidicus, Leptochromus, Papusus, and later also several extinct genera. Jałoszyński et al. (2018) redefined this tribe and restricted it only to Clidicus, with the problematic Middle Miocene Palaeoleptochromus placed in this tribe tentatively and pending further study. The same applies to the Upper Eocene Aenictosoma known from Baltic amber, which was originally described as possibly belonging to Cerambycidae (Schaufuss 1891), and later, solely on the basis of the original description, transferred to Scydmaeninae: Clidicini (Vitali 2006). See Remarks for each of these taxa below.	en	Paweł Jałoszyński (2018): World genera of Mastigitae: review of morphological structures and new ecological data (Coleoptera: Staphylinidae: Scydmaeninae). Zootaxa 4453 (1): 1-119, DOI: 10.11646/zootaxa.4453.1.1
2161879CFFF28A2EFF7A358C633CDF8F.taxon	diagnosis	Diagnosis. Leptochromus differs from all remaining Leptochromini in unique apomorphies: maxillary palpomere II with a long and slender cuticular projection with bifurcate apex and a pair of thick apical bristles (Fig. 121); and maxillary palpomere IV strongly elongate, nearly as long as palpomere III, only slightly broadened in subapical region and with subconical apex. Characteristics. Adults. Body (Figs 107 – 109) moderately large, 2.70 ̄ 4.20 mm in length, from yellowish to dark brown, strongly convex or slightly flattened, dorsally densely setose, setae long and suberect to erect, unmodified except for several long and thick bristles on protrochanters, postgenae, and maxillary palpomere II. Head capsule (Figs 114 – 116, 119 – 120, 124, 127 – 129) divided into large and exposed anterior part and much smaller, subcylindrical ' neck' region retracted into prothorax and demarcated by distinct occipital constriction; ' neck' region about as broad as half width of head. Anterior part of head strongly flattened, subequal in width with prothorax, strongly transverse, broadest at eyes, slightly in front of middle. Composite eyes dorsolateral, near anterior margin of head, moderately large, composed of numerous small ommatidia, strongly convex. Vertex strongly transverse and posteriorly impressed at middle, convex at sides, with posterior margin concave. Tempora long and rounded. Frons between antennal insertions with straight margin, not forming a demarcated ' platform', anteriorly demarcated by a deep and complete frontoclypeal groove. Clypeus very short and broad, with rounded or straight sides. Antennal insertions located anterodorsally, very broadly separated. Gular plate (Fig. 129; gp) lacking sutures, nearly smooth; posterior tentorial pits (Figs 124, 129; ptp) arcuate, in front of transverse impression demarcating ' neck' region ventrally; hypostomal ridges (Figs 124, 129; hr) nearly straight or arcuate, posteriorly reaching middle between anterior submental margin and posterior tentorial pits. Postgenae (Figs 115 – 116, 119 – 120, 124, 127 – 129) in most species with a tubercle or projection (postgenal or subocular process) with 2 – 3 thick apical bristles. Head with or without punctures, densely setose (Fig. 115). Antennae (Figs 107 – 109, 114 – 116, 127) shorter or slightly longer than body, slender; scape (Figs 114 – 116, 127; sc) 6 to more than 10 times as long as broad, usually slightly broadening distad, longer than head, with very deep lateroventral emargination; pedicel (Figs 114, 115, 127; pd) subequal in length to antennomere III or shorter and broadening from narrow base to apex; antennomeres III – XI elongate (usually strongly, rarely weakly) and weakly thickening distad, basal stalks not exposed, basal rings absent or indistinct; antennomere XI elongate and slightly asymmetrical. Antennomeres covered with variously dense, long setae; surface of antennomeres smooth. Mouthparts. Labrum strongly transverse, with lateral margins divergent anterad, nearly straight or weakly rounded, and with anterior margin concave, with a pair of broad and short submedian teeth separated at middle by a relatively shallow emargination, and with a transverse dorsoanterior row of long setae. Mandibles symmetrical, subtriangular and robust, each with two subapical teeth, at least one of them above the coronal plane of mandible, setose prostheca present and long. Maxilla (Figs 124, 129) with large but relatively short cardo (Fig. 124; cd); basistipes (Fig. 124; bst) subtriangular and elongate; mediostipes (Fig. 124; mst) large and sharply demarcated from lacinia and galea, which are both elongate and curved mesally and each with dense row of distal setae; palpifer (Fig. 124; ppf) broad and elongate; maxillary palp (Figs 114, 116, 121, 127, 129) conspicuously long, much longer than head capsule, composed of minute palpomere I (Fig. 121; mxp 1), extremely elongate, slender, only slightly broadening distad palpomere II (Fig. 121; mxp 2), which is distinctly recurved and bearing a long, slender bifurcate cuticular process with two apical bristles, palpomere III (Fig. 121; mxp 3) strongly elongate, slender, broadened only slightly in its distal third or fourth, with transverse distal margin, palpomere IV (Fig. 121; mxp 4) nearly rod-like, strongly elongate, slender, only slightly shorter than III, slightly broadened distad up to its distal third and then narrowed to subconical and blunt apex. Palpomeres II ̄ IV round or nearly round in crosssection, covered with moderately long setae. Labium (Figs 124, 129) with broad and short submentum (Fig. 124; smn) posteriorly not demarcated from gular region, bearing a pair of long thin setae in its subanterior region; mentum (Fig. 124; mn) subtrapezoidal and strongly transverse, with anterior margin weakly concave; prementum (Fig. 124; pm) long, subtrapezoidal, broadest distally, lacking demarcated ligula, with one or two pairs of anterolateral setae, with broadly separated bases of labial palps; lateral hypopharyngeal lobes (Fig. 124; lhl) moderately large; labial palp (Fig. 124; lp) composed of three palpomeres: palpomere I small, weakly elongate, broadening distad, palpomere II largest, strongly elongate and barrel-shaped, palpomere III narrow, long and pointed. Porous fields on mouthparts and clypeus not found. Prothorax (Figs 107 – 108, 125, 131) elongate, strongly convex, broadest near anterior third or at least distinctly in front of middle. Pronotum with anterior and posterior margins arcuate or nearly straight, sides rounded in anterior half and sinuate in posterior half; anterior and posterior corners obtuse-angled; pronotal base with a short posterior ' collar' (Fig. 131; prcl) demarcated by a dorsal transverse row of several variously distinct pits connected by a groove. Prosternum (Figs 125, 130) with basisternal part (Fig. 125; bstr) much longer than coxal part (Fig. 125; cxst). Prosternum laterally completely fused with hypomera. Coxal region demarcated anteriorly by carina extending laterally up to apices of very weakly developed adcoxal hypomeral lobes. Procoxal cavities broadly open. Prosternal intercoxal process indistinct or absent. Ventral surface of prothorax densely setose. Mesoventrite (Figs 126, 132) subtrapezoidal, broadening posteriorly. Prepecti (Fig. 126; pre) long and together with anteromedian mesoventral area forming a massive ' collar', which bears an indistinct transverse groove just behind its anterior ridge, posterior margin of collar broadly subtriangular, only slightly projecting posterad at middle, tip of this median projection is broadly separated from a similar anteromedian projection of posterior margin of setose impression (Fig. 126; si), margins of impression are obscured by setae (Fig. 132). Region just behind collar strongly and abruptly constricted, ventrally forming setose impression (Figs 126, 132; si) filled with dense, unmodified setae. Mesoventral intercoxal process (Figs 126, 132; msvp) long, narrow and weakly elevated, nearly parallel-sided or indistinctly broadened near middle of mesocoxal cavities, fully separating mesocoxae, posteriorly with indistinct posterior tip, which is fused with metaventrite. Mesanepisterna large and subtriangular; mesepimera partly exposed in ventral view. Mesonotum with subtriangular, strongly elongate mesoscutellum (Fig. 122; scl 2) with pointed or rounded apex, in intact specimens only its very tip visible between elytral bases; scutoscutellar suture absent. Metanotum and hind wings fully developed. Metaventrite (Figs 123, 126, 132) short, subquadrate or subrectangular and slightly transverse, with lateral margins rounded; mesocoxal cavities not carinate; posterior margin of metaventrite deeply bisinuate laterally (in front of each metacoxa) and with a broad metaventral intercoxal process (Fig. 126; mtvp) with variously deeply concave posterior margin and subtriangular posterolateral corners; metaventrite with lateral mesocoxal (Figs 123, 126; lmcf) and postmesocoxal (Figs 123, 126; pmcf) foveae. Posterior metaventral margin lacking adcoxal carinae. Metanepisterna (Fig. 126; aest 3) broad and partly visible in ventral view, broadened posteriorly; metepimera broader than metanepisterna, with indistinctly demarcated inner and outer component, posteriorly extending far behind metacoxae. Metendosternite (metafurca) with stem much broader than long and broadly separated divergent lateral furcal arms (Fig. 126; lmfa), additionally with median longitudinal projection. Legs (Figs 107 – 109, 117 – 118, 126, 130, 132) conspicuously long, slender. Pro- and mesocoxa short subconical, metacoxa with nearly hemispherical basal part and subconical distal part. Mesocoxa lacking coxal bristles, but usually with a row or patch of thin long setae. All trochanters short and subtriangular; protrochanters (Figs 118, 125, 130) in both sexes with a row of several thick bristles (in one species reduced to 1 – 2). Femora weakly clavate, profemur in both sexes with a ventral row of several conspicuously thick bristles, usually with papillate insertions (Figs 118, 130). Tibiae slender, protibiae in both sexes curved, only rarely nearly straight. Tarsi long and slender, nearly subcylindrical, tarsomeres I – V subequal or reducing in length, tarsomere V strongly elongate, with curved and slender claws lacking elongate costae; empodial region was not studied. Elytra (Figs 107 – 108, 131) oval, strongly convex, with prominent humeral calli (Fig. 131; hcl), lacking basal impressions, with rounded apices; elytral disc with distinct large and deep punctures arranged in six nearly complete and typically regular longitudinal rows. Elytra variously densely setose, setae long and erect. Abdomen (Fig. 132) with sternite III not fused with metaventrite, about as long as sternites IV and V together; sternite VIII in male distinctly emarginate (Fig. 134). Aedeagus (illustrated in Jałoszyński et al. (2018), Lord et al. (2014) and O’Keefe (2002 )) strongly elongate, with symmetrical median lobe, parameres heavily sclerotized, symmetrical, with arrowhead-shaped or spatulate and pointed apices; flagellum simple, not coiled. Ejaculatory duct with elongate and narrow sperm pump lacking funnel-like structures. Aedeagus in repose positioned symmetrically inside abdomen, with basal orifice facing up. Spermatheca (illustrated by O’Keefe (2002 )) globular, nearly spherical, with relatively large accessory gland. Larva. Unknown.	en	Paweł Jałoszyński (2018): World genera of Mastigitae: review of morphological structures and new ecological data (Coleoptera: Staphylinidae: Scydmaeninae). Zootaxa 4453 (1): 1-119, DOI: 10.11646/zootaxa.4453.1.1
2161879CFFF28A2EFF7A358C633CDF8F.taxon	description	Composition and distribution. Leptochromus comprises five extant species distributed in Brazil, Colombia, Costa Rica and Panama (Fig. 113), and one extinct species (Fig. 110) found in the Middle Miocene Chiapas amber. Natural history. Little is known about natural history of Leptochromus, except collecting circumstances for some species. Adults were collected from moist leaf litter, moss and lichens in subtropical forests (Figs 135 – 137). Numerous individuals of Leptochromus laselva were collected " lifting dead palm fronds lying on the floor of secondary-growth forest (...) and beating them over a white sheet during late afternoon and early evening hours " (Lord et al. 2014). The latter authors also noted that although adults of L. laselva are fully winged, no individuals were collected at light (including UV) for two weeks at the collecting site. Leptochromus fulvescens and L. laselva were also collected by canopy fogging, demonstrating that members of this genus are active not only on the forest floor (Lord et al. 2014). The unusual modifications of the maxillary palpomere II and postgenae may be used by beetles as a sensory and / or prey-capture apparatus, but hunting and feeding-related behavior of Leptochromus have not been observed.	en	Paweł Jałoszyński (2018): World genera of Mastigitae: review of morphological structures and new ecological data (Coleoptera: Staphylinidae: Scydmaeninae). Zootaxa 4453 (1): 1-119, DOI: 10.11646/zootaxa.4453.1.1
2161879CFFF28A2EFF7A358C633CDF8F.taxon	discussion	Remarks. All nominal extant and extinct species of Leptochromus were adequately described or redescribed by O’Keefe (2002) and Lord et al. (2014).	en	Paweł Jałoszyński (2018): World genera of Mastigitae: review of morphological structures and new ecological data (Coleoptera: Staphylinidae: Scydmaeninae). Zootaxa 4453 (1): 1-119, DOI: 10.11646/zootaxa.4453.1.1
2161879CFFFB8A2DFF7A330860EAD857.taxon	diagnosis	Diagnosis. Euroleptochromus differs from all remaining Leptochromini in the following unique apomorphies: maxillary palpomere II with short and broad submedian protuberance bearing apical bristles; and maxillary palpomere IV suboval and only about twice as long as broad. Additionally, a long cuticular process bearing apical bristles is present on each postgena (shared with some Leptochromus, but not with Rovnoleptochromus). Composition and distribution. Euroleptochromus (Fig. 111) comprises two extinct species known from the Upper Eocene Baltic amber.	en	Paweł Jałoszyński (2018): World genera of Mastigitae: review of morphological structures and new ecological data (Coleoptera: Staphylinidae: Scydmaeninae). Zootaxa 4453 (1): 1-119, DOI: 10.11646/zootaxa.4453.1.1
2161879CFFFB8A2DFF7A330860EAD857.taxon	discussion	Remarks. Euroleptochromus, except for characters listed in the diagnosis, closely resembles Leptochromus in all remaining structures.	en	Paweł Jałoszyński (2018): World genera of Mastigitae: review of morphological structures and new ecological data (Coleoptera: Staphylinidae: Scydmaeninae). Zootaxa 4453 (1): 1-119, DOI: 10.11646/zootaxa.4453.1.1
2161879CFFF88A2DFF7A31BB6375DE6C.taxon	diagnosis	Diagnosis. Rovnoleptochromus differs from all remaining Leptochromini in one unique apomorphy: maxillary palpomere II lacking cuticular projection but bearing one long bristle; and a combination of characters that, in a different set, occur in other genera: postgenae bearing long bristles but lacking cuticular postgenal processes; maxillary palpomere III broadening from base to apex and with strongly oblique apex; maxillary palpomere IV indistinctly subtriangular, broadest at base; mesocoxal cavities with carinate posterior margins; and metaventrite with a median longitudinal carina. Composition and distribution. Rovnoleptochromus (Fig. 112) comprises one extinct species known from the Upper Eocene Baltic (Rovno) amber.	en	Paweł Jałoszyński (2018): World genera of Mastigitae: review of morphological structures and new ecological data (Coleoptera: Staphylinidae: Scydmaeninae). Zootaxa 4453 (1): 1-119, DOI: 10.11646/zootaxa.4453.1.1
2161879CFF808A55FF7A307C627FDFC0.taxon	diagnosis	Diagnosis. Papusus is the only genus of Papusini; diagnosis and characteristics as for tribe, vide supra. Composition and distribution. Papusus comprises nine species distributed in Mexico and USA (Arizona, California, Colorado, Nevada, Utah) (Fig. 140).	en	Paweł Jałoszyński (2018): World genera of Mastigitae: review of morphological structures and new ecological data (Coleoptera: Staphylinidae: Scydmaeninae). Zootaxa 4453 (1): 1-119, DOI: 10.11646/zootaxa.4453.1.1
2161879CFF808A55FF7A307C627FDFC0.taxon	description	Natural history. Species of Papusus are unique among all Scydmaeninae in inhabiting the driest deserts of North America (Great Basin Desert, including the Death and Saline Valleys, Mojave Desert, Colorado Desert and Sonora Desert); moreover, adults were collected only during the hottest seasons of the year (O’Keefe 2003). Beetles were collected predominantly by pitfall traps set in the Shedscale Scrub and Creosote Bush Scrub communities, and in deserts with Agave, Amrosia, Atriplex, Yucca, Fouquieria, Pachycereus, Lysiloma and Stenocereus. Adults were found most often in alkali scrub or sand habitats with scattered vegetation, but not on open sand dunes. O'Keefe (2003) observed beetles walking on sand at night and found individuals under volcanic rocks 10 – 20 cm in diameter. He concluded that " in the Great Basin Desert adults of Papusus are active during the warmest time of the year and are most likely nocturnal " (O'Keefe 2003).	en	Paweł Jałoszyński (2018): World genera of Mastigitae: review of morphological structures and new ecological data (Coleoptera: Staphylinidae: Scydmaeninae). Zootaxa 4453 (1): 1-119, DOI: 10.11646/zootaxa.4453.1.1
2161879CFF808A55FF7A307C627FDFC0.taxon	discussion	Remarks. All species were revised by O'Keefe (2003).	en	Paweł Jałoszyński (2018): World genera of Mastigitae: review of morphological structures and new ecological data (Coleoptera: Staphylinidae: Scydmaeninae). Zootaxa 4453 (1): 1-119, DOI: 10.11646/zootaxa.4453.1.1
2161879CFF808A41FF7A335E6062DADE.taxon	description	Mastigini Reitter, 1882 b: 142. Type genus: Mastigus Latreille, 1802. Tribe was redefined by Jałoszyński et al. (2018).	en	Paweł Jałoszyński (2018): World genera of Mastigitae: review of morphological structures and new ecological data (Coleoptera: Staphylinidae: Scydmaeninae). Zootaxa 4453 (1): 1-119, DOI: 10.11646/zootaxa.4453.1.1
2161879CFF808A41FF7A335E6062DADE.taxon	diagnosis	Diagnosis. Mastigini differ from all remaining Mastigitae in unique autapomorphies: head with sharply marked median longitudinal groove on posteriorly impressed vertex; maxillary palpomere IV inversely suboval or indistinctly subtriangular, slightly asymmetrical, broadened from base to about distal third and with broadly rounded subtriangular apex, longer than broad; submentum with many setae, lacking one outstanding pair of anterolateral setae (but this character was not possible to see in † Baltostigini and in extinct Mastigini); mentum anteriorly very deeply emarginate, so that anterolateral corners form triangular and usually pointed lobes projecting anterad (also this character was not possible to examine in † Baltostigini and extinct Mastigini); first visible abdominal sternite firmly fused with metaventrite; and aedeagus with asymmetrical median lobe and asymmetrical parameres, with one paramere distinctly shorter than the other one (in some cases only one paramere is visible, the other one is either vestigial or completely obliterated), aedeagus in repose asymmetrically rotated inside abdomen, flagellum very long, with several coils, endophallus permanently everted on tip of elongate copulatory piece; larva with setose frontal impression; larval antennomere II subdivided into three sections; larval antennomere III vestigial, developed as a barely discernible papilla near much longer accessory appendage of palpomere III; larval palpomeres I and II with conspicuously long spines; and larva lacking urogomphs (but larva unknown in one extant and one extinct genus). Additionally, Mastigini share with † Baltostigini synapomorphies not known in other Mastigitae: enlarged and ventrally spiny scape and pedicel; elongate head; narrowly separated antennal insertions; and pronotum lacking posterior collar and antebasal pits. Characteristics. Adults. Body (Figs 153 – 163) large, 3.10 ̄ 7.50 mm in length, yellowish-brown to black, in some species head or head and pronotum dark brown to nearly black and elytra testaceous, brown or reddish, strongly convex, dorsally densely but finely setose, setae short (often extremely so) and recumbent, unmodified except for long and thick bristles on scape and pedicel. Head capsule (Figs 167 – 168, 172, 176 – 177, 221 – 224) divided into large and exposed anterior part and much smaller, subcylindrical ' neck' region retracted into prothorax and demarcated by distinct occipital constriction; ' neck' region much broader than half width of head. Anterior part of head flattened, subequal in width with prothorax, distinctly elongate, broadest near middle. Composite eyes dorsolateral, moderately large, composed of numerous small ommatidia, not projecting or weakly projecting from silhouette of the head, broadly separated from mandibular bases. Vertex and frons divided by a distinct median longitudinal groove (Figs 167, 176, 223; mg); vertex strongly transverse, convex at sides, with posterior margin nearly straight or slightly concave. Tempora much longer than eyes, weakly rounded. Frons between antennal insertions not forming a demarcated ' platform', weakly convex or flattened, anteriorly demarcated by a deep and very short frontoclypeal groove largely obliterated at sides. Clypeus very short and broad, with nearly straight sides slightly convergent anterad or parallel. Antennal insertions (Figs 176, 223; ia) located anterodorsally, relatively narrowly separated. Gular plate (Figs 172, 177, 224; gp) lacking sutures, indistinctly transversely reticulate; posterior tentorial pits (Figs 172, 177, 224; ptp) circular or oval, in front of broad and diffuse transverse impression demarcating ' neck' region ventrally; hypostomal ridges (Figs 177, 224; hr) arcuate, posteriorly reaching middle between anterior submental margin and posterior tentorial pits. Head finely to strongly punctate, densely setose (e. g. Fig. 176). Antennae (Figs 153 – 163, 168 – 170, 178 – 179, 221 – 222) long and slender, shorter than body or subequal in length; scape (Figs 168 – 169, 178, 222; sc) 5 – 10 or even more times as long as broad, thickened, much longer than head, with lateroventral (more ventral than lateral) emargination; pedicel (Figs 168 – 169, 178, 222; pd) also conspicuously enlarged, slightly to much shorter and narrower than scape, typically 5 – 8 times as long as broad, broadening from narrow base to subapical region. Both scape and pedicel with two ventral longitudinal rows of several long and thick bristles with papillate insertions, area between bristles with variously densely distributed porous fields (Figs 170, 179; pf; unknown in fossils); antennomeres III – XI distinctly narrower than pedicel, elongate (often strongly so), each slightly thickened distad, basal stalks not exposed in intact beetles, basal rings absent or indistinct; antennomere XI elongate and indistinctly asymmetrical. Antennomeres covered with variously dense, long setae; surface of antennomeres smooth. Mouthparts. Labrum (Figs 176, 180 – 181, 225 – 226; lbr) strongly transverse, with lateral margins slightly convergent anterad or parallel and weakly rounded, and with anterior margin weakly concave, with a pair of broad and short sublateral teeth broadly separated at middle by a very shallow emargination, and with two partly irregular transverse rows of long and short setae; epipharynx (Fig. 181; eph) smooth, with dense lateral trichia. Mandibles (Figs 173, 176, 180, 182 – 183, 222 – 226) symmetrical, subtriangular and robust, each with one dorsal and a group of 2 – 3 ventral mesal teeth, setose prostheca (Figs 182 – 183; pst) present and long, its setae extending to dorsal and ventral surface of basal half of mandible. Maxilla (Figs 172 – 173, 177, 184 – 185, 224, 229) with large, long cardo (Fig. 185; cd); basistipes (Fig. 185; bst) subtriangular and elongate; mediostipes (Fig. 185; mst) large and sharply demarcated from lacinia (Fig. 185; lac) and galea (Fig. 185; gal), which are both elongate and each with conspicuously dense group of thin distal setae; palpifer (Fig. 185; ppf) broad and elongate; maxillary palp slightly to much longer than head capsule, composed of minute palpomere I (Figs 172 – 173, 177, 185, 229; mxp 1), slender, curved, distinctly but only slightly broadening distad palpomere II (Figs 167 – 168, 172, 177, 187, 222; mxp 2), palpomere III (Figs 16 7 – 168, 172, 177, 187, 222; mxp 3) strongly elongate, strongly and gradually broadened distad, with transverse or (rarely) slightly obtuse distal margin, palpomere IV (Figs 167 – 168, 172, 177, 187, 222; mxp 4) slightly asymmetrical, shorter, about as long as, or longer than III, suboval or indistinctly subtriangular, broadening from base to about distal third and with subtriangular or rounded apex, or broadly boomerang-shaped (with one margin convex and the other one concave), elongate. Surface of palpomere IV, and sometimes also III, with sparsely distributed porous fields (Fig. 188; pf). Palpomeres III and IV slightly (sometimes indistinctly) flattened, II – IV covered with relatively long and dense setae. Labium (Figs 172 – 173, 177, 184, 186, 224, 229) with broad and short submentum (Figs 172, 177, 224; smn) posteriorly not demarcated from gular region, densely setose and lacking an outstanding pair of anterior or subanterior lateral setae; mentum (Figs 173, 184, 229; mn) subtrapezoidal and strongly transverse, with anterior margin very deeply emarginate, so that anterolateral corners form triangular and usually pointed lobes projecting anterad; prementum (Figs 173, 184, 229; pm) long, subtrapezoidal, broadest distally, lacking demarcated ligula, with several pairs of submedian anterior setae, with broadly separated bases of labial palps; lateral hypopharyngeal lobes (Fig. 186; lhl) moderately large, with conspicuously sparse, thick mesal setae; hypopharynx (Fig. 186) with two lateral groups of long trichia; labial palp composed of three palpomeres: palpomere I (Figs 173, 184, 229; lp 1) small, elongate, strongly broadening distad, palpomere II (Figs 173, 184, 229; lp 2) largest, conspicuously enlarged, long and broad, approximately barrelshaped, palpomere III (Figs 173, 184, 229; lp 3) very small and narrow in relation to III, as long as about half length of III and very narrow, pointed. Prothorax (Figs 153 – 162, 174, 189, 230 – 231) elongate (usually strongly so), strongly convex but usually with flattened dorsum, broadest near anterior third. Pronotum with anterior and posterior margins arcuate (anterior margin sometimes nearly straight), sides rounded in anterior half and sinuate or (rarely) nearly straight in posterior half; anterior and posterior corners obtuse-angled or broadly rounded; pronotal base lacking pits and groove. Prosternum (Figs 174, 189, 230) with basisternal part (Figs 174, 189, 230; bstr) usually slightly shorter than or equal in length to coxal part (Fig. 174, 189, 230; cxst). Prosternum laterally completely fused with hypomera. Coxal region anteriorly and laterally without marginal carina; postcoxal hypomeral lobes (Fig. 174, 189, 230; pchl) conspicuously large, rounded and strongly projecting mesad or anteromesad and overlapping with (but not fused to) posterolateral lobes of prosternum, so that procoxal cavities are not open, but entirely delimited posterioly by hypomeral lobes. Prosternal intercoxal process developed as a narrow and weakly elevated carina in intact beetles hidden between procoxae (character not studied in extinct taxa). Ventral surface of prothorax densely setose. Mesoventrite (Figs 171, 175, 191 – 192, 232, 233) subtrapezoidal, broadening posteriorly. Prepecti (Fig. 191; pre) moderately long and together with anteromedian mesoventral area forming a relatively short ' collar', which is weakly impressed just behind its anterior ridge, anterior margin of impression typically with a short subtriangular posteromedian projection. Impressed area can be shallow, short and diffuse, posteriorly gradually becoming shallower to indistinguishably fuse with median precoxal portion of mesoventrite (Figs 175, 232), or forming a deep and broad, well-defined setose impression densely filled with long setae, with its posterior margin abruptly elevated and sharply demarcated from median precoxal portion of mesoventrite (Fig. 191; si). Mesoventral intercoxal process (Figs 171, 175, 191 – 192, 232, 233; msvp) reaching middle of mesocoxal cavities; in extant taxa short and very broad, subtriangular or subtrapezoidal, weakly convex, broadly separating mesocoxae, posteriorly fused with metaventrite or separated from the latter by transverse groove, and then with its posterior margin subtriangular, rounded or nearly straight; in one extinct genus mesoventral process relatively narrow, longer than broad, broadest between anterior margins of mesocoxae, slightly narrowing caudad and with truncate apex. Mesanepisterna (Fig. 233; aest 2) relatively narrow and strongly elongate, demarcated from median part of mesoventrite by a distinct ridge and from mesepimera by complete suture; mesepimera (Fig. 233; epm 2) elongate, indistinctly demarcated from metepimera, not exposed in ventral view. Mesonotum with cordiform, broad mesoscutellum (Fig. 190; scl 2) with pointed apex, in intact specimens not visible between elytral bases, or only its very tip discernible; scutoscutellar suture absent (mesonotum not studied in fossils). Metanotum (Fig. 193) partly reduced, with lightly sclerotized mesoscutum, but only slightly shortened alacristae (Fig. 193; ala); hind wings absent (not studied in fossils). Metaventrite (Figs 171, 175, 191, 194, 232, 233) short, subrectangular and usually strongly transverse, with lateral margins rounded; mesocoxal cavities with all margins non-carinate; posterior margin of metaventrite deeply bisinuate laterally (in front of each metacoxa) and with a broad metaventral intercoxal process (Figs 191, 232; mtvp) with shallowly concave or straight posterior margin; anteriorly metaventrite forming a short (in extant forms) or elongate (in extinct species) anterior metaventral process (Figs 175, 191, 232; amvp) similar in shape to mesoventral intercoxal process and meeting the latter at middle of mesocoxal cavities; metaventral foveae absent. External admetacoxal part of posterior metaventral margin lacking adcoxal carinae, but with adcoxal expansions (Figs 191, 232; ade). Metanepisterna (Fig. 233; aest 3) relatively narrow, partly visible in ventral view, narrowing posterad; metepimera (Fig. 233; epm 3) 2 – 3 times as broad as metanepisterna, with inner and outer components not demarcated, posteriorly extending far behind metacoxae. Metendosternite (metafurca) with stem much broader than long and with broadly separated, divergent lateral furcal arms (Fig. 194; lmfa), lacking median longitudinal projection (metafurca not studied in fossils). Legs (Figs 153 – 163, 171, 174, 191, 221, 232) very long and slender (often extremely so). Pro- and mesocoxa short subconical, metacoxa with nearly hemispherical basal part and subconical distal part. Mesocoxa lacking coxal bristles. All trochanters short and subtriangular (sometimes with various male secondary sexual characters); femora weakly clavate; tibiae slender; tarsi long and slender, nearly subcylindrical, tarsomeres I – V reducing in length, tarsomere V strongly elongate, with curved and slender claws lacking elongate costae; empodial region was not studied. Elytra (Figs 153 – 163, 2 – 235) oval, strongly convex, lacking humeral calli and basal impressions, with rounded or pointed apices; elytral disc in extant forms with superficial, indistinct and incomplete longitudinal rows of fine punctures, often barely discernible, obscured by chaotic secondary punctures or coarse microscultpure; in one extinct genus elytra with complete and deeply impressed longitudinal striae. Elytra densely setose, setae short and nearly recumbent. Abdomen (Figs 171, 195 – 196) with sternite III firmly fused with metaventrite (so that during disarticulation it is almost impossible to separate intact abdomen), much longer than sternite IV, but shorter than IV – VI together; sternite VIII in male distinctly, often deeply emarginate (Fig. 195). Aedeagus (illustrated e. g. in Bordoni & Castellini (1973), Jałoszyński (2012 c; d; e), Jałoszyński et al. (2015), Leleup (1968), Lhoste (1936, 1937 )) elongate, sometimes extremely so, with asymmetrical median lobe and asymmetrical parameres, one shorter than the other, sometimes vestigial or completely obliterated; flagellum very long and forming several coils (Fig. 199; fl). Ejaculatory duct with elongate and narrow sperm pump lacking funnel-like structures. Aedeagus with elongate copulatory piece (Fig. 199; cp) with membranous endophallus permanently everted, and only inflated during copulation, with distal end of flagellum permanently fixed, so that flagellum is not extricable. Aedeagus in repose positioned asymmetrically inside abdomen, with basal orifice lateral or dorsolateral. Spermatheca (Figs 197, 198; sp) subglobose, with relatively large accessory gland. Characteristics. Larvae. Larvae (Figs 200 – 201, 236) are known for two extant genera. Campodeiform, subparallel or with strongly narrowing abdomen, slightly flattened; head, tergal and sternal plates brown to nearly black, remaining areas whitish or yellowish, or nearly entire larva orange. Body sparsely covered with long unmodified setae and dense asperities forming patterns among smooth areas of tergal plates, additionally with sparse short leaf-like setae with elongate ribs; frontal impression with setae covered with irregular convexities. Head prognathous and slightly tilted ventrad, lacking ' neck', with one stemma at each side; epicranial stem and frontal sutures distinct but short, together with antennal insertions shifted to posterior half of head capsule; nasale with a row of several short setae with papillate insertions. Head with large glandular impression at the junction of epicranial stem and frontal sutures, impression surrounded by or filled with short modified setae (first instar larvae without impression). Antenna 2 – 4 times longer than head, very slender, antennomeres subcylindrical and not broadened distad, antennomeres I and II very long and similarly broad, antennomere II subdivided into three sections, antennomere III vestigial, developed as a barely discernible papilla adjacent to base of strongly elongate, slightly asymmetrical, subconical and pointed accessory appendage. Mandibles falciform, moderately slender, pointed, each with one submedian mesal tooth; stipital projection of maxilla divided into two very short and broad, densely setose lobes; maxillary palp longer than head, with palpomere I short and II and III strongly elongate; labial palp with palpomere I longer than II, both subconical. Thoracic tergites with ecdysial line visible as a smooth elongate median stripe among lateral fields of dense asperities. Abdomen with ten segments, all except X (or IX and X) transverse; segment X elongate; urogomphi absent. Sternal plates on thorax and abdomen reduced to small, paired (2 – 4) and setose sclerites. Legs conspicuously long and slender, with particularly densely setose tibiotarsi. Spiracles annular, lateral, nine pairs: one on mesothorax and eight pairs on abdominal segments I – VIII.	en	Paweł Jałoszyński (2018): World genera of Mastigitae: review of morphological structures and new ecological data (Coleoptera: Staphylinidae: Scydmaeninae). Zootaxa 4453 (1): 1-119, DOI: 10.11646/zootaxa.4453.1.1
2161879CFF808A41FF7A335E6062DADE.taxon	description	Composition and distribution. Mastigini include three extant and one extinct genera, comprising 56 extant species and subspecies distributed in southern Europe and south Africa (Figs 164 – 166), and three extinct species known from Cenomanian Myanmar amber.	en	Paweł Jałoszyński (2018): World genera of Mastigitae: review of morphological structures and new ecological data (Coleoptera: Staphylinidae: Scydmaeninae). Zootaxa 4453 (1): 1-119, DOI: 10.11646/zootaxa.4453.1.1
2161879CFF808A41FF7A335E6062DADE.taxon	discussion	Remarks. Mastigini and † Baltostigini form a monophyletic and distinct group characterized, among some less conspicuous structures, by strongly enlarged and spinose scape and pedicel. All known Mastigini, including the extant genus Clidicostigus Jałoszyński et al., 2017, have asymmetrical aedeagi twisted inside abdomen in repose and lack wings and humeral calli. † Baltostigini differ in symmetrical aedeagus, which in repose is symmetrically positioned inside the abdomen, and are winged, with prominent humeral calli. Moreover, Mastigini comprise large and strikingly elongate beetles, whereas adults of † Baltostigini are small and stout. The shape of maxillary palpomere IV also differentiates these tribes; it is elongate in Mastigini, and broader than long, axe-shaped in † Baltostigini.	en	Paweł Jałoszyński (2018): World genera of Mastigitae: review of morphological structures and new ecological data (Coleoptera: Staphylinidae: Scydmaeninae). Zootaxa 4453 (1): 1-119, DOI: 10.11646/zootaxa.4453.1.1
2161879CFF948A46FF7A302D623FD8BB.taxon	diagnosis	Diagnosis. Unclearly diagnosed genus; within Mastigini identifiable on the basis of a combination of subtriangular mesoventral intercoxal process with narrow and pointed or narrowly rounded tip (Fig. 171; msvp) and the aedeagus with the short paramere distinct, always delimited from the long paramere by at least subtriangular or rounded emargination (but see Remarks). Characteristics. Adults. Body (Figs 153 – 154) large, 3.50 ̄ 6.80 mm in length, with dark brown or nearly black head, similarly dark or yellowish-brown pronotum, and elytra brown, often with reddish or yellowish hue, strongly convex, dorsally densely but finely setose, setae unmodified except for long and thick bristles on scape and pedicel. Head capsule (Figs 167 – 168, 172) divided into large and exposed anterior part and much smaller, subcylindrical ' neck' region retracted into prothorax and demarcated by distinct occipital constriction; ' neck' region much broader than half width of head. Anterior part of head flattened, subequal in width with prothorax, distinctly elongate, broadest near middle. Composite eyes dorsolateral, moderately large, composed of numerous small ommatidia, weakly projecting from the silhouette of the head, broadly separated from mandibular bases. Vertex and frons divided by a distinct median longitudinal groove (Fig. 167; mg); vertex strongly transverse, convex at sides, with posterior margin slightly concave. Tempora much longer than eyes, weakly rounded. Frons between antennal insertions not forming a demarcated ' platform', weakly convex or flattened, anteriorly demarcated by a deep and very short frontoclypeal groove largely obliterated at sides. Clypeus very short and broad, with nearly straight sides, slightly convergent anterad or parallel. Antennal insertions located anterodorsally, relatively narrowly separated. Gular plate (Fig. 172; gp) lacking sutures, indistinctly transversely reticulate; posterior tentorial pits (Fig. 172; ptp) oval, in front of broad and diffuse transverse impression demarcating ' neck' region ventrally; hypostomal ridges (Figs 177, 224; hr) arcuate, posteriorly reaching middle between anterior submental margin and posterior tentorial pits. Head finely to strongly punctate, densely setose. Antennae (Figs 153 – 154, 168 – 170) long and slender, shorter than body; scape (Figs 168 – 169; sc) 5 – 6 times as long as broad, thickened, broadest in front of middle, much longer than head, with lateroventral (more ventral than lateral) emargination; pedicel (Figs 168 – 169; pd) enlarged, slightly to much shorter and narrower than scape, 5 – 8 times as long as broad, broadening from narrow base to subapical region. Both scape and pedicel with two ventral longitudinal rows of several long and thick bristles with papillate insertions, area between bristles with variously densely distributed porous fields (Fig. 170; pf); antennomeres III – XI distinctly narrower than pedicel, elongate, each slightly thickened distad, basal stalks not exposed in intact beetles, basal rings absent or indistinct; antennomere XI elongate and indistinctly asymmetrical. Antennomeres covered with variously dense, long setae; surface of antennomeres smooth. Mouthparts. Labrum (Fig. 167) strongly transverse, with lateral margins slightly convergent anterad and weakly rounded, and with anterior margin weakly concave, with a pair of broad and short sublateral teeth broadly separated at middle by a very shallow emargination, and with two partly irregular transverse rows of long and short setae. Mandibles symmetrical, subtriangular and robust, each with one mesal dorsal tooth and a group of 3 mesal ventral teeth, setose prostheca long. Maxilla (Figs 172 – 173) with large cardo (Fig. 173; cd); basistipes (Fig. 173; bst) subtriangular and elongate; mediostipes (Fig. 173; mst) large and sharply demarcated from lacinia and galea (Fig. 173; gal), which are both elongate and each with dense group of thin distal setae; palpifer (Fig. 173; ppf) broad and elongate; maxillary palp slightly to much longer than head capsule, composed of minute palpomere I (Fig. 172 – 173; mxp 1), slender, curved, distinctly but only slightly broadening distad palpomere II (Figs 167 – 168, 172; mxp 2), palpomere III (Figs 167 – 168, 172; mxp 3) strongly elongate, strongly and gradually broadened distad, with transverse distal margin, palpomere IV (Figs 167 – 168, 172) shorter than III, approximately subtriangular and broadening distad. Labium (Figs 172 – 173) with broad and short submentum (Fig. 172; smn) posteriorly not demarcated from gular region, densely setose and lacking an outstanding pair of anterior or subanterior lateral setae; mentum (Fig. 173; mn) subtrapezoidal and strongly transverse, with anterior margin very deeply emarginate, so that anterolateral corners form triangular and usually pointed lobes projecting anterad; prementum (Figs. 173; pm) long, subtrapezoidal, broadest distally, lacking demarcated ligula, with several pairs of submedian anterior setae, with broadly separated bases of labial palps; lateral hypopharyngeal lobes moderately large; labial palp composed of three palpomeres: palpomere I (Fig. 173; lp 1) small, elongate, strongly broadening distad, palpomere II (Fig. 173; lp 2) largest, conspicuously enlarged, long and broad, approximately barrel-shaped, palpomere III (Fig. 173; lp 3) very small and narrow in relation to III, as long as about half length of III and very narrow, pointed. Prothorax (Figs 153 – 54, 174) elongate, strongly convex but with flattened dorsum, broadest near anterior third. Pronotum with anterior and posterior margins arcuate, sides rounded in anterior half and sinuate in posterior half; anterior and posterior corners obtuse-angled; pronotal base lacking pits and groove. Prosternum (Fig. 174) with basisternal part (Fig. 174; bstr) indistinctly shorter than coxal part. Prosternum laterally completely fused with hypomera. Coxal region anteriorly and laterally without marginal carina; postcoxal hypomeral lobes (Fig. 174; pchl) conspicuously large, rounded and strongly projecting anteromesad and overlapping with (but not fused to) posterolateral lobes of prosternum, so that procoxal cavities are not open, but entirely delimited posteriorly by hypomeral lobes. Prosternal intercoxal process developed as a narrow and weakly elevated carina in intact beetles hidden between procoxae. Ventral surface of prothorax densely setose. Mesoventrite (Figs 171, 175) subtrapezoidal, broadening posteriorly. Prepecti moderately long and together with anteromedian mesoventral area forming a relatively short ' collar', which is weakly impressed just behind its anterior ridge, anterior margin of impression with a short subtriangular posteromedian projection. Impressed area shallow, short and diffuse, with median sternal area behind it becoming gradually convex caudad. Mesoventral intercoxal process (Figs 171, 175; msvp) reaching middle of mesocoxal cavities; short and very broad, subtriangular, weakly convex, broadly separating mesocoxae, posteriorly separated from anterior metaventral process, with its posterior margin subtriangular. Mesanepisterna relatively narrow and strongly elongate, demarcated from median part of mesoventrite by a distinct ridge and from mesepimera by complete suture; mesepimera elongate, indistinctly demarcated from metepimera, not exposed in ventral view. Mesonotum with cordiform, broad mesoscutellum pointed at apex, in intact specimens not visible between elytral bases; scutoscutellar suture absent. Metanotum partly reduced, with lightly sclerotized mesoscutum, but only slightly shortened alacristae; hind wings absent. Metaventrite (Figs 171, 175) short, subrectangular and strongly transverse, with lateral margins rounded; mesocoxal cavities with all margins non-carinate; posterior margin of metaventrite deeply bisinuate laterally (in front of each metacoxa) and with a broad metaventral intercoxal process with shallowly concave posterior margin; anteriorly metaventrite forming a short anterior metaventral process (Figs 171, 175; amvp) similar in shape to mesoventral intercoxal process and meeting the latter at middle of mesocoxal cavities; metaventral foveae absent. External admetacoxal part of posterior metaventral margin lacking adcoxal carinae, but with adcoxal expansions. Metanepisterna relatively narrow, partly visible in ventral view, narrowing posteriorly; metepimera 2 – 3 times as broad as metanepisterna, with not demarcated inner and outer components, posteriorly extending far behind metacoxae. Metendosternite (metafurca) with stem much broader than long and broadly separated, divergent lateral furcal arms, lacking median longitudinal projection. Legs (Figs 153 – 154, 171, 174 – 175) long and slender. Pro- and mesocoxa short subconical, metacoxa with nearly hemispherical basal part and subconical distal part. Mesocoxa lacking coxal bristles. All trochanters short and subtriangular; femora weakly clavate; tibiae slender; tarsi long and slender, nearly subcylindrical, tarsomeres I – V reducing in length, tarsomere V strongly elongate, with curved and slender claws lacking elongate costae; empodial region was not studied. Elytra (Figs 153 – 154) oval, strongly convex, lacking humeral calli and basal impressions, with rounded apices; elytral disc with barely discernible incomplete and superficial longitudinal rows of fine punctures, in some species obscured by very fine transverse striae; microsculpture fine. Elytra densely setose, setae short and nearly recumbent. Abdomen (Fig. 171) with sternite III firmly fused with metaventrite, much longer than sternite IV, but shorter than IV – VI together; sternite VIII in male distinctly, often deeply emarginate. Aedeagus (illustrated in Jałoszyński (2012 c), Jałoszyński et al. (2018), Leleup (1968 )) moderately elongate, with asymmetrical median lobe and asymmetrical parameres, one shorter than the other; flagellum very long and forming several coils. Ejaculatory duct with elongate and narrow sperm pump lacking funnel-like structures. Aedeagus with elongate copulatory piece with membranous endophallus permanently everted, and only inflated during copulation, with distal end of flagellum permanently fixed, so that flagellum is not extricable. Aedeagus in repose positioned asymmetrically inside abdomen, with basal orifice lateral or dorsolateral. Spermatheca subglobose, with relatively large accessory gland. Sexual dimorphism distinct, females distinctly larger and with relatively broader elytra than males; males with emarginate, females with rounded sternite VIII. Larva. Unknown.	en	Paweł Jałoszyński (2018): World genera of Mastigitae: review of morphological structures and new ecological data (Coleoptera: Staphylinidae: Scydmaeninae). Zootaxa 4453 (1): 1-119, DOI: 10.11646/zootaxa.4453.1.1
2161879CFF948A46FF7A302D623FD8BB.taxon	description	Composition and distribution. Mastigus comprises five species distributed in west-southern part of the Republic of South Africa (western West Cape Province) (Fig. 164). Natural history. Almost nothing is known about natural history of these beetles. They are rarely collected, typically by using pitfall traps, often near the coastline, on sandy places with sparse vegetation. Judging from a similar body form and structures, Mastigus species may resemble Palaeostigus and Stenomastigus in diurnal, open life style. However, all species were described on the basis of a few specimens only, and Leleup (1968), who in his large monograph of South African Mastigini included hundreds of specimens of Palaeostigus and Stenomastigus, also found only a small number of adult Mastigus during his field studies. It seems that species of Mastigus do not live in such large populations as Palaeostigus and Stenomastigus.	en	Paweł Jałoszyński (2018): World genera of Mastigitae: review of morphological structures and new ecological data (Coleoptera: Staphylinidae: Scydmaeninae). Zootaxa 4453 (1): 1-119, DOI: 10.11646/zootaxa.4453.1.1
2161879CFF948A46FF7A302D623FD8BB.taxon	discussion	Remarks. Mastigus is currently the most problematic genus of Mastigini, comprising species that show the greatest diversity in structures of the aedeagus. Species of Mastigus have a general appearance more similar to that of Palaeostigus than Stenomastigus; the latter genus includes species very slender and with extremely long legs and antennae, with coarse microgranulation of the elytra, whereas Mastigus and Palaeostigus share a stouter body form, less elongated appendages and either shiny or matt but always not coarse elytral surface. Consequently, Mastigus can be distinguished from Stenomastigus solely by its general body shape. However, fine morphological structures of these two genera are nearly identical and a major difference can be found only in the aedeagus. In Mastigus the aedeagus is distinctly stouter and shorter than that in Stenomastigus, the short paramere is always developed and demarcated from the long paramere by a subtriangular or rounded emargination, so that the apex of short paramere is distant from the lateral margin of the long paramere. Also the capsular, basal part of the aedeagus that contains flagellar coils is larger in relation to the parameral part in Mastigus than in Stenomastigus. In the latter genus the aedeagus is typically very narrow and long, with one paramere strongly elongate and the other one usually absent. However, in few cases the short paramere is present, but its apex overlaps with the lateral margin of the long paramere or it is very indistinctly demarcated. Aedeagi of Mastigus species are diverse in shapes and structures and this genus must be revised to clarify its separate placement. Palaeostigus, although very similar to Mastigus, differs clearly in the shape of the mesoventral intercoxal process, which is not subtriangular but subtrapezoidal and its posterior margin is nearly straight and widely separates mesocoxae; and in a deep mesoventral setose impression, which is absent in Mastigus. All species were treated by Leleup (1968) and Jałoszyński (2012 c); two species originally placed in Mastigus were treated as incertae sedis within Mastigini by Leleup (1968).	en	Paweł Jałoszyński (2018): World genera of Mastigitae: review of morphological structures and new ecological data (Coleoptera: Staphylinidae: Scydmaeninae). Zootaxa 4453 (1): 1-119, DOI: 10.11646/zootaxa.4453.1.1
2161879CFF938A4DFF7A35C46552DC3E.taxon	description	Mastigus of Leleup, 1968: 10 (misidentified, not Latreille, 1802). Type species: Mastigus palpalis Latreille, 1804 (des. Leleup, 1968: 7; invalid designation, species not originally included).	en	Paweł Jałoszyński (2018): World genera of Mastigitae: review of morphological structures and new ecological data (Coleoptera: Staphylinidae: Scydmaeninae). Zootaxa 4453 (1): 1-119, DOI: 10.11646/zootaxa.4453.1.1
2161879CFF938A4DFF7A35C46552DC3E.taxon	diagnosis	Diagnosis. Adults of Palaeostigus differ from remaining Mastigini in subtrapezoidal mesoventral intercoxal process with broad and nearly straight posterior tip (Figs 191 – 192); and distinct setose impression of mesoventrite (Fig. 191; si). Mature larva: antennae about twice as long as head capsule (Figs 200 – 201). Characteristics. Adults. Body (Figs 155 – 156, 159 – 160) large, 3.40 ̄ 7.50 mm in length, uniformly black or nearly black or with dark brown or nearly black head and pronotum and reddish-brown or testaceous elytra; strongly convex, dorsally densely but finely setose, setae unmodified except for long and thick bristles on scape and pedicel. Head capsule (Figs 176 – 177) divided into large and exposed anterior part and much smaller, subcylindrical ' neck' region retracted into prothorax and demarcated by distinct occipital constriction; ' neck' region much broader than half width of head. Anterior part of head flattened, subequal in width with prothorax, distinctly elongate, broadest near middle. Composite eyes dorsolateral, moderately large, composed of numerous small ommatidia, not projecting or weakly projecting from the silhouette of the head, broadly separated from mandibular bases. Vertex and frons divided by a distinct median longitudinal groove (Fig. 176; mg); vertex strongly transverse, convex at sides, with posterior margin nearly straight or slightly concave. Tempora much longer than eyes, weakly rounded. Frons between antennal insertions not forming a demarcated ' platform', weakly convex or flattened, anteriorly demarcated by a deep and very short frontoclypeal groove (Fig. 176; fcg) largely obliterated at sides. Clypeus very short and broad, broadly subtrapezoidal, with nearly straight sides, which are slightly convergent anterad or parallel. Antennal insertions (Fig. 176; ia) located anterodorsally, relatively narrowly separated. Gular plate (Fig. 177; gp) lacking sutures, indistinctly transversely reticulate; posterior tentorial pits (Fig. 177; ptp) oval, in front of broad and diffuse transverse impression demarcating ' neck' region ventrally; hypostomal ridges (Fig. 177; hr) arcuate, posteriorly reaching middle between anterior submental margin and posterior tentorial pits. Head finely to strongly punctate, densely setose (Fig. 176). Antennae (Figs 155 – 156, 159 – 160, 178) long and slender, shorter than body; scape (Fig. 178; sc) 5 – 7 times as long as broad, thickened, much longer than head, with lateroventral (more ventral than lateral) emargination; pedicel (Fig. 178; pd) also conspicuously enlarged, much shorter and narrower than scape, typically 4 – 6 times as long as broad, broadening from narrow base to subapical region. Both scape and pedicel with two ventral longitudinal rows of several long and thick bristles with papillate insertions, area between bristles with variously densely distributed porous fields (Fig. 179; pf); antennomeres III – XI distinctly narrower than pedicel, elongate, each slightly thickened distad, basal stalks not exposed in intact beetles, basal rings absent or indistinct; antennomere XI elongate and indistinctly asymmetrical. Antennomeres covered with variously dense, long setae; surface of antennomeres smooth. Mouthparts. Labrum (Fig. 180; lbr) strongly transverse, with lateral margins slightly convergent anterad and weakly rounded, and with anterior margin weakly concave, with a pair of broad and short sublateral teeth broadly separated at middle by a very shallow emargination, and with two partly irregular transverse rows of long and short setae; epipharynx (Fig. 181; eph) smooth, with dense lateral trichia. Mandibles (Figs 182 – 183) symmetrical, subtriangular and robust, each with one dorsal mesal tooth and a group of 2 – 3 ventral mesal teeth, setose prostheca (Figs 182 – 18; pst 3) present and long, its setae extending to dorsal and ventral surface of basal half of mandible. Maxilla (Figs 184 – 185) with large, long cardo (Fig. 185; cd); basistipes (Fig. 185; bst) subtriangular and elongate; mediostipes (Fig. 185; mst) large and sharply demarcated from lacinia (Fig. 185; lac) and galea (Fig. 185; gal), which are both elongate and each with conspicuously dense group of thin distal setae; palpifer (Fig. 185; ppf) broad and elongate; maxillary palp slightly to much longer than head capsule, composed of minute palpomere I (Fig. 185; mxp 1), slender, curved, distinctly but only slightly broadening distad palpomere II (Figs 185, 187; mxp 2), palpomere III (Fig. 187; mxp 3) strongly elongate, strongly and gradually broadened distad, with transverse or indistinctly obtuse distal margin, palpomere IV (Fig. 187; mxp 4) about as long as III, broadening to distal third and with rounded or slightly subtriangular apex. Surface of palpomere IV, and sometimes also III, with sparsely distributed porous fields (Fig. 188; pf). Palpomeres III and IV slightly (sometimes indistinctly) flattened, II – IV covered with relatively long and dense setae. Labium (Figs 177, 184, 186) with broad and short submentum (Fig. 177; smn) posteriorly not demarcated from gular region, densely setose and lacking an outstanding pair of anterior or subanterior lateral setae; mentum (Fig. 184; mn) subtrapezoidal and strongly transverse, with anterior margin very deeply emarginate, so that anterolateral corners form triangular and usually pointed lobes projecting anterad; prementum (Fig. 184; pm) long, subtrapezoidal, broadest distally, lacking demarcated ligula, with several pairs of submedian anterior setae, with broadly separated bases of labial palps; lateral hypopharyngeal lobes (Figs 184, 186; lhl) moderately large, with conspicuously sparse, thick mesal setae; hypopharynx (Fig. 186) with two lateral groups of long trichia; labial palp composed of three palpomeres: palpomere I (Fig. 184; lp 1) small, elongate, strongly broadening distad, palpomere II (Figs 184, 186; lp 2) largest, conspicuously enlarged, long and broad, approximately barrel-shaped, palpomere III (Figs 184, 186; lp 3) very small and narrow in relation to III, as long as about half length of III and very narrow, pointed. Prothorax (Figs 155 – 156, 159 – 160, 189) elongate, strongly convex but usually with flattened dorsum, broadest near anterior third. Pronotum with anterior and posterior margins arcuate (anterior margin sometimes nearly straight), sides rounded in anterior half and sinuate or (rarely) nearly straight in posterior half; anterior and posterior corners obtuse-angled or broadly rounded; pronotal base lacking pits and groove. Prosternum (Fig. 189) with basisternal part (Fig. 189; bstr) subequal in length to coxal part (Fig. 189; cxst). Prosternum laterally completely fused with hypomera. Coxal region anteriorly and laterally without marginal carina; postcoxal hypomeral lobes (Fig. 189; pchl) conspicuously large, rounded and strongly projecting anteromesad and overlapping with (but not fused to) posterolateral lobes of prosternum, so that procoxal cavities are not open, but entirely delimited posterioly by hypomeral lobes. Prosternal intercoxal process developed as a narrow and weakly elevated carina in intact beetles hidden between procoxae. Ventral surface of prothorax densely setose. Mesoventrite (Figs 191 – 192) subtrapezoidal, broadening posteriorly. Prepecti (Fig. 191; pre) moderately long and together with anteromedian mesoventral area forming a relatively short ' collar', which is strongly impressed just behind its anterior ridge, anterior margin of impression with a short median subtriangular projection. Impressed area deep and broad, forming a well-defined setose impression (Figs 191 – 192; si) densely filled with long setae, with its posterior margin sharply demarcated from median precoxal portion of mesoventrite (but posterior margin usually obscured by dense setae). Mesoventral intercoxal process (Figs 191 – 192; msvp) reaching middle of mesocoxal cavities; short and very broad, subtrapezoidal, weakly convex, broadly separating mesocoxae, posteriorly separated from metaventrite by a transverse groove (often obscured by setae but discernible in transparent mounts (Fig. 192 )), posterior margin of mesoventral process nearly straight. Mesanepisterna relatively narrow and strongly elongate, demarcated from median part of mesoventrite by a distinct ridge and from mesepimera by complete suture; mesepimera elongate, indistinctly demarcated from metepimera, not exposed in ventral view. Mesonotum with cordiform, broad mesoscutellum (Fig. 190; scl 2) with pointed apex, in intact specimens not visible between elytral bases, or only its very tip discernible; scutoscutellar suture absent. Metanotum (Fig. 193) partly reduced, with lightly sclerotized mesoscutum, but only slightly shortened alacristae (Fig. 193; ala); hind wings absent. Metaventrite (Figs 191, 194) short, subrectangular and transverse, with lateral margins rounded; mesocoxal cavities with all margins non-carinate; posterior margin of metaventrite deeply bisinuate laterally (in front of each metacoxa) and with a broad metaventral intercoxal process (Fig. 191; mtvp) with shallowly concave or straight posterior margin; anteriorly metaventrite forming a short anterior metaventral process (Figs 191 – 192; amvp) similar in shape to mesoventral intercoxal process and meeting the latter at middle of mesocoxal cavities; metaventral foveae absent. External admetacoxal part of posterior metaventral margin lacking adcoxal carinae, but with adcoxal expansions (Fig. 191; ade). Metanepisterna relatively narrow, partly visible in ventral view, narrowing posteriorly; metepimera 2 – 3 times as broad as metanepisterna, with not demarcated inner and outer components, posteriorly extending far behind metacoxae. Metendosternite (metafurca) with stem much broader than long and with broadly separated, divergent lateral furcal arms (Fig. 194; lmfa), lacking median longitudinal projection. Legs (Figs 155 – 156, 159 – 160, 191) very long and slender. Pro- and mesocoxa short subconical, metacoxa with nearly hemispherical basal part and subconical distal part. Mesocoxa lacking coxal bristles. All trochanters short and subtriangular; femora weakly clavate; tibiae slender; tarsi long and slender, nearly subcylindrical, tarsomeres I – V reducing in length, tarsomere V strongly elongate, with curved and slender claws lacking elongate costae; empodial region was not studied. Elytra (Figs 155 – 156, 159 – 160) oval, strongly convex, lacking humeral calli and basal impressions, with rounded apices; elytral disc with superficial, indistinct and incomplete longitudinal rows of fine punctures, often barely discernible; microsculpture fine. Elytra densely setose, setae short and nearly recumbent. Abdomen (Figs 191, 195 – 196) with sternite III firmly fused with metaventrite; much longer than sternite IV, but shorter than IV – VI together; sternite VIII in male distinctly, often deeply emarginate (Fig. 195). Aedeagus (illustrated in Bordoni & Castellini (1973), Jałoszyński et al. (2015, 2018), Lhoste (1936 )) elongate, but not conspicuously long, with asymmetrical median lobe and asymmetrical parameres, one shorter than the other, both always well-developed; flagellum very long and forming several coils (Fig. 199; fl). Ejaculatory duct with elongate and narrow sperm pump lacking funnel-like structures. Aedeagus with elongate copulatory piece (Fig. 199; cp) with membranous endophallus permanently everted, and only inflated during copulation, with distal end of flagellum permanently fixed, so that flagellum is not extricable. Aedeagus in repose positioned asymmetrically inside abdomen, with basal orifice lateral or dorsolateral. Spermatheca (Figs 197, 198; sp) subglobose, with relatively large accessory gland. Sexual dimorphism distinct, females distinctly larger and with relatively broader elytra than males; males with emarginate, females with rounded abdominal sternite VIII; males often with slightly curved apices of protibiae. Characteristics. Larvae. De Marzo (1983, 1984) described eggs, all larval instars and the pupa of the European Palaeostigus pilifer (Kraatz, 1879); Newton (1991) illustrated some structures for the Turkish Palaeostigus ruficornis schimitscheki (Machulka, 1944); and Grebennikov & Newton (2009) illustrated some larval structures of the South African Palaeostigus bifoveolatus (Boheman, 1851). Larvae (Figs 200 – 201) campodeiform, subparallel or with strongly narrowing abdomen, slightly flattened; head, tergal and sternal plates in living mature larvae dark brown to nearly black, remaining areas creamy white; young instars orange. Body sparsely covered with long unmodified setae and dense asperities forming patterns among smooth areas of tergal plates, additionally with sparse short leaf-like setae with elongate ribs; frontal impression with setae covered with irregular convexities. Head prognathous and slightly tilted ventrad, lacking ' neck', with one stemma at each side; epicranial stem and frontal sutures distinct but short, together with antennal insertions shifted to posterior half of head capsule; nasale with a row of several short setae with papillate insertions. Head with large glandular impression at the junction of epicranial stem and frontal sutures, impression surrounded by or filled with short modified setae (but first instars without impression). Antenna about twice as long as head or slightly shorter, antennomeres subcylindrical and not broadened distad, antennomeres I and II very long and similarly broad, antennomere II subdivided into three sections, antennomere III vestigial, developed as a barely discernible papilla adjacent to base of strongly elongate, slightly asymmetrical, subconical and pointed accessory appendage. Mandibles falciform, moderately slender, pointed, each with one submedian mesal tooth; stipital projection of maxilla divided into two very short and broad, densely setose lobes; maxillary palp longer than head, with palpomere I short and II and III strongly elongate; labial palp with palpomere I longer than II, both subconical. Thoracic tergites with ecdysial line visible as a smooth elongate median stripe among lateral fields of dense asperities. Abdomen with ten segments, all except X (or IX and X) transverse; segment X elongate; urogomphi absent. Sternal plates on thorax and abdomen reduced to small, paired (2 – 4) and setose sclerites. Legs conspicuously long and slender, with particularly densely setose tibiotarsi. Spiracles annular, lateral, nine pairs: one on mesothorax and eight pairs on abdominal segments I – VIII.	en	Paweł Jałoszyński (2018): World genera of Mastigitae: review of morphological structures and new ecological data (Coleoptera: Staphylinidae: Scydmaeninae). Zootaxa 4453 (1): 1-119, DOI: 10.11646/zootaxa.4453.1.1
2161879CFF938A4DFF7A35C46552DC3E.taxon	description	Composition and distribution. Palaeostigus comprises 11 species and 14 subspecies distributed in Asia (Turkey), Europe (Albania, Austria, Bosnia & Herzegovina, Croatia, France, Greece, Italy, Portugal, Slovenia, Spain), and in South Africa (Fig. 166). Natural history. Most species (both Mediterranean and South African) live in large and relatively dense populations restricted to small areas. Adults are diurnal, they can be found near streams or rivers in warm places that provide some rocks, logs and vegetation for the adults to climb and moist leaf litter microhabitats where the larvae develop (Figs 202 – 207). In places inhabited by these beetles it is often possible to observe hundreds of individuals; adults walk on the ground, on mosses, twigs and logs, climb rocks and bushes. Frequently such places are relatively small and can be restricted to a 20 - m stripe of ground along a stream, outside of which beetles cannot be found (Jałoszyński, pers. obs.). In Europe, females of Palaeostigus lay eggs in autumn and the larvae develop during winter (De Marzo 1983); the last-instar larvae can still be found in the beginning of April (Jałoszyński, pers. obs.). Later during summer, at least in Europe, places where beetles live frequently dry and beetles disappear; it is unclear whether they die, enter a diapause, or migrate to unknown shelters. Pérez Fernández et al. (2013) reported a common occurrence of the Iberian P. palpalis (Letreille, 1804) at mouths of caves in the Sistema de la Murcielaguina and even used this species as an indicator to locate new entrances to this cave system. It is possible that during the hot and dry season beetles enter caves or underground microhabitats. Adults of Palaeostigus kept for a long time in captivity and offered a broad spectrum of potential prey, readily attacked and devoured small caterpillars (tentatively identified as Lecithoceridae) that were obtained by sifting leaf litter from which also numerous beetles were collected (Figs 208 – 213). During attack, beetles were not using the antennal spines to catch their prey. Once a caterpillar was detected by palpating with tips of antennae and then with maxillary palps, the antennae were spread so that during the attack and during nearly entire feeding the spines on scape and pedicel did not touch the prey. Beetles used their mandibles to catch the caterpillar in any place on its body, and they ate captured prey while it was still alive. When the attack started from the tip of the abdomen, as in Fig. 208, prey's movements ceased only when about half or even more of its body was consumed. Beetles continuously moved their mandibles to damage prey's cuticle and increase the surface for digestive juices. The feeding process documented in Figs 208 – 213 took 1 hour and seven minutes; the abandoned remains were hardly recognizable as a lepidopterous larva. When given large, dead caterpillars of noctuids or geometrids with their cuticle pierced or cut, so that some body fluids spilled outside, beetles quickly detected this source of food and gathered around to feed on the liquids (Figs 214 – 216). Live and undamaged large caterpillars were not attacked. During about two months of laboratory observations, adults of P. palpalis did not catch any living springtails; beetles showed some interest toward any moving objects, including Collembola, but were not able to get close enough to any springtails for a successful attack. The touching with antennae caused springtails to jump away and avoid being captured. Only when dead or half-dead (pierced with a needle; still alive, but not able to jump or walk) tomocerid springtails were found by beetles, they were readily eaten (Fig. 217). Beetles also fed on living enchytraeids found between leaves or on the surface of soil in their containers (Fig. 218), attacking them and eating in a similar way as small caterpillars. Beetles also readily scavenged on a variety of dead arthropods, including large freshly killed flies (Fig. 219); they also used as food any raw or processed meat provided, including pieces of ham or mortadella (Fig. 220). They gathered around large pieces of solid food and used their mandibles to chew on it. It was previously hypothesized that long spines on the scape and pedicel of Mastigini may be used as a ' springtail-trap', a catching device that may work in a similar way as spiny antennae of springtail-eating carabid Loricera Latreille (Jałoszyński 2016 a; Yin et al. 2017 a). These hypotheses were based solely on the antennal structure and required verification. Field observations and laboratory experiments (not only with Palaeostigus, but also with Stenomastigus, which has a similar antennal structure) carried out during the present project in South Africa and Spain, did not provide any evidence to support this view. In a contrary, not even once the beetles were observed using their antennal spines to catch (or to try to catch) any prey organisms, and during attack and feeding on various arthropods beetles kept their antennae spread in such a way that the spines did not touch the prey. Interestingly, Yin et al. (2017 a) published their study on Mastigini fossils focused on the catching antennal apparatus knowing about my results clearly falsifying the initial ' springtail trap' hypothesis (correspondence with Z. - W. Yin). Function of the enlarged and spiny scape and pedicel in Palaeostigus and other Mastigini remains unclear. The large groups of porous fields between the rows of spines suggest a sensory function, but it does not explain the spines, which in fact would interfere with moving the ventral surface of scape and pedicel close to any objects. The only occasions when the spines possibly directly touch objects in front of beetles are frequent encounters between individuals of the same species, under laboratory conditions forced by keeping many individuals on a relatively small surface, but observed to occur also in nature, as Palaeostigus lives in large groups and beetles continuously walk around and often meet each other. When two beetles meet face to face, they engage in a rapid palpating each other with antennae; it lasts only a second or two, then beetles stop palpating each other and each walks away. The action of antennae is very quick; beetles resemble two ant workers meeting, but the interaction is very quickly terminated. It seems that the antennal spines of beetles engaged in such a brief interaction touch, but movements are too fast to observe any details (my attempts to photograph such meetings failed). Beetles walking on a ground, gathering around dead arthropods or pieces of meet, and ' communicating' among themselves appear similar to ants. The body form, geniculate antennae and the short and dense vestiture of setae that appears silverish on black European adults of Palaeostigus enhance this impression, as Formica ants that are similar in size, shape, pigmentation and behavior can be commonly found in places where Palaeostigus lives (Jałoszyński, pers. obs.). It is possible that beetles benefit from their apparent ' myrmecomorphy', as it was demonstrated that some predators avoid not only ants but also ant-like prey (e. g. Taniguchi et al. (2005 )). Copulation in South African Palaeostigus was observed (Jałoszyński et al. 2015); males have relatively short aedeagi, and they mount females with both individuals facing the same direction, with the tip of copulatory piece inserted into the female's gonopore, and parameres touching female's terminal abdominal segments; because of the different length of parameres, the aedeagus is twisted and positioned asymmetrically.	en	Paweł Jałoszyński (2018): World genera of Mastigitae: review of morphological structures and new ecological data (Coleoptera: Staphylinidae: Scydmaeninae). Zootaxa 4453 (1): 1-119, DOI: 10.11646/zootaxa.4453.1.1
2161879CFF938A4DFF7A35C46552DC3E.taxon	discussion	Remarks. Palaeostigus and Mastigus are morphologically very similar; see Remarks for the latter genus. All nominal species were treated by Bordoni & Castellini (1973) and Leleup (1968).	en	Paweł Jałoszyński (2018): World genera of Mastigitae: review of morphological structures and new ecological data (Coleoptera: Staphylinidae: Scydmaeninae). Zootaxa 4453 (1): 1-119, DOI: 10.11646/zootaxa.4453.1.1
2161879CFF988A75FF7A31786595DAAA.taxon	description	Acanthostigus Leleup, 1968: 95 (as subgenus of Stenomastigus). Type species: Stenomastigus basilewskyi Leleup, 1968 (des. orig.). Synonymized by Jałoszyński (2012 d).	en	Paweł Jałoszyński (2018): World genera of Mastigitae: review of morphological structures and new ecological data (Coleoptera: Staphylinidae: Scydmaeninae). Zootaxa 4453 (1): 1-119, DOI: 10.11646/zootaxa.4453.1.1
2161879CFF988A75FF7A31786595DAAA.taxon	diagnosis	Diagnosis. Adults of Stenomastigus differ from remaining Mastigini in extremely elongate, conspicuously slender aedeagus with one paramere extremely long, much longer than copulatory piece, and the other paramere either completely obliterated or vestigial; in microsculptured, granulose surface of elytra (Fig. 235); and in elytra in females strongly elevated along suture in posterior half, forming a steeply pitched gable roof-like structure (Fig. 162). Mature larva: antennae about 3 – 4 times as long as head capsule (Fig. 236). Characteristics. Adults. Body (Figs 157 – 158, 161 – 162) moderately large, 3.10 ̄ 4.90 mm in length, brown to nearly black, in some cases with elytra lighter than head and prothorax, testaceous or reddish-brown; body strongly convex, dorsally densely but finely setose, setae very short and recumbent, unmodified except for long and thick bristles on scape and pedicel. Head capsule (Figs 221 – 224) divided into large and exposed anterior part and much smaller, subcylindrical ' neck' region retracted into prothorax and demarcated by distinct occipital constriction; ' neck' region much broader than half width of head. Anterior part of head flattened, subequal in width with prothorax, distinctly elongate, broadest near middle. Composite eyes dorsolateral, moderately large, composed of numerous small ommatidia, not projecting or weakly projecting from the silhouette of the head, broadly separated from mandibular bases. Vertex and frons divided by a distinct median longitudinal groove (Fig. 223; mg); vertex strongly transverse, convex at sides, with posterior margin nearly straight or slightly concave. Tempora much longer than eyes, weakly rounded. Frons between antennal insertions not forming a demarcated ' platform', weakly convex or flattened, anteriorly demarcated by a deep and very short frontoclypeal groove largely obliterated at sides. Clypeus very short and broad, broadly subtrapezoidal, with nearly straight sides, which are slightly convergent anterad or parallel. Antennal insertions (Fig. 223; ia) located anterodorsally, relatively narrowly separated. Gular plate (Fig. 224; gp) lacking sutures, indistinctly transversely reticulate; posterior tentorial pits (Fig. 224; ptp) typically circular, rarely oval, in front of broad and diffuse transverse impression demarcating ' neck' region ventrally; hypostomal ridges (Fig. 224; hr) arcuate, posteriorly reaching middle between anterior submental margin and posterior tentorial pits. Head finely to strongly punctate, densely setose (Fig. 223). Antennae (Figs 157 – 158, 161 – 162, 221 – 222) very long and slender, subequal to body length or distinctly longer; scape (Fig. 222; sc) 6 – 8 times as long as broad, thickened, much longer than head, with lateroventral (more ventral than lateral) emargination; pedicel (Fig. 222; pd) enlarged, slightly to much shorter and narrower than scape, typically 5 – 8 times as long as broad, broadening from narrow base to submedian or subapical region. Both scape and pedicel with two ventral longitudinal rows of several long and thick bristles with papillate insertions, area between bristles with variously densely distributed porous fields; antennomeres III – XI distinctly narrower than pedicel, strongly elongate, each indistinctly thickened distad, basal stalks not exposed in intact beetles, basal rings absent or indistinct; antennomere XI elongate and indistinctly asymmetrical. Antennomeres covered with variously dense, long setae; surface of antennomeres smooth. Mouthparts. Labrum (Figs 223, 225; lbr) strongly transverse, with lateral margins slightly divergent anterad or parallel and weakly rounded, and with anterior margin weakly concave, with a pair of broad and short sublateral teeth broadly separated at middle by a very shallow emargination, and with two partly irregular transverse rows of long and short setae. Mandibles (Figs 225 – 226) symmetrical, subtriangular and robust, each with one dorsal mesal tooth and a group of 2 – 3 ventral mesal teeth, setose prostheca present and long. Maxilla (Fig. 229) with large, moderately long cardo (Fig. 229; cd); basistipes (Fig. 229; bst) subtriangular and elongate; mediostipes (Fig. 229; mst) large and sharply demarcated from lacinia and galea, which are both elongate and each with conspicuously dense group of thin distal setae; palpifer (Fig. 229; ppf) broad and elongate; maxillary palp slightly to much longer than head capsule, composed of minute palpomere I (Fig. 229; mxp 1), slender, curved, distinctly but only slightly broadening distad palpomere II, palpomere III strongly elongate, strongly and gradually broadened distad, with transverse or (rarely) slightly obtuse distal margin, palpomere IV (Fig. 227; mxp 4) subequal in length to III, suboval or indistinctly subtriangular and broadening from base to distal third, with rounded apex, elongate. Surface of palpomere IV, and sometimes also III, with sparsely distributed porous fields (Fig. 188; pf). Palpomeres III and IV slightly (sometimes indistinctly) flattened, II – IV covered with relatively long and dense setae. Labium (Figs 224, 229) with broad and short submentum (Fig. 224; smn) posteriorly not demarcated from gular region, densely setose and lacking an outstanding pair of anterior or subanterior lateral setae; mentum (Fig. 229; mn) subtrapezoidal and strongly transverse, with anterior margin very deeply emarginate, so that anterolateral corners form triangular and usually pointed lobes projecting anterad; prementum (Fig. 229; pm) long, subtrapezoidal, broadest distally, lacking demarcated ligula, with several pairs of submedian anterior setae, with broadly separated bases of labial palps; lateral hypopharyngeal lobes moderately large, with conspicuously sparse, thick mesal setae; labial palp composed of three palpomeres: palpomere I (Fig. 229; lp 1) small, elongate, strongly broadening distad, palpomere II (Fig. 229; lp 2) largest, conspicuously enlarged, long and broad, approximately barrel-shaped, palpomere III (Fig. 229; lp 3) very small and narrow in relation to II, about half length of II, pointed. Prothorax (Figs 157 – 158, 161 – 162, 230 – 231) strongly elongate, strongly convex but usually with flattened dorsum (Fig. 231), broadest near anterior third. Pronotum with anterior and posterior margins arcuate (anterior margin sometimes nearly straight), sides rounded in anterior half and sinuate in posterior half; anterior and posterior corners obtuse-angled or broadly rounded; pronotal base lacking pits and groove. Prosternum (Fig. 230) with basisternal part (Fig. 230; bstr) subequal in length to coxal part (Fig. 230; cxst). Prosternum laterally completely fused with hypomera. Coxal region anteriorly and laterally without marginal carina; postcoxal hypomeral lobes (Fig. 230; pchl) conspicuously large, rounded and strongly projecting mesad or anteromesad and overlapping with (but not fused to) posterolateral lobes of prosternum, so that procoxal cavities are not open, but entirely delimited posterioly by hypomeral lobes. Prosternal intercoxal process developed as a diffuse, barely discernible longitudinal elevation, which in intact beetles is hidden between procoxae. Ventral surface of prothorax densely setose (Fig. 230). Mesoventrite (Figs 232 – 233) subtrapezoidal, broadening posteriorly. Prepecti (Fig. 232; pre) moderately long and together with anteromedian mesoventral area forming a relatively short ' collar', which has a narrow transverse groove just behind its anterior ridge; setose impression absent or very shallow and diffuse. Mesoventral intercoxal process (Fig. 232; msvp) reaching middle of mesocoxal cavities; short and very broad, subtriangular with pointed or narrowly rounded apex, weakly convex, broadly separating mesocoxae, posteriorly separated from metaventrite. Mesanepisterna (Fig. 233; aest 2) relatively narrow and strongly elongate, demarcated from median part of mesoventrite by a distinct ridge and from mesepimera by complete suture; mesepimera (Fig. 233; epm 2) elongate, indistinctly demarcated from metepimera, not exposed in ventral view. Mesonotum with cordiform, broad mesoscutellum with pointed apex, in intact specimens not visible between elytral bases, or only its very tip discernible; scutoscutellar suture absent. Metanotum partly reduced, with lightly sclerotized mesoscutum, but only slightly shortened alacristae; hind wings absent. Metaventrite (Figs 232 – 233) short, subrectangular and strongly transverse, with lateral margins rounded; mesocoxal cavities with all margins non-carinate; posterior margin of metaventrite deeply bisinuate laterally (in front of each metacoxa) and with a broad metaventral intercoxal process (Fig. 232; mtvp) with shallowly concave or straight posterior margin; anteriorly metaventrite forming a short anterior metaventral process (Fig. 232; amvp) similar in shape to mesoventral intercoxal process or with its tip concave (when the tip of mesoventral process is rounded, as in Fig. 232) and meeting the latter at middle of mesocoxal cavities; metaventral foveae absent. External admetacoxal part of posterior metaventral margin lacking adcoxal carinae, but with adcoxal expansions (Fig. 232; ade). Metanepisterna (Fig. 233; aest 3) relatively narrow, partly visible in ventral view, narrowing posteriorly; metepimera (Fig. 233; epm 3) 2 – 3 times as broad as metanepisterna, with not demarcated inner and outer components, posteriorly extending far behind metacoxae. Metendosternite (metafurca) with stem much broader than long and broadly separated, with divergent lateral furcal arms, lacking median longitudinal projection. Legs (Figs 157 – 158, 161 – 162, 221, 232) extremely, strikingly long and slender. Pro- and mesocoxa short subconical, metacoxa with nearly hemispherical basal part and subconical distal part. Mesocoxa lacking coxal bristles. All trochanters short and subtriangular; femora weakly clavate; tibiae slender; tarsi long and slender, nearly subcylindrical, tarsomeres I – V reducing in length, tarsomere V strongly elongate, with curved and slender claws lacking elongate costae; empodial region was not studied. Elytra (Figs 157 – 158, 161 – 162, 234 – 235) oval, strongly convex, lacking humeral calli and basal impressions, with rounded or pointed apices; elytral disc with superficial, very indistinct and incomplete longitudinal rows of fine punctures obscured by coarse granulose microscultpure (Figs 234 – 235). Elytra densely setose, setae short and nearly recumbent. Abdomen with sternite III firmly fused with metaventrite (so that during disarticulation it is almost impossible to separate intact abdomen), much longer than sternite IV, but shorter than IV – VI together; sternite VIII in male distinctly, deeply emarginate. Aedeagus (illustrated in Jałoszyński et al. (2018 )) strongly, often extremely elongate, with asymmetrical median lobe and asymmetrical parameres, one paramere extremely long, much longer than copulatory piece, the other one obliterated or (rarely) present but vestigial; flagellum very long and forming several coils. Ejaculatory duct with elongate and narrow sperm pump lacking funnel-like structures. Aedeagus with elongate copulatory piece with membranous endophallus permanently everted, and only inflated during copulation, with distal end of flagellum permanently fixed, so that flagellum is not extricable. Aedeagus in repose positioned asymmetrically within abdomen, with basal orifice lateral or dorsolateral; in species with extremely long paramere aedeagus cannot be completely retracted into abdomen and in repose a short apical portion of paramere is permanently outside. Spermatheca subglobose, with relatively large accessory gland. Sexual dimorphism distinct, females distinctly larger and with much broader elytra, which have different shape than those in males, with a pair of variously large and deep impressions in anterior third (to receive protrochanters of males during mating) and posteriorly steeply elevated along suture, forming a ' gable roof', which, through a gap between separately rounded elytral apices, receives the long paramere during copulation. Slender males have evenly convex elytra and the apical portion of each protibia curved and / or provided with a variously developed tooth, in some species protrochanters are modified, angulate or with variously long ventral projection. Larva. Very similar to larva of Palaeostigus, but lightly pigmented (Fig. 236), orange when alive, and with extremely elongate antennae, which in mature larvae are more than three times as long as head capsule.	en	Paweł Jałoszyński (2018): World genera of Mastigitae: review of morphological structures and new ecological data (Coleoptera: Staphylinidae: Scydmaeninae). Zootaxa 4453 (1): 1-119, DOI: 10.11646/zootaxa.4453.1.1
2161879CFF988A75FF7A31786595DAAA.taxon	description	Composition and distribution. Stenomastigus comprises 17 species and 6 subspecies distributed in eastern part of South Africa (Fig. 165). Natural history. Species of Stenomastigus live in large groups on relatively restricted areas in South Africa; adults are diurnal and they can be found along streams, rivers or ditches with trees, bushes and long grasses (Figs 237 – 239) providing some shade and surface for beetles to climb; or in forests, also close to water (Figs 241 – 242) or in swampy areas. They seem to be more active, or at least can be encountered in greater numbers, during cloudy or even rainy days. Beetles can be found on ground, on mosses and among low vegetation (Figs 234 – 246), but they also climb tall grasses, bushes and trees (Fig. 240) and can be collected by using an entomological umbrella and beating twigs at 2 m and higher. Larvae live in moist leaf litter close to water. During short field observations and a few days of keeping living adults of two species in plastic containers, I was not able to observe any feeding-related behavior. An unusual way of copulation in South African Stenomastigus was described (Jałoszyński et al. 2015); males have extremely long aedeagi, in some cases nearly reaching 3 / 4 of the body length, and they mount females in such a way that both individuals face the same direction, with the tip of copulatory piece inserted into the female's gonopore, and the long paramere inserted into the female's subelytral space, the apical portion of paramere is positioned along suture between anterior impressions of female's elytra, which provides an additional stabilization for the aedeagus. Retracting the aedeagus after copulation takes some time, during which male lays on its back twisting and positioning its enormously long aedeagus. Not only the copulation is remarkably different that that in externally similar Palaeostigus; the male terminal abdominal segments in Stenomastigus have an additional, asymmetrical group of muscle fibers, presumably to assist the rotation of the aedeagus during its retraction (Jałoszyński et al. 2015). It was hypothesized (Jałoszyński et al. 2015) that the elongation of the aedeagus during evolution was possible because of an already existing preadaptation, a broadly separated arms of the metendosternite, between which the median lobe of aedeagus can be placed, and in extreme cases its base can reach the mesoventrite. In genera with Y-shaped metendosternite and especially in those with a long metafurcal stem, this structure forms a structural obstacle between the abdomen and thorax, and the aedeagus is always shorter than the abdomen. Stenomastigus is so far the only staphylinid genus, and one of a few among beetles, in which the architecture of extrinsic and intrinsic aedeagal muscles was studied in detail, with three-dimensional reconstructions of the postabdomen and also some other internal abdominal organs (Jałoszyński et al. 2015).	en	Paweł Jałoszyński (2018): World genera of Mastigitae: review of morphological structures and new ecological data (Coleoptera: Staphylinidae: Scydmaeninae). Zootaxa 4453 (1): 1-119, DOI: 10.11646/zootaxa.4453.1.1
2161879CFF988A75FF7A31786595DAAA.taxon	discussion	Remarks. Although Stenomastigus can be easily recognized on the basis of enormously long antennae and legs, the slender body form of males and the modified elytra in females, diagnostic features of this genus are relatively weak, and the classification of Mastigini may require further study. All nominal species were treated by Franz (1984), Leleup (1968) and Jałoszyński (2012 d; e).	en	Paweł Jałoszyński (2018): World genera of Mastigitae: review of morphological structures and new ecological data (Coleoptera: Staphylinidae: Scydmaeninae). Zootaxa 4453 (1): 1-119, DOI: 10.11646/zootaxa.4453.1.1
2161879CFFA18A74FF7A348061B7DA11.taxon	description	Cascomastigus Yin & Cai (in Yin et al. (2017 a )): 2. Type species: Cascomastigus monstrabilis Yin & Cai, 2017 (orig. des.). Placed as junior synonym of Clidicostigus by Jałoszyński et al. (2018).	en	Paweł Jałoszyński (2018): World genera of Mastigitae: review of morphological structures and new ecological data (Coleoptera: Staphylinidae: Scydmaeninae). Zootaxa 4453 (1): 1-119, DOI: 10.11646/zootaxa.4453.1.1
2161879CFFA18A74FF7A348061B7DA11.taxon	diagnosis	Diagnosis. Clidicostigus differs from all remaining Mastigini in deep, regular, continuous longitudinal elytral grooves; mesoventral and anterior metaventral processes both much longer than broad and together forming nearly parallel-sided narrow carina separating mesocoxae; and elongate maxillary palpomere IV with one side strongly convex and the other sinuate (approximately boomerang-shaped). Composition and distribution. Clidicostigus (Fig. 163) comprises three extinct species known from the Cenomanian Myanmar amber.	en	Paweł Jałoszyński (2018): World genera of Mastigitae: review of morphological structures and new ecological data (Coleoptera: Staphylinidae: Scydmaeninae). Zootaxa 4453 (1): 1-119, DOI: 10.11646/zootaxa.4453.1.1
2161879CFFA18A74FF7A348061B7DA11.taxon	discussion	Remarks. Clidicostigus has an asymmetrical aedeagus, with its distal region (reconstructed by microcomputer tomography by Jałoszyński et al. (2017 )) closely resembling that of the extant Mastigus spinicornis (Fabricius, 1787). The elytral base devoid of humeral calli suggests that adults were wingless; extremely long legs, antennae and maxillary palps and other structures closely resembling those in extant Mastigini suggest a similar mode of life, i. e. possibly diurnal activity and relaying on walking and running to locate a source of food and to disperse.	en	Paweł Jałoszyński (2018): World genera of Mastigitae: review of morphological structures and new ecological data (Coleoptera: Staphylinidae: Scydmaeninae). Zootaxa 4453 (1): 1-119, DOI: 10.11646/zootaxa.4453.1.1
2161879CFFAE8A7BFF7A36E463C6DAD5.taxon	diagnosis	Diagnosis. Baltostigus is the only genus of † Baltostigini; diagnosis and characteristics as for tribe, vide supra. Composition and distribution. Baltostigus comprises three species known from Upper Eocene Baltic amber (Jałoszyński 2016 a; Jałoszyński et al. 2018).	en	Paweł Jałoszyński (2018): World genera of Mastigitae: review of morphological structures and new ecological data (Coleoptera: Staphylinidae: Scydmaeninae). Zootaxa 4453 (1): 1-119, DOI: 10.11646/zootaxa.4453.1.1
