Description of Kashmirobroscus gen. nov. with two new species from the Northwest Himalaya, and remarks on the East Asian genus Eobroscus Kryzhanovskij, 1951 (Coleoptera: Carabidae: Broscini) Author Schmidt, Joachim Lindenstrasse 31, 18211 Admannshagen, Germany; Author Wrase, David W. Dunckerstrasse 78, 10437 Berlin, Germany; Author Sciaky, Riccardo Via Fiamma 13, 20129 Milano, Italy text Journal of Natural History 2013 J. Nat. Hist. 2013-09-03 47 41 - 42 2671 2689 http://dx.doi.org/10.1080/00222933.2013.791952 journal article 55814 10.1080/00222933.2013.791952 857d6372-de50-463d-8471-717e72bdd3ba 1464-5262 4608237 1775013D-FE87-41CA-87AF-D730E809331B Kashmirobroscus gen. nov. Type species: Kashmirobroscus ortrudae sp. nov. urn:lsid:zoobank.org:act: 304B2D40-25B9-4FD1-9A55-4CDABAA86101 Derivation of generic name A masculine noun that combines the name of the geographical origin of the new taxon, the Kashmir Himalaya, with the name of the nominotypical genus of the tribe Broscini , the genus Broscus Panzer, 1813 . Description Body length: 11.6–13.6 mm . Colour: Body surface reddish brown to piceous, glossy. Appendages reddish brown with mouthparts somewhat lighter and femora and tibiae somewhat darker. Micro-sculpture: Entire surface covered with micro-puncturation (magnification 100 ×). Surface of head with mesh pattern isodiametric, very indistinctly engraved, visible under high magnification (100 ×) except on labrum which has micro-meshes distinctly engraved (magnification 40 ×). Pronotum with mesh pattern somewhat irregular, very shallowly impressed (magnification 100 ×). Elytra with mesh pattern isodiametric, somewhat more deeply impressed than on head and pronotum. Head: Large, moderately convex, somewhat narrower than pronotum, with surface smooth. Eyes relatively small, slightly prominent, with one (exceptionally with two) supraorbital seta at level of hind margin of eye ( Figure 3A, C ). Tempora distinct, indistinctly swollen, smooth, about as long as length of eye diameter (viewed dorsally), temporal ridge incomplete. Frontal impressions shallow, divergent posteriorly. Vertex with an impunctate transverse impression that is deepened in middle ( Figure 3A, C ). Mandibles long and slender with seta in scrobe. Mentum with two setae each side and with pair of deep paramedian foveae; apical tooth truncate, shallowly emarginate, hence shallowly bifid; submentum with three setae each side, the middle one longer and broader ( Figure 3B ). Ligula in central part swollen, bisetose apically, paraglossa almost as long as ligula. Cardo with one, stipes with two or three setae. Gula with two deep transverse grooves on each side. Antennae relatively long (ninth or tenth antennomere extended to pronotal base), with scape glabrous apart from the dorsoapical seta; pedicel with one or two ventroapical setae; third antennomere glabrous apart from the ring of obligatory apical setae; fourth antennomere apart from the ring of long obligatory apical setae with numerous fine and short setae in apical third; 5th to 11th antennomeres pubescent on all surfaces. Pronotum: Cordate, distinctly narrower than elytra (EW / PW 1.54–1.69), as wide as long or somewhat wider than long (PW / PL 0.99–1.10), widest at apical third, strongly contracted towards base (PW / PBW 1.50–1.64), dorsally fairly convex. Apical margin somewhat wider than basal margin, with apical angles weakly protruded. Basal margin markedly convex. Lateral margin finely beaded, rounded in anterior four-fifths and rectilinearly divergent towards basal angles; latter obtuse, blunt at tip. Basal portion of epipleuron near basal angles visible in dorsal view ( Figures 1A , 2A ). Lateral margin each side with three to five setae in apical half and one seta slightly anteriad basal angle. Median line moderately deep, not reaching apical and basal margin; anterior transverse impression indistinct, posterior transverse impression very shallow but distinct, laterobasal foveae narrow, shallow. Base more or less coarsely punctuate or transversely wrinkled. Elytra: Long-ovate, flattened on disc; humeri widely rounded, with apical portion of lateral border not reflexed, terminated at level of fifth stria. Parascutellar seta and parascutellar stria absent. Striae on disc slightly impressed, distinctly punctured, almost absent at sides and near apex, intervals on disc weakly convex. Lateral umbilical series consists of one posthumeral, one preapical and two apical setae. Hind wings: Reduced to about one-third of elytral length. Sterna: Prosternal process furrowed, not margined; prosternum, pro- and mesepisternum, mesepimeron and metepisternum smooth or, in some specimens, with few shallow punctures. Metepisternum moderately long, with ventral margin approximately 1.5 times longer than anterior margin, moderately narrowed posteriad. Abdomen: Smooth in middle, somewhat wrinkled laterally and with a fine irregular puncturation ( Figure 1B ), sternites IV–VI each with one pair of ambulatory paramedian setae, sternite VI of male apically with a button-like, round elevation at middle ( Figure 1B, C ), last abdominal sternite apically usually with one seta on each side in male, two in female (but chaetotaxy sometimes irregular). Legs: Slender, with protrochanters bisetose and with mesotrochanters and metatrochanters unisetose. Male protarsomeres very indistinctly dilated, tarsomeres 1 and 2 with adhesive vestiture ventrally; mesotarsomeres without adhesive vestiture. Dorsal side of mesotarsomere and metatarsomere 1 (less on metatarsomeres 2) with a blunt longitudinal carina in middle that is bordered by a distinct longitudinal furrow on each side ( Figure 4J, K ). Tarsomere 5 on ventral side with one couple of setae behind middle ( Figure 4J , exceptionally with an additional seta in middle). Male genitalia: Sternum IX with sclerotized ring complete, with right margin markedly narrowed ( Figure 5J ). Median lobe of aedeagus with basal orifice partially open, with basal third of dorsal surface sclerotized, without basal expansions, but with wing-like expansions on lateral and ventrolateral surfaces ( Figure 5G, I ). Internal sac with sclerite X wide, with a distinct constriction in middle and with two latero-apical projections ( Figure 5 G–I); sclerite Y present; apical plate composed of free spiculae, without tooth. Both parameres setiferous ( Figure 5I ). Figure 1. Kashmirobroscus ortrudae sp. nov. , paratype, male. (A) Dorsal view; (B) ventral view; (C) left lateral view. Abbreviation: bt, button-like elevation on sternite VI. Female genitalia ( Figure 6A ): Helminthoid sclerite of accessory gland of bursa copulatrix long and thin, U-shaped. Phylogenetic relationships Evidence for inclusion of Kashmirobroscus gen. nov. in the subtribe Broscina is based on the following two diagnostic features, which are considered synapomorphies (RoigJuñent 2000; Liebherr et al. 2011 ): Figure 2. Kashmirobroscus and Eobroscus in dorsal view. (A) Kashmirobroscus mirabilis sp. nov. , holotype; (B) Eobroscus bhutanensis Morvan, 1982 , female specimen from Mt Manaslu, Nepal. Sclerite X in the internal sac of aedeagal median lobe with upturned latero-apical projections (see Figure 5 G–I). Helminthoid sclerite of accessory gland of bursa copulatrix U-shaped (see Figure 6A ). Within Broscina , Kashmirobroscus gen. nov. , together with Eobroscus Kryzhanovskij, 1951 and Zacotus LeConte, 1869 , seems to form a natural group that is defined by the following two derived character states ( Roig-Juñent 2000 ; Liebherr et al. 2011 ): Figure 3. Head in dorsal (A, C, D) and ventral (B) views of Kashmirobroscus and Eobroscus . (A, B) Kashmirobroscus ortrudae sp. nov. , paratype, male; (C) Kashmirobroscus mirabilis sp. nov. , holotype; (D) Eobruscus bhutanensis Morvan, 1982 , female specimen from Mt Manaslu, Nepal. Abbreviation: ms, setae of mentum; sms, setae of submentum. Gula on each side with two deep transverse grooves or oblique foveae ( Figure 3B ). Helminthoid sclerite of the female genitalia thin and long (see Figure 6A, B ). Within this group the East Asian genus Eobroscus and the Northwest Himalayan genus Kashmirobroscus gen. nov. have six characters in common that are considered synapomorphies and so, allow for the assumption of a sister-group relationship: Transverse impression on vertex markedly deep ( Figure 3A, C, D ). Head with vertex smooth seems to represent the plesiomorphic character state as it is developed in most Broscini genera and in the outgroup taxa Apotomini and Melaenini. A transverse impression is also developed in the Broscina genera Broscodera Lindroth, 1961 (incl. Sinobrosculus Deuve, 1990 ), Broscosoma Rosenhauer, 1846 and Craspedonotus Schaum, 1863 ; however, it is distinctly shallower in these taxa than in Eobroscus and in Kashmirobroscus gen. nov. In Zacotus matthewsii LeConte, 1869 , which is a supposed sister taxon of Eobroscus + Kashmirobroscus , a deep short longitudinal furrow exists in the middle of the vertex beside a shallow transverse impression. Hence, the markedly deep transverse impression of vertex in Eobroscus + Kashmirobroscus seems to represent a character state that evolved independently from that of the Broscina genera named above. Roig-Juñent (2000: 27 , fig. 72) figured misleadingly a transverse impression also on vertex of Miscodera arctica (Paykull, 1798) . Instead, this species has the dorsal surface of head completely smooth and rounded in basal half but has a distinct transverse impression on frons. Sternite VI of male apically with a button-like elevation at middle ( Figure 1B, C ). This character is unique within Broscini and outgroups. Protarsi of male weakly dilated ( Figure 4D, G ). Zacotus matthewsii possesses the plesiomorphic character state with male protarsomere 1–3 markedly dilated ( Figure 4A ). Dorsolateral surfaces of tarsi with longitudinal furrows, at least on first hind tarsomere (see Figure 4F , I–K). Tarsi with dorsal side completely smooth (see Figure 4C for Zacotus matthewsii ) seems to represent the plesiomorphic character state as it is developed in other Broscini (except for Broscodera ) and outgroups ( Apotomini , Melaenini). In contrast to what is developed in Eobroscus and Kashmirobroscus , which have dorsal surfaces of all tarsomeres glabrous (except for the pair of dorsoapical setae on fifth tarsomere), the longitudinal furrows on dorsal surfaces of the tarsomeres of Broscodera are bordered by fine keels which each terminate in a fine seta. It is therefore most likely that longitudinal furrows in Broscodera evolved independently from those of Eobroscus + Kashmirobroscus . Fifth tarsomere with one or two pairs of ventral seate ( Figure 4 D–K). Three or more pairs of setae (see Figure 4 A–C for Zacotus matthewsii ) seem to represent the plesiomorphic character state as it is developed in other Broscini and outgroups. Aedeagal median lobe with wing-like lateral expansions in middle portion ( Figure 5 A–G, I). Except for some Himalayan species of the genus Broscosoma ( B. monticola group), which have a single sail-like expansion on the ventral surface this character is unique within Broscini and outgroups. Independent evolution of the ventral expansion within a terminal lineage of Broscosoma is most likely. In addition, the monotypic genus Zacotus from North America can be distinguished from Eobroscus + Kashmirobroscus by the following characters: Figure 4. Protarsi (A, D, G) and mesotarsi (B, E, H) in ventral view, and metatarsi in lateral view (C, F, I–K) of Broscini . (A–C) Zacotus matthewsii LeConte, 1869 , male specimen from Vancouver, Canada; (D–F) Eobroscus lutshniki (Roubal, 1928) , male specimen from Ussurisk, Russia; (G–I), Eobroscus bhutanensis Morvan, 1982 (G, H, male specimen from Qingcheng Houshan, China; I, female specimen from Mt Manaslu, Nepal); (J) Kashmirobroscus ortrudae sp. nov. , paratype, male; (K) Kashmirobroscus mirabilis sp. nov. , holotype. Abbreviation: lf, longitudinal furrow on dorsolateral surface of first metatarsomere. Figure 5. Aedeagal median lobe in left lateral view (A, C, E, G; H shows a cut-out of G) and in dorsal view (B, D, F, I; B and I additionally with parameres), and sternum IX (J) of Kashmirobroscus and Eobroscus . (A, B) Eobroscus lutshniki (Roubal, 1928) , specimen from Ussurisk, Russia; (C–F) Eobroscus bhutanensis Morvan, 1982 (C, D, specimen from Qingcheng Houshan, China; E, F, specimen from Mt Fansipan, Vietnam); (G–J), Kashmirobroscus ortrudae sp. nov. , paratype, male. Abbreviations: lap, latero-apical projections of internal sac; pl, left paramere; pr, right paramere. Dorsal side of body with splendid metallic coloration. Fourth antennomere glabrous in basal three-quarters. Dorsal side of fifth tarsomere with numerous fine setae. The character of incomplete temporal ridge, given by Roig-Juñent (2000: 13) for separating Zacotus from Eobroscus , is not useful, as the ridge is also incomplete in Eobroscus species, though the area of interruption is not completely smooth and convex (as in Zacotus ), i.e. it is somewhat angled between the upper and lower part. Differential diagnosis In Eobroscus the transverse neck constriction is much more sharply incised (as if made with an axe, autapomorphy of Eobroscus , see Figure 3D ) than in Kashmirobroscus gen. nov. ( Figure 3A, C ). In Eobroscus longitudinal furrows are developed on dorsal surfaces of all tarsomeres, at least near base (derived character state, see Figure 4F, I ), whereas in Kashmirobroscus longitudinal furrows exist only on dorsolateral surfaces of the first mesotarsomeres and metatarsomeres ( Figure 4J, K ). The distal portion of the tibiae is longitudinally wrinkled in Eobroscus (autapomorphy, see Figure 4I ) and smooth in Kashmirobroscus gen. nov. (plesiomorphic character state, see Figure 4K ). The adhesive vestiture on the ventral side of male mesotarsomeres 1 + 2 is developed in Eobroscus (plesiomorphic character state, see Figure 4E, H ) and absent in Kashmirobroscus gen. nov. (autapomorphy). The parascutellar seta is present in Eobroscus (plesiomorphic character state) and absent in Kashmirobroscus gen. nov. (autapomorphy). In Kashmirobroscus gen. nov. the number of lateral setae in the pronotal apical half is increased to three to five (autapomorphy) in contrast to a single pair of lateral setae in Eobroscus (plesiomorphic character state). Additionally, differences of Kashmirobroscus gen. nov. (autapomorphies) are developed in the degree of adaption to the high-altitude environment: body slender ( Figures 1A , 2A ), eyes smaller ( Figure 3 A–C), elytra flattened ( Figure 1C ), hind-wings reduced, humeri rounded ( Figures 1A , 2A ) and metepisterna shortened. Geographical distribution Figure 7 shows the geographical distribution. Only two species are known from different localities in the Northwest Himalaya of Pakistan (for details see below).