Revision of the East Mediterranean Orthomus (Coleoptera, Carabidae, Pterostichini), with description of Parorthomus gen. n. socotranus sp. n. from Socotra Island and key to the Old World genera of subtribe Euchroina

Abstract The East Mediterranean species of Orthomus Chaudoir, 1838 are revised. The type series of Feronia longula Reiche & Saulcy, 1855, F. berytensis Reiche & Saulcy, 1855, F. proelonga Reiche & Saulcy, 1855, Orthomus longior Chaudoir, 1873, O. sidonicus Chaudoir, 1873, and O. berytensis akbensis Mateu, 1955 were studied and lectotypes for the first four are designated. Also, the following nomenclatural acts are proposed: Feronia proelonga Reiche & Saulcy, 1855, syn. n. of Orthomus berytensis (Reiche & Saulcy, 1855); Feronia elongata Chaudoir, 1859, syn. n. of Orthomus berytensis (Reiche & Saulcy, 1855); Orthomus sidonicus Chaudoir, 1873, syn. n. of Orthomus longior Chaudoir, 1873; Orthomus velocissimus andalusiacus Mateu, 1957, syn. n. of Orthomus velocissimus akbensis Mateu, 1955, new assignment for Orthomus berytensis akbensis Mateu, 1955. As a result, three species of the genus inhabit the East Mediterranean biogeographical region: O. berytensis, O. longior, and O. longulus. A key to these three species is given. O. longior is recorded for Turkey and Syria for the first time. In addition, a new synonymy of two West Mediterranean taxa is proposed: O. szekessyi (Jedlička, 1956), syn. n. of O. balearicus (Piochard de la Brûlerie, 1868), and a new genus and a species are described: Parorthomus gen. n. socotranus sp. n. (type locality: Republic of Yemen, Socotra Archipelago, Socotra Island, Fimihin env., 530 m.a.s.l.). Illustrations of the species dealt with here are provided including external characters, habitus, mentum and submentum, and genitalia are provided. Nine genera of the “African Series” of subtribe Euchroina Chaudoir, 1874 are keyed for the first time. Checklists of the species of Orthomus and of the Old World euchroine genera are given.


Introduction
Orthomus Chaudoir, 1838 is a Palaearctic genus of pterostichine carabid beetles with Circum-Mediterranean distribution, which includes 23 species (Bousquet 2003, Lorenz 2005, Wrase and Jeanne 2005, Pupier and Coulon 2013. The greatest diversity of species is concentrated in the Western Mediterranean, mostly in the Iberian Peninsula and North-West Africa. Only a few species live in the Eastern Mediterranean. The group is often ranked as nominotypical subgenus of Orthomus s.l., together with the Macaronesian Eutrichopus Tschitschérine, 1897 (three species), Gietopus Machado, 1992 (one species), Nesorthomus Bedel, 1899 (eight species), Wolltinerfia Machado, 1985 (two species), and the North-West African Trichopedius Bedel, 1899 (one species) (Bousquet ibid., Lorenz ibid., Serrano et al. 2009, Serrano et al. 2012), but some authors accepted a generic status (Mateu 1954, Machado 1992, Ortuño 1996, Wrase and Jeanne 2005, Donabauer 2008. A recent work on mitochondrial DNA variations among species of Eutrichopus, Wolltinerfia and Orthomus, found relatively high genetic divergence between them and suggested dealing with these as distinct genera (Moya et al. 2004: 3163). Based on established phyletic distance, the last taxon is treated here as an independent genus.
All the afore-enumerated taxa together with the Afrotropical genera Abacillius Straneo, 1949 andAbacillodes Straneo, 1988 were referred as to the "African Series" of subtribe Euchroina Chaudoir, 1874(Will 2006. The species included in Euchroina share several derived characters that is exceedingly short or no coronal suture (i), ventrally extended membranous band on the maxillary stipes (ii) in the larvae (Bousquet and Liebherr 1994), well-impressed frontal furrows of head, not or hardly attaining the level of anterior supra-orbital puncture (iii), hind trochanters without setae (iv), and "gooseneck" shaped bursa copulatrix (v) in the imago (Will 2002, 2006, Frania and Ball 2007, which suggest monophyletic origin of the subtribe. Besides, most Old World euchroines have relatively large and prominent eyes compared to the size of the head, submentum without lateral setae, distinct parascutellar stria, sternites V-VII with transverse sulci (complete or not), segment 5 of tarsomeres setose beneath and median lobe of aedeagus with dorsal ostium.
The genus Cedrorum Borges & Serrano, 1993 from the Azores does not belong to this series as its only species, C. azoricus Borges & Serrano, 1993, exhibits a set of features atypical of the euchroines, such as: superficial frontal furrows, submentum with lateral setae, left paramere with a slight transverse aphophysis, falcate right paramere, no transverse sulci on sternites V-VII (contrary to the statement of Borges and Serrano 1993: 320) and segment 5 of tarsomeres glabrous ventrally.
The East Mediterranean species of Orthomus were the object of taxonomic reviews quite a while ago (Mateu 1955). Since then, however, new data and new material have accumulated. Investigations of material in several European museum and private collections demonstrated that the identifications of the species from the area in question were often incorrect, our investigations proved the occurrence of only three species there. For example, a review of material from Greece, formerly published or identified as O. barbarus and O. berytensis, revealed that only the latter species inhabits this country (Arndt et al. 2011: 58). Hence, the necessity to improve our taxonomic knowledge of the East Mediterranean representatives of the genus was the main reason to start this work. Besides, a few years ago one of us (DWW) received a series of Orthomus-like specimens from Socotra Island. Subsequent examination has proven that it belongs to a species and a genus new to science.
Male specimens were boiled in water and their genitalia were extracted, put in 10% KOH solution, then washed and neutralized and then stored in glycerine. The figures were made with Zeiss transmitted-light microscope. After that, the aedeagus and parameres were embedded in Euparal either on the same card with specimen from which they were extracted or on a separate transparent label beneath the specimen from whom they were removed. The measurements and other drawings were made with the aid of an Olympus SZX10 stereobinocular. The photos of habitus and pronotum were made with a Zeiss Stemi 2000 microscope equipped with an AxioCam ERc 5s camera.
Measurements: body length from the apex of the longer mandible to the apex of the longer elytron (BL); maximum linear distance across the head, including the eyes (HW); maximum width of pronotum (PW); length of pronotum, measured from the apical margin to the basal margin along the midline (PL); width of the pronotal base, between the tips of the hind angles (PbW); length of elytra, from a line connecting the apices of the humeral angles to the apex of the longer elytron (EL); maximum width of elytra (EW). The surface of the paramere close-fitting to the distal part of the median lobe of the aedeagus is denoted as internal face, on the contrary the second surface is the external face.
The material examined is housed in the collections listed below:  Straneo, 1949 Republic of South Africa Abacillodes Straneo, 1988 Malawi Eutrichopus Tschitschérine, 1897

BMNH
Canary Islands Gietopus Machado, 1992 Canary Islands Nesorthomus Bedel, 1899 Madeira Orthomus Chaudoir, 1838 Canary Islands, Mediterranean Parorthomus gen. n. Socotra Trichopedius Bedel, 1899 Algeria Wolltinerfia Machado, 1985 Canary Islands  : Mateu, 1955: 56, 76 Orthomus muluyensis Antoine, 1957: 205 (type locality: Guercif ) Pterostichus (Orthomus) haligena Wollaston, 1860: 87 (type locality: "Great Salvage") Note. The taxon haligena, described from the Salvage Islands belongs to O. berytensis (see Machado 1992: 260, Lorenz 1998: 249, 2005 and not to O. barbarus where it was quoted by Bousquet (2003: 477), as a subspecies, erroneously with the distribution data Madeira. The examination of 13 specimens (Ilhas Selvagens, Selvagem Grande, SE. sublittoral zone, 65-90 m (under stones), XI/XII 2006, D. Putzer leg., DWBG) in comparison with specimens from the Canaries and from Morocco revealed some differences, namely a smaller body size and a strong elytral reticulation in males, almost as strong as in females. These differences could suggest that we deal with a distinct taxon but further investigations are needed. The taxonomic status of the names atlantica and muluyensis, both partly considered in the literature as a "forma" or a subspecies of berytensis or as a synonym to this name must be cleared up by further investigation of a larger sample of material.  (Reiche & Saulcy, 1855), male, "Tel-Aviv" 2 O. longior Chaudoir, 1873, male, "Upper Galilee, Ya'ar Bar'am" 3 O. longulus (Reiche & Saulcy, 1855), male, "Upper Galilee, Ha Khula Valley, Ma'agar Einan lake". Scale bar = 2 mm. male paralectotype from MHNG were examined and glued to cards, pinned beneath the specimens from which they had been removed. The specimen from MNHP is a male, with previously extracted genitalia and glued on a separate card pinned beneath the specimen. This sample bears two old labels equal in size and type: "berytensis type Reiche" [handwritten on white label by Mateu] We got the information that the part of the Reiche collection housed in MHNG was integrated into the general collection (which was build up essentially from the collection of André Melly) with all specimens with: "Coll. Reiche" but usually there are neither locality, nor identification labels attached to individual specimens, and that identification and locality data figure only on the 'taxa labels' (handwritten by Melly) pinned to the bottom of the drawer, which have thus to be considered as pertaining collectively to the specimens pinned above (Cuccodoro in litt.). The species was described from "Beyruth", taking into consideration the above mentioned facts concerning labelling of the types we suppose the above mentioned typical specimens as coming from the type locality.

I. Revision of the East Mediterranean
Type material. Feronia longula Reiche & Saulcy, 1855 (specimens belonging to O. berytensis). The type series in MHNG contains two specimens belonging to O. berytensis: 1 ♀, "Jaffa, Syrie" [Reiche's handwriting on yellow label], "Coll. Reiche" paralectype from MHNG examined by ourselves (the apices of both median lobes were found damaged!) and glued to cards together with the respective specimens. The specimen from MNHP, designated as paralectotype, was with previously extracted genitalia. Its aedeagus and parameres are glued to a separate card pinned beneath the specimen. Additionally, this paralectotype bears two old labels equal in size and type: "proelongus The study of the type material of Feronia proelonga found that this taxon is to be removed from synonymy with O. barbarus barbarus (cfr. Bousquet 2003: 477) and treated as conspecific with O. berytensis. Originally, it was separated by the authors in the relatively shorter size of body and pronotum seemingly more transversal, with two basal impressions distinct and strongly punctured. We found such variations in the populations of O. berytensis coming from modern-day Israel. Type material. Feronia elongata Chaudoir, 1859. We were unable to find any type specimen(s) of this taxon, but the description and especially the distribution Chaudoir gave for his species (Chaudoir 1859: 116) Wollaston (1864: 47), and Wollaston (1865: 39), which concerns, according to Machado (1992: 259), O. berytensis. On the other side, the first locality listed by the author from among the type one is "Moräa". Morea was the name of the Peloponnese Peninsula in South Greece during the early modern period (ca. 1500-1800). At present, based on recent data (Arndt et al. 2011: 58, present Chaudoir, 1873 Figs 2, 5, 9, 13, 17 Othomus [sic!] longior Chaudoir, 1873: 105 (type locality: "Sidon"), (as locality of LT), part Feronia (     Pronotum with hind angles obtuse at tip, often with small denticles protruding laterally (Fig. 16). All discal setiferous punctures of elytra as a rule situated close to or in stria 3. Elytral striae smooth, sometimes with a shallow punctuation. Mesepisternum smooth or with shallow punctuation only. Elytral microsculpture in females consisting of isodiametric meshes almost regularly arranged (as in O. longior). Median lobe of aedeagus toward apex distinctly shifted to the left, apical lamella narrowed distally, almost round at tip (dorsal aspect) (Figs 4, 8)  Remarks. Orthomus berytensis akbensis was described from "Akbes, Siria", based on a male and a female specimen. Mateu characterized it as having the pronotal base not bordered bilaterally, the pronotum widest at about middle, and with basal fovea punctured, the elytral striae hardly punctured, the metatibia in male crenulate at internal side, and the median lobe (Mateu 1955: 75, Fig. 9) somewhat differing from O. barbarus berytensis.

Orthomus longior
The study of the structure of the median lobe of aedeagus and the parameres, as well as selected external features in the holotype of O. barbarus akbensis demonstrated that it is really different from the other three East Mediterranean species. Its comparison with various taxa of the genus revealed that it is identical with specimens of O. velocissimus andalusiacus Mateu, 1957(compare Figs 4, 8, 12 and Mateu 1957, Lámina IV, Figs 5-8, Pupier and Coulon 2013 Fig. 1d). Hence, we synonymize the latter with O. barbarus akbensis which becomes the senior synonym and therefore the name of a valid subspecies of O. velocissimus (Waltl, 1835). Lorenz (2005: 265) formally declared Feronia hesperica as a nomen oblitum, in spite of the fact that a junior name has never been declared as nomen protectum. This statement is incorrect, since after 1899 this taxon was cited at least twice in the coleopterological literature. Heyden et al. (1906: 86) and Csiki (1930: 612) recorded it as synonym of Pterostichus (Orthomus) barbarus. The type locality cannot be fixed exactly geographically, Motschulsky (1849: 52) wrote that the coleopterological yields the collector Handschuh made in 1847 in southern Spain came from "principalement aux environs de Carthagène", which means that other parts of southern Spain cannot be excluded. Thus, we accept the view of Mateu (1957: 103) and list F. hesperica as a questionable senior synonym of O. velocissimus akbensis, before the identity of the former can be settled.

Diagnosis.
A Euchroina genus of beetles that are medium-sized (8.2-10 mm), black coloured, brachypterous, with the following combination of characters: convex eyes; mentum with bifid tooth and large labial pits; pronotum sides posteriorly straight to slightly convex; elytra with 3-4 (rarely 2 or 5) discal setiferous puncture in stria 3/interval 3, with last puncture in posterior third of elytron; intercoxal process of prothorax subquadrate, distinctly bordered at sides and backwards; metaepisterna as wide as long; abdominal sternites V-VII with transverse basal sulci complete and wellimpressed; mesotibia and metatibia straight in both sexes, mesotibia distally with slight inner callus in males; tarsomeres glabrous dorsally, with segment 5 setose ventrally; distal part of median lobe of aedeagus considerably curved to left in dorsal aspect; spermatheca with appended gland spherical and elongate diverticulum.
Description. None required because the genus is monobasic, and its characters are the same as those of its type species.
Etymology. A prefix in apposition (masculine), formed from the Greek παρά-, meaning "beside", "near", "alongside", and the name of the Mediterranean pterostichine genus Orthomus to which the new taxon is related.
Microsculpture distinct on the whole dorsal and ventral surfaces (including coxae, trochanters and femora), consisting of isodiametric and slight transversal meshes, more apparent in females (female specimens almost matt on dorsal surface, males somewhat shiny), reduced on the most part of the clypeus and gula.
Head noticeably longer and narrower with respect to the pronotum, frons smooth, frontal furrows well-marked, divergent posteriorly, reaching the level of anterior supraorbital punctures; neck without constriction posteriorly; eyes fairly large, convex, moderately prominent, with diameter as long as the combined length of segment I-II of the antennae, temporae short, as long as or shorter than half of eye diameter; paraorbital sulci moderately deep, encircling eyes behind; clypeus trapezoidal, separated from frons by fine suture, with anterior margin slightly concave; labrum rectangular; antennae moderately long, pubescent from second fourth of segment IV, the apex of terminal segment not reaching basal margin of pronotum; mentum transverse, deeply emarginate, with large labial pits, median tooth slightly bifid at tip, epilobes narrow, slightly projecting beyond lobes; submentum with medial setae, without lateral ones (Fig. 28).
Pronotum wide, transverse, sub-trapezoid, widest about middle, with margins distinctly, narrowly bordered (the bordering reduced in the middle quarter of apical margin, and sometimes in the middle of basal margin, just between the internal basal impressions); sides somewhat more constricted apically than basally, with two pairs of setiferous punctures, lateral punctures situated at about end of apical third, posterolateral ones situated near hind angles, near to lateral margin and close to basal margin; apical margin moderately emarginate, narrower than basal margin, fore angles rounded, moderately projecting; basal margin nearly straight, slightly concave in the middle, hind angles almost rectangular, rounded at tip; basal impressions somewhat variable in extension and size, internal ones always present, linear, narrow and falcate, diverging toward base, impunctate, deeper and longer than the outer ones, outer impression present or reduced becoming evanescent, when present then mostly faint, foveolate, somewhat punctate; disc slightly convex, midline well-impressed, long, not reaching both anterior and posterior margins.
Elytra sub-elongate, moderately convex, widest at about the second third, fused at suture; shoulders well-marked, obtusely angulate; basal margin complete, reaching stria 1 inwards, forming a very minute denticle at humerus; discal striae moderately impressed, impunctate, parascutellar striae distinct, striae 1-8 joining basal margin; intervals slightly flat, smooth, interval 3 with three to four (rarely two or five) setiferous punctures adjoining stria 3, with last puncture in posterior third of elytron, rarely in about middle of interval 3 (see also Variability); scutellar setiferous puncture present; hind wings reduced to small scales.
Abdominal sternites IV-VI with transverse basal sulci complete (continuous) and well-impressed, abdominal sternum VI with posterior margin rimmed throughout, with one pair of foveate setigerous punctures medially in males and two pairs in females.
Male genitalia (5 specimens dissected). Median lobe of aedeagus slender , narrower at middle, with basal part long, almost rectangularly bent behind apical part, narrowest at middle, from there toward apical lamella rectilinear, right external angle of apical lamella somewhat bent down, left external angle elevated (left lateral view), median lobe from middle part shifted to left, with right margin moderately convex, lengthwise appreciably elevated over left margin, left margin concave towards apex, apical lamella wide, rounded on left side, obtusely angled on right side, with a slight front concavity (dorsal view), ostium slightly deflected to right; right paramere narrow and elongate, smaller than left one, with a slanting lateral process (Fig. 21); left paramere conchoid (Fig. 22).
Variability. Interval 3 with three to four (rarely two or five) setiferous punctures adjoining stria 3, with last puncture in posterior third of elytron, rarely in about middle of interval 3. The number of punctures can increase to five or decrease to two, often the number of punctures of the left and the right elytron is unequal. Also the position of the punctures can somewhat vary. While the first two discal punctures always adjoin stria 3 (and so also the majority of the following punctures), sometimes the third puncture is located on the middle of interval or adjoins stria 2, rarely, the fourth discal pore is located on the middle of interval 3 or adjoins stria 2.
For variability of body size and indices see 'Description' and Table 1. Etymology. The specific epithet is an adjective, referring to Socotra, the island where the new species was collected.
Distribution. Up to present only known from Socotra. Ecology. A mesotopic to eurytopic epigeic beetle, collected from the end of September to the first ten days of February, and on higher ground (Hagher Mts., Scand Mt. env., 1450 m) some specimens were also found in June. From the list of localities, the species seems to be quite widespread across Socotra from the mouths of wadis (near or far off the water) till the highest mountains. Referring on Bezděk et al. (2012), the dominant habitat at most of the mentioned localities is high shrubland with dominant Croton socotranus and Jatropha unicostata (in higher altitude with intermixed Boswellia spp., Dracaena cinnabari, Euphorbia arbuscula, etc.) (Jiří Hájek, personal communication).
Systematic and biogeographic considerations. The Socotra Archipelago is a Gondwanan continental fragment, which has experienced a long period of geological isolation. This landmass was separated from the Arabian plate during the rifting which began to open the Gulf of Aden in the Oligocene to Miocene epochs (d' Acremont et al. 2006). It is supposed that Socotra was isolated from Arabia at least 16 million years ago (d' Acremont et al. 2010). The high level of endemism found among the insects in Socotra (Batelka 2012) is in accordance with the estimated geological age and the supposedly continuous stability of its ecosystems.
At present, it is difficult to ascertain whether Parorthomus gen. n. socotranus sp. n. derived from ancestral populations on the Arabian mainland that probably reached Socotra by transoceanic dispersal in relatively recent geological times, or it Figures 23-24. Parorthomus gen. n. socotranus sp. n., paratype, female genitalia, ventral view: 23 left ovipositor 24 female reproductive tract (spermathecal complex and ovipositor). Legend: as apical stylomere; bc bursa copulatrix; bs basal stylomere; co common oviduct; di diverticulum; es ensiform setae; ns nematiform setae; ov ovipositor; rc receptaculum of spermatheca; smc seminal canal of spermatheca; sg appended gland of spermatheca; spc spermathecal canal; vf valvifer. Scale bars = 0.2 mm (Fig. 23), 0.5 mm (Fig. 24). is a descendent of an ancestor and evolved in situ in the course and after the separation of the island. Notwithstanding, a few taxonomic and biogeographic facts are consistent with the hypothesis that the species is not a phyletically young descendant of continental populations.
Combinations of distinguishing features (see 'Diagnosis', Key to the genera of the "African Series" of Euchroina) clearly distinguish the new genus from the other related genera. However, some character states: 1/ mentum tooth bifid (Fig. 28); 2/ sides of pronotum straight or slightly convex posteriorly (Fig. 18); 3/ elytra with setiferous punctures in interval 3 (Fig. 18); 4/ intercoxal process of prothorax bordered; 5/ abdominal sternites V-VII with transverse sulci, complete and well-impressed; 6/ tarsomeres 1-5 of all legs glabrous dorsally; 7/ segment 5 of tarsomeres setose ventrally; 8/ aedeagus with sides nearly equally broadened in the distal half, with apical lamella wide, nearly rounded at tip (Fig. 20); 9/ appended spermathecal gland with diverticulum (Fig. 24), show that the new species may be related to two geographically "close" genera, the Afrotropical Abacillodes and the Mediterranean Orthomus. The two species from the first genus and the new species share characters 1-3 and 5-8. Some species from the second genus and Parorthomus gen. n. socotranus sp. n. divide states 1-7 and 9 between, while the last taxon and Orthomus velocissimus s.l. possess all the listed character states.
The median lobe with sides nearly equally broadened along the distal half and apical lamella wide, rounded or semi-rounded at tip in the new species looks alike the median lobe in the Afrotropical Abacillodes (see Straneo 1988: 482, Fig. 1d, 484, Fig.  2b), as well as those in some Mediterranean taxa of Orthomus (see Mateu 1957, laminas I-IV, Machado 1992: 262, Fig. 100, Wrase and Jeanne 2005: 894-896, Figs 1b, 2b, 3b, 4b, 5b, 6b, 7b, Pupier and Coulon 2013. In contrast, the median lobe of Parorthomus gen. n. socotranus sp. n. has the distal half considerably curved to the left in dorsal aspect. Hitherto all the representatives of Orthomus and Abacillodes have the median lobe of the aedeagus straight or nearly straight. Based upon the condition in the other continental, African and Eurasian euchroines (aedeagus of Abacillius and Trichopedius not yet known or described), we consider the bent aedeagus to be an apotypic character in Parorthomus gen. n. The median lobes in the "gracilipes" group of Nesorthomus, with N. gracillipes (Wollaston, 1854) and N. berrai Battoni, 1987, have also the above discussed character state, more pronounced in the former and less pronounced in the second species (see Sciaky 1988: Figs 1b, 2b, Machado 1992: 264, Fig. 101A, Donabauer 2008: 111, Figs 1b, 2b, Serrano et al. 2009. This case is an instance of convergency. Change in this state has taken place independently in Nesorthomus, since the other six species from the genus have a straight median lobe of the aedeagus. Besides, we infer that the presence of three to four discal setiferous punctures (by exception, two or five punctures on one elytron only) in the elytral interval 3 or stria 3, with the last puncture in posterior third of elytron, is another clear apotypic feature in the new taxon. This state occurs in no other species among the Old World Euchroina, except Parorthomus gen. n. socotranus sp. n. The most species of the subtribe have two discal punctures in the elytral interval 3, as the second one lies at the posterior third of elytron. The species of Abacillius have no discal punctures on the elytron.
Parorthomus gen. n. socotranus sp. n. has a unique combination of two apotypic characters, distal third of the aedeagus considerably curved to the left in dorsal aspect (i), and presence of 3-4 discal setiferous puncture in elytral stria 3/interval 3, with the last puncture situated in the posterior third of elytron (ii), which is indication for a long-time existing isolation and merit surely the erection of an own genus. The absence of close relative/s sharing together with the new species these two marked structural features exclude a close relationships and suggests that we deal with an ancient lineage which most probably arisen within the basic stem of the "African Series" of Euchroina (according to Will 2006) long time ago. As well, the lack of extant relatives, akin to the new species, in the Arabian Peninsula or somewhere else is a strong biogeographic argument, which certainly excludes geologically recent migration.
In spite of all, special character states and main ecologic preference in Parorthomus gen. n. socotranus sp. n. suggest that it is phylogenetically closer to Orthomus and Abacillodes than to any other genus. The exact phylogenetic position within the euchroines can be disclosed only after investigation of more taxa, especially from the Old World, including also genetic technics and providing cladistic analysis, this could probably also identify its sister taxon.  Straneo, 1962, holotype 27 Abacillodes jocquei Straneo, 1988, holotype 28 Parorthomus gen. n. socotranus sp. n., male paratype, "Fimihin". Scale bar = 0.5 mm.
The presence of large labial pits on the mentum is a trait in the new taxon that is worth noting. Each pit has a distinct, deep aperture, its diameter wider than the diameter of the labial pore, and both are situated more medially (Fig. 28). The distinct labial pits, destined to a particular function of use, seem to be a plesiotypic condition in Pterostichini (Bousquet 1999: 33), as well in the Nearctic euchroines (Frania and Ball 2007: 120). The species of Abacillius, Abacillodes and Orthomus possess no or small labial pits (Figs 25-27). In the second case, they have indistinct, shallow apertures, diameters similar to or smaller than the diameters of the labial pores, and both are situated more basally.

Remarks.
A striking characteristic in the type species of Abacillodes, A. jocquei, are the elytral intervals 2, 4, and 6, significantly wider than the adjacent uneven intervals. But, this character state is not present in A. malawianus, thus it can not be used as a generic distinguishing mark. Straneo, 1988 Abacillodes malawianus Straneo, 1988 Remarks. Straneo (1988: 483) stated the holotype is a male. Actually, it is a female specimen having protarsomeres not dilated and last abdominal sternite with four marginal setiferous punctures.