Nesaeoduvalius (gen. n.) vailatii sp. n., from Euboia island (Eastern Greece) (Coleoptera: Carabidae, Trechinae) (**)

Nesaeoduvalius (new genus) vailatii Casale & Giachino, new species (Coleoptera: Carabidae: Trechini) is described and illustrated from the Euboia (Evvia) island (eastern Greece). The new taxon belongs to the Duvalius “isotopic” trechine lineage, but is markedly characterized by the peculiar combination of several morphological features in the general shape, chaetotaxy of head, serrate lateral margins of elytra, and features of male genitalia (both in median lobe of aedeagus and endophallus). Its putative position amongst the trechines of the Duvalius phyletic lineage is discussed. Data on the distribution and habitat of this new, insular and unexpected taxon are also provided.


Introduction
The discoveries of new subterranean taxa in the Euro-Mediterranean area are fre quent, and their descriptions are a current, every-day taxonomical practice. Nevertheless, some of these taxa are particularly interesting, because their morphological features and geographical patterns of distribution per mit postulating the history and evolution of some lineages and better under standing the taxonomic position of previously described relatives (Casale et al. 2011(Casale et al. b, 2012. The diversity of subterranean genera of Trechini in Greece is low, when compared to the Dinaric chain and the Balkan peninsula as a whole, which are an impressive hotspot of subterranean biodiversity. Amongst Carabidae Trechini, as recently showed by Lohaj & Lakota (2010),  and Casale et al. (2012), ten genera of so-called "anisotopic" aphaenopsoid Trechini beetles of the former « série phylétique d'Aphaenops » sensu Jeannel (1922Jeannel ( , 1928Jeannel ( , 1930 and Casale & Laneyrie (1982) are currently known from the territory of the Dinaric range, from northern Croatia up to the north and central Albania in the south: Scotoplanetes Absolon, 1913, Adriaphaenops Noesske,1928, Dalmataphaenops Monguzzi, 1993, Al bano trechus Casale & Guéorguiev, 1994, Croatotrechus Ca sale & Jalzic, 1999, Minosaphaenops Quéinnec 2008, De rossiella Quéinnec, 2008, Jalzicaphaenops Lohaj & La kota, 2010, Acheroniotes Lohaj & Lakota, 2010, and Ve lebitaphaenops Casale & Jalzic, 2012. Besides these, several genera of different lineages are represented in the area: the "anisotopic" genus Neotrechus G. Müller, 1913and the "isotopic" genera Anophthalmus Sturm, 1844and Aphaenopsis G. Müller, 1913 in Dinarides, the isolated genus Pheggomisetes Knirsch, 1923 in the Balkan, while the genus Duvalius Delarouzée, 1859 (in the widest sense of authors) is widely spread everywhere.
On the contrary, in Greece, the genus Duvalius only is represented, which shows however an impressive specific diversity in all areas of the country and a wide adaptive radiation both in caves and in upper hypogean zone (Casale 2011a;Casale et al. 1996Casale et al. , 2013. Therefore, it is particularly interesting the discovery in the Euboia island (south-eastern Greece) of a blind, specialized trechine species, which is here attributed to a new genus for the peculiar combination of its morphological features.

Material and methods
The morphological features of the holotype of the new taxon were examined using Wild M5, Wild M3 and Olympus eISSN: 2284-4880 (online version) pISSN: 0429-288X (print version) ble with one, acute tooth at base; right mandible with a large sized, elongate retinaculum, with traces of two teeth. Submentum (Fig. 4) not transversally furrowed, with eight setae along the anterior margin.
Mentum free, not fused with the submentum along the suture, with a pair of long labial setae, without a pair of sensory foveae; mentum tooth wide, very prominent, widely emarginated at apex, largely bifid, with acute teeth; ligula elongate, not bifid at apex, with two long apical and several lateral setae; paraglossae very long and slender, pubescent on the inner margin. Penultimate palpomere of the labial palpi with three setae, two on the inner margin and one, sub-apical, on the outer one.
Pronotum transverse, markedly cordiform, with deep, wide marginal furrows and with lateral margins deeply sinuate anteriorly to the basal angles. Median furrow impressed, reaching the basal margin. Anterior angles obtusely rounded, basal angles acutely prominent outside. One pair of antero-lateral and one pair of baso-lateral setae present, the anterior pair at the anterior sixth of pronotum, the basal one moved anteriorly to the basal angles.
Elytra elongate, widened in the middle; humeral angles rounded but evident. Lateral margins deeply sinuate posteriorly to the humeral angles and finely but distinctly serrate in the basal two/thirds, from the humeral angles to the level of pores 5-6 of the lateral umbilicate series. Apical lobe, pre-apical sinuation and apical recurrent stria very reduced. Juxta-scutellar striola and inner striae 1-4 evident, the outer striae 5-8 vanished. Chaetotaxy: one pair of basal setiferous pores near scutellum; two discal setiferous pores and one preapical pore on interval 3 in each elytron; marginal umbilicate series of 8 punctures aggregated, i.e. punctures 1-8 (4 of the humeral group, 4 of the apical group) situated along the putative stria 8; latero-apical pore close to the apical stria; inner apical pore present, so that the so-called apical triangle (in the sense of Jeannel, 1928) is complete.
Abdominal sterna IV-VII with a pair of setae (male), longer on VII.
Legs long but robust; protibiae thickened, pubescent, without external furrow; two basal tarsomeres of male protarsi dilated and denticulate inwards. Male genitalia (Figs 3, 4): median lobe of aedeagus very elongate and slender, with apical lamina axe-shaped, hooked on both the dorsal and ventral sides. Endophallus without copulatory piece, but armed with a series of small, sclerotized spines. Parameres long and slender, each with 4 apical long setae.
Etymology. Named after the combination of the Latinized adjective Nesaeus (insular) (from the ancient Greek Nísaios), and Duvalius, to stress the relationships of this new taxon with the genus Duvalius in the widest sense, and the insular, isolated distribution of its type species. Gender is masculine.
SZ 60 stereo-microscopes. Male genitalia were dissected, cleaned and mounted in Canada balsam on transparent slide under the examined specimen. Drawings of aedeagus were obtained using Leitz Dialux transmitted-light microscopes with attached drawing tube.

Measurements:
TL: total body length (measured from the anterior margin of clypeus to the apex of elytra) L: overall length, from apex of mandibles to apex of elytra, measured along the suture PL/PW: ratio length of pronotum, as linear distance from the anterior to the basal margin, measured along the midline/maximum width of pronotum. EL/EW: ratio length of elytra/maximum width of elytra.

Diagnosis.
A genus of medium-sized trechine beetles with the character states of the "isotopic" lineage of Duvalius Delarouzée, 1859 in the widest sense of Jeannel (1928) and Casale & Laneyrie (1982), recently confirmed as monophyletic unit in the molecular phylogeny of Alpine trechines by Faille et al. (2013), but markedly characterized by the peculiar combination of the following apomorphic features: genae pubescent, tempora each with a long seta in the posterior half; elytra with lateral margins serrate from the humeral angles to the median group of the umbilicate series; apical stria almost vanished and apical carina very reduced; median lobe of aedeagus hooked on the dorsal and ventral sides, endophallus without copulatory piece but armed with small, dense spines in dorsal position.
Head glabrous with tempora pubescent, each with some short lateral erected setae and one additional long seta in the posterior half, shorter than the supraorbital setae (Figs 1, 3). Eyes absent. Frontal furrows very deep, complete.
Two supraorbital setae present on each side. Clypeus ( Fig. 1) with two pairs of setae. Labrum with a prominent median tooth on the anterior margin and three pairs of setae (Fig. 2).
Genae swollen, convergent and markedly narrowed to the neck. Labrum (Fig. 2) slightly emarginate at the anterior side, with obtuse, prominent tooth in the middle; mentum tooth (Fig. 4) wide, prominent, widely emar ginate at apex. Antennae elongate but robust, exceeding backwards the level of the fourth humeral setiferous puncture.
Pronotum markedly cordiform, transverse (ratio PL/ PW: 0.83), wider than head, widest at the anterior fifth; disc moderately convex; lateral sides curved in front, shortly and deeply sinuate towards hind angles, which are acute and prominent outside; marginal furrows wide and deep; anterior margin gently arcuate, front angles effaced; base curved in the centre; basal foveae very deep.
Legs long but robust; protibiae dilated, pubescent on the anterior side, not grooved on the external side; protarsi in male each with two basal tarsomeres dilated and denticulate inwards. Tarsal claws long and curved.
Male genitalia as in Figs 7-9. Median lobe of aedeagus long (1.26 mm), slender, in lateral aspect straight, sinuate on the dorsal side, hooked on both the dorsal and ventral sides at apex. Basal bulb and basal sagittal carina not distinguishable in the holotype (immature, damaged male genitalia). Parameres long and slender, each with 4 apical long setae. Endophallus without copulatory piece, but armed with a series of small, dense, sclerotized spines in dorsal-apical position.
Specific epithet. We wish to dedicate this species to our good friend Dante Vailati (Brescia, Italy), excellent speleologist and explorer of subterranean environments in Greece and everywhere, who sampled the only specimen known so far of this interesting new taxon.
Geographical distribution and habitat. The only male specimen of the new species here described was sampled on the Eastern slope of Mt. Dirfis, near the place named Lirí, along the road from S. Dirfios to A. Iríni (Euboia Island, Central-Eastern Greece) at 1080 m of altitude, un- 1 mm der stones near the snow (Figs 10-11). Like many Trechini species of the Euro-Mediter ranean area, it is an upper hypogean species in the alpine zone. This specimen was sampled during the first survey to the island by P.M. Giachino and D. Vailati in 1998. Later, in the following years (1999, 2000, 2002, 2003, 2004, 2005, 2006, 2010, 2011, 2014, they investigated with care the same biotope, and caves and subterranean superficial habitat (M.S.S.) of the same and close mountains, and in different environmental conditions (presence or absence of snow, more or less dried substrate), but were not able to collect any further specimen of this species.
Morphological features (body flattened, eyes fully absent, serrate lateral margins of elytra, legs robust, protibiae without external furrows), its extreme rarity in investigated environments, and its absence in caves of the island, indicate a high grade of specialization of this species to life in fis-sures of superficial subterranean compartments (Giachino & Vailati 2010), where only in particular environmental (climatic) conditions it is likely possible to meet it.

Relationships
Nesaeduvalius (new genus) vailatii sp. nov. is particularly interesting for both taxonom ical and biogeographical aspects. Its peculiar, apomorphic features, and its isolation on an island merit the institution of a new genus. We have to stress that the Euboia (Evvia) island, despite its position relatively close to the eastern coast of continental Greece, hosts some peculiar, very isolated subterranean genera and species which don't show apparent relationships with any continental taxon. As examples, amongst carabids, we should cite the exceptional, big-sized spho- drine Hystrichosphodrus vailatii Casale & Giachino, 2004 and, amongst cholevids Leptodirini, Henrotiella eubeensis Perreau, 1999, from the cave of Agia Triada near Káristos (South Euboia). The genus Duvalius as a whole in the Balkan peninsula includes, in the current taxonomic treatment, a high number of species (Moravec et al. 2003;Lorenz 2005). The validity of some so-called "genera" recently described has been widely debated by Lohaj & Mleinek (2013) and Ćurčić et al. (2013). Personally, we are inclined to preserve this genus as a monophyletic unit , as perfectly established by Jeannel (1928) and confirmed by molecular data (Faille et al. 2013), despite the heterogeneity of morphological features of species ascribed to the genus (epigean or hypogean, pigmented or depigmented, winged or micropterous, oculate, microphthalmous or blind). As for aphaenopsoid Dinaric and Balkan genera of the "anisotopic" lineages, in which the diagnostic features are mostly based on plesiomorphic characters, or derived by different grades of specialization, relationships amongst the different species of Duvalius in this area remain unclear, and hidden by features corresponding to different grades of evolution, radiation, and adaptation to the subterranean way of life (Quéinnec 2008).
We proposed however a generic rank for this new taxon of the Euboia island because the combination of the unique, peculiar characters described above, and for its geographical isolation by the closest continental relatives of the genus Duvalius (in the widest sense), which are very numerous: 47 species described so far in Greece (Crete island included), and many others not yet described (Casale, Giachino & Vailati, in progress).
Conversely, in Nesaeoduvalius some other informative apomorphic features are absolutely unique, i.e. the general shape of body, the occurrence of an additional temporal seta, elytra with serrate humeral and lateral margins and vanished both apical lobe and apical stria. In particular, serrate humeral angles and lateral margins of elytra are a character present in endogean taxa of Trechini of the Cimmerites lineage and, amongst other tribes of carabid beetles, in Clivinini Reicheiina and Bembidiini Anillina, but absent in all genera of the Duvalius phyletic lineage. These characters, associated with its insular, isolate geographical distribution (no Duvalius species is known from Euboia island), support the institution of Nesaeduvalius as a new genus distinct from all described taxa currently attributed to Duvalius and related "isotopic" genera.
In this case, we could hypothesize an ancient origin of Nesaeoduvalius vailatii as an extant taxon derived by a common ancestor (with Biharotrechus, Hungarotrechus and Platyduvalius), now extinct, of a phyletic lineage of continental (Carpathian and Balkan) Duvalius lineage in isolated massifs of the eastern Aegean coast.
Molecular phylogenies, like those recently performed in different areas (see Faille et al. 2010Faille et al. , 2013 and further discoveries of new taxa should provide in the future new and interesting information on these lineages of Dinaric and Balkan subterranean carabids, speciated and radiated in mountains and islands of a very complex geographical area.