Aphaenops

nodes showing incongruence (Figure 3) (see detailed comments below). In all cases the monophyly of Trechina was strongly supported, with three main clades that were recovered in all analyses. Clade 1 (Figures 3a-c and 4) included most of the anisotopic species: the Pyrenean hypogean lineage (clade 1.1, Figures 3a and 4), the genusEpaphius and relatives (Blemusand some species ofTrechus, clade 1.2) and a large group including most species of the genusTrechusplus a number of subterranean gen- eranestedwithinit(clade1.3).Theseincludedallthe Alpine and Dinaric subterranean anisotopic genera with the only exception ofDoderotrechus,plussome other genera from nearby areas (Speotrechus,Jeannelius, Typhlotrechus,SardaphaenopsandParaphaenops, Figures 1 and 3). Figure 3Phylogram of Alpine Trechina obtained in RAxML using the combined data matrix.Number in nodes, Bayesian posterior probability obtained in the two alternative states of MrBayes (see main text)⁄ML bootstrap obtained in RAxML. Subterranean genera and lineages are indicated by asterisks (see Additional file 3: Table S1). -: unresolved or poorly supported node (bt≤70%, Bpp≤70); x: contradictory node. 3a: clade 1; 3b: clade 1.3.2.2; 3c: clade 1.3.2.1; 3d: clades 2 and 3. Failleet al. BMC Evolutionary Biology2013,13:248Page 5 of 16 The clade 1.3.2.1 was well-supported (bt = 80; pp = 0.99/1) and included the subterranean generaOrotrechus,Bol- doriella,Lessinodytes,Allegrettia,Italaphaenopsand thetwoDinaricgeneraAdriaphaenopsandNeotrechus (Figure 3c). No epigean species were nested within this clade. The monophyly of both generaOrotrechusand Boldoriellawas confirmed, whereas the position ofLes- sinodytes,AllegrettiaandItalaphaenops,aswellasthat of the Dinaric species, was uncertain. A sister relationship was suggested in the ML analysis betweenAllegrettiaand Italaphaenops, but without support and not recovered in the Bayesian analyses. The sampled species of the genusOrotrechuswere re- covered as monophyletic, and divided in two groups of species: the first, poorly supported (pp = 0.54/0.97; bt = 66) includingO. vicentinus,O. fabianii,O. giordaniiandO. montellensis, and the second, well-supported (pp = 1; bt = 99), with all other sampled species (Figure 3c). The two aphaenopsian speciesO. jamaeandO. messai,cur- rently attributed to different species groups [20], were in- cluded in the same clade. The genusBoldoriellawas found monophyletic with low support (pp = 0.86/0.92; bt = 54), but the separation of two subgenera withinBoldoriella (BoldoriellaandInsubrites[26]) was not supported by our analyses, asB.(Insubrites)focarileiwas closer toB. (Boldoriella)carminatiithan toB.(Boldoriella)humeralis. The clade 1.3.2.2 was also well-supported (pp = 0.97/ 0.99; bt = 81), and included most of the species of Trechussensu stricto (including the type species of the genus). Seven sub-clades were recovered with general good support, together with some isolated species of uncertain position (T. karadenizusandT. lepontinus, Figure 3b): (1) a Pyrenean-Cantabrian clade (pp = 1; bt = 77), which included 16 species, some of them hypogean (T. beusti ,T. navaricus,T. bouilloni,T. escaleraeand Apoduvalius alberichae); (2) the group ofT. fair- maireiand closely related species; (3) two species of the Maritime and Ligurian Alps,T. liguricusandT. putzeysi, as sister (with low support) to a species from the Eastern Alps,T. gracilitarsis;(4)thetwospeciesoftheT. strigipen- nisspecies group as currently defined [14,39] (T. pochoni andT. piazzolii); (5) two species ofApoduvalius,withun- certain affinities; (6) theTrechus fulvusgroup, here repre- sented by the speciesT. fulvusandT. gigoni(the latter described asAntoinella[40]); and (7) a clade containingT. quadristriatusandT.obtusus, plus some Alpine and Pyren- ean endemics as well as species from Turkey. Clade 2, the isotopic Trechina The isotopic Trechina in the traditional sense [2,30] were confirmed as monophyletic with strong support (pp = 1; bt = 100), alt