Figure 1.
The showing main parts of the Himalayan Tibetan mountain system, main ecological divisions of the High Himalayan mountain arc, and distribution of five species groups of primary wingless Pterostichus ground beetles (areas framed by coloured lines): red, Ethira clade, a Himalayan endemic with an extensive distributional gap across the Kumaon Himalaya; green, Pterostichus (unplaced) leviculus group, a Northwest Himalayan endemic; light blue, subgenus Sinosteropus, with more than 30 species and subspecies on the eastern Tibetan Plateau, with a single species in the Western Himalaya, and with two species in a restricted area of the eastern Tibetan Himalaya; dark blue, subgenus Pseudohaptoderus, endemic to the eastern Tibetan Plateau; yellow, Pterostichus (unplaced) migliaccioi group, an East Himalayan endemic (distributional data after [14,30,60,61,62 and unpublished data]). General habitus and median lobe of the male genital of typical representatives of the relevant species groups are figured above. Abbreviations: al = apical lamella; os = ostium of aedeagal median lobe.
Figure 2.
Results of the combined 28S + 18S rDNA + COI mtDNA data analysis.
The 50% majority rule consensus tree of the Bayesian analysis. of the combined 28S + 18S rDNA + COI mtDNA sequence data set of Pterostichini (Ethira clade, 10 additional Pterostichus subgeneric taxa, and 5 putatively related genera) using Abax parallelepipedus as outgroup taxon. Numbers on or beside branches are Posterior Probabilities ≥90%/Bootstrap support values. Numbers in brackets refer to the internal specimen identification code and are added in cases when more than one specimen of a morphological species was added to the analysis. The deflected median lobe of male genital of the “modern Pterostichus” with ostium turned to the left is exemplary figured beside the relevant subtree; the presumably plesiomorphic shape with ostium in dorsal position is figured above. Beside the Ethira clade four of its basal lineages and six terminal species groups of clade A are designated and are discussed in the text. Abbreviations: balach. = balachowskyi; imm. = immarginatus; nepal. = nepalensis; P. = Pterostichus; Ps. = Pseudethira.
Figure 3.
Comparison of phylogeny and distribution of the basal lineages of the Ethira clade.
Below: The 50% majority rule consensus tree of the Bayesian analysis of the Rib + COI data set (see Figure 2). The four basal lineages are designated by different colors. The same colors are used to frame the distributional areas of each of the lineages in the map above the tree. The black arrows in the tree mark six events of wing reduction which necessitate from the assumption that the separated distributional area of the harmandi group originated from migration of a winged ancestor. The white arrow points to the single event of wing reduction that follows from the assumption of a Tibetan origin of the Ethira clade. For details see chapter discussion.
Figure 4.
Comparison of phylogeny and distribution of the terminal lineages of sub-clade A of the Ethira clade as followed from the Bayesian analyses of the Rib + COI data set (see Figure 2).
Figure 5.
Comparison of mitochondrial phylogeny and geographical distribution of the balachowskyi species group.
Below: The 50% majority rule consensus tree of the Bayesian analysis of the COI sequences using Pterostichus immarginatus and P. matsumurai as outgroups. Numbers on or beside branches are Posterior Probabilities ≥90%/Bootstrap support values. Numbers in brackets refer to the internal specimen identification code. Private haplotypes are numbered in colored boxes. The collecting localities of these haplotypes are shown in the map above.