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DEUVE ET AL.: CARABID BEETLE FAUNA OF THE GAOLIGONG MOUNTAINS 433<br />
ranges of these species, three general range patterns are apparent. The first pattern (1) is shown by<br />
one species, Perileptus imaicus, with a geographical range that includes only a narrow swath along<br />
the southern base of the Himalayan Range from Himachal Pradesh, India in the west to the<br />
Gaoligong Shan region, where it reaches its eastern distributional limit. Among the zabrines,<br />
Amara elegantula Tschitschérine shares this same pattern but in higher elevation habitats<br />
(Kavanaugh et al. 2014). The second (2) pattern is shown by two species, Agonotrechus wuyipeng<br />
and Eocnides fragilis, both of which have a geographical range that extends from central or northern<br />
Sichuan, respectively, southwest to the northern half of the Gaoligong Shan region, where they<br />
reach their western distributional limit. Finally, the fourth species, Trechus indicus, shows the third<br />
pattern (3), which is a combination of the first two. The range of T.indicus extends from eastern<br />
Afghanistan eastward along the southern edge of Himalayan range and the Qinghai-Xizang<br />
(Tibetan) Plateau to northcentral Sichuan, with its southern limit in the mountains of the Gaoligong<br />
Shan region. Several zabrine species, including Amara sikkimensis Andrewes, A. chalciope<br />
(Bates), A.dissimilis Tschitschérine, A.latithorax Baliani, and A.birmana Baliani share this same<br />
pattern (Kavanaugh et al. 2014, Fig. 27 ), although with varied eastern and western extents. Deuve<br />
(1997, 2013b) recognized what he called subzones within both the southern Palearctic and northern<br />
Oriental Regions based mainly on his biogeographic analysis of the Carabus fauna of China<br />
and adjacent areas. Our general ranges patterns (1) to (3) correspond well to the southern parts of<br />
his Sichuano-Tibetan subzone (“Sous-zone Sichuano-Tibétaine”, subzone III in Deuve 2013b, Fig.<br />
13); and pattern (2) is similar to his Yunnan Plateau subregion (“Plateau du Yunnan”, subzone IIIa<br />
in the same figure).<br />
It is likely no coincidence that all four of the trechine species with known ranges extended<br />
beyond the Gaoligong Shan region have adults that are fully-winged. The ability to fly undoubtedly<br />
supports the maintenance of larger occupied ranges as well as greater potential for dispersal<br />
to new areas. within the trechine fauna of the study area, four additional species have fully-winged<br />
adults: Perileptus pusilloides, Agonotrechus fugongensis, A. xiaoheishan, and A. yunnanus.<br />
Although all of these species are currently known only from the study area, the discovery of one<br />
or more of them in additional, adjacent areas is possible or even likely in the future. However, it<br />
would be unexpected to discover populations of any of the remaining 21 species, all of which have<br />
flightless adults, outside the Gaoligong Shan region.<br />
with such a large proportion of the trechine fauna of the region known from nowhere else (25<br />
of 29 known species [86%] and four of eight genera [50%, plus one additional subgenus]), a look<br />
at the overall distributions of the genera or subgenera to which these trechines belong may provide<br />
a broader geographic context for understanding the composition of fauna. Perileptus is widely distributed<br />
in tropical to temperate portions of all continents in the Eastern Hemisphere and in the<br />
Caribbean portion of the Neotropical Region, probably introduced into the latter from Africa with<br />
commercial trade. Agonotrechus is restricted to the southern portion of the Sichuano-Tibetan subzone<br />
and the Subtropical China subzone (“Sous-zone de Chine subtropicale”, subzone V in Deuve<br />
2014b, Fig. 13) including Japan. Eocnides is restricted to the southcentral portion of the Sichuano-<br />
Tibetan subzone and Queinnectrechus to the Yunnan Plateau subregion of that subzone. Trechus,<br />
with its present taxonomic inclusiveness, is Holarctic in distribution, and species in the study area<br />
likely have their closest relatives to the north. Three of the remaining genera, Trechepaphiopsis,<br />
Epaphiotrechus, and Trechepaphiama, are members of the Epaphiopsis complex of genera, which<br />
mainly occupy the Subtropical China subzone. The phyletic affinity of Minutotrechus remains<br />
unclear based on morphological features. As noted above, it is likely related to either Uenoites, and<br />
therefore has a Sichuano-Tibetan affinity, or to Hubeitrechus,and hence has a Subtropical China<br />
affinity.