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434 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES<br />

Series 4, Volume 63, No. 12<br />

Kavanaugh et al. (2014) suggested that the Gaoligong Shan region may have been an area of<br />

differentiation, speciation and origin of montane elements from which, rather than to which, at least<br />

some of the species that now range more broadly subsequently spread. This hypothesis was based<br />

on meager evidence provided by the zabrine fauna of the Gaoligong Shan, but also on geologic evidence.<br />

The Hengduan Mountains date their origins to the late Mesozoic, whereas the uplift of the<br />

Himalayan Ranges and Qinghai-Xizang Plateau began later, in the early Cenozoic (Chaplin 2005).<br />

Hence the biota of the Gaoligong Shan region probably predates that of these other areas as well.<br />

The trechine fauna of the region supports this hypothesis even more strongly. The occurrence of<br />

four precinctive genera and one apparently precinctive subgenus of small, flightless beetles, two of<br />

which are represented by two and seven species, respectively, suggests differentiation and diversification<br />

within the region. The occurrence of another nine likely precinctive species in other genera<br />

suggests speciation in situ. Finally, even the four species with ranges extended beyond the<br />

Gaoligong Shan region all have those ranges either centered on the region of anchored there. As<br />

with the zabrines, a better understanding of phylogenetic relationships among the Gaoligong Shan,<br />

Eurasian and Oriental trechinespecies and genera is required in order to test this hypothesis, and<br />

such an analysis has not yet been undertaken.<br />

REGIONAL GEOGRAPHICAL AND ALTITUDINAL DISTRIBUTION PATTERNS.— within the<br />

Gaoligong Shan study area, most of the trechine species represented are narrowly distributed, both<br />

geographically and altitudinally. This is not surprising given the high percentage (72%) of species<br />

with flightless adults and their preferences for moist, undisturbed habitats. Such areas are restricted<br />

within the region mainly to remaining forested areas at low to middle elevations and alpine<br />

meadows, moist tundra, stable talus slopes, stream edges and bamboo and Rhododendron thickets<br />

at higher elevations. Many such areas in the region are separated from each other by deep valleys<br />

and, increasingly, by disturbance associated with agriculture and human habitation. while human<br />

disturbance is relatively recent, topographic diversity of the region has been developing since the<br />

Miocene (Chaplin 2005).<br />

The chart in Fig. 46 summarizes the recorded regional distributions of the species with respect<br />

to our project-designated Core Areas (see Fig. 3); and the recorded altitudinal ranges for each<br />

species are shown in Fig. 47. These charts clearly demonstrate the relatively narrow geographical<br />

and altitudinal ranges of most of the trechine species occurring in the region. This is especially<br />

apparent from comparisons with the ranges of zabrine species in the same area (see Kavanaugh et<br />

al. 2014, Figs. 28 and 29). Among the 13 zabrine species in the fauna, one is recorded from all 7<br />

Core Areas, one from 6 Core Areas (all except 5) and three from four or five Core areas; and most<br />

of these species are likely to occur in all seven Core Areas. In contrast, only one species, Trechus<br />

indicus, is recorded from as many as four Core Areas (1, 3, 6 and 7) and also has a relatively broad<br />

known altitudinal range (1230 to 2486 m). Given that it is recorded from the northernmost and<br />

southernmost core areas, as well as from both eastern and western slopes of the Gaoligong Shan,<br />

it is likely to be found in additional, if not all, core areas with further sampling. Only one additional<br />

species, Perileptus imaicus, is recorded from as many as three Core Areas (2, 6 and 7) and also has<br />

a broad known altitudinal range (from 680 to 2030 m). Like T.indicus, it is recorded from both<br />

northernmost and southernmost core areas and from both slopes of the mountain range, so it is likely<br />

to be more widespread in the region than is presently known, at low to middle elevations. Five<br />

additional species are recorded from as many as two core areas: Perileptus pusilloides (Core Areas<br />

6 and 7, known altitudinal range from 680 to 1105 m); Agonotrechus fugongensis (Core Areas 2<br />

and 3, known altitudinal range from 2300 to 2530 m); A.wuyipeng (core areas 2 and 3, known altitudinal<br />

range from 2687 to 2770 m); Queinnectrechus balli (Core Areas 1 and 2, known altitudinal<br />

range from 3300 to 3750 m); and Trechepaphiopsis unisetosa (Core Areas 4 and 5, known altitu-

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