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Cover image 

Species in the southern Black Mountains that occur in both Mohave and Sonoran deserts 

(Fig. 4) 

(See Butterneck & Daniel, Vegetation and Flora of the Southern Black Mountains 

of W est-Central Arizona) 

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Published by the California Academy of Sciences 
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PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES 


Series 4, Volume 63, No. 8, pp. 269-320, 12 fig. pis., 3 tables 


October 14, 2016 


Vegetation and Flora of the Southern Black Mountains 

ofWest-Central Arizona 


Mary L. Butterwick and Thomas F. Daniel 

D epartment of B otany, C alifornia Academy of Sciences, 55 Music Concourse Drive, Golden Gate Park, 
San Francisco, CA 94118 U.S.A.; Email: m Ibutterwick@ gm ail.com 

This study documents the botanical resources of the southern Black Mountains 
along the western edge of Mohave County, Arizona. The flora area covers approxi¬ 
mately 883 sq. km (341 sq. mi) in the southern third of the range from A rizona H w y. 

68 to Sacramento Wash along U.S. Interstate 40. Eight plant communities are iden¬ 
tified and described and an annotated catalog of the vascular plants of the southern 
Black Mountains is provided. Seventy-two families, 276 genera, 471 species, and 
seven additional infraspecific taxa are documented from this region of the Mohave 
Desert (478 total taxa at rank of species or below). Plant families with the largest 
presence in the flora area include Asteraceae (89 species), Poaceae (53), Boragi- 
naceae (35),Fabaceae (24), Brassicaceae (19), Cactaceae (16), and Polygonaceae (16). 

Of the 205 taxa of ephemeral herbs in the flora area, 182 (89% ) a re win ter ephemer- 
als. Thirty-eight taxa (8% of the flora) are not native to the flora of A rizona. Twelve 
species of biogeographic significance are noted, including Moncit'della eplingii , which 
is endemic to the Black Mountains. 

Keywords: Mohave Desert, Arizona, plant communities, floristics, ethnobotany 

The Black Mountains are an extensive desertic mountain range along the western edge of 
Mohave County, Arizona, paralleling the Colorado River for approximately 156 km (97 mi), from 
Lake Mead southward to Sacramento Wash. The range was previously called the Ute Mountains, 
in reference to the Ute people who lived in the vicinity (Barnes 1960). The names Ute Mountains 
or Black Ute Mountains occur on the labels of several early plant collections from the area. This 
flora covers the southern third of the range from Union Pass along Arizona Hwy. 68 to Sacramen¬ 
to Wash along U.S. Interstate 40. This portion of the range is approximately 24 km (15 mi) west of 
Kingman and 13 km (8 mi) east of the Colorado River (Fig. 1). The flora area comprises approxi¬ 
mately 883 sq. km (341 sq. mi). 

The flora area (Fig. 1) is bounded on the north by Arizona Hwy. 68 through Union Pass, and 
on the northeast by the main mass of the mountains, generally along the 1036 m (3400 ft) to 853 
m (2800 ft) elevation contours from north to south. South of Mohave Co. 10 (Oatman Road), the 
flora area extends from Meadow Creek southeastward down Sacramento Valley to Sacramento 
Wash, and follows the wash as it curves to the west near its junction with Franconia Wash. From 
there the southwest boundary generally follows the 305 m (1000 ft) elevation contour to Mohave 
Co. 10 (Topock—Oatman road), then continues northward along the eastern edge of Mohave Val¬ 
ley on the west side of McHeffy Butte and Boundary Cone to Mohave Co. 153 (Boundary Cone 
Road). From there, the boundary continues northward to encompass the hills on the west side of 
the Black Mountains generally following the 610 m (2000 ft) to 670 m (2200 ft) elevation contours 
from south to north, and reconnects with Arizona Hwy. 68 in Union Pass. 

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Figure 1 . Satellite image of the southern Black Mountains from Google Earth showing boundary (red) of floristic 
study area, and collecting locales (green) of the authors. The Colorado River, which forms the border between Arizona and 
California, is at left, and the Sacramento Valley is on the right. Nearby towns outside of the study area are indicated. With¬ 
in the study area, Oatrnan (orange diamond) and Black Mesa are shown. Major roads in the vicinity are Interstate Hwy. 40 
(=Arizona Hwy. 95) in purple, Arizona Hwy. 68 (traversing the Black Mountains through Union Pass) in yellow, Mohave 
Co. 10 (=old U.S. Hwy. 66 or “Oatrnan Road” northward/eastward from Oatman and traversing the eastern Black Moun¬ 
tains through Sitgreaves Pass, and “Oatman Topock Hwy.” southward/westward from Oatman to the Colorado River) in 
blue, and Mohave Co. 153 in pink. Inset map shows state of Arizona with Mohave County shaded in green, and the loca¬ 
tion of the satellite image. 


BUTTERWICK & DANIEL: FLORA OF SOUTHERN BLACK MTS., ARIZONA 


271 


Most of the flora area is public land administered by the U.S. Department of Interior’s Bureau 
of Land Management (BLM), and includes two wilderness areas: Mount Nutt Wilderness, a 112- 
sq. km (27,660-acre) area north of Mohave Co. 10 through Sitgreaves Pass, and Warm Springs 
Wilderness, a 455-sq. km (112,400-acre) area at the southern end of the Black Mountains (U.S. 
Department of Interior 2016a, 2016b). Although land ownership is primarily BLM, there are some 
private and state of Arizona holdings within the flora area, particularly in the town of Oatman and 
along Mohave Co. 10 (Oatman Road) in Sitgreaves Pass. The Black Mountains are known for their 
mineral resources. Numerous mines throughout the Oatman mining district produced significant 
quantities of gold (2.2 million troy oz) and silver (0.8 million troy oz) between 1897 and 1942 
(DeWitt et al. 1986). 

The present study was undertaken to document the botanical resources of the southern Black 
Mountains. Based on Moore and Cole (2004), this is the first published flora for a region within 
the Mohave Desert of Arizona. Certain plant communities such as Great Basin Conifer Woodland 
and both Desert Oasis Woodland and Streamside Marsh at perennial springs are very restricted in 
the Black Mountains, and thus are particularly vulnerable to increased warming and drought antic¬ 
ipated from climate change in the region (Garfm et al. 2014). Our results can serve as a basis for 
examination of future changes in the flora of this range and other areas of Mohave Desert in south¬ 
ern Nevada and southeastern California. 

Topography and Geology 

The Black Mountains are located in the Basin and Range Physiographic Province (Wilson 
1962) and form a north-northwest-trending mountain range with wide valleys on each side—Sacra¬ 
mento Valley to the east and Mohave Valley to the west. The range is known for its extremely 
rugged terrain. The northern portion of the flora area has a total relief of about 914 m (3000 ft). 
The highest points in the flora area are an unnamed peak (“Benchmark Nutt”) at 1590 m (5216 ft) 
and Mt. Nutt at 1543 m (5062 ft). These peaks are surrounded by precipitous cliffs, small mesas, 
steep and rocky slopes, and several steep-walled canyons such as Grapevine Canyon, Cottonwood 
Canyon, and several unnamed canyons (Gray et al. 1990b), An area that is underlain by Protero¬ 
zoic granite, schist, and gneiss is exposed near Arizona Hwy. 68 between Thumb Butte and Secret 
Pass Arch (Nations and Stump 1996). However, most of the remainder of the northern portion of 
the flora area is overlain by a variety of Tertiary volcanic rocks including rhyolite, andesite, dacite, 
basalt, and tuff (Gray et al. 1990b). In fact, most of the Black Mountains, including our flora area, 
are volcanic in nature and in this respect differ from surrounding mountains and basins in Arizona, 
California, and Nevada—in effect forming a geologic/edaphic “sky island” in the Mohave Desert 
of western Arizona (Elvin et al. 2013). West of the main mass of the mountains is a complex of vol¬ 
canic and granitic hills in the vicinity of Pheland Butte, Cathedral Rock, Hardy Mountain, Finger 
Butte, and Thumb Butte (Nations and Stump 1996; U.S. Department of Agriculture 2016). 

Meadow Creek, a major east-west-trending tributary of Sacramento Wash, forms a prominent 
divide in the southern Black Mountains. Mohave Co. 10 (Oatman Road) east of Sitgreaves Pass 
generally follows Meadow Creek. The southern portion of the flora area, south of Meadow Creek, 
is dominated by Black Mesa (Fig. 2), a northwest-trending plateau of Tertiary basalt (Gray et al. 
1990a). The summit of Black Mesa ranges in elevation from approximately 975 m (3200 ft) to 1158 
m (3800 ft). Topographic relief is abrupt particularly along the east front of the mesa with drops in 
elevation of 305 m (1000 ft) or more to the surrounding pediment, alluvial fan, and piedmont sur¬ 
faces that slope gradually across the Sacramento Valley toward Sacramento Wash. The mesa is 
deeply incised by Warm Springs Creek and several unnamed tributaries (Gray et al. 1990a), and it 


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¥ ¥ ¥ 


Figure 2. Southern Black Mountains and plants communities-1. Panoramic view of western escarpment from Nee¬ 
dles, California showing (arrows, from right): Black Mesa, Boundary Cone, and Mt. Nutt area (top). Mixed Desert Scrub 
on Black Mesa (southern portion of the flora area) with Bromus rubens (abundant ground cover), Cylindropuntia spp., 
Ephedra aspera, Nolina bigelovii, and Yucca schidigera (center). Mixed Desert Scrub at Secret Pass (northern portion of 
the flora area) with Juniperus califomica (bottom). Photos by the authors. 


BUTTERWICK & DANIEL: FLORA OF SOUTHERN BLACK MTS., ARIZONA 


273 


is surrounded by a variety of Tertiary volcanic rocks (Gray et al. 1990a; Nations and Stump 1996). 
The topography is more complex on the west side of Black Mesa where it consists of lower-eleva¬ 
tion buttes, ridges, and rocky hills in the vicinity of McHeffy Butte, Wrigley Peak, and Boundary 
Cone, a prominent landmark in the area (Fig. 2). 

Perennial springs occur occasionally throughout the flora area on slopes (e.g., Shaffer, Fig, and 
Cave springs), in narrow, boulder-strewn canyons (e.g., Cottonwood, Grapevine Springs, and Twin 
Springs canyons), and along a few of the major tributaries including Meadow Creek, Warm Springs 
Creek, and Silver Creek Wash. 


Climate 

Climatic patterns in Arizona are affected by elevational gradients and by large-scale meteoro¬ 
logical systems that affect weather in western North America. As a result of the latter, a biseason- 
al pattern characterizes precipitation in Arizona (Sellers and Hill 1974). Winter rains generally 
occur from October to April when the westerlies move moist Pacific air masses eastward. These 
rains take the form of widespread gentle showers that may last for several days. Summer monsoon 
rains, concentrated from July to September, are created when moist air masses from the Gulf of 
California and the Gulf of Mexico move northward into the state. Rapid cooling and condensation 
occur as moist superheated air rises over mountainous terrain. Summer rains tend to form as local¬ 
ized heavy thunderstorms of short duration. 

The character of the biseasonal rainfall pattern varies across the state. In general, as one moves 
westward, a decreasing proportion of the annual precipitation occurs during the summer. In south¬ 
eastern Arizona, dependable summer rains contribute up to 70% of the annual total (Hastings and 
Turner 1965). However, descriptions of mapped ecological sites indicate that summer rains in the 
flora area account for only 35% of the annual precipitation (U.S. Department of Agriculture 2016). 
Thus, most of the annual precipitation (ca. 65%) is in the form of whiter rains. 

There are no site-specific climatic data for the Black Mountains. The nearest climate stations 
are in Bullhead City to the west, which has hotter and drier conditions, and in Kingman to the east, 
which has cooler and wetter conditions. However, the average annual precipitation in the portion 
of the Desert Region of the Basin and Range Province, which includes our flora area, ranges from 
12.7 cm (5 inches) to 25.2 cm (10 inches) (Sellers and Hill 1974). Increases in elevation generally 
correlate with greater average amounts of precipitation. This general pattern is reflected in the eco¬ 
logical site descriptions for the flora area with the lowest elevation sites being in the 7.5-15 cm (3- 
6 inch) precipitation zone, the mid-elevation sites being in the 15-23 cm (6-9 inch) precipitation 
zone, and the highest-elevation sites being in the 25-33 cm (10-13 inch) precipitation zone (U.S. 
Department of Agriculture 2016). 

According to Sellers and Hill (1974), average daily temperatures for the region that includes 
our flora area range from 7° to 10° C (45° to 50° F) during January and from 29° to 32° C (85° to 
90° F) during July. Like precipitation, temperatures also follow elevational gradients. Thus, high¬ 
er areas tend to be cooler and more mesic. Average daily temperatures for January and July are 
10.5° and 35° C (51° and 95° F) in the 7.5-15 cm (3-6 inch) precipitation zone, 7° and 30.5° C 
(45° and 87° F) in the 15-23 cm (6-9 inch) precipitation zone and 6° and 28° C (43° and 82° F) in 
the 25-33 cm (10-13 inch) precipitation zone (U.S. Department of Agriculture 2016). 


274 


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Series 4, Volume 63, No. 8 


Early Inhabitants and Ethnobotany 

Excavations at Bighorn Cave, a large and deeply stratified site in the southern Black Moun¬ 
tains, documented an occupational sequence extending back to 1800 BC (Geib and Keller 2002). 
Bighorn Cave served as a seasonal foraging basecamp with diverse artifact and native plant assem¬ 
blages. Such base camps tend to be situated near springs or other perennial water sources (Fryman 
et al. 1977). A scries of perennial springs currently exists near the Bighorn Cave site. According to 
Geib and Keller (2002), native plants (with names in parentheses corresponding to species we doc¬ 
umented from the flora area) recovered from Bighorn Cave include: agave ( Agave mckeiveyana ), 
beargrass ( Nolina b igelovii), Cactus ( Opuntia basilaris , O. chlorotica , O. polyacantha, O. xcur- 
vospina ), catclaw ( Senegalia greggii), creosotebush ( Larrea tridentata), gourd {C ucurbita palma- 
ta), grass (38 native species), hackberry ( Celtis reticulata ), juniper (Ju niperus californica\ milk¬ 
weed (Asclepias albicans, A. erosa,A. nyctaginifolia, A. subulata ), lUUStard (D escurainea pinna- 
ta ), Oak (Q ue reus turbinella ), pine ( Pinus monophylla'), reed (A hragmites australis ), SCrewbean 
mesquite (p rosopis pubescens\ and yucca (Yucca sc hi dig era , Y. baccata). Several plant taxa 
reported from Bighorn Cave either as pollen or plant fragments have not been otherwise collected 
in the flora area or elsewhere in the Black Mountains, based on our collections and on occurrence 
data in SEINet (2016). These taxa include Acer negun do, A Inns Sp Asclepias latifolia, Canotia 
holacantha, C ucurbita foetidissim a, Jugtans major , Nolina microcarpa, and Sarcobatus sp. (Geib 

and Keller 2002). 

During the late and terminal Archaic period (ca. 1200^400 BC and 200 BC-100 AD, respec¬ 
tively) Bighorn Cave was used as a hunting camp, primarily for deer and bighorn sheep, although 
plants were exploited as well. As farming communities became established along the Colorado 
River (perhaps during 200 BC-100 AD), the traditional hunting/foraging settlement-subsistence 
practices gave rise to a “valley centric” settlement-subsistence strategy, which became that of the 
Mohave people here (Kroeber 1925). Later, during the Formative period (550-1200 AD), plant har¬ 
vesting and processing became significant. By 1600 AD, Bighorn Cave appears to have been 
almost abandoned as an important foraging basecamp (Geib and Keller 2002). 

During early historic times, the Black Mountains were within the ranges of two peoples, the 
Upland Yuman group (or Hualapai people) on the east side of the mountains and the River Yuman 
group (or Mohave people) on the west side (Stone 1987; McGuire 1983). The Hualapai were able 
to successfully exploit wild resources by using an organizational flexibility that could effectively 
respond to the patchy distribution of resources and to the seasonal and year-to-year variability and 
unpredictability of the natural environment (Kniffen 1935; Manners 1974). As a result, camps 
could change in membership composition, periodically disperse into separate families, or aggre¬ 
gate into larger groups as the availability of resources dictated. Their staples of existence consist¬ 
ed of agave leaves, amaranth, Mentzelia seeds, grass seeds, prickly pear fruits, mesquite beans, 
banana yucca fruits, piny on nuts, and juniper berries, all of which we documented from the flora 
area. While most of these wild food plants produce reliable yields from year to year, the various 
species occur in different areas and became available for harvesting at different times of the year. 
For instance, grass seeds were obtained in the valleys during the summer months. Juniper berries 
and pinyon nuts are less dependable. Good yields of juniper berries are produced every two to five 
years (Jeter 1977). The Hualapai harvested the pinyon nuts every four years (Stone 1987). Hunts 
to pursue deer and bighorn sheep took place in desert ranges, such as the Black Mountains, during 
winter months (Castetter and Bell 1951). Additional animals hunted for food included antelope and 
rabbits (McKennan 1935). Portions of the wild harvest were stored at winter base camps to survive 
the lean months during winter and early spring (Stone 1987). The excavation of Bighorn Cave doc¬ 
umented several storage pits, some of which were lined with juniper bark (Geib and Keller 2002). 


BUTTERWICK & DANIEL: FLORA OF SOUTHERN BLACK MTS., ARIZONA 


275 


The Yurnan peoples used many additional plant resources, not only for food but also for fiber, 
construction, implements, basketry, water proofing, adhesives, dyes, medicine and fuel, among 
other uses (Stone 1987). We documented the following plant species, which serve as sources for 
many of these uses, in the flora area: Allium nevadense, Baccharis salicrfolia, Celtis reticulata, 
Chenopodium berlandieri, Chenopodium fremontii, Chilopsis linearis, Fouquieria splendens, 
Fraxinus anomala, Larrea tridentata, M entzelia albicaulis, Physalis c ra ssifo lia, Pluchea sericea, 
Poputus fremontii, Rhus aromatica, Saiix exigua, Salix gooddingii, Salvia columbariae, Typha lat- 
ifo Ha, and Viti s arizonica . 

The Hualapai in the Black Mountains interacted frequently with the Mohave people, trading 
wild products for agricultural produce from the floodplain of the Colorado River (Stone 1987). The 
Mohave practiced a generalized settlement-subsistence pattern, which included farming, fishing, 
hunting, and gathering wild plants. Their settlements shifted in arrangement, composition, and 
location from year to year and season to season to cope with the unpredictable variations in the tim¬ 
ing and volume of annual floods (Bee 1981). Crops they cultivated included com, beans, squash, 
melons, and grasses. However, gathered resources were not mere supplements to cultivated crops 
but constituted a diverse and important component of the Mohavean diet. According to Castetter 
and Bell (1951:179-180), mesquite beans continued to be a staple resource into the 20 th century 
and was “more important than maize . . . and virtually supplied the living through the winter and 
until the next cultivated crop was ready.” Because large game was scarce along the river, fish, such 
as the humpbacked sucker and Colorado salmon or squawfish, and small game were the major 
sources of protein. During winter, small hunting parties pursued deer and bighorn sheep in the 
desert mountain ranges, including the Black Mountains. An annual lean period of reduced supplies 
occurred in the spring, when few wild plant resources were available and fish were relatively 
scarce. The duration and severity of this lean time depended on the amount and rate of consump¬ 
tion of stored foods from the previous seasons (Stone 1987). 

The remote and rugged terrain of the Black Mountains provided the Yuman peoples respite 
from the effects of early Spanish contact. However, by the 1800s Anglo-American explorers ven¬ 
tured into the area. The California Gold Rush in the 1850s led to increased travel and settlement in 
western Arizona. In 1863 gold was first discovered in the Black Mountains at the Moss vein north¬ 
west of Oatman (Gray et al. 1990b; Ransome 1923). Varying levels of mineral production contin¬ 
ued in the Oatman mining district until 1942. Numbers of feral burros, originally released by the 
miners working in the area, increased significantly, resulting in degradation of the vegetation and 
perennial springs and competition with the desert bighorn sheep for limited resources. After the 
introduction of cattle in Mohave County, army officers consistently reported that livestock grazing 
reduced the quantity of wild game and the supply of edible seed (United States Senate 1936). Davis 
(1973) documented that Sacramento Valley, now desertscrub dominated by Larrea tridentata and 
Ambrosia dumosa, was once an open grassland. In 1865 the Colorado River Indian Reservation 
was established, thereby providing the Mohave people with only a portion of their former prime 
farming and gathering lands. In 1883 the Hualapai Indian Reservation was established at the north 
end of the Hualapai people’s historic area of occupation. A profound impact to the Mohave peo¬ 
ple’s way of life resulted from construction of major dams on the Colorado River between 1930 
and 1970. These changed the water flows and aquatic temperature, tamed the annual floods, and 
reduced transport of sediment, all of which contributed to the loss of marshes and mesquite bosques 
as well as the native fish populations (Cole 1981). Given the magnitude of changes within the past 
century, it is difficult to see how a traditional subsistence economy could be sustained under cur¬ 
rent conditions in and around the Black Mountains. 


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Previous Plant Collectors 

Botanical exploration in the Black Mountains apparently began during the 19 th century with 
collections of Ericameria cuneata var. spathulata and Ericam eria teretifolia (Chrysothamnus 
teretifolius ) from Union Pass by Edward Palmer in 1870 (Kearney et al. 1960). According to 
McVaugh (1956), in 1870, Palmer traveled from Fort Whipple, in the vicinity of Prescott, across 
the desert to Fort Mohave on the Colorado River. Presumably he followed the Fort Mohave- 
Prescott Road, which crosses the Black Mountains at Union Pass (Paulsen 2010). Between 13 and 
21 May 1884, Marcus E. Jones made 70 collections in the vicinity of Yucca just outside the south¬ 
east boundary of the flora area (Lenz 1986:397); that year he described the desert around Yucca as 
“. . . a perfect garden in all directions, never was as good for 20 years afterwards.” 

A search of the Southwest Environmental Information Network (SEINet 2016) indicates that 
the early 20 th century was a rather active period for collecting even in as remote an area as the 
Black Mountains. The majority of collections from this period were from the Oatman-Sitgreaves 
Pass area. Frank A. Thackery, an agriculturist with the U.S. Department of Agriculture’s Bureau of 
Plant Industry, collected 8 specimens from Oatman in 1927 and 1928. On 22 March 1931, George 
John Harrison, with the U.S. Department of Agriculture Field Station in Sacaton, Arizona, made 18 
additional collections near Oatman. That same year, Alice Eastwood, curator of botany at the Cal¬ 
ifornia Academy of Sciences, also collected in the Oatman area. According to her collection notes 
in the archives at the Academy, she collected 83 specimens in Oatman during 20-21 April and 23 
specimens on the road to Oatman on 16 May. Thomas H. Kearney and Robert H. Peebles, botanists 
with the U.S. Department of Agriculture, collected 24 specimens from the “Black Ute Mountains” 
in the vicinity of Oatman on the following dates: 23 March 1931, 19 April 1935, 18-19 September 
1935, and 16 April 1937. On 14 April 1937, Bassett Maguire, then at Cornell University, and 
Andrew Alexander Nichol, with the Agricultural Experiment Station at the University of Arizona, 
collected 30 specimens in the Cool Springs area on the east side of the Black Mountains. On 1 
October 1940, Nichol made a few additional collections in Union Pass at the northern boundary of 
the flora area. On 1 and 15 April 1941, Joseph F. Arnold, also with the Agricultural Experiment Sta¬ 
tion at the University of Arizona, collected in Oatman, Secret Pass, and on the east side of the Black 
Mountains. On 16 October 1943, botanist Charlotte Goodding Reeder collected pin die a o do rata 
in the Goldroad Region just east of Sitgreaves Pass. Allen R. Phillips, a well-known ornithologist 
and co-author of The Birds of Arizona (Phillips et al. 1964) collected Quercus tu rb in ell a below 
Union Pass on 26 November 1947. John Thomas Howell, a curator of botany at the California 
Academy of Sciences, collected Larrea tridentata and Stephanomeria pauciflora in the vicinity of 
Oatman on 29 May 1950. 

During the latter half of the 20 th century numerous botanists (other than the authors) made 
additional collections from the Black Mountains, including: J. Anderson, R. J. Barr, J. Biggs, 
C. Carter, C. Christy, C. Edgar, S. Ferricr, R. Haberle, H. Hammond, G. Helmkamp and 
F. Helmkamp, R. Hevly, W. Hodgson, M. Hovezak, S. Ickert-Bond, E. Lehto, B. Marshall, 
C. Michaels, B. Parfitt, J. Parrott, K. Price, A. Phillips, B. Phillips, C. Reeder, J. Reeder, G. Rink, 
and J. Rominger. 

At least 390 collections from the Black Mountains have been made between 2000 and 2016 
(SEINet 2016). The vast majority of these specimens was collected north of our flora area and are 
deposited at ASU, ASC, and/or GMDRC. Within the flora area, the main collectors since 2000 have 
been G. Goodwin (ASC) and J. Andre (GMDRC). 


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Vegetation 

Mohave Desertscrub Biome.— We have relied on Brown et al. (1979) and Turner (1982) 
for the description of the Mohave Desertscrub Biome. However, in the absence of a single com¬ 
prehensive system for classifying specific plant communities in Arizona, we generally followed 
that of Thome et al. (1981). Their flora covers several mountain ranges in the Mohave Desert of 
southeastern California; all of these ranges are in close proximity to the Black Mountains, and they 
include several plant communities that are comparable in scale and composition to those found in 
our flora area. The plant communities represented in the southern Black Mountains are: Blackbrush 
Scrub, Creosote Bush Scrub, Desert Oasis Woodland, High-Gradient Desert Wash Scrub, Low- 
Gradient Desert Wash Scrub, Mixed Desert Scrub, Pinyon-Juniper Woodland, and Streamside 
Marsh. 

The vast majority of our flora area is within the Mohave Desertscrub Biome. Mohave 
Desertscrub is considered to be intermediate between the Great Basin and Sonoran desertscrub bio- 
mes, both geographically and floristically (Brown et al. 1979; Turner 1982). The pattern is well 
represented in the southern Black Mountains. The following species occur in both the Mohave and 
Sonoran deserts and are found in the southern Black Mountains: Ambrosia saisoia, Crossosoma 
bigelovii , Dudleya arizonica, Encelia farinosa, Eriogonum fasciculatum, Larrea tridentata, 
M entzelia invo luc rata , M ohavea confertiflora, Molina bigelovii, Opuntia bigelovii., Prosopis glan- 
dulosa Var. torreyana, Psorotham nus spinosa, Senegalia greggii, Trixis ealiforniea, Yucca bacca- 
ta, and Y, schidigera (Figs. 3-5). Coleogyne ramosissima (Figs. 6, 7) can be a dominant species in 
either the Mohave or Great Basin deserts. Other species in the southern Black Mountains that occur 
in both Mohave and Great Basin deserts include: Echinocereus engelmannii var. chrysocentrus, 
Ericam eria teretifo lia , Grayia spinosa , K ra scheninnikovia lanata, and Opuntia polyacantha var. 
erinacea (Figs. 6, 8). Monardella eplingii (Fig. 6), a recently described species, known only from 
the Black Mountains, is endemic to the Mohave Desert. Yucca brevifolia, another Mohave Desert 
endemic, occurs just outside of the flora area in the vicinity of Yucca. 

The Mohave Desertscrub Biome is known for its diversity of whiter ephemerals (annuals), pre¬ 
sumably due to the preponderance of precipitation between September and December (Beatley 
1974). A significant percentage (89%) of the ephemeral species documented in the flora area are 
winter ephemerals, many of which show a strong geographic affinity with those of the Sonoran 
Desert. In the flora area, we documented 63 of the 109 species of winter ephemerals listed by 
Shreve and Wiggins (1964) as characteristic of both the Mohave and Sonoran deserts. They include 
species of Amsinckia, Chaenactis, Chorizanthe, C ryptantha, Eriogonum, Eschscholzia, Euphorbia, 
Gilia, Linanthus, Lupinus, Malacothrix, M entzelia, Naina, and Phacelia, ill addition to Achyrony- 
chia cooperi, Acmispon brachycarpus, Bowlesia incana, Bromus arizonicus, Calycoseris wrightii, 
Chylismia claviformis, Daucus pusillus, Draba cuneifolia var. integrifolia, D escurainea pinnata, 
Eriophyllum lanosum, Erodium texanum, Eucrypta chrysanthem ifolia var. bipinnatifida, Eulobus 
cham aenerioides, Harpagonella palm eri, Lepidium lasiocarpum , Logfia filaginoides, Monoptilon 
bellio ides, M uhlenbergia microsperma, Myosurus cupulatus, N emacladus g la nd u life ru s, 
Oligomeris linifolia, Pectocarya hete rocarpa, Petunia parviflo ra, Pholistoma auritum, Pla- 
giobothrys arizonicus, Plantago ovala, Poa bigelovii, Pterostegia drytn arioides, Rafinesquia 
neomexicana, Salvia colum bariae, Thysanoca rpus curvipes, Vicia exigua , and Vulpia octoflora. 

Additional winter ephemerals are identified in the species list. Among ephemeral species, a much 
lower percentage (9%) of summer ephemerals occurs in the flora area, probably as a result of the 
relatively limited rainfall in August or September. These include Am a ran thus fimbriatus, Boer- 
havia wrightii, Bouteloua a ristido ides, Kallstroemia ealiforniea, Kallstroemia parviflo ra, Muhlen- 


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Figure 3. Species in the southern Black Mountains that occur in both Mohave and Sonoran deserts-1. Yucca schidigera 
(top left). Carnegiea gigantea (top right; photo by E. Butler, provided by and used with permission of J. Pebworth, Arizona 
Game and Fish Department). Mohavea confertiflora (bottom left; photo by L. Makings, SEINet). Larrea tridentata (bottom 
right; photo by S. Carnahan, SEFNet). Photos by the authors, except as noted 


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Figure 4. Species in the southern Black Mountains that occur in both Mohave and Sonoran deserts-2. Fouqueria spien- 
dens (top). Janusia gracilis (bottom left). Mentzelia involucrata (middle Center; photo by P. Alexander, SEINet). Trixis Cal¬ 
ifornia a (bottom center). Dudleya arizonica (bottom right). Photos by the authors, except as noted. 


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bergia fragilis, and Pectis papposci . Six species of ephemeral herbs flower throughout the year in 
response to rainfall: Aristida adscensionis, Baileya pleniradiata, C ham aesyce setiloba, Conyza 
canadensis, E riogonum hookeri, and Eriogon urn palmerianum. 

Creosote Bush Scrub.— This plant community (Fig. 7) is generally comparable to the Cre- 
osotebush Series of Brown et al. (1979) and Turner (1982). It occurs on the lower hills, upper and 
lower bajadas, and valley basins (generally between 335 m (1100 ft) and 914 m (3000 ft) eleva¬ 
tion), from the base of the mountains toward Sacramento Wash on the south and east sides of the 
flora area and toward the Colorado River on the west side of the flora area. The dominant species 
Ambrosia dumosa, Encelia farinasa, and Larrea tridentata are characteristic of this plant commu¬ 
nity. Common associates include: Cylindropuntia acanthocarpa, Cylind ropuntia bigelovii , 
Ephedra aspera, Fouquieria splendens, Krameria erecta, Opuntia basilaris, Psorothamnus fre- 

montii, and Sphaeralcea ambigua (Figs. 4, 6, 8). Ephemeral species can be abundant in response 
to rainfall. They include: A cmispon rigidus, A cm ispon maritimus, Acmispon strigosus. Amsinckia 
intermedia. Am sinckia tessel lata. At richo se ris platyphylla, B o utelou a barbata, Bouteloua aristi- 
doides, C hylism ia brevipes, C hylism ia claviform is, C haenactis carphoclinia, Chaenactis fremontii, 
Cham aesyce arizonica, Cham aesyce m elanadenia, Cham aesyce polycarpa, Chorizanthe brevicor- 
nu, C ho rizanthe rigida, Cryptantha barbigera, Cryptantha maritima, Cryptantha pterocarya, 
C ryptantha utahensis, Erem olhera refracta, E riastrum diffusum, E ria strum eremicum, E riogonum 
deflexum, E riogonum inflatum, Eriogonum thorn asi't, E riogonum trichopes, E rio phyllum lanosum, 
Eschscholzia calif'arnica, Eschscholzia glyptosperm a, Eschscholzia minutiflora, Gilia scopulorum, 
Gilia stellatci, Langloisia setosissima, Linanthus bigelovii, Linanthus demissus, Lupinus concinnus, 
M alacothrix glabrata, M entzelia albicaulis, M entzelia involucrata, M onoptilon bellioid es, Narna 
demissa, Nem acladus o rientalis, P ectoca rya platycarpa, P ectoca rya recu rvata, Phacelia crenula- 
ta, Phacelia d is tans, P Uinta go ovata, Rafine squia California a, R a fine squia n earn exic an a, 
Stephanomeria pauciflora, Syntrichopappus fremontii, and Vulpia octoflora, 

M ix ed Desert Scrub. —Mixed Desert Scrub dominates the upper mountain slopes and rock 
outcrops (generally between 850 m (2800 ft) and 1370 m (4500 ft) elevation) and features a greater 
diversity of species than Creosote Bush Scrub. This catch-all plant community (Fig. 2) occurs 
throughout much of the flora area and exhibits various combinations of associated species. Sever¬ 
al of the desert growth forms are represented including: stem-succulent cacti (Fig. 8) such as Cylin¬ 
dropuntia echinocarpa, Ech inocereus coccineus, Echinocereus engelm annii, Echinom astus john- 
sonii, Ferocactus cylind race u s, M amm illaria graham ii, M ammiliaria tetrancistra, Opuntia 
ch lorotica, Opuntia x curvospina, and Opuntia polyacantha j leaf semisucculents like Agave mck- 
elveyana, Nolina bigelovii, Yucca baccata, and Yucca schidigera (Figs. 3, 5); leaf-SUCClllentS (i.e.. 
Dudleya arizonica; Fig. 4); shrubby, often microphyllous, species, such as Brickella atractyloides, 
B ricke Ilia incana, Encelia resinifera, Encelia virginensis, Ephedra aspera, Ericamerici laricifolia, 
Ericameria linearifolia, E riogonum J'ascicu latum, Eriogonum heermannii, Gutierrezia sarothrae, 
Keckiella antirrhinoides. Salvia dorrii, Salvia mohavensis, Thamnosma montana , and Xylorhiza 
tortifolia (FigS. 6, 9); SubsllRlbs and perennial herbs, such as Acmispon rigidus, Artemisia ludovi- 
ciana. Baccharis brachyphylla, Boechera perennans, Cryptantha racemosa, Mirabilis laevis, 
Mirabilis multiflo ra, N icotiana obtusifolia, P enstemon eatonii, Physalis crassifolia, P leurocoronis 
plu ris eta, p o ro phyllum gracile, and p eucephyllum schottii; perennial grasses including 
Acnatherum co ronatum, Acnatherum hymenoides, Aristida purpurea, Bothriochloa barbinodis, 
Bouteloua curtipendula, Hilaria rigida, Jarava speciosa, Melica frutescens, Melica imperfecta, 
Muhienbergia porteri, and Poa fendleriana; and numerous ephemerals, many of which are listed 
above. In more mesic sites on north-facing slopes and at the base of cliff faces at higher elevations 
are found species characteristic of woodland and chaparral plant communities, including: Ceanoth- 


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us vestitus, Forestie ra pubescens, Fraxinus anom ala, G arrya flavescens, P runus fascicu lata, Quer- 
cus turbinella , and R h us arom atica. 

Blackbrush Scrub. —Geographically interspersed within the widespread Mixed Desert 
Scrub are areas of Blackbrush Scrub, which are dominated by Coleogyne ram osissima (Figs. 6, 7). 
This plant community is generally comparable to the Blackbrush Series, which is ecotonal between 
Great Basin and Mohave desertscrub communities (Turner 1982). Here, perennial grasses such as 

Acnatherum hymenoides, Dasyochloa pulchella, Hilaria rigida, Jarava speciosa, M uhlenb ergia 
porteri, Sporobolus cryptandrus, and Tridens muticus are commonly prevalent in unbumed stands 
that are in good condition. According to Bradley (1964), these areas are important for Desert 
Bighorn Sheep (ovis canadensis nelsoni) because of the occurrence of these grasses. Indeed, 
extensive areas of high value habitat and lambing grounds for Desert Bighorn Sheep exist in the 
flora area (U.S. Department of Interior 1996). Other species commonly associated with Blackbrush 
Scrub include: Encelia virginen sis, Ephedra aspera, Eriogonum fasciculatum, Krameria bicolor, 
Krameria erecta , , Larrea trident,ata , Opuntia basilaris , and Yucca schidigera. Juniperus califor- 
n ica , an indicator species of the southern portion of the Great Basin Conifer Woodland (Brown 
1982), is frequently found on dry slopes generally above 914 m (3000 ft.) elevation both with 
Blackbrush Scrub and Mixed Desert Scrub. 

Great Basin Conifer w oodland Biome, —A small area of the Great Basin Conifer Wood¬ 
land Biome was mapped by Brown et al. (2007) in the southern Black Mountains. We collected two 
of the characteristic species, Juniperus califo mica and Pin us monop hylla (Fig. 10), of this biome 
at higher elevations (1341-1433 m) north of Mt. Nutt. We treat this area as a relict Pinyon-Juniper 
Woodland plant community due to the rarity of p in u s m on op hylla in the southern Black Mountains. 
Characteristic species associated with this community in the flora area include: Artemisia ludovi- 

ciana, Coleogyne ramosissima, Echinocereus coccineus, Echinocereus engelmannii, Fraxinus 
anom ala, G arrya flavescens, Krascheninnikovia lanata, Nolina bigelovii, Opuntia basilaris, Pur- 
shia stansburyana, Quercus turbinella, Rhamnus ilicifolia, Rhus arom atica, aild Yucca baccata. 
Locally common grasses here are Achnatherum hymenoides, Aristida purpurea, Bouteloua cur- 
tipendula, Hilaria rigida , Jarava speciosa, Muhlenbergia porteri, Sporobolus cryptandrus, and 
Tridens mu ticus. We collected the following species only in this plant community in the flora area: 
Gilia flavocincta Subsp. australis, Hymenopappus filifotius, Linanthus pungens, Penstemon 
palmeri, Ptelea trifoliata, mdsilene verecunda. The majority ofthese associated species also occur 
with the Interior Chaparral Biome, as described by Pase and Brown (1982). Although not present 
in our flora area, Interior Chaparral is widespread in the Hualapai Mountains to the east across the 
Sacramento Valley (Butterwick et al. 1992). 

Desert Wash Scrub.— Numerous ephemeral streams occur throughout the flora area. The 
Desert Wash Scrub plant community varies with stream gradient, which is defined as the change in 
elevation per unit distance along the stream channel and expressed as a percent (Society of Amer¬ 
ican Foresters 1998). The low-gradient (generally < 3%) tributaries with predominantly sand and 
gravel substrates form incised arroyos and open, shallow washes at the lower elevations. These 
tributaries support a Low-Gradient Desert Wash Scrub plant community in and along the channels 
comprised Ot small trees such as Chilopsis linearis, Pa rkinsonia florida, Prosopis glandulosa var. 
torreyana, Prosopis pubescens, and Psorothamnus spinosus. Representative shrub species include 
Ambrosia monogyra, Ambrosia salsola, Bebbia juncea, Lycium exsertum, P sorothamnus fremontii, 
Scutellaria mexicana, Senegalia greggii, and Ziziphus obtusifolia . Vines SUCh 3S Cucurbita digita- 
ta, Funastrum cynanchoides, Funastrum hirtellum, Marah gilensis, Maurandella antirrhiniflora, 
and Neogaerrhinum filipes are occasionally found twining over the shrubs and trees or Railing 
along the streambed. These tributaries occur primarily within the Creosote Bush Scrub plant com- 


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Figure 5. Some conspicuous plants in the southern Black Mountains- 1. Nolina bigelovii (top). Calochortus kennedyi 
(bottom left). Eschscholzia glyptosperma (bottom right). 


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Figure 6. Species in the southern Black Mountains and their floristic affinities. Coleogyne ramosissima (top left; photo 
by Keir Morse, SElNet) and G rayia spinosa (top right; photo by C. Webber, © California Academy of Sciences, used with 
permission), occur in both Mohave and Great Basin deserts. Monardella eplingii, the only known plant species endemic to 
the Black Mountains (middle left; plant photo by and copyright reserved to M. Elvin, used with permission; inset photo 
showing inflorscence with flower by J. Anderson, used with permission), psorothamnus fremontii var. attenuatus (bottom 
left; photo by Daniela Roth, SElNet) and Salvia mohavensis (bottom right; photo by A. Schusteff, with permission), occur 
in both Mohave and Sonoran deserts. 


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Figure 7. Southern Black Mountains, plant communities-2. Creosote Bush Scrub near Wami Springs Wash (with Black 
Mesa in background) with Cylindropuntia spp., Encelia farinosa , and Larrea tridentata (top). Blackbrush Scrub SOUth of 
Sitgreaves Pass with Coleogyne ramosissima (bottom). Photos by the authors. 


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Figure 8. Some Cactaceae (stem succulents) that occur in the southern Black Mountains. Cylindropuntia acanthocarpa 
(top left). Cylindropuntia bigelovii (top Center). Opuntia polyacantha var. erinacea (top right). Mam m iliaria grahamii (cen¬ 
ter left). Ferocactus cylindraceus (center right). Opuntia basilaris (bottom left). Echinocereus coccineus (bottom right). 

Photos by the authors. 


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munity and, thus, feature many of the same species including: Ambrosia dumosa, Ephedra asp era, 
Eriogonum fasciculaium , Hilaria rigida, Jarava speciosa, Krameria erecta , and Larrea tridentata. 

With sufficient late winter rains, a high diversity of winter ephemerals are found in the Low-Gra¬ 
dient Desert Wash Scrub, many of which are listed above in the description of the Mohave 
Desertscrub Biome. 

Sacramento Wash is the largest low-gradient drainage near the southern Black Mountains, has 
multiple channels, and forms the southeastern boundary of the flora area for approximately 32 km 
(20 mi). The Low-Gradient Desert Wash Scrub here is dominated by Ambrosia monogyra, Baccha- 
ris salisifolia, Parkinsonia microphylla, Psorothamnus spinosus, and Senegalia greggii, with an 
herbaceous Stratum consisting of Cryptantha decipiens, Cryptantha micrantha, Descurainea pin- 
nata, Pectocarya platycarpa, Pectocarya recurvata, Senecio Jiaccidus, and Stilling ia linearifolia. 

The sand substrate provides suitable habitat for several species not found elsewhere in the flora 

area including Bailey a pleniradiata, Brickellia incana , Cryptuntha circum scissa, Dim orphocarpa 
w islizenii, Oenothera deltoides Sllbsp. deltoides, P alafoxia arida, and Tiquilia plica fa. An undeter¬ 
mined species of what appears to pertain to Petalonyx (Daniel & Butterwick 6042 ) has leaves with 
a dense pubescence of soft, white, and flexuose hairs, unlike the scabrous herbage of Petalonyx 
nitidus, which occurs elsewhere in the flora area. 

Upstream reaches of these tributaries, generally over 914 m (3000 ft) in elevation, are high- 
gradient (generally >3%) streams often with cobble, boulder, or bedrock substrates and steep side 
slopes that in places narrow into canyons. The relatively cooler and moister conditions at these sites 
support a more species-rich variant of Desert Wash Scrub, referred to here as High-Gradient Desert 
Wash Scrub, and they include several woodland and chaparral species (Fig. 11). We observed this 
plant community in several tributaries north of Mohave Co, 10 (Oatman Road). In this communi¬ 
ty species collected on canyon bottoms or adjacent slopes include: Acetmptopappus sphaero- 

cephalus, Amsonia palmeri, Artemisia ludoviciana, Baccharis sergiloides, Bothriochloa barbin- 
odis, Eucnide urens, Forestiera pubescens, Fraxinus anomala, Hyptis emoryi, Keckiella antirrhi- 
noides, Melicci imperfecta, Nolina bigelovii, Quercus turbinella, Rhamnus ilicifolia, SLUG. Salvia 
mohavensis. In shaded crevices of the canyon walls are found Brickellia atractyloides, Cheilan thes 
parryi, Crossosoma bigelovii, Erigeron oxyphyllus , Ivesia arizonica. Pellaea truncata, Perityle 
emoryi, Phacelia perityloides, Phacelia rotundifolia, Phacelia saxicola, P leu rocoron is pluriseta, 
and P terosteg ia drytnarioides. Among these, Erigeron oxyphyllus, Ivesia arizonica, Phacelia per¬ 
ityloides, Phacelia rotundifolia, and Phacelia saxicola are apparently rare, and thus far each one 
has been documented from a single locale in the flora area. 

Wetlands. —Year-round sources of water are restricted to perennial springs that occur on 
slopes, in narrow and boulder-strewn canyons, or in the more open, sand and gravel beds of tribu¬ 
taries at lower elevations. Of the 23 springs we sampled, the linear extent of surface and shallow 
subsurface flows varies from approximately 0.8 km (0.5 mi), at Cool Spring, to the immediate 
vicinity of the spring itself. The more extensive springs support a Desert Oasis Woodland plant 
community with a tree and shrub stratum (Fig. 11) and/or a Streamside Marsh plant community 
comprised of herbaceous species along the wet streambed and banks (Fig. 11). Several springs with 
limited flows support very little distinct vegetation, often consisting of single trees, such as 
Prosopis glandulosa var. torreyana at Metate Spring and Celtis reticulata at Whiskey Spring. In 
total, the vegetation associated with these perennial springs covers a very small area within this arid 
landscape and, consequently, many of the species associated with these plant communities are rare 
in the flora area. 

The Desert Oasis Woodland is dominated by the following trees: Celtis reticulata, Parkinso- 
nia florid a, P opulus fremontii, Prosopis glandulosa var. torreyana, Prosopis pubescens, Quercus 


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turbinella , and Salix gooddingii. Common shrub Species include Ambrosia monogyra, Baccharis 
salicifolia, Baccharis sergiloides, Salix exigua, and Pluchea sericea. The Stl'eamside Marsh may 
occur as an herbaceous stratum of the Desert Oasis Woodland or alone in more open drainages such 
as the series of step pools at Columbine Spring (Fig. 11). Species characteristic of the Streamside 
Marsh are numerous and include: A diantum capHlis-veneris, Polypogon viridis, Apocynum 
cannabinum, Aquile gia chrysantha, Carex alma, C astilleja minor subsp. spiralis, C henopodium 
berlandieri, Cyperus laevigatas, Eleocharis ovata, Eleocharis parishii, Eleocharis rostellata, 
Pseudognaphalium luteoalbum, Heliotropium curassavicum , Juncus acutus, Juncus bufonius, Jun- 
cus ensifolius, Lythrum californicum, Mimulus guttatus, Mimulus verbenaceus, Muhlenbergia 
rigens, Nasturtium officinale, P aspalum dilatatum, Petunia parviflora, Phragmites australis, Poly¬ 
pogon interruptus, Polypogon m onsp eliensis, Rumex hymeno sepalus, Solanum am ericanum, Sper- 
gularia marina, Typha latifolia, Veronica anagallis-aquatica, Vitis a rizonica, and W islizenia 
refra c ta (Fig. 12). 

A few of the herbaceous wetland species are currently known from a single spring. For 
instance, Columbine Spring, near the headwaters of an ephemeral tributary to Warm Springs Wash, 
has the only documented occurrence of Sisyrinchium demissum and Zeltnera arizonica (Fig. 12) in 
the flora area. Two grass species, Andropogon glomeratus and imperata brevifolia, have been doc¬ 
umented only from Cottonwood Spring in Cottonwood Canyon in the southern Black Mountains. 


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Figure 9. Some conspicuous plants in the southern Black Mountains-2. Castnieja chromosa (top left). Faiiugia para- 
doxa (top right). Euc aide urens (center left). Mimulus bigelovii (center right). Thamnosma montana (bottom left; photo by 
P. Alexander, SEINet). Oenothera caespitosa subsp. marginata (bottom right). Photos by the authors, except as noted. 


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Figure 10. Southern Black Mountains, plant communities-3. Pinyon-Juniper Woodland on saddle north of Mt. Nutt with 
Juniperus californica (top). Juniperus californica with female Cones (center right). Pinus monophylla (bottom left). Branch 
with female cones of P. monophylla (bottom right). Photos by the authors 


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Figure 11 . Southern Black Mountains, plant communities-4. Desert Oasis Woodland in Bighorn Cave Canyon with 
overstory of Populus fremontii and Baccharis sergiloides in understory (top left). Streamside Marsh at Columbine Spring 
(looking downslope with series of step pools) with Aqu Hegia chrysantha and M im ulus verbenaceus (top right). Desert Oasis 
Woodland with Streamside Marsh at “Twin Spring,” with M im ulus guttatus and Baccharis sergiloides in marsh understory 
(bottom left). High-Gradient Desert Wash Scrub above “Twin Spring” with Quercus turbinella (bottom right). Photos by 
the authors. 


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Figure 12. Plants restricted to springs in the southern Black Mountains. Plants of M im ulus verbenaceus and Aquilegia 
chrysantha at Columbine Spring (top). Aquilegia chrysantha (center left; photo by Max Licher, SEINet). Castilleja minor 
subsp. spiralis (middle center). Plant of Zeltnera arizonica (bottom right). Flower of Zeltnera arizonica (bottom left). Flow¬ 
ers of m im ulus verbenaceus (bottom center). Photos by the authors, except as noted. 


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Annotated Catalog of the Vascular Plants in the 
Southern Black Mountains 

Our list of plants is based in large part on extensive collection and examination of specimens 
deposited at ARIZ, ASC, ASU, CAS, DES, GMDRC, MNA, and UCR. We conducted 11 field trips 
of from one to six days each between 1986 and 1993. Eight of the trips involved sampling the 
springtime flora between February and May; two of the trips were in June and August; and one trip 
was in November. An effort was made to sample the diversity of habitats represented. Some sites 
were revisited in order to obtain a more complete inventory and adequate material for identifica¬ 
tion. Figure 1 shows the boundary of our flora area and the sites we sampled. Our collections from 
the southern Black Mountains consist of 1040 numbers, with a complete set of voucher specimens 
deposited at CAS and duplicates (when available) distributed to ASU, MO, and NY. A query of 
SEINet (2016) identified several species that might be expected to occur in the southern Black 
Mountains but which were not encountered by us. We examined these specimens or in some cases 
images of them in order to verify their localities and identifications. 

The vascular flora of the study area consists of 471 species in 276 genera representing 72 fam¬ 
ilies. In at least 44 species infraspecific taxa are recognized, and in seven of these species more 
than one inffaspecific taxon occurs in the southern Black Mountains. Thus, 478 taxa at the rank of 
species or below are recognized in our study.The flora is summarized based on major taxonomic 
categories (Table 1), numbers of genera and species in the largest families and numbers of species 
in the largest genera (Table 2), and life-form (Table 3). The life-form spectrum (Table 3) is char¬ 
acteristic of arid regions—the 205 total ephemeral herbs comprise 43% of the total flora. Thirty- 
eight taxa (8% of the flora) are not native to the flora of Arizona; most of them have been intro¬ 
duced from Europe. This is nearly the same percentage that was noted by Kearney et al. (7%; 1960) 
for the non-native flora of the state.The non-native species, b ramus rubens and Erodium cicutari- 
um , are abundant and conspicuous throughout much of the flora area (Fig. 2). 

Occurrences of at least 12 taxa in the southern Black Mountains are noteworthy. Monardella 
epling n (Fig. 6), a recently described species that is endemic to the Black Mountains (Elvin et al. 
2013), is found on various volcanic substrates in cracks of bedrock, among boulders along tribu¬ 
taries, and on rock outcrops, cliffs, and open rocky slopes. In our flora area, the species occurs in 
association with Mixed Desert Scrub and High-Gradient Desert Wash Scrub plant communities at 
elevations between 850-1100 m (2789-3608 ft). Six taxa appear to occur at a geographic limit of 
their respective distributions in the southern Black Mountains: Asclepias albicans (northern), 
Astra galis new be rryi var. blyae (western), Carnegiea giganlea (northwestern), Packera querceto- 
rum (western), Phacelia suxicola (southern), and Parkinsonia microphylla (northern). Five addi¬ 
tional taxa occur at a geographic limit of the Arizona portion of their respective distributions in our 
study area: Coreopsis califomica (northern and western), Parkinsonia florid a (northern), 
P sorothain nus sp inosus (northern), Typha latifolia (western), and Yucca schicligera (southeastern). 

In the following list, nomenclature and taxonomic circumscriptions generally follow Flora of 
North America North of Mexico (Flora of North America Editorial Committee 1993-2016), The 
Jepson Manual: Vascular Plants of California (Baldwin et al. 2012), Vascular Plants of Arizona 
Project (Vascular Plants of Arizona Editorial Committee 1992-2015), and /\ rizona Flora (Kearney 
et al. 1960) in that order and as published. In some cases, more recent and/or widely used syn¬ 
onyms are provided. In the following catalog, the name and author of each taxon are followed by 
an indication of whether the species is native or introduced, life form, plant community(ies) of 
occurrence, and reference to at least one voucher collection studied. In the collection numbers, “B” 
refers to a collection by Butterwick (or Butterwick et al.), “DB” refers to a collection by Daniel and 


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Table 1. Summary of the flora of the southern Black Mountains by major tax¬ 
onomic categories. Infraspecific taxa refers to species with occurrences of more 
than one variety or subspecies in the flora area. 


Families Genera 

Species Infraspecific Taxa 

Ferns 

1 3 

5 

0 

Gymnosperms 

3 3 

3 

0 

Flowering Plants 

68 270 

463 

7 

Totals 

72 276 

471 

7 

Table 2. Numbers of genera and species 

in the largest families and numbers 

of species in the largest genera. 


Largest families 

Genera/Species 

Largest Genera 

Species 

Asteraceae 

55/89 

C ryptan iha 

12 

Poaceae 

29/53 

Eriogonum 

12 

Boraginaceae 

12/35 

Phacelia 

7 

Fabaceae 

13/24 

B rom us 

6 

Brassicaceae 

14/19 

Chylism ia 

5 

Cactaceae 

7/16 

Cylindropuntia 

5 

Polygonaceae 

4/16 

Eleocharis 

5 


Ericam eria 

5 


G ilia 

5 


June us 

5 


M uhlenbergia 

5 


Table 3. Spectrum of life forms in the flora of the southern Black Mountains. WA = winter ephemeral 
herb; SA = summer ephemeral herb; YRA = year-round ephemeral herb; PH = perennial herb; PV = perenni¬ 
al vine; S = shrub; T = tree; SS = stem succulent; LSS = leaf semisucculent; LS = leaf succulent. Total flora 
refers to the 478 total taxa at the rank of species or below. 


WA 

SA 

YRA 

PH 

PV 

S 

T 

SS 

LSS 

LS 

Number of species 

182 

17 

6 

155 

7 

75 

15 

16 

4 

1 

Percent of total flora 

38 

3.6 

1.2 

32.6 

1.5 

15.7 

3.1 

3.4 

0.8 

0.2 


Butterwick, “DBH” refers to a collection by Daniel, Butterwick, and Deborah Hillyard, “DBa” 
refers to a collection by Daniel and Bruce Bartholomew, and “DBP” refers to a collection by 
Daniel, Butterwick, Arthur Phillips, and Barbara Phillips. The location of these collections is noted 
above. For all other collections, the herbarium in which the specimen is deposited is noted. Plant 
communities in the southern Black Mountains are designated with the abbreviations listed below. 


BBS 

Blackbrush Scrub 

CBS 

Creosote Bush Scrub 

DOW 

Desert Oasis Woodland 

HDWS 

High-Gradient Desert Wash Scrub 

LDWS 

Low-Gradient Desert Wash Scrub 

MDS 

Mixed Desert Scrub 

PJW 

Pinyon-Juniper Woodland 

SSM 

Streamside Marsh 


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Ferns 

Pteridaceae 

Adiantum capillus-veneris L. Native perennial herb; DOW, SSM. db 4518. 

Cheiianthes coviliei Maxon Native perennial herb; HDWS, MDS. DB 6645, 6682; dbh 
5704; DBP 5954. 

Cheiianthes parryi (D.c. Eaton) Domin Native perennial herb; LDWS, HDWS. B 8954; DB 
5919. 

Cheiianthes xparishii Davenp. Native perennial herb; HDWS. Windham 347 (ASU). 
Peiiaea truncata Goodd. Native perennial herb; MDS, HDWS. b 8957; dbh 5695. 

Gymnosperms 

C upressaceae 

juniperus californica Carriere Native shrub (sometimes treelike); BBS, HDWS, MDS, 
PJW. B 8992; DB 4495, 6006; Hale 5045 (ARIZ); Kearney & Peebles 1263 0 (ARIZ); Phillips 74- 

266 (ARIZ). Fig. 10. 


Ephedraceae 

Ephedra aspera Engelm. ex S. Watson Native shrub; BBS, CBS, HDWS, LDWS, MDS. 

B 8887; DB4623, 6483; Barr 67-104 (ARIZ), 67-105 (ARIZ); Carter s.n. (ASC); Currie 5 (ASC); 
Helmkamp 1555 (ARIZ); H uis in g a s.n. (ASC); Ickert-Bond 571 (ASU, ARIZ); N elson 9 (ASC); 
Nichol s.n. (ARIZ); Reeder & Reeder 8981 (ARIZ), 8981a (ARIZ); Stevens-Rumann 11 (ASC). 

Goodwin 3320 (ASC) was determined as Ephedra nevadensis S. Watson. Because the speci¬ 
men lacks female cones, we were unable to confim this determination. 

Pinaceae 

Pinus monophyiia Torr. & Frem. Native tree; PJW; saddle above Grapevine Spring Canyon, 
north facing slopes, db 6702, Hovezak 14 (MNA-not seen). Fig. 10. 

We observed and collected this species at a single locale, the saddle above Grapevine Spring 
Canyon on north-facing slopes at 1357 m elevation. Fewer than 10 individuals were noted at this 
site. Our collection was from a plant about six meters tall with a trunk diameter (DBH) of 40 cm. 
Cones were present on the ground. Hovezak’s collection appears to have been made about 0.8 km 
northwest of ours at 1433 meters. Notes on his collection indicate the presence of mature, cone¬ 
bearing trees with an average trunk diameter of 46 cm. 

Flowering Plants 

A doxaceae 

Sambucus nigra L. subsp. caerulea (Raf.) Bolli (5. m exicana C. Presl. ex DC.) Native 
shrub; MDS. db 5933. 


Agavaceae 

Agave mckelveyana Gentry Native leaf semisuCClllent; MDS, HDWS. DB 4507; Gentry 
9961 (ARIZ), 21979 (ARIZ, DES), Hovezak s.n. (MNA). 

The type of this name, Gentry 21979 (ARIZ, DES), is from Sitgreaves Pass in the southern 
Black Mountains. Based on a mixed collection of Marcus Jones ( 25167 at CAS, POM) from Oat- 
man, Gentry (1982) included a. Utah en si 5 Engelm. subsp. Utah e ns is as occurring in the flora 


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area.The -4. make Ivey ana portion of the collection was likely collected near Oatman, but subse¬ 
quent collecting activities have failed to locate A. u taken sis in the southern Black Mountains. We 
tentatively exclude it from the currently known flora. 

Noiina bigelovii (Torn) s. Watson Native leaf semisucculent; BBS, HDWS, MDS, PJW. 
B 8942. Fig. 5. 

Yucca baccata Torr. Native leaf semisucculent; MDS, PJW. DB 5673, 6667; DBH 5744. 

Yucca schidigera Ortgies Native leaf semisucculent; BBS, MDS. B 9349\ DB 5638. Fig. 3. 

The southeastern extent of the native distribution of this species in Arizona occurs in the south¬ 
ern Black Mountains (ca. 35.037651° lat.; based on Hodgson et al. 3536 at DES; SEINet 2016). 


A m aranthaceae 

Amaranthus fimbriatus (Torr.) Benth. ex S. Watson Native Summer ephemeral herb; 
LDWS, HDWS. DB 4598. 

Tidestromia lanuginosa (Nutt.) Standi. Native Slimmer ephemeral herb; CBS. Andre 32087 
(GMDRC). 


A nacardiaceae 

Rhus aromatica Aiton (r. triiobata Nutt. var. anisophylla (Greene) Jeps.) Native shrub; 
HDWS, MDS, PJW. DB 5916, 5937; DBH 5715. 

A piaceae 

Bowlesia incana Ruiz & Pav. Native winter ephemeral herb; CBS, LDWS. db 4686. 
Daucus pusilius Michx. Native winter ephemeral herb; CBS, LDWS. db 5594, 6423, 6473; 
DBH 5750. 

Lomatium nevadense (S. Watson) J.M. Coult. & Rose var. parishii (J.M. Coult. & Rose) 
jeps. Native perennial herb; HDWS, PJW. db 5973, 6707. 

Yabea microcarpa (Hook. & Am.) Koso-Poi. Native winter ephemeral herb; HDWS, MDS. 
DBH 5690. 


A pocynaceae 

Amsonia palm eri A. G ray Native perennial herb; LDWS, HDWS. b 6821, 8932; dbh 5724; 
DBP 5963; H ovezak s.n . (MNA). 

Apocynum cannabinum L. Native perennial herb; DOW, SSM, local at springs, db 4519, 
6451; B 8922. 

Asciepias albicans S. Watson Native perennial herb; CBS. DB 6466. 

This collection from the southern portion of our flora area (34.861486° lat.) and one collected 
in the same region ( Power s.n. at ASC), represent the northernmost known occurrence of this 
species (Consortium of California Herbaria 2016; SEINet 2016). 

Asciepias erosa Torr. Native perennial herb; CBS, LDWS. db 6044. 

Asciepias nyctaginifolia A. Gray Native perennial herb; HDWS. DBH 5735. 

Asciepias subuiata Dccne. Native perennial herb; CBS. db 450i;dbh 5751. 

Funastrum cynanchoides (Decne) Schltr. var. hartwegii (Vail) Krings (Sarcostemma 

cynanchoides Decile, subsp. hartwegii (Vail) R.W. Holm.) Native perennial vine; HDWS, LDWS. 
B 8990; DB 5616, 5912; Hodgson et al. 9203 (DES), 9200 (DES). 

Funastrum hirtellum (A. Gray) Schltr. ( Sarcostemma hirtellum (A. Gray) R.W. Holm) 
Native perennial vine; LDWS. b 9330; db 5600, 6001. 

Matelea parvifolia (Torr.) Woodson Native perennial vine; LDWS, MDS. DB 4695; DBa 


5113. 


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A steraceae 

Acamptopappus sphaerocephalus A. Gray Native shrub; HDWS, MDS. fi 8935. 

Acourtia wrightii (A. Gray) Reveal & R.M . King Native perennial herb; HDWS. DB 4562, 
5 985; DBH 5733. 

Adenophyllum porophylloides (A. Gray) Strother (Dyssodia porophylloid.es A. Gray) 
Native perennial herb; HDWS, LDWS. b 8949, 9354; db 4646, 6388. 

Ambrosia confertiflora (DC.) Rydb. Native perennial herb; HDWS. DBP 5942. 

Ambrosia dumosa (A. Gray) W.W. Payne Native shrub; CBS, LDWS, MDS. B 9337; 
DB 6662. 

Am brosia eriocentra (A. G ray) W.W. Payne Native shrub; HDWS. B 8974; DB 5 981; DBH 

5725. 

Ambrosia monogyra (Torr. & A. Gray) Strother & B.G. Baldwin (Hymenoclea m ono gyra 

Torr. & A. Gray) Native shrub; DOW, HDWS, LDWS. db 4608, 6316. 

Ambrosia salsola (Torr. & A.Gray) Strother & B.G. B a Id win (Hymenoclea s also la Torr. & 
A. Gray) Native shrub; LDWS. b 8877, 9329, 933 4. 

Amphipappus fremontii Torr. & A. Gray var. spinosus (A. Nelson) Ced. Porter Native 
shrub; MDS. Weaver & Web be s.n. (DES). 

Artemisia dracuncuius l. Native perennial herb; HDWS. Db 4551. 

Artemisia ludoviciana Nutt. Native perennial herb; HDWS, MDS, PJW.fi 8981. 
Atrichoseris piatyphyiia A. Gray Native winter ephemeral herb; CBS, LDWS. db 5607. 
Baccharis brachyphylla A. Gray Native shrub; DOW, HDWS, MDS. DB 4541,6675; DBH 
5717. 

Baccharis salicifolia (Ruiz & Pav.) Pers. Native shrub; DOW, LDWS. DB 5993, 6330. 

B accharis ser giloides A. Gray Native shrub; DOW, HDWS. fi 8923; DB 451 7; Hovezak s.n. 

(DES, MNA). 

Bahiopsis parishii (Greene) E.E. Schill. & Panero (Vig uiera deltoidea A. Gray var. parishii 
(Greene) Vasey & Rose) Native shrub; HDWS, LDWS, MDS. fi 8889, 8952; db 5595; Anderson 
473 0 (ASU); H ovezak s.n . (MNA); P arfitt & Clark 2182 (ASU)j Rominger 307-65 (ASC), 312-65 
(ASC). 

Baileya muitiradiata Harv. & A. Gray Native perennial (biennial) herb; CBS, HDWS, 
LDWS. fi 9360; DB 5685. 

Baileya pieniradiata Harv. & A. Gray Native year-round ephemeral herb; LDWS. DB 6035. 
Bebbia juncea (Benth.) Greene var. aspera Greene Native shrub; HDWS, LDWS. fi 8885; 
DB 4513. 

Brickellia atractyloides A. Gray Native shrub; HDWS, MDS. fi 8891; DB 5665. 

B rickellia desertorum Coville Native shrub; HDWS, MDS. fi 8919, 8982; DB 4522, 5991, 
6328. 

B rickellia incana A. G ray Native shrub; LDWS. DB 6021, 

Brickellia microphylla (N u tt.) A. G ray Native slimb; HDWS. DB 4521. 

Caiycoseris parryiA. Gray Native winter ephemeral herb; HDWS, LDWS, MDS. fi)fi 63 99; 
DBH 5741. 

Caiycoseris wrightii A. Gray Native winter ephemeral herb; HDWS, LDWS, MDS. fi 8985; 
DB 4665; DBa 5142. 

Centaurea meiitensis l. Introduced weedy winter ephemeral herb; LDWS. db 5910. 
Chaenactis carphoclinia A. Gray var. carphoclinia Native winter ephemeral herb; CBS, 
LDWS. fi 9332; DB 4697, 6439. 


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Chacnactis fremontii A. Gray Native winter ephemeral herb; CBS, HDWS, LDWS, MDS. 
B 8934; DB 4626, 4632, 5589, 5975; DBa 5107,5141. 

Chaenactis raacrantha D.C. Eaton Native winter ephemeral herb; MDS. DB 6660. 
Chaenactis stevioides Ilook. & Arn. Native winter ephemeral herb; MDS. DBH 5740. 
Cirsium neomexicanum A. Gray Native perennial (biennial) herb; HDWS, MDS. b 8964, 
9351; DB 6400. 

Conyza canadensis (L.) Cronquist Native weedy, year-round ephemeral herb; HDWS. db 

4565. 

Coreopsis californica (Nutt.) H. Sharsm. ( Leptosyne californica Nutt.) Native winter 
ephemeral herb; MDS; Kearney & Peebles 11281 (ARIZ). 

This collection, presumably from Sitgreaves Pass, represents the northern- and westernmost 
occurrence of this species in Arizona, and it is apparently the only known occurrence of the species 
in Mohave County (SEfNet 2016). 

Dieteria asteroides Torr. var. asteroides (M ciehaeranthera asteraides (Ton*.) Greene var. 
asteroides ) Native perennial herb; HDWS. db 4559 ; Hevly & Hovezak 62 (MNA). 

Dieteria canescens (Pursh) Nuttaii Native perennial (biennial) herb; HDWS. b 6817. 
Encelia farinosa A. Gray ex Torr. Native shrub; CBS, MDS. DB 4620. 

Encelia resinifera C. Clark Native shl'llb; MDS. DB 5629; Lehto 21055 (ASU). 

Encelia virginen sis A. N els. Native shrub; BBS, MDS. B 9348. 

Ericameria cuneata McClatchie var. spathulata (A. Gray) H.M. Ha 11 Native shrub; MDS. 
Paim er s.n . (US-image). 

Ericameria laricifolia (A. Gray) Shinners Native shrub; HDWS, MDS. B 8997; Hovezcik 
s.n . (MNA). 

Ericameria linearifolia (DC.) Urbatsch & Wussow ( Stenotopsis linearifolia (DC.) Rydb. 
var. interior (Coville) J.F. Macbr.) Native shrub; MDS. db 5672, 5927. 

Ericameria paniculata (A. Gray) Rydberg (Chrysothamnus paniculatus (A. Gray) H.M. 
Hall) Native shrub; HDWS, LDWS. B 8971 ;DB 5913, 6020, 6302; Hovezak & Hevly s.n. (MNA). 

Ericameria teretifolia (Durand & Hilgard) Jeps. ( Chrysothamnus teretifolius (Durand & 
Hilgard) H.M. Hall) Native shrub; MDS. Anderson 93-28 (ASU); Palmer s.n. (US-image). 

Erigeron divergens Torr. & A. Gray Native perennial herb; HDWS, MDS. D B 4566, 5926. 
Erigeron lobatus A. Nelson Native winter ephemeral herb; HDWS. b 8958. 

Erigeron oxyphyllus Greene Native perennial herb; HDWS. B 8939. 

Eriophyllum lanosum A. Gray Native winter ephemeral herb; CBS, HDWS, LDWS, MDS. 

B 8968; DB 4639; DBa 5096. 

Eriophyllum wallacei (A. Gray) A. Gray Native winter ephemeral herb; MDS. Arnold s.n. 
(DES); Kearney & Peebles 11279 (ARIZ); Kearney et al. 7617 (ARIZ). 

Geraea canescens Torr. & A. Gray Native winter ephemeral herb; CBS. DB 6469. 
Gutierrezia microcephala (DC.) A. Gray Native shrub (subshrub); HDWS. Hovezak s.n. 
(MNA) 

Gutierrezia sarothrae (Pursh) Britton & Rusby Native shrub (subshrub); HDWS, MDS. 

B 8945; DB 4499, 6304; Hovezak s.n. (MNA). 

H ym enopappus filifolius H ook. var. lugens (G reene) Jeps. Native perennial herb; PJW. DB 

6697. 

Hymenothrix loomisii S.F. Blake Native Summer ephemeral herb; MDS. Rominger 312-65 
(ARIZ, ASC). 

Layia gianduiosa (Hook.) Hook. & Arn. Native winter ephemeral herb; MDS. DB 5650, 


6690. 


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Logfia filaginoidcs (Hook & Arn.) Morefield ( Filago californica Nutt.) Native winter 
ephemeral herb; HDWS, MDS. b 9007; db 6665. 

M alacothrix coulteri Harv. & A. Gray Native winter ephemeral herb; MDS. DB5631,6658; 
DB a 5108. 

M alacothrix glabrata (A. Gray ex D.C. Eaton) A. Gray (M alacothrix californica DC. var. 
g lab rata Eaton) Native winter ephemeral herb; CBS, HDWS, LDWS. b 8986; db 5630, 6386; 
DBH 5747; D B a 5131. 

M alacothrix sonorae w.s. Davis & p.h. Raven Native winter ephemeral herb; MDS. DB 
6689; Hodgson H-757 (DES); Price s.n. (ASC). 

Monoptiion beiiioides (A. Gray) H.M. Hall Native winter ephemeral herb; CBS, LDWS. 

DB 4661; DBa 5153. 

Pack era m u Itilo ba ta (Torr. & A . G ray) W .A . W eber & A . L ove ( Senecio multilobatus Torr. 
& A. Gray) Native perennial herb; HDWS, MDS. db 4548, 5656, 5920; dbh 5713. 

Packera quercetorum (Greene) C. Jeffrey Native perennial herb; DOW, LDWS. Maguire 
17814 (UTC). 

This collection from Cool Springs on the east side of the Black Mountains (-114.309949° 
long.), represents the westernmost known occurrence of this species (SEINet 2016). 

Paiafoxia arida b.l. Turner & M.1. Morris Native winter ephemeral herb; LDWS. DB 

6034. 

Parthenium incanum Kunth Native shrub; MDS. D B 6695. 

Pectis papposa Harv. & A. Gray Native summer ephemeral herb; HDWS. Hovezak s.n. 
(MNA). 

Perityie emoryi Torr. Native winter ephemeral herb; HDWS, LDWS. b 8929;db 469i;DBa 
5156. 

Peucephyllum schottii A. G ray Native shrub; MDS. DB 5611. 

Pluchea sericea (Nutt.) Coville ( Tessaria sericea (Nutt.) Shinners) Native shrub; DOW. db 

5676, 6462. 

Pleurocoronis pluriseta (A. Gray) R.M. King & H. Rob. Native shrub; CBS, HDWS, 
MDS. B 8943; DB 4644. 

Porophyiium graciie Benth. Native perennial herb; LDWS, MDS. b 8880; db 5609; DBa 
5100. 

Prenantheiia exigua (A. Gray) Rydb. Native winter ephemeral herb; CBS, LDWS. B 9338; 
DB 5601,6428. 

Pseudognaphalium canescens (DC.) Anderb. (G naphalium wrightii A. Gray) Native peren¬ 
nial herb; HDWS, SSM. db 4569; dbh 5731,5737. 

Pseudognaphalium luteoalbum (L.) Hilliard & B.L. Burtt. (G naphalium luteoalbum L.) 

Introduced winter ephemeral herb; DOW, SSM. db 5995 , 6325, 6464. 

Pseudognaphalium stramineum (K until) A nderb. {G naphalium chilense Spreng.) Native 
perennial herb; SSM. dbh 5771. 

Psilostrophe cooperi (A. G ray) G reene Native perennial herb; HDWS. B 8996. 
Rafinesquia californica Nutt. Native winter ephemeral herb; CBS, HDWS, MDS. b 8928, 
9006; DB 5636. 

Rafinesquia neomexicana A. Gray Native winter ephemeral herb; CBS, LDWS, MDS. db 

4633, 5592; DBa 5106. 

Senecio flaccidus Less. var. flaccidus (5. douglasii DC. var. douglasii) Native perennial 
herb; DOW, HDWS, LDWS, MDS. b 8879; db 4500, 6465. 


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Senecio flaccidus Less. var. monocnsis (Greene) B.L.Turner & T.M. Barkley ( S. douglasii 
DC. var. monoensis (Greene) Jeps.) Native perennial herb; HDWS. Hovezak s.n. (MNA). 
Sonchus asper (L.) Hill Introduced winter ephemeral herb; SSM. db 5684. 

Sonchus olcraceus L. Introduced winter ephemeral herb; HDWS, SSM. b 8909; dbh 5756. 
Stephanom eria pauciflora (Torr.) A. Nelson Native perennial herb; CBS, HDWS. B 9336; 
Christy 1482 (ASU); Hovezak s.n. (MNA); Howell 2 6615 (CAS), 

Christy 1482 was previously determined as Stephanom eria exigua Nutt. Based on characters 
of the pappus, length of involucre, and branching pattern, it conforms to S. pauciflora. 
stephanom eria exigua has been collected ( White 527 at CAS) ca. 5 miles north of Union Pass (the 
northern border of our flora area) and likely also occurs in the southern Black Mountains. 

Stylociine micropoides A. Gray Native winter ephemeral herb; CBS, LDWS. db 5681, 
6460; DBa 5167. 

Sym p hy otrichum subulatum (Michx.) G .L. Nesom var. parviflorum (Nees) S.D. Sundb. 

Native winter ephemeral herb; MDS. db 5677. 

Syntrichopappus fremontii A. Gray Native winter ephemeral herb; CBS, MDS. DB5649, 
6004, 6698; DBP 5949. 

Trichoptiiium incisum (A. G ray) A. Gray Native winter ephemeral herb; CBS, LDWS. d b 

4690, 5603,6381, 6437. 

Trixis californica Kellogg Native shrub; CBS, LDWS. DB 4655; DBH 5748. Fig. 4. 
Uropappus lindleyi (DC.) Nutt. ( Microseris lindleyi (DC.) A. Gray) Native winter ephemer¬ 
al herb; MDS. b 9370; db 5667, 6687. 

Verbesina encelioides (Cav.) Benth. & Hook.f. ex A. Gray Native winter ephemeral herb; 
CBS, LDWS. DB 6033. 

Xanthisma spinulosum (Pursh) D.R. Morgan & R.L. Hartman var. gooddingii (A. Nel¬ 
son) D.R. Morgan & R.L. Hartman (M achaeranthera pinnatifida (Hook.) Shimiers) Native 
perennial herb; CBS, MDS. db 4641,4648. 

X ylorhiza tortifolia (Torr. & A. Gray) Greene Native perennial herb (subshmb); MDS. D B a 

5101. 


B ignoniaceae 

C hilopsis linear is (Cav.) Sweet Native tree; LDWS. db 4 502. 

B oraginaceae 

Amsinckia intermedia Fisch. & c.A. Mey. Native winter ephemeral herb; CBS, HDWS, 
LDWS. B 9003; DB 4588; DBa 5133. 

Amsinckia tesseiiata A. Gray Native winter ephemeral herb; CBS, LDWS, MDS. db 4650, 
5608; DBa 5089, 5175. 

Cryptantha angustifolia (Torr.) Greene Native winter ephemeral herb; CBS, HDWS, 
LDWS. DB 6028, 6029; DBa 5129. 

Cryptantha barbigera (A. Gray) Greene Native winter ephemeral herb; CBS, HDWS, 
LDWS, MDS. B 8882; DB 4613, 5588, 6420, 6663, 6668; DBa 5088, 5120, 5128, 5136, 5162; 
Lehto 21048 (ASU). 

Cryptantha circumscissa (Hook. & Am.) i.m. johnst. Native winter ephemeral herb; 
LDWS. DB 6025. 

Cryptantha decipiens (M.E. Jones) A. Heller Native winter ephemeral herb; CBS, HDWS, 
LDWS. B 8966; DB 6026. 

Cryptantha gracilis Osterh. Native winter ephemeral herb; CBS. db 4635. 


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Cryptantha maritima (Greene) Greene Native winter ephemeral herb; CBS, LDWS. DB 

4640, 4645; DBa 5161, 5/73. 

Cryptantha micrantha (Torr.) i.m. johnst. Native winter ephemeral herb; CBS, LDWS. 
DB 6027. 

Cryptantha nevadensis A. Nelson & P.B. Kenn. var. nevadensis Native winter ephemeral 
herb; LDWS, MDS. db 5661 ;Dba 5160. 

Cryptantha pterocarya (Torr.) G reene var. cycloptera (G reene) J.F. M acbr. Native win¬ 
ter ephemeral herb; MDS. db 6488. 

Cryptantha pterocarya (Torr.) Greene var. pterocarya Native winter ephemeral herb; 
CBS, HDWS, LDWS, MDS. B 8970; DB 4581,4602, 5619, 6664; DBa 5087, 5117, 5135. 

Cryptantha racemosa (S. Watson ex A. Gray) Greene Native perennial herb; HDWS, 
MDS. B 8916, 8924. 

Cryptantha utahensis (A. Gray) Greene Native winter ephemeral herb; CBS, HDWS, 
LDWS, MDS. B 8895, 8983; DB 5587, 5622, 5669, 6481; DBa 5163. 

Emmenanthe penduliflora B enth. var. penduliflora Native winter ephemeral herb; HDWS, 
MDS. B 8933; DBa 5111. 

Eucrypta chrysanthem ifolia (Benth.) Greene var. bipinnatifida (Torr.) Constance Native 
winter ephemeral herb; HDWS. b 890 7 . 

Eucrypta micrantha (Torr.) A. Heller has been collected ( Fugate 6 at ASC) ca. 10 miles north 
of Union Pass and likely also occurs in the southern Black Mountains. 

H arpagonella palmeri A. Gray Native winter ephemeral herb; MDS. Andre 28622 
(GMDRC). 

H eliotropium curassavicum L . var. oculatum (A . H eller) Tidestr. Native perennial herb; 
SSM. DB 6450. 

Nama demissa A. Gray Native winter ephemeral herb; CBS, LDWS. b 9342; db 4688, 
6407; DBa 5132. 

Nama hispida A. Gray var. spathulata (Torr.) C .L. Hitchc. Native winter ephemeral herb; 
LDWS. DB 6041. 

Pectocarya heterocarpa (I.M. johnst.) i.m. johnst. Native winter ephemeral herb; CBS, 
LDWS. DB 4643b, 4710. 

Pectocarya platycarpa (Munz & I.M. Johnst.) Munz & I.M. Johnst. Native winter 
ephemeral herb; CBS, HDWS, LDWS, MDS. db 4540, 4603, 4643, 4710a, 6031; DBa 5098, 
5134. 

Pectocarya recurvata i.m. johnst. Native winter ephemeral herb; CBS, HDWS, LDWS, 
MDS. DB 4538, 4603a, 4643a, 4678, 6030; DBa 5097, 5174. 

Pectocarya setosa a. Gray Native winter ephemeral herb; MDS. db 5972,6691; d bp 5943. 
Phacelia crenuiata Torr. ex s. Watson Native winter ephemeral herb; CBS, HDWS, MDS. 
B 892 5, 8925a; DB 4671, 4717, 63 79; DBa 5092, 5122. 

Phacelia cryptantha Greene Native winter ephemeral herb; HDWS, LDWS. b 6822; db 
5647, 5660, 6482; Edgar s.n. (ASC); Hodgson etal.9143 (DES), 9150 (DES). 

Phacelia distans Benth. Native winter ephemeral herb; CBS, HDWS, LDWS. b 8910, 8973; 
DB 4590, 4708, 5646, 6478, 6486; DBa 5159; Goodwin 3314 (ASC). 

Phacelia fremontii Torr. Native winter ephemeral herb; MDS. db 6709. 

Phacelia perityloides Coville var. laxiflora (J.T. Howell) Cronquist Native perennial herb; 
HDWS. B 8956; Hodgson et al. 9185 (DES). 

Phacelia rotundifolia Torr. ex s. Watson Native winter ephemeral herb; HDWS. DB 5617; 
Hodgson 9198 el al. (DES). 


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Phaceiia saxicoia A. Gray Native winter ephemeral herb; HDWS. b 8917. 

This collection (at 35.119416° lat.) represents the southernmost geographic extent of the 
species. 

Pholistom a auritum (Lindl.) Lilja var. arizonicum (M .E . Jones) Constance Native winter 
ephemeral herb; HDWS. b 8913. 

Plagiobothrys arizonicus (A. Gray) Greene ex A. Gray Native winter ephemeral herb; 
HDWS, MDS. DB 5659, 6487; DBP 5947. 

Plagioboth rys jonesii A. Gray Native winter ephemeral herb; MDS. b 9357 

Tiquilia canescens (DC.) A. Richardson var. canescens Native perennial herb; CBS, MDS. 

DB 5632, 6471; DBa 5099. 

Tiquilia plicata (Torr.) A. Richardson Native perennial herb; LDWS. DB 6037. 

Brassicaceae 

Athysanus pusilius (Hook.) Greene Native winter ephemeral herb; MDS. DB 6693. 
Boechera percnnans (S. Watson) W.A. Weber ( Arabis perennans S. Watson) Native peren¬ 
nial herb; HDWS, MDS. b 9001 ; db 4612, 6671 . 

Brassica tournefourtii Gouan. Introduced winter ephemeral herb; DOW, SSM. db 6447. 
Cauianthus cooperi (S. Watson) Payson Native winter ephemeral herb; CBS, HDWS, 
LDWS, MDS. B 8896; DB 4649, 5593; DBa 5119. 

Cauianthus lasiophyllus (Hook. & Arn.) Payson var. lasiophyllus Native winter ephemer¬ 
al herb; LDWS, MDS. db 4679, 6655. 

Descurainia pinnata (Walter) Britton subsp. glabra (Wooton & Standi.) Detling Native 
winter ephemeral herb; HDWS, LDWS, MDS. db 4601,5613,6002,6032,6656,67 12;db a 5086, 
5139. 

Dimorphocarpa wislizenii (Engiem.) Rollins Native winter ephemeral herb; LDWS. DB 

6023. 

Draba cuneifolia Nutt, ex Torr. & A. Gray var. integrifolia S. Watson Native winter 
ephemeral herb; HDWS, LDWS. db 4580, 47u. 

Lepidium fremontii S. Watson Native perennial herb; BBS, MDS. DB 5925, 6670; DBP 
5948. 

Lepidium iasiocarpum Nutt. var. iasiocarpum Native winter ephemeral herb; BBS, CBS, 
HDWS, LDWS, MDS. DB 4552, 4584, 4642, 4685, 6492; DBa 5116, 5169. 

Nasturtium officinale w.t. A iton Introduced perennial herb; DOW, SSM. b 9367;db 5769, 
5992. 

Physaria tenella (A. Nelson) O’Kane & Al-Shehbaz ( Lesquerella tenella A. Nelson) Native 
winter ephemeral herb; MDS. db 5743 , 6674; Hammond 11118 (ASU); m arshaii & Tall s.n. 
(ASC); Nichol 2983 (ARIZ). 

Sisym brium aitissim um l. Introduced winter ephemeral herb; DOW, HDWS, LDWS, MDS. 

B 9368; DB 5612,5621,6651. 

Sisym brium irio L. Introduced winter ephemeral herb; CBS, LDWS. db 4662, 4712 , 6446. 
Sisymbrium orientate l. Introduced winter ephemeral herb; MDS. db 5930; Carter s.n. 
(ASC); Christy 1493 (ASU); Hodgson et al. 9163 (DES); Parfitt 21 78 (ASU). 

Stanleya pinnata (Pursh) Britton var. pinnata Native shrub; HDWS, MDS. B 8920; DB 
5618, 6479. 

Streptanthella longirostris (S. Watson) Rydb. Native winter ephemeral herb; MDS. Isobe 
et al.ll ( ASC). 


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Thysanocarpus curvipes Hook. Native winter ephemeral herb; HDWS, LDWS, MDS. db 

6646;DBa 5114, 5140. 

Thysanocarpus laciniatus Nutt. var. laciniatus Native winter ephemeral herb; HDWS, 
MDS. B 8897; DB 4553, 4582, 4604, 5664. 


C actaceae 

Carnegiea gigantea (Engelm.) Britton & Rose Native Stem Succulent; MDS. Fig. 3. 

The iconic columnar cactus of the Sonoran Desert, Carnegiea gigantea, attains the northwest¬ 
ern extent of its distribution in the southern Black Mountains. Jeff Pebworth and Erin Butler sight¬ 
ed and photographed (Fig. 3) a single plant on the east side of the Black Mountains south of Sitg- 
reaves Pass (at ca. 35° lat., -114.35° long.). 

Cylindropuntia acanthocarpa (Engelm. & J.M. Bigelow) F.M . Knuth var. acanthocarpa 

Native stem succulent; BBS, CBS, HDWS, LDWS, MDS. db 5627; Eastwood 18220 (CAS); 
Hodgson et at. 9126 (DES), 9183 (DES). Fig. 8. 

Some of the collections of this species from the southern Black Mountains have been identi¬ 
fied as var. coloradensis (L.D. Benson) Pinkava. For those noted above, the number of spines per 
areole varies between 12 and 22, which conforms to var. acanthocarpa (Benson 1982). The num¬ 
ber of “abaxial spines deflcxed” varies from five to 11, which also suggests the nominate variety 
(Pinkava 2003). Eastwood’s collection was determined by M. Baker as var. acanthocarpa in 2015. 
Thus, we treat these collections as more readily identifiable with var. acanthocarpa. 

Cylindropuntia bigelovii (Engelm .) F.M . K nuth var. bigelovii Native Stem SUCClllent; CBS, 
HDWS, MDS. DB 5988, 6438a; Hovezak & Hevly s.n (MNA). Fig. 8. 

Cylindropuntia echinocarpa (Engelm. & J.M. Bigelow) F.M. Knuth Native Stem SUCCU- 
lent; CBS, MDS. db 6438, 6467. 

Cylindropuntia leptocaulis (DC.) F.M . Knuth Native Stem SUCClllent; CBS. DB 5976. 
Cylindropuntia ramosissima (Engelm.) F.M. Knuth Native Stem Succulent; CBS. DBP 

5970. 

Echinocereus coccineus Engelm. Native Stem SUCClllent; MDS, PJW. D B H 56 87; DBP 5938. 
Fig. 8. 

Echinocereus engelm annii (Parry ex Engelm.) Lem. Native Stem SUCClllent; MDS, PJW. 
B 93 5 5 ; Eastw ood 18215 (CAS); Hovezak s.n. (MNA). 

Echinomastusjohnsonii (Parry ex Engelm.) E.M. Baxter Native Stem SUCClllent; MDS. D B 

5908. 

Ferocactus cylindraceus (Engelm.) Orcutt Native Stem Succulent; CBS, MDS. DB 6474. 
Fig. 8. 

A collection ( Hodgson 15527 at DES) cultivated at the Desert Botanical Garden (Phoenix, Ari¬ 
zona) from material collected by P. Quirk in the southern Black Mountains was identified as Fero¬ 
cactus wislizeni (Engelm.) Britton & Rose. The plant has perianth segments (all similar) that con¬ 
form to F. wislizeni, but the color and architecture of the spines are ambiguous with respect to dis¬ 
tinguishing that species from f. cylindraceus. Because f. wilsizeni, in general a more southerly 
occurring species, is not otherwise known from Mohave County, we feel further studies and/or col¬ 
lections are needed to document the presence of F. wislizeni in the Black Mountains. 

Mammillaria grahamii Engelm. (M. microcarpa Engelm. subsp. grahamii (Engelm.) Mot- 
tram Native stem succulent; MDS. db 6644, 6656.5. Fig. 8. 

Mammillaria tetrancistra Engelm. Native Stem SUCClllent; FIDWS, MDS. D B 5648.5. 
Opuntia basilaris Engelm. & J.M. Bigelow 7 var. hasilaris Native Stem Succulent; BBS, 
CBS, HDWS, MDS, PJW. B 9353; Hovezak & Hevly s.n. (MNA). Fig. 8. 


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Opuntia chlorotica Engelm. & .J.M. Bigelow Native Stem SUCClllent; HDWS, MDS. DB 

4624.5, 5662. 

Opuntia xcurvispina Griffiths Native Stem SUCClllent; MDS. DBH 5732. 

Opuntia polyacantha Haw. var. erinacea (Engelm. & J.M. Bigelow) B .D. Parfitt Native 
stem succulent; CBS, MDS. db 5934, 601 7, 6018. Fig. 8. 

C am panulaceae 

Nemacladus gianduliferus jeps. Native winter ephemeral herb; LDWS. DBa 5147. 
Nemaciadus orientalis (McVaugh) Morin Native winter ephemeral herb; CBS, LDWS. DB 

5586,6425. 

C aryophyllaceae 

Achyronychia cooperi A. Gray Native winter ephemeral herb; CBS. Harrison et al. 7552 
(ARIZ, not seen; fide SEINet (2016)). 

Silene antirrhina L. Native winter ephemeral herb; HDWS, LDWS. b 8993; DBa 5151. 
Siiene verecunda s. Watson Native perennial herb; PJW. DB 6710. 

Sperguiaria saiina j. Presl & c. Presi Native winter ephemeral herb; DOW, SSM.De 6459. 
steiiaria nitens Nutt. Native winter ephemeral herb; HDWS, MDS. db 5658, 6684; dbh 
5714. 

C henopodiaceae 

Atriplex canescens (Pursh) Nutt. var. canescens Native shrub; MDS. DB 5932, 6008. 
Atriplex elegans (Moq.) D. Dietr. var. elegans Native winter ephemeral herb; LDWS. A n dre 
3081 7 (GMDRC). 

Atriplex polycarpa (Torr.) S. Watson Native shrub; MDS. DB 6652. 

Chenopodium berlandieri Moq. var. zschackei (Murr) Graebn. Native Summer ephemer¬ 
al herb; DOW, SSM. db 6449, 6461. 

Chenopodium fremontii S. Watson Native summer ephemeral herb; MDS. db 6489. 
Grayia spinosa (Hook.) Moq. Native shrub; BBS, MDS. DB 5653. Fig. 6. 
Kraseheninnikovia lanata (Pursh) A. Meeuse & A. Smit {C eratoides lanata (Pursh) J.T. 
Howell) Native shrub; BBS, MDS, PJW. db 64 75,6476. 

Saisoia tragus l. Introduced winter ephemeral herb; MDS. db 6009. 

C leom aceae 

w isiizenia refracta Engelm. Native winter ephemeral herb; DOW, LDWS, SSM. db 5 749, 
6317. 

Convolvulaceae 

Cuscuta caiifornica Hook. & Am. Native ephemeral parasitic vine; CBS. DB 4680, 6427, 
6468. 

Plants were parasitic on Encelia farinosa aild Psorothamnus fremontii. 

C rassulaceae 

Dudleya arizonica Rose (D. pulverulenta (Nutt.) Britton & Rose subsp. arizonica (Rose) 
Moran) Native leaf succulent; HDWS, MDS. db 4570, 6669. Fig. 4. 

Crossosomataceae 

Crossosoma bigelovii S. W atson Native shrub; HDWS, MDS. B 6820, 8890; DB 4616; DBP 
5960. 


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C ucurbitaceae 

Cucurbita palmata S. Watson Native perennial vine; HDWS, LDWS. B 9328', DB 5682, 
6398. 

Marah gilensis Greene Native perennial vine; HDWS, LDWS. B 9002‘, DB 4585, 5935, 
6391,6650; DBH 5728. 

Cyperaceae 

Carex alma l.h. Bailey Native perennial herb; DOW, SSM. B 8948; DB 4528, 4628, 5986; 
DBH 5700, 5702, 5703; DBF 5952. 

Cyperus laevigatus L. Native perennial herb; DOW, SSM. db 6314, 6320. 

Cyperus odoratus l. Native summer ephemeral herb; SSM. db 6312. 

Eleocharis montevidensis Kunth Native perennial herb; DOW, SSM. B 8963. 

Eleocharis ovata (Roth) Roem. & Schult. Native summer ephemeral herb; DOW, SSM. db 

6458. 

Eleocharis palustris (L.) Roem. & Schult. Native perennial herb; SSM. DB 6315. 
Eleocharis parishii Britton Native perennial herb; DOW, SSM. DB 4511,5679, 5994. 
Eleocharis rostellata (Torr.) Torr. Native perennial herb; SSM. DBH 5764, 5770. 

Euphorbiaceae 

Chamaesyce albom arginata (Torr. & A. Gray) Small Native perennial herb; LDWS, MDS. 

DB 5628, 6319. 

Chamaesyce arizonica (Engeim.) Arthur Native perennial herb; CBS, HDWS, LDWS, 
MDS. B 8946; DB 4629, 5758, 6318. 

Chamaesyce melanadenia (Torr.) Millsp. Native perennial herb; CBS, MDS. DB 6014; 
DBH 5739. 

Chamaesyce poiycarpa (Benth.) Millsp. Native perennial herb; BBS, CBS, HDWS, MDS. 

DB 4651, 4666 , 4670; Hovezak s.n. (MNA). 

Chamaesyce setiloba (Engeim. ex Torr.) Millsp. Native ephemeral herb; HDWS. D B 4589. 
Ditaxis lanceolata (Benth.) Pax & K. H offm. Native perennial herb; CBS, MDS. DB 4707. 
Ditaxis neom exicana (M ueii. Arg.) A. Heiier Native perennial herb; HDWS, LDWS, MDS. 

DB 4539, 6411; D B a 5118. 

Euphorbia eriantha Benth. Native ephemeral herb; CBS. DB 6470. 

Euph orbia incisa Engeim. Native perennial herb; HDWS, MDS. db 4563,6708; dbh 5734 . 
Stillingia linearifolia S. Watson Native perennial herb; HDWS, LDWS. DB 4542, 6024. 
Tragia ramosa Torr. Native perennial herb; HDWS. DB 5696. 

F abaceae 

Acmispon brachycarpus (Benth.) D .D. Sokoloff ( Lotus hum istratus Greene) Native winter 
ephemeral herb; MDS. DBa 5093. 

Acmispon maritim us (N utt.) D .D. Sokoloff var. brevivexillus (Ottley) Brouillet ( Lotus sal- 
suginosus Greene var. brevivexillus Ottley) Native winter ephemeral herb; CBS, HDWS, LDWS, 
MDS. B 8936; DB 4560, 4660, 6384; D B H 5755; DBa 5112. 

Acmispon rigidus (Benth.) Brouillet ( Lotus rigidus (Benth.) Greene) Native perennial herb; 
CBS, HDWS, MDS. B 8969; DB 4622. 

Acmispon strigosus (Nutt.) Brouillet (Lotus strigosus (Nutt.) Greene var. tom entellus 
(Greene) Isley) Native winter ephemeral herb; CBS, HDWS, LDWS, MDS. b 8886 , 89 11 ,8961; 
DB 4595, 4636, 4653, 4657, 4677, 4715, 4716;DBa 5094. 

Astragalus lentiginosus Dougl. var. yuccanus M.E. Jones Native perennial herb; LDWS, 
MDS. B 9371; DB 5977, 5997, 6040. 


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Astragalus newbcrryi A. Gray var. blyae (Rydb.) Barneby Native perennial herb; MDS, 
PJW. DB 5671,6012, 6706; Pinkava et al. 10268 (ASU). 

These collections (from -114.348005° to -114.387081° long.) represent the westernmost geo¬ 
graphic extent of the variety. 

Astragalus nuttallianus DC. var. imperfectus (Rydb.) Barneby Native winter ephemeral 
herb; CBS, HDWS, MDS. B 8978 ; DB 4654, 4668 ; DBa 5095. 

Astragalus tephrodes Gray var. chloridae (M.E. Jones) Barneby Native perennial herb; 
HDWS. DB 4567. 

Dalea mollis Benth. Native winter ephemeral herb; CBS, LDWS. db 4658, 4687, 4714, 
6038. 

Lupinus brevicaulis s. Watson Native winter ephemeral herb; MDS. b 9359. 

Lupinus eoncinnus J. Agardh Native winter ephemeral herb; CBS, HDWS, MDS. db 4594, 
6394, 6683 ; DBM 5727. 

Lupinus sparsifiorus Benth. Native winter ephemeral herb; HDWS, LDWS. b 8975; DBa 
5123. 

Marina parryi (Torr. & A. Gray) Barneby Native perennial herb; CBS, LDWS, MDS. 
B 6813, 9344; DB 6385, 6429; DBa 5105. 

Medicago sativa L. Introduced perennial herb; MDS. Parfitt 2197 (ASU). 

Meliiotus indicus (L.) All. Introduced winter ephemeral herb; DOW, LDWS. b 9361; db 
5918,6442. 

Parkinsonia florida (Benth. ex A. Gray) S. Watson ( Cercidium floridum Benth.) Native 
tree; DOW, HDWS, LDWS. b 8991. 

Both species of Parkinsonia are at or very near the northern extent of their respective native 
distributions in the southern Black Mountains. Parkinsonia florida reaches its northern extent in 
Arizona in the southern Black Mountains (35.111714° lat., based on the collection noted above), 
but the species appears to occur slightly farther north in San Bernardino County, California (ca. 
35.14333° lat., based on Harris 23 72 at UVSC, not seen; Consortium of California Herbaria 2016; 
SEINet 2016). 

Parkinsonia microphylla Torr. ( Cercidium microphyllum (ToiT.) Rose & I.M. JohllSt.) 
Native tree; CBS, LDWS. db 6036. 

This collection from the southeastern portion of our flora area (34.926794° lat.), represents the 
northernmost known occurrence of this species (Consortium of California Herbaria 2016; SEINet 
2016). 

Prosopis glandulosa Torr. var. torreyana (L.D. Benson) M .C. Johnst. Native tree; DOW, 
HDWS, LDWS. B 8977; DB 6453; DBH 5753. 

Prosopis pubescens Ben th. Native tree; DOW, LDWS. db 4531, 63li. 

Psorothamnus fremontii (Torr.) Benth. var. attenuatus Barneby Native shrub; CBS, 
LDWS. B 9345, DB 6435. Fig. 6. 

Psorothamnus spin os us (A. Gray) Barneby Native tree; LDWS. DB 6016. 

This collection (at 35.082278° lat.), and one collected in the same region ( Rinks.n . at ASC), 
represent the northernmost known occurrences of the species in Arizona (SEINet 2016). These col¬ 
lections are also veiy near the northernmost native geographical extent of the species, which 
appears to be in San Bernardino County, California at 35.118363° lat. (based on Gross et al. 1203 
at UCR, not seen; Consortium of California Herbaria 2016). 

Senegalia greggii (A. Gray) Britton & Rose Native shrub; HDWS, LDWS. DB 4520. 

Senna covesii (A. Gray) H.S. Irwin & Barneby Native perennial herb; CBS, HDWS, MDS. 
DB 4587; Eastwood s. n. (ARIZ). 


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vicia ludoviciana Nutt. Native winter ephemeral herb; HDWS. b 8984; db 4617; dbh 
5701 . 

F agaceae 

Quercus turbinella Greene Native shrub; BBS, DOW, HDWS, MDS, JPW. D B 4504, 4506, 
5936, 6657, 6700; Hevly & Hovezak 45 (MNA); H odgson et al. 9141 (DES). 

Fou q u ieriaceae 

Fouquieria splendens Engelm. subsp. splendens Native shrub; CBS, MDS. DB 6661; Hov¬ 
ezak s.n. (MNA). Fig. 4. 


G arryaceae 

Garrya flavescens S. Watson Native shrub; BBS, HDWS, MDS, PJW. DBH 5709; DBP 
5965, 5966, 5967. 

G entianaceae 

Zeltnera arizonica (A. Gray) G. Mans. (Centaurium calycosum (Buckley) Fernald) Native 
ephemeral herb; SSM. dbh 5759. Fig. 12. 

G eraniaceae 

Erodium cicutarium (L.) L’Her. ex Aiton Introduced winter ephemeral herb; CBS, HDWS, 
LDWS. DB 4596. 

Erodium texanum A. Gray Native winter ephemeral herb; LDWS. db 4676. 


Iridaceae 

Sisyrinchium demissum Greene Native perennial herb; SSM. dbh 5 760. 

This collection appears to have simple stems and the outer spathe bract slightly longer than the 
inner one, thus showing some affinity with s. idahoense E.P. Bicknell of northeastern Arizona. 

Juncaceae 

juncus acutus l. subsp. acutus Native perennial herb; SSM. db 4558 

Juncus acutus L. subsp. leopoldii (Parlatore) Snogerup (j. acutus var. sphaerocarpus 

Engelm.) Native perennial herb; SSM. db 5909; dbh 5766. 

juncus bufonius L. Native summer ephemeral herb; SSM. db 4578, 5678, 6455. 

Juncus ensifolius Wikstrom var. montanus (Engelm.) C .L. Hitchcock (j. ensifolius var. 
brunnescens (Rydb.) Cronquist) Native perennial herb; SSM. b 6823; db 4510, 6323; Hovezak 
s.n. (MNA). 

juncus torreyi Coviiie Native perennial herb; SSM. db 6015. 


Kram eriaceae 

Krameria bicolor s. Watson (k. grayi Rose & Painter) Native shrub; BBS, CBS, MDS. DB 

4494, 4496, 6432. 

Krameria erecta Willd. Native shrub; BBS, CBS, LDWS, MDS. DB 4497, 5597, 5987. 


L am iaceae 

Hedeoma nana (Torr.) Briq. Native perennial herb; HDWS, MDS. B 8898; DB 6010; DBH 
5775. 

Hyptis emoryi Torr. Native shrub; CBS, HDWS, LDWS, MDS. B 8960; DB 4647. 
Marrubium vuigare l. Introduced perennial herb; DOW, HDWS. b 9369; dbh 5745. 

M onardella eplingii Elvin, A .C . Sanders & J.L. A nderson Native shlTlb (perennial herb to 


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subshrub); HDWS, MDS. B 6819, 8927', DB 4550, 4574', Elvin 6292 (UCR), 6295 (UCR), 6300 
(UCR); M. Hovezak s.n. (MNA). Fig. 6. 

This is the only plant species currently recognized as endemic to the Black Mountains. 
Although the type is from the southern Black Mountains, the species also occurs to the north of 
Union Pass (Elvin et al. 2013). 

Salvia columbariae Benth. Native winter ephemeral herb; HDWS, LDWS, MDS. b 8987; 
DB 4673, 6654; DBa 5091. 

Salvia dorrii (Kellogg) Abrams var. pilosa (A. Gray) Strachan & Reveal Native shrub; 
MDS. DBH 5746. 

Salvia mohaven Sis Greene Native shrub; CBS, HDWS, MDS. B 8926; DB 4546, 4664. 
Fig. 6. 

Scutellaria mexicana (Torr.) A.J. Paton (Salazaria m exicana Torr.) Native shrub; HDWS, 
LDWS, MDS. B 8876; DB 5599, 5982, 6485; DBH 5686. 

L iliaceae 

Allium nevadense s. Watson Native perennial herb; MDS. Elvin 6298 (UCR, not seen; col¬ 
lector’s identification verified by A. Sanders in 2016). 

Calochortus kennedyi Porter var. kennedyi Native perennial herb; MDS. DB 5637. Fig. 5. 
Dichelostemma capitatum (Benth.) Alph. Wood subsp. pauciflorum (Torr.) Keator 
(d. pule he Hum (Salisb.) A. Heller) Native perennial herb; MDS. DBa 5102; Edgar s.n. (ASC). 

Loasaceae 

Eucnide urens Parry Native perennial herb (subshrub); HDWS. dbh 5767 . Fig. 9. 
Mentzelia albicaulis (Douglas) Douglas ex Torr. & A. Gray Native winter ephemeral herb; 
CBS, HDWS, LDWS, MDS. B 9341; DB 4702, 4706, 5165, 5623, 6713; DBH 5693, 5742; DBa 
5150, 5171a. 

Mentzelia involucrata s. Watson Native winter ephemeral herb; CBS, LDWS. b 9339; db 
5591,6414; DBa 5148. Fig. 4. 

Mentzelia nitens Greene Native winter ephemeral herb; LDWS. DBa 5171. 

Mentzelia tricuspis A. Gray Native winter ephemeral herb; LDWS. DBa 5170. 

Petaionyx nitidus S. Watson Native perennial herb (subshrub); LDWS, MDS. db 5989, 
6403, 6672; Anderson 95-16 (ASU). 

Petaiony x sp. Native perennial herb (subshrub); LDWS. db 6042. 

The leaves of this collection have atypical pubescence consisting of dense, soft, white, and 
flexuose hairs, unlike the scabrous herbage of Petalonyx nitidus, which occurs elsewhere in the 
flora area. 


Lythraceae 

Lythrum californicum Torr. & A. Gray Native perennial herb; SSM. DB 4556, 6321 ; DBH 
5752. 


M alpighiaceae 

janusia gracilis A. Gray Native perennial vine; CBS, HDWS, LDWS. B 8950; db 6043, 
6436; Hovezak s.n. (MNA). 

This species has been treated recently as Cottsia gracilis (A. Gray) W.R. Anderson & C. Davis 
(Anderson and Davis 2007). 


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M alvaceae 

Abutilon incanum (Link) Sweet subsp. incanum Native perennial herb; HDWS. DB 4554. 
Abutilon parvulum A. Gray Native perennial herb; HDWS. B 8998’, DB 4512’, D BH 572 3. 
Maiva parvifiora l. Introduced winter ephemeral herb; LDWS. Andre 28583 (GMDRC). 
Sphaeralcea am bigua A. G ray Native perennial herb; CBS, HDWS, LDWS, MDS. B 9343’, 
DB 5674, 5914, 6390. 

Sphaeralcea emoryi Torr. ex A. Gray Native perennial herb; HDWS. B 8915', Hovezak 5 

(MNA). 

Sphaeralcea parvifolia A. Nelson Native perennial herb; BBS, MDS. Reeder & Reeder 8982 
(ARIZ). 

M oraceae 

Ficus carica L. Introduced tree, possibly naturalized; DOW. Hodgson et al. 9146 (DES). 


M yrsinaceae 

Anagaiiis arvensis L. ( Lysimachia arvensis (L.) U. Manns & Anderb.) Introduced winter 
ephemeral herb; SSM. db 5978. 


N yctaginaceae 

A llionia incarnata L. Native perennial herb; CBS, LDWS, MDS. D B 4694,5663,6472’, Hov¬ 
ezak s.n. (MNA). 

Boerhavia coccinea Mill. Native perennial herb; HDWS. Hovezak s.n. (MNA). 

Boerhavia wrightii a. Gray Native summer ephemeral herb; HDWS. db 4606. 

Mirabilis laevis (Benth.) Curran var. villosa (Kellogg) Spellenb. Native perennial herb; 
HDWS, MDS. B 8962; DB 4621; Eastwood 18221 (CAS); Hovezak & Iievly 37 (MNA). 

M irabilis m ultiflora (Torr.) A . G ray var. m ultiflora Native perennial herb; MDS. D B 6480. 

O leaceae 

Forestiera pubescens Nutt. var. parvifolia (A. Gray) Nesom Native shrub; HDWS, MDS. 

B 8930; DB 4523; D BH 5707. 

Fraxinus anomala Torr. ex s. Watson Native shrub (small tree); DOW, HDWS, MDS, PJW. 
DB 6007.5, 6703; DBH 5710; D BP 5961,5950. 

Menodora scabra A. Gray var. scabra Native perennial herb (subshrub); MDS. B 9347. 

M enodora scabra A. G ray var. glabrescens A . Gray ex S. Watson Native perennial herb 
(subshrub); HDWS. B 8972; Hovezak s.n. (MNA). 

Onagraceae 

Chylismia arenaria A. Nelson (Camissonia arenaria (A. Nelson) P.H. Raven) Native winter 
ephemeral herb; HDWS, LDWS, MDS. B 8906; db 5639, 6647. 

Chylismia brevipes (A. Gray) Small (Camissonia brevipes (A. Gray) P.H. Raven subsp. bre- 
vipes ) Native winter ephemeral herb; CBS, LDWS, MDS. db 4625, 4659, 4674,4713, 6659; DBa 
5115, 5143. 

C hylism ia claviform is (Torr. & Frem.) A. H eller subsp. aurantiaca (Munz) W.L. Wagner 
& Hoch (C amissonia claviform is (Tori'. & Frem.) P.H. Raven subsp. aurantiaca (S. Watson) P.H. 
Raven) Native winter ephemeral herb; LDWS. db 6045,6410; DBa 5154.5. 

C hylism ia claviform is (Torr. & F rem .) A . Heller subsp. peeblesii (Munz) W.L. Wagner & 
Hoch ( Camissonia claviform is (Torr. & Frem.) P.H. Raven subsp. peeblesii (Munz) P.H. Raven) 


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Native winter ephemeral herb; CBS, LDWS. db 4672, 4700 ; dB a 5137, 5154. 

Chylismia multijuga (S. Watson) Small ( Camissonia multijuga (S. Watson) RH. Raven) 
Native winter ephemeral herb; CBS, LDWS. db 6405, 6424. 

Ereraothera boothii (Douglas) W.L. Wagner & Hoch (C amissonia boothii (Dougl.) P.H. 
Raven) Native winter ephemeral herb; CBS, LDWS. db 6426. 

Eremothera refracta (S. Watson) W.L. Wagner & Hoch (Cam issonia refracta (S. Watson) 
P.H. Raven) Native winter ephemeral herb; CBS, HDWS, LDWS. b 9004 ; db 4689, 6406, 6415; 
DBa 5130, 5146. 

Eulobus californicus Nutt, ex Torr. & A. Gray ( Camissonia californica (Nutt, ex Torr. & 
A. Gray) P.H. Raven) Native winter ephemeral herb; HDWS, LDWS. b 8884. 

Eulobus cham aenerioides (A. Gray) W.L. Wagner & Hoch ( Camissonia chain aenerioides 

(A. Gray) P.H. Raven) Native winter ephemeral herb; HDWS, LDWS. B 9005, 8883A. 

Oenothera caespitosa Nutt, subsp. marginata (Nutt, ex Hook. & Arn.) Munz (O. caespi- 
tosa var. m a rg in a la (Nutt.) Munz) Native perennial herb; HDWS, LDWS. b 898o;db 5921 ,6395. 
Fig. 9. 

O enothera deitoides Torr. & F rem. subsp. deitoides Native winter ephemeral herb; LDWS. 

DB 6019, 6022. 

Orobanchaceae 

Castilleja chrom osa A. Nelson Native perennial herb; BBS, HDWS, MDS. db 5654, 5670, 
5922, 6673; DBP 5953; Biggs s.n. (ASC); Edgar s.n. (ASC); LehtoL 21047 (ASU). Fig. 9. 

Castilleja minor (A. Gray) A. Gray subsp. spiralis (Jeps.) T.I. Chuang & Heckard 
(c. stenantha A. Gray) Native summer ephemeral herb; SSM. db 4561 , 5675. Fig. 12. 

Orobanche ludoviciana Nutt. Native perennial herb; DOW, HDWS. db 4583; dbp 5951. 

O xalidaceae 

Oxalis albicans Kunth subsp. pilosa (Nutt.) G. Eiten ( Oxalis pilosa Nutt.) Native perenni¬ 
al herb; HDWS. B 8901; Hovezak 7 (MNA). 

Papaveraceae 

Eschscholzia californica Cham, subsp. mexicana (Greene) C. Clark Native winter 
ephemeral herb; CBS, HDWS, LDWS, MDS. DB 4586, 5640; Hovezak & Hevly s.n. (MNA). 

Eschscholzia glyptosperma Greene Native winter ephemeral herb; CBS, LDWS. db 4696, 
5606, 6i82. Fig. 5. 

Eschscholzia minutiflora S. Watson Native winter ephemeral herb; CBS, HDWS, LDWS. 

B 89 76; DB 4631,5598, 5605, 6413; DBa 5144,5158. 

Phrym aceae 

Mimulus guttatusDC. Native perennial herb; SSM. B 8994, 93 64; DB 5917; Hevly & Hov¬ 
ezak 51 (MNA). 

This species has been treated recently as Erythranthe guttata (DC.) G.L. Nesom (Barker et al. 
2012 ). 

Mimulus bigeiovii (A. Gray) A. Gray Native winter ephemeral herb; CBS, LDWS. db 
4703, 6417; DBa 5152. Fig. 9. 

This species has been treated recently as Dipiacus bigeiovii (A. Gray) G.L. Nesom (Barker et 
al. 2012). 

Mimulus cordatus Greene Native peremiial herb; MDS. Barr 67-97 (ARIZ). 

This species has been treated recently as Erythranthe cordata (Greene) G.L. Nesom (Barker 
et al. 2012). 


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Mimulus vcrbenaceus Greene (M. cardinalis Benth. var. verbenaceus (Greene) Keamey & 
Peebles) Native perennial herb; SSM. db 452 7, Fig. 12. 

This species has been treated recently as Erythranthe verbenacea (Greene) Nesom & Fraga 
(Barker et al. 2012). Other collections of Mimulus with red flowers have been made in the south¬ 
ern Black Mountains (e.g., Christy 1494, Goodding & Reeder 22, and Walker & Prigge s.n.', all at 
ASU and seen on SEINet 2016) and show morphological similarities to both M. eardinalis andM. 
verbenaceus. These two species have been treated as conspecific and the latter has been recognized 
as a variety of the former. Molecular studies (Beardsley et al. 2003) reveal the species to be dis¬ 
tinct and not phylogenetically sister to one another. Examination of materials at CAS/DS show that 
most specimens can be distinguished only by the orientation of the anther thecae: parallel to sagit¬ 
tate (i.e., those of the pair spreading from the filament at an angle < 45° and thus arrowhead shaped) 
in m , verbenaceus versus sagittate (but with the pair spreading from the filament at an angle > 45°) 
to explanate (i.e., those of a pah oriented apex to apex and ± perpendicular to filament) in m. car- 
din a Hs. Other characters sometimes used to distinguish these species (e.g., corolla tube length from 
base of sinus between calyx lobes to separation of limb from tube) varies among specimens from 
the flora area, and do not correlate with anther orientation. The collections from the flora area cited 
above all have anther thecae as noted for m . verbenaceus. 

Plantaginaceae 

Keckiella antirrhinoides (Benth.) Straw subsp. microphylla (A. Gray) N.H. Holmgren 

Native shrub; HDWS, LDWS, MDS. b 8931; db 5644, 5990. 

Maurandella an tirrhiniflora (Willd.) Rothm. ( M aurandya antirrhiniflo ra Willd.) Native 
perennial vine; HDWS, MDS. B 8967; dbh 5736. 

Mohavea confertiflora (Benth.) A. Heller Native winter ephemeral herb; CBS, LDWS. DB 

4663, 6404; DBa 5168 , Fig. 3. 

Neogaerrhinum filipes (A. Gray) Rothm. Native winter ephemeral herb (vine-like); CBS, 
LDWS. DB 4656, 4692; DBa 5145. 

Penstemou eatouiiA. Gray subsp. undosus (M .E. Jones) D.D. Keck Native perennial herb; 
HDWS, MDS. B 9000, 9358; DB 5923, 6494. 

Penstemon palm eri A . G ray Native perennial herb; PJW. D B 6705. 

Although this fruiting collection lacks flowers, the plant’s overall size (to 2 m) is more char¬ 
acteristic of P, palm eri than of P. pseudospectabilis or P. bicolor. 

Piantago ovata Forssk. Native winter ephemeral herb; CBS, HDWS, LDWS, MDS. db 

4634, 4669; DBa 5121. 

Piantago major l. Introduced perennial herb; SSM. db 5998. 

Veronica anagaiiis-aquatica L. Introduced perennial herb; SSM. B 9366; DB 6448. 

Poaceae 

Achnatherum xbloomeri (Bol.) Barkworth Native perennial herb; MDS. Michaels 2091 

(ARIZ). 

Achnatherum coronatum (Thurb.) Barkworth ( Stipa coronata Thurb. var. depauperata 
(M.E. Jones) Hitchc.) Native perennial herb; HDWS, MDS. db 4543 ; dbh 5688. 

Achnatherum hymenoides (Roem & Schult.) Barkworth (O ryzopsis hym enoides (Roem. & 

Schult.) Ricker) Native perennial herb; BBS, MDS, PJW. db 6477. 

Andropogon glomeratus (Walter) Britton, Sterns & Poggenb. Native perennial herb; 
SSM. DB 4555. 

Aristida adscensionis L. Native year-round ephemeral herb; HDWS, MDS. db 4549, 6307. 
Aristida purpurea Nutt. var. parishii (H itchc.) A llred (A. parishii Hitchc.) Native perenni- 


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al herb; HDWS, MDS. b 8999; db 4600, 5722 . 

Aristida purpurea Nutt. var. nealleyi (Vasey) Allred (A. purpurea var. glauca (Nees) A.H. 
Holmgren & N.H. Holmgren) Native perennial herb; CBS, HDWS, LDWS, MDS, PJW. b 8979, 
9333; DB 4505, 4593, 5602, 5643, 6308. 

Avena barbata Pott ex Link Introduced winter ephemeral herb; MDS. Reeder & Reeder- 
8986 (AR1Z). 

Avena fatua l. Introduced winter ephemeral herb; CBS. b 9346; db 6011 . 

Bothriochloa barbinodis (Lag.) Herter Native perennial herb; HDWS, MDS. B 8944; DB 
4575, 4627. 

Bouteloua aristidoides (Kunth) Griseb. Native summer ephemeral herb; CBS, MDS. 

Rominger 319-65 (ASC). 

Bouteloua barbata Lag. var. barbata Native winter ephemeral herb; CBS, HDWS. DB 

4619, 5983. 

Bouteloua curtipendula (M ichx.) Torr. var. caespitosa Gould & Kapadia Native perenni¬ 
al herb; HDWS, MDS, PJW. db 4571, 5974 ; dbh 5697. 

Bromus arizonieus (Shear) Stebbins Native winter ephemeral herb; HDWS. B 8903; DBP 
5956. 

Bromus berteroanus Coiia (s. trinii Desv.) Native winter ephemeral herb; HDWS, MDS. 
DB 5625, 5648; DBH 5774; Reeder & Reeder 8984 (ARIZ, ASU). 

Bromus carinatus Hook. & Am. Native perennial herb; HDWS. DBH 5694. 

Bromus diandrus Roth (b. rigidus Roth) Introduced winter ephemeral herb; DOW, MDS. 

B 9356, 9362. 

Bromus rubens L. Introduced winter ephemeral herb; common throughout, db 46 10 . 
Bromus tectorum L. Introduced winter ephemeral herb; MDS. Reeder & Reeder 8987 
(ARIZ). 

Cynodon dactylon (L.) Pers. Introduced perennial herb; SSM. db 4529, 6454. 

Dasyochloa pulchella (Kunth) W illd. ex Rydb. ( Erioneuron pulchellum (Kunth) Tateoka) 
Native perennial herb; BBS, HDWS, MDS. db 4615. 

Digitaria californica (Benth.) Henrard (Trichachne californ ica (Benth.) Chase) Native 
perennial herb; HDWS, MDS. DB 4573. 

Echinoehloa colona (L.) Link Introduced summer ephemeral herb; SSM. db 6313. 

Elymus elymoides (Raf.) Swezey subsp. elymoides ( Sitanion hystrix (Nutt.) J.G. Sm.) 
Native perennial herb; HDWS, MDS. db 6393, 6484. 

Elymus elymoides subsp. brevifolius (J.G. Sm.) Barkwor th (S itanion hystrix var. brevifoli- 

um (J.G. Sm.) C.L. Hitchc.) Native perennial herb; HDWS, MDS. db 63 10 ; dbh 5716. 

Elymus multisetus (J.G. Sm.) Burtt Davy (Sitanion jubatum J.G. Sm.) Native perennial 
herb; BBS, MDS. Gould tfe D arrow 4 294 (ARIZ). 

Hilaria rigida (Thurb.) Benth. Native perennial herb; BBS, HDWS, LDWS, MDS, PJW. db 

4684; DBH 5691. 

Hordeum murinum L. subsp. glaucum (Steud.) Tzvelev (H. glaucum Steud.) Introduced 
winter ephemeral herb; SSM. b 9363; db 5680, 6493. 

Hordeum pusiiium Nutt. Native winter ephemeral herb; SSM. dbh 5768. 

Imperata brevifolia Vasey Native perennial herb; SSM. DB 4568. 

Jarava speciosa (Trin. & Rupr.) Penail. ( Stipa speciosa Trin. & Rupr.) Native perennial 
herb; BBS, HDWS, LDWS, MDS, PJW. B 9350; DB 4524, 4611, 5655; Hevly & Hovezak 58 
(DES); Reeder & Reeder 8985 (ARIZ). 


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Leptochloa panicea (Retz.) Ohwi subsp. brachiata (Steud.) N. Snow (L. filiform is (Lam.) 
Beauv.) Native winter ephemeral herb; SSM. db 5984. 

Meiica fruteseens Seribn. Native perennial herb; HDWS, MDS. b 6818; db 6648; dbh 
5651. 

Meiica imperfecta Trin. Native perennial herb; HDWS, MDS. B 8892, 8899, 8900, 8989; 
Hovezak & Hevly 59 (MNA). 

Muhlenbergia appressa C.O. Goodd. Native winter ephemeral herb; HDWS, MDS. db 

6007, 6306; DBP 5941. 

Muhlenbergia fragili s Swalien Native summer ephemeral herb; HDWS, MDS. dbp 5945. 
Muhlenbergia microsperma (DC.) Kunth. Native winter ephemeral herb; HDWS, SSM. 

DB 4605; DBH 5757. 

Muhlenbergia porteri Seribn. Native perennial herb; BBS, HDWS, MDS, PJW. DB 4618. 
Muhlenbergia rigens (Benth.) Hitehc. Native perennial herb; DOW, SSM. B 8947. 
Panicum capiliare L. Native summer ephemeral herb; BBS, MDS. Baker ill33 (ASU). 
Paspaium diiatatum Poir. Introduced perennial herb; DOW, SSM. db 6322. 

Phragmites australis (Cav.) Trin. Introduced perennial herb; DOW, SSM. db 4515, 4557 ; 
Hovezak s.n. (MNA). 

Poa annua l. Native winter ephemeral herb; SSM. db 5999 . 

Poa bigeiovii Vasey & Seribn. Native winter ephemeral herb; HDWS. db 4576. 

Poa fendleriana (Steud.) Vasey subsp. longiligula (Seribn. & T.A. Williams) Soreng 

Native perennial herb; HDWS, MDS, PJW. db 67 11 ; dbh 5708; dbp 5955 . 

Polypogon interruptus Kunth Introduced perennial herb; DOW, SSM. db 6324, 6327. 

Poiypogon monspeiiensis (L.) Desf. Introduced winter ephemeral herb; DOW, SSM. b 8959, 
9365; DB 6444. 

Polypogon viridis (Gouan) Breistr. ( Agrostis semiverticillata (Forssk.) C. Chr.) Introduced 
perennial herb; SSM. db 6326, 6443, 6452; dbh 5763. 

Schism us arabicus Nees Introduced winter ephemeral herb; HDWS, MDS. db 4577 , 4607; 
DBH 5773. 

Schismus barbatus (L.) Then. Introduced winter ephemeral herb; HDWS, MDS. db 6418. 
Sporobolus cryptandrus (Torr.) A. Gray Native perennial herb; BBS, LDWS, HDWS, 
MDS, PJW. DB 6408; DBP 5957. 

Tridens muticus (Torr.) Nash Native perennial herb; BBS, HDWS, MDS, PJW. dbh 5738; 
DBP 5940. 

Triticum acstivum l. Introduced winter ephemeral herb; MDS. db 6491. 

This cultivated species occurs along the roadside and is a probable waif here. 

Vulpia microstachys (Nutt.) Benth. var. pauciflora (Seribn. ex Beal) Lonard & Gould 
Native winter ephemeral herb; HDWS, MDS. db 4579, 5620; dbh 5698, 5721; dbp 5944. 

Vulpia octofiora (Waiter) Rydb. Native winter ephemeral herb; CBS, HDWS, MDS. dbh 
5772. 

Polem oniaceae 

Eriastrum diffusum (A. Gray) H. Mason Native winter ephemeral herb; CBS, HDWS, 
MDS. DB 6003, 6401; DBH 5726. 

Eriastrum eremicum (jeps.) H. Mason Native winter ephemeral herb; CBS, HDWS, 
LDWS, MDS. B 8875; DB 5683, 6402, 6416. 

G ilia flavocincta A. N elson subsp. australis (V.E. Grant) A.G. Grant & V.E. Grant Native 
winter ephemeral herb; PJW. db 6696. 


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Cilia minor A.D. Grant & V.E. Grant Native winter ephemeral herb; LDWS. DBa 5126, 
5127; Andre 28644. 

Plants in the alia inconspicua (J.E. Sm.) Sweet complex are sometimes distinguishable only 
with considerable difficulty. None of the color patterns for corollas used by Porter (2014) in the 
portion of his key that includes these species are evident on the collections noted above. Both 
Daniel & Bartholomew 5126 (pinkish corollas) and 5127 (white corollas) were annotated as 
G. transm on tan a (H. Mason & A.D. Grant) A.D. Grant & V.E. Grant by A. Day in 1989. Based on 
characters that are present on these plants (i.e., leaf lobes mostly less than 1 mm wide that are point¬ 
ed toward the leaf apex and ellipsoid capsules that are only slightly exserted beyond the calyx) and 
using the keys of Wilken and Porter (2005) and Day (2002), these collections are either more con¬ 
sistent with features of G. minor (leaf lobes), or are intermediate between G. minor and o. trans- 
montana (capsules). Andre & La Doux 28644, originally determined as G. minor, shares these 
same characters and has more mature capsules with valves that are not detaching (as reported for 
G. minor). Based on the specimens and data available, we tentatively treat these collections from 
the southern Black Mountains as G. minor. Cronquist (1984) treated Gilia minor, G. sinuata, and 
G. transm on tan a as synonyms of a broadly delimited G. inconspicua . While such treatment 
appears reasonable and appealing based on morphological mtermediacics present, it has not been 
followed in the more recent accounts (e.g., Porter 2014; Wilken and Porter 2005) on which our 
species concepts and nomenclature are based. 

G ilia scopuiorum M .E. Jones Native winter ephemeral herb; CBS, HDWS, LDWS. b 8883, 
8953; DB 4709, 5125b; DBa 5138, 5155; Hovezak s.n. (MNA). 

Gilia sinuata Douglas ex Benth. (G. inconspicua Sweet var. sinuata (Douglas ex Benth.) 
Brand) Native winter ephemeral herb; CBS. Heimkamp& Heimkamp 15 74 (DES). 

Gilia steiiata a. Heller Native winter ephemeral herb; CBS, HDWS, LDWS, MDS. b 8881; 
DB 5615; DBa 5104, 5110, 5125, 5172; Hevly & Hovezak 65 (MNA). 

Langloisia setosissima (Torr. & A. Gray) Greene subsp. setosissima Native winter 
ephemeral herb; CBS, LDWS, MDS. db 5604, 5635, 6430. 

Leptosiphon aureus (Nutt.) J.M. Porter & L.A. Johnson {Lilianthus aureus (Nutt.) Greene) 
Native winter ephemeral herb; HDWS, MDS. dbh 5720 . 

Linanthus bigeiovii (A. Gray) Greene Native winter ephemeral herb; CBS, LDWS. DB 
4704; DBa 5149. 

Linanthus demissus (A. Gray) Greene Native winter ephemeral herb; CBS, LDWS. DB 

4705, 6433; DBa 5164. 

Linanthus dichotomus Benth. Native winter ephemeral herb; MDS. Hodgson et al. 3551 
(DES, UCR). 

Linanthus pungens (Torr.) J.M. Porter & L.A. Johnson Native perennial herb; PJW. DB 

6704. 

Loeseliastrum schottii (Torr.) Tim brook Native winter ephemeral herb; CBS, LDWS. Beck 
s.n. (ASC); Heimkamp & Heimkamp 1582A (UCR). 

Both collections determined as Loeseliastrum schottii share certain corolla and calyx charac¬ 
teristics withL. mathewsii (A. Gray) Timbrook. 

Microsteris gracilis (Hook.) Greene Native winter ephemeral herb; MDS. DB 6692; Biggs 
s.n.( ASC). 

Polygonaceae 

Chorizanthe brevicornu Torr. var. brevicornu Native winter ephemeral herb; CBS, LDWS. 


DB 4652, 4681,6383. 


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Chorizanthe rigida (Torr.) Torr. & A. Gray Native winter ephemeral herb; CBS, LDWS. 

DB 4675, 4693, 6380. 

Eriogonum deflexum Torr. var. defiexum Native winter ephemeral herb; CBS, HDWS, 
LDWS. D B 4682', D ha 5 / 66 ', Eastwood 18211 (CAS); H ovezak s.n . (MNA). 

Eriogonum fasciculatum Benth. var. polifolium (Benth.) Torr. & A. Gray Native shrub; 
BBS, HDWS, LDWS, MDS. b 8878; db 4599. 

Eriogonum hcermannii Durand & Hiig. var. argense (M.E. Jones) Munz Native shrub; 
HDWS, MDS. DB 4545, 6005. 

Eriogonum hookeri s. Watson Native year-round ephemeral herb; LDWS. DBa 5157. 
Eriogonum inflatum Torr. & Frem. Native perennial herb; CBS, LDWS. DB 463 7, 6303. 
Eriogonum maculatum A. Heller Native winter ephemeral herb; HDWS. DB 5624, 6392. 
Eriogonum niduiarium Coviiie Native winter ephemeral herb; CBS, LDWS. db 5610, 
6421. 

Eriogonum paimerianum Reveal Native year-round ephemeral herb; HDWS, LDWS. db 

6412; Hovezak & Iievly s.n. (MNA). 

Eriogonum reniforme Torr. & Frem. Native winter ephemeral herb; CBS, LDWS. B 9340; 
DB 4699. 

Eriogonum thomasii Torr. Native winter ephemeral herb; CBS, LDWS. b 9335; db 4667, 
4683,4698,6419. 

Eriogonum trichopes Torr. var. trichopes Native winter ephemeral herb; CBS, LDWS. 

B 9331; DB 6422. 

Eriogonum wrightii Torr. ex Benth. var. wr ightii Native shrub; HDWS. db 4544. 
Pterostegia drymarioides Fisch. & Mey. Native winter ephemeral herb; HDWS. B 8894; 
Hovezak & Hevly 24 (MNA), 

Rumex hymenosepaius Torr. Native perennial herb; HDWS, SSM. DB 4547; DBH 5730; 
DBP 5939, 5962, 5979. 

Portulacaceae 

Claytonia perfoliata Donn ex W illd. subsp. mexicana (Rydb.) John M. Mill. & K .L. 
Chambers Native winter ephemeral herb; HDWS, MDS. b 8914; db 5652. 


Prim ulaceae 

Androsace occidentaiis Pursh Native winter ephemeral herb; MDS. dbp 5946. 

R anunculaceae 

Anemone tuberosa Rydb. Native perennial herb; HDWS, MDS. db 4614. 

Aquilegia chrysantha A. Gray Native perennial herb; SSM. B 8921; DBH 5761 . Fig. 12. 
Delphinium parishiiA. Gray subsp. parishii Native perennial herb; HDWS, LDWS, MDS. 

B 9352; DB 5590, 6397. 

Delphinium scaposum Greene Native perennial herb; CBS, LDWS, MDS. db 6434; dbh 
5689. 

Myosurus cupulatus s. Watson Native winter ephemeral herb; MDS. db 6685. 

R esed aceae 

Oiigomeris linifoiia (Vahi) j.F. Macbr. Native winter ephemeral herb; SSM. db 644i;dbh 
5754. 


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R ham naceae 

Ceanothus vestitus Greene (C. greggii A. Gray) Native shrub; MDS. DB 5712, 5931. 
Rhamnus ilicifolia Kellogg ( R. crocea Nutt. var. ilicifolia (Kellogg) Greene) Native shrub; 
HDWS, MDS, PJW. B 8902; DB 4516. 

Ziziphus obtusifolia (Hook, ex Torr. & A. Gray) A. Gray var. canescens (A. Gray) M.C. 
johnst. Native shrub; LDWS, MDS. db 5633. 

R osaceae 

Coleogyne ram osissima Torr. Native shrub; BBS, MDS, PJW. DB 5668, 5 924; DB a 5103; 
DBP 5968. Fig. 6. 

Falliigia paradoxa (D. Don) Endl .ex Torr. Native shrub; HDWS. DBH 5706. Fig. 9. 

Ivesia arizonica (Eastw. ex J.T. Howell) Ertter var. arizonica ( Purpusia arizonica Eastw. 
ex J.T. Howell) Native perennial herb; HDWS. B 8 908; db 4514. 

This species occurs in isolated populations throughout its range in Arizona, California, 
Nevada, and Utah, and is considered to be of conservation concern because of its rarity (Ertter and 
Reveal 2014). 

Prunus fasciculata (Torr.) A. Gray var. fasciculata Native shrub; HDWS, MDS. B 8937; 
DB 5634; DBH 5718; DBP 5964. 

Purshia stansburyana (Torr.) Henrickson (P. mexicana (D. Don) S.L. Welsh var. stansburi- 
ana (Torr.) S.L. Welsh) Native shrub; MDS, PJW. db 5928, 6701. 

R u biaceae 

Galium aparine l. Native winter ephemeral herb; HDWS. b 8912; dbh 5719. 

Galium microphyllum A. Gray ( Relbunium microphyllum Hemsl.) Native perennial herb; 
HDWS. DB 4564; DBH 5699. 

Galium stellatum Kellogg (G. stellatum Kellogg var. eremicum Hilend & J.T. Howell) 
Native shrub; LDWS. db 5596. 

R utaceae 

Ptelea trifoliata L. Native shrub (small tree); PJW. db 6699. 

Thamnosma montana Torr. & Frem. Native shlTlb; CBS, MDS. DB 4638. Fig. 9. 

Sa licaceae 

Pop ulus frem ontii S. W atson subsp. fremontii Native tree; DOW. DB 4508, 6653. 

Salix bonplandiana Kunth Native tree; DOW. B 6816. 

Salix exigua Nutt.var. exigua Native tree (shrub); DOW, SSM. B 8965 ; D B 4509,4526; DBH 
5692. 

Salix gooddingii C.R. Ball Native tree; DOW. DB 6456, 6457; DBH 5765. 

Saururaceae 

Anemopsis ealifornica (Nutt.) Hook. & Arn. Native perennial herb. 

The inclusion of this species is based on an observation by the authors on 26 March 1989 of 
numerous plants growing in an impoundment at Fig Spring, on the eastern side of the range. 

Solanaceae 

Datura wrightii Regel Native perennial herb (shrub); LDWS. DB 4503, 6389; Hovezak s.n. 
(MNA). 

Lycium andersonii A. Gray Native shrub; HDWS, MDS. DB 5626, 5657; DBa 5090. 
Lycium cooperi A. G ray Native shlTlb; HDWS. DB 5971 ; DBP 5969. 


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Lycium exsertum A. Gray Native shrub; HDWS, LDWS, MDS. B 8951', DB 4597, 4609, 
4630, 5641,5911 . 

Daniel & Butterwick 5911 from a spring south of Thumb Butte represents the westernmost 
occurrence for this species both in Arizona and the United States (at -114.434590° long.; Consor¬ 
tium of California Herbaria 2016; SEINet 2016) for this otherwise primarily Sonoran Desert 
species. 

Lycium fremontii A. Gray Native shrub; MDS. Andre 28618 (GMDRC). 

This specimen differs from Lycium fremontii as treated in Chiang and Landrum (2009) in that 
the fruits appear to have far fewer than 40 seeds. 

Nicotiana glauca Graham Introduced tree; LDWS. DB 5614, 5996, 6039,6329. 

Nicotiana obtusifolia M. Martens & Galeotti var. obtusifolia (N. trigonophylla Dunal) 
Native perennial herb; HDWS, MDS. b 8904. 

Petunia parvifiora juss. Native winter ephemeral herb; SSM. db 6445. 

Physaiis crassifoiia Benth. Native perennial herb; HDWS, LDWS, MDS. b 8995', db 4592, 
6409', DBa 5124. 

Physaiis hederifolia A. Gray Native perennial herb; MDS. Christy 1498 (ASU). 

Solanum americanum Mill. Native perennial herb; HDWS, LDWS, SSM. DB 4530, 6463. 

Solanum douglasii Dunal Native perennial herb; MDS. M ichaels 2094 (ARIZ). 

Solanum elaeagnifolium Cav. Native perennial herb; SSM. DB 6686. 


Tam aricaceae 

Tamarix chinensis Lour. Introduced tree; LDWS. DB 4532, 6000, 6387. 

Typhaceae 

Typha latifoiia l. Native perennial herb; SSM. db 5980, 6396. 

Daniel and Butterwick 5980 from Fig Spring on the east side of the Black Mountains has near¬ 
ly contiguous spikes and carpellate flowers that lack bracts. Both of these characters support its 
placement into this species. This collection would appear to represent the westernmost occurrence 
of T. latifoiia ill Arizona (at -114.307630° long.; SEINet 2016;. Daniel and Butterwick 6396 from 
the west side of the range is sterile; however, the flat leaf blades suggest that this collection may 
also pertain to t. latifoiia. At least one other collection from the northern Black Mountains (i.e., 
Cagney et al. 9 at ASC, not seen; SEINet 2016) has been determined as t. domingensis Pers. 


Ulmaceae 

Celtis reticulata Torr. Native tree; DOW, HDWS. DB 5642, 6688; DBH 5705; DBP 5959. 

U rticaceae 

Parietaria hespera Hinton Native winter ephemeral herb; HDWS. B 8893, 8941. 
Parietaria pensyivanica Muhi.ex w iiid. Native winter ephemeral herb; HDWS. B 8940; DB 
5645. 

Verbenaceae 

Aloysia w rightii (A . G ray) A . H eller Native shrub; HDWS. B 8938. 

Verbena ciliata Benth. Native perennial herb; HDWS, MDS. Hodgson 796 (DES); Hovezak 
s.n. (MNA); Pa rfitt 2191 (ASU). 

Verbena gooddingii Briq. Native perennial herb; HDWS, MDS. B6815, 8905, 8952; DB 
4591,5666, 6013; DBa 5085;DBP 5958. 


BUTTERWICK & DANIEL: FLORA OF SOUTHERN BLACK MTS., ARIZONA 


317 


Viscaceae 

Phoradendron californicum Nutt. Native parasitic perennial herb; BBS, HDWS, MDS. db 

4498,4624, 5915, 6649. 

Plants were parasitic on Senegalia greggii. 

Phoradendron coryae Trcl. (P. villosum (Nutt.) Nutt, ex Engelm. subsp. coryae (Trel.) 
Wiens) Native parasitic perennial herb; MDS. db 6694. 

Plants were parasitic on Quercus turbinellu. 

Vitaceae 

Vitis arizonica Engelm. Native perennial vine; DOW. db 4525. 

Although rare in the southern Black Mountains, this species is locally common in portions of 
Grapevine Canyon. 

Z ygophyllaceae 

Fagonia laevis Standi. Native shrub; CBS. Richardson & Ayers 1 (ASC). 

Kaiistroem ia californica (S . W atson) Vail. Native summer ephemeral herb; MDS. DB 6490. 
Kallstroemia parviflora Norton Native Summer ephemeral herb; HDWS. Hovezak s.n. 
(MNA). 

Larrea tridentata (DC.) Coville (l. divaricata Cav. subsp. tridentata (DC.) Felger) Native 
shrub; BBS, CBS, LDWS, HDWS, MDS. db 4701,6431. Fig. 3. 

Tribuius terrestris l. Introduced winter ephemeral herb; CBS, MDS. Denver 2814 (ASC). 


Acknowledgments 

We are grateful to the Phoenix District Office of the Bureau of Land Management for logisti¬ 
cal support for fieldwork in 1980 and 1986. Subsequent field studies were funded, in part, by the 
California Academy of Sciences. For their assistance in the field, we are most thankful to our co- 
collectors: B. Bartholomew, W. Gehres, D. Hillyard, W. Hodgson, A. Phillips, B. Phillips, D. Simo- 
nis, and B. Welsh. 

The following individuals provided invaluable support: T. Columbus (identification of an East- 
wood collection of miaria rigid a ), A. Day (identification of numerous collections of G Uia),V. 
Funk and C. Kelloff (images of Palmer specimens at US from Union Pass), L. Makings (informa¬ 
tion on specimens of M im ulus at ASU, J. Pebworth (information about and a photograph of the only 
known saguaro in the Black Mountains), R. Peck (logistics and information), and A. Sanders (ver¬ 
ifying the identification of a specimen of Allium at UCR). We thank the following photographers 
who kindly made their images available for our use, either by granting us permission or by mak¬ 
ing them available on the internet via CC BY-NC 3.0 license at http://swbiodiversity.org/seinet/: 
Patrick Alexander {M entze lia involucrata and Thamnosma in o n tan a), El'in Butler (C arnegiea 
gigantea ), S. Carnahan ( Larrea tridentata), Mark Elvhl and John Anderson ( Salvia eplingii), C. 
Webber (c rayia sp in os a), Max Licher (a quite g i a c h ry s a n th a ), Keir Morse ( Coieog yne ram osissi- 
nia ), Liz Makings ( Mohavea confertiflora), Daniela Roth (Psorothamnus fremontii), and Aaron 
Schusteff (5 a l v ia moh a v e ns ii }). We thank the curators and staff of the following herbaria for loans, 
images, and/or hospitality during our visits: ARIZ, ASC, ASU, CAS, DES, GMDRC, MNA, UCR, 
US. 


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PROCEEDINGS OF THE CALIFORNIA ACADEM Y OF SCIENCES 


Series 4, Volume 63, No. 9, pp. 321-328, 6 figs. 


October 14, 2016 


Chauligenion Camelopardalis, aNe w G e n u s a n d species 

of D eepw a te r Snake Eel (A nguilliform es: O phichthidae) 

from the East C hina Sea 


John E.M cCosker 1 f * and M akoto Okamoto 2 

1 California Academy of Sciences, San Francisco, California 94118; E-mail: jmccosker@calacademy.org 

2 Seikai National Fisheries Research Institute, Nagasaki 851-2213, Japan; E-mail: jitabagu@yahoo.co.jp 

Corresponding author 

A new genus and species of ophichthid eel, ChCtltllgetTlOTl C CfflielOp CirdCtfaS M cCosker 
and Okamoto, is described based on a specimen trawled at 150 m off western Oki¬ 
nawa. It differs from other ophichthids in its elongation, its pointed tail and snout, 
low er jaw extension, anterior nostril condition, elongate jaw s, gill arch reduction, 
vertical gill openings, numerous small closely packed teeth, lack of pectoral fins, and 
its yellow ish-tan coloration overlain w ith row s of brow n spots. It is sim ilar to genera 
of the tribe O phichthini; how ever, w e are unable to assign it w ith confidence. 

Keywords: Fish taxonomy, Ophichthidae, Chauligenion Camelopardalis , genus and sp. 
nov, East China Sea. 

The snake eels of the family Ophichthidae are the most diverse and speciose family of anguil- 
liform fishes, occupying tropical and subtropical habitats including nearshore sand and mud bot¬ 
toms, rivers and streams, and estuaries and coral reefs, ranging from the sandy intertidal to mid¬ 
water depths of 800 m and to more than 1000 m in the benthos. Most, however, live shallower than 
200 m. The family was revised on the basis of its osteology, morphology and meristics by 
McCosker (1977), who recognized 49 genera and more than 200 species. Subsequent discoveries 
and generic revisions have elevated those numbers to 61 genera and nearly 300 valid species. The 
capture of a remarkably distinctive individual by a bottom trawl at 150 m in the East China Sea has 
resulted in an additional genus and species. It is so unique that we are unable to assign it a tribal 
rank, and will be unable to do so until additional material is collected. 

Methods and Materials 

Measurements are straight-line, made either with a 300 mm ruler with 0.5 mm gradations (for total 
length, trunk length, and tail length) and recorded to the nearest 0.5 mm, or with dial calipers (all other 
measurements) and recorded to the nearest 0.1 mm. Body length comprises head and trunk lengths. 
Head length (HL) is measured from the snout tip to the posterodorsal margin of the gill opening; trunk 
length is taken from the end of the head to mid-anus; maximum body depth does not include the medi¬ 
an fins. Head-pore terminology follows that of McCosker et al. (1989:257) such that the supraorbital 
pores are expressed as the ethmoid pore + pores in supraorbital canal, e.g., 1 + 3, and the infraorbital 
pores are expressed as pores along the upper jaw + those in vertical part of canal behind eye (the 
“postorbital pores’'), e.g., 4 + 2, in that frequently the last pore included along the upper jaw is part of 
the postorbital series. Osteological examination of the gill arches involved clearing and counterstain- 

321 


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ing with alcian blue and alizarin red dyes (Dingerkus and Uhler 1977). Other osteological examination 
was made from radiographs. Vertebral counts (which include the hypural) were taken from a radi¬ 
ograph. Radiographic techniques are described in Boh Ike (1989). The vertebral formula (VF) is 
expressed as the predorsal/preanal/total vertebrae (Bohlke 1982). Institutional abbreviations are as 
listed at http://asih.org/codons.pdf. The holotype of the new species is deposited in the fish collec¬ 
tion of the National Museum of Nature and Science, Tsukuba, Japan (NSMT-P). 


F am ily O phichthidae 


Genus 


Chauligenion M cCosker and Okamoto, 

(N e w Japanese nam e: K irin-uni ihebi-zoku) 


novum 


Type species: Chauligenion Camelopardalis McCosker & Okamoto 2016, by original designation. 

(Figures 1-6) 

d . * c n » s , s .— An elongate ophichthine with tail longer than head and trunk, body cylindrical, 
becoming laterally compressed in tail region; snout and tail tip pointed; lower jaw slightly extended; 
anterior nostril a hole above outer lip, without a tube; posterior nostrils along upper lip; jaws elongate, 
capable of closing completely; orbit large, in posterior half of upper jaw, its rear margin above rictus; 
gill openings low lateral, vertical; median fins elongate, not elevated; dorsal-fin origin above gill open¬ 
ing; pectoral fins absent; teeth small, conical, numerous and densely packed; preopercular pores 3; col¬ 
oration yellowish-tan overlain with rows of brown spots. 

d e s c r , t i <> n (other characters those of the single species).— An elongate (Fig. 1) ophichthine, 
with tail longer than head and trunk, cylindrical in head and trunk, becoming laterally compressed in 
tail region. Snout acute, pointed, its underside not split. Lower jaw slightly in advance of snout tip. 
Branchial basket moderately expanded. Anterior nostrils above outer lip, without a tube; posterior nos¬ 
trils along upper lip, not opening into mouth, not visible externally. Jaws elongate, capable of closing 
completely; orbit large, in posterior half of upper jaw, its rear margin in advance of the rictus. Lips 
smooth, without crenulae or barbels. Gill openings low lateral (Fig. 2), vertical, not converging for¬ 
ward, opening about twice eye diameter. Median fins elongate, not elevated, entering tail well in 
advance of pointed tail tip. Dorsal-fin origin above gill opening. Pectoral fins absent. Cephalic and lat¬ 
eral-line pores developed, supraorbital (SO), infraorbital (TO), preoperculomandibular (POP) and medi¬ 
an supratemporal pore (ST) and interorbital pores present. Teeth small, conical, numerous and dense¬ 
ly packed. Anterior etlunovomerine teeth the largest, slightly retrorse, followed by 6-7 irregular rows 
of smaller teeth, followed by a row of 25 smaller vomerine teeth. Teeth of maxillary and mandibular 
in patches, becoming smaller and more numerous posteriorly. Gill arches (Fig. 3) mostly cartilaginous: 
basibranchials 1-4 cartilaginous; hypobranchials 1—4 cartilaginous; ceratobranchial 1 cartilaginous, 
2-4 ossified, 5 minute and cartilaginous; epibranchial 1 cartilaginous, 2-4 ossified; infrapharyngob- 
ranchials 2-3 ossified; upper and lower pharyngeal tooth plates with 5 nearly regular rows of small, 
conical, densely packed and slightly retrorse teeth, the upper pharyngeal tooth plate with 5 longitudi¬ 
nal rows of ca. 15 teeth, plates closely sutured but not fused; the lower pharyngeal tooth plate with 
20-25 conical teeth in 5 irregular longitudinal rows. Teeth comparable in size and appearance to those 
of jaws.Neurocranium depressed (Fig. 4). Supraoccipital condyle developed. Suspensorium nearly ver¬ 
tical; maxillae not tapering posteriorly. Opercle and preopercle weakly developed. Branchiostegal rays 
slender, condition typically ophichthine (sensu McCosker 1977: 28-30), ca. 20-25 (as seen by radi¬ 
ograph to be) attached to outer face of each epihyal. Pectoral girdle reduced to cleithrum and thin supr- 
acleithrum.Epipleural ribs attached to precaudal vertebrae 5-11 are elongate. Other characters those of 
the single species. 


MCCOSKER & OKAMOTO: NEW SNAKE EEL FROM EAST CHINA SEA 


323 


Figure l.Holotype of Chauligenion Camelopardalis sp. nov., NSMT-P 125489, female, 407 mniTL, photographed soon 
after capture and before preservation. Arrows indicate origin of dorsal and anal fins. 


d , S t r , b u t , o n .— Known only from the type specimen, collected by trawl in 150 m from the East 
China Sea. 

etymology .— From the Greek chaulios (prominent) and genion (chin), neuter, in reference to 
its obtrusive lower jaw tip. 

remarks.— In that the new genus and species is known only from a single specimen, we were 
unable to prepare a cleared-and-stained specimen and therefore were unable to make a complete oste- 
ological examination. The holotype was preserved in formalin soon after capture and fresh tissue was 
not removed, making a genomic comparison impossible at this time. We look forward to additional 
fresh material so that those studies can be performed. 

We were able however, based on its morphometry, meristics, and the removal of its gill arches, to 
make some conclusions as to its similarity to other ophichthids. It is clearly within the subfamily 
Ophichthinae based on its hard-pointed tail tip (without caudal rays), its frontal and temporal canals, 


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Figure 2 . Head of holotype of Chauligenion Camelopardalis sp. nov., NSMT-P 125489, female, 407 mm TL, pho¬ 
tographed soon after capture and before preservation. 


5 mm 


Figure 3 . Gill arches (interior view, cut longitudinally along dorsal surface and spread laterally) of holotype of Chauli¬ 
genion Camelopardalis sp. nov., NSMT-P 125489, female, 407 mm TL. Bone is stained red and cartilage is blue. 


MCCOSKER & OKAMOTO: NEW SNAKE EEL FROM EAST CHINA SEA 


325 


Figure 4 . Radiograph of head of holotype of Chauligenion Camelopardalis sp. nov., NSMT-P 125489, female, 407 mm 
TL. 


Figure s . Schematic illustration of head of holotype of Chauligenion Camelopardalis sp. nov., NSMT-P 125489, female, 
407 mm TL. 


Figure <, . Schematic illustration of dentition (upper jaw left, lower jaw right) of holotype of Chauligenion 
Camelopardalis sp. nov., NSMT-P 125489, female, 407 mm TL. 


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its gill arch condition, and the location of its branchiostegal rays (McCosker 1977). Its other charac¬ 
teristics are so unique that we are unable to confidently place Chauligenion Camelopardalis within 
any of the currently recognized tribes. We would exclude it from the tribe Callelchelyini which has a 
body greater than its tail length, low lateral gill openings, a dorsal fin arising above the nape, and a 
short and posterior-sloping neurocranium. They too lack pectoral fins, however their pectoral girdle is 
somewhat simpler. The species of Sphagebranchini are similar in body/tail proportions to the new 
species, however their gill-opening location is entirely ventral and converging forward, their neuro¬ 
cranium is elongate, and depressed and truncate posteriorly, and their coloration is generally without 
markings and nearly uniform. The species of Bascanichthyini are similar in their gill-opening location, 
their body is usually longer than the tail, their dorsal fin usually arises on the head, they lack the third 
preopercular pore and the fifth ceratobranchial, and their coloration is nearly uniform. The species of 
the tribe Ophichthini, the most numerous within the family, are the most variable of ophichthids. The 
new species is similar to many ophichthines in some of its characteristics, but it is unique among them 
in lacking a pectoral fm and in the location and shape of its posterior nostril. The condition of its 
extended lower jaw is also unique. Whereas species of other ophichthin genera ( Echiophis , Brachyso- 
mophis, and Aplatophis) also have lower jaws extending beyond their snouts, those species all possess 
pectoral fins, slender, fewer, and longer teeth, and are more robust. Species of Herpetoichthys and 
Quassiremus are similar in general appearance (boldly spotted), but they too are stouter and have small 
pectoral fins and have fewer and more slender teeth. The new species is similar to species of Xyrias in 
the size and distribution of its maxillary dentition, and in its snout shape, anterior nostril condition, and 
spotted appearance, however it differs from all of them in lacking a pectoral fm and in other anatomi¬ 
cal characters (cf McCosker et al. 2009). 

The gill arch condition of Chauligenion Camelopardalis deserves special mention. It is clearly that 
of an ophichthid (Nelson 1966; McCosker 1977), albeit some elements are reduced from bone to car¬ 
tilage. Ophichthids are unique among anguilliforms (McCosker et al. 1989) in having: a cartilaginous 
connection between the proximal ends of the dorsal parts of the first and second gill arches; the first 
basibranchial either ossified or absent, all others are cartilaginous, rudimentary, or absent; hypo- 
branchials 1-2 ossified; the second infrapharyngobranchial is ossified; and, members of the subfami¬ 
ly Ophichthinae possess or have reduced or lost the fifth ceratobranchial, whereas those of the 
Myrophinae have lost it. Chauligenion Camelopardalis satisfies all of those conditions. 

Chauligenion Camelopardalis m c c „ * k , ,■ & okam 0 to )SP .n 0 Y. 

New English name: Giraffe- spotted snake eel; New Japanese name: Kirin-umihebi 
Figures 1-6 


M a x e » ,a L examined .— Holotype, N SMT-P 125489, 407 mm TL, female, from 27°15'00"N, 
125°45 W'E, East China Sea, west of Okinawa, captured by trawl at 150 m, on 25 Nov. 2014. 

n iMi.gsu .— An elongate ophichthine with the following characteristics: tail 53% of TL; snout 
and tail tip pointed; chin in advance of snout tip; dorsal fin arising above gill opening; median fins low; 
pectoral fin absent; eye moderate, 10.8 in head, behind middle of upper jaw; cephalic pores small but 
apparent, 3 preopercular and 3 supratemporal; teeth small, conical, numerous and densely packed on 
jaws, irregularly biserial on vomer; coloration yellowish-tan overlain with 2 rows of 70-75 eye-sized 
brown spots; vertebral formula 9/82/186. 

counts and m e a s v r e m e n t s o p i h e u o l o t v p e (in mm). Total length 407, head length 33.6, 
trunk length 158.4; tail length 215; body depth at gill openings ca. 9.3; body width at gill openings ca. 
6.9; body depth at branchial basket ca. 7; body width at branchial basket ca. 8; body depth at anus ca. 
10; body width at anus ca. 8; origin of dorsal fin 32.6; gill-opening length ca. 2.8; isthmus width ca. 


MCCOSKER & OKAMOTO: NEW SNAKE EEL FROM EAST CHINA SEA 


327 


5; snout length 7.3; tip of snout to tip of lower jaw 0.3; upper-jaw length 13.2; eye diameter 3.1; interor¬ 
bital distance 1.6. Total left-lateral pores 171, 8-10 above branchial basket, 83 before anus, last pore 
13 mm before tail tip. Vertebral formula 9/82/186. 

d e s (■ k 11 * t i o n (other characters those of the genus).— Body elongate, depth at gill openings 44 in 
TL, cylindrical in head and trunk, laterally compressed in tail region. Head and trunk 2.2 and head 12.1 
in TL. Snout acute, without a median groove on its underside. Jaws elongate, capable of closing com¬ 
pletely; center of eye above posterior ; of upper jaw. Lower jaw extended slightly beyond tip of snout. 
Anterior nostrils open well above upper lip with a small barbel in anterodorsal comer, without a tube 
or marginal extensions; posterior nostril in upper lip, not entering mouth, covered by a flap and not 
obvious, its posterior margin abutting anterior margin of orbit. Eye moderate, 10.8 in head and 4.3 in 
upper jaw, its center above posterior v, of upper jaw, its rear margin in advance of rictus. Interorbital 
region narrow, slightly elevated. Gill openings tow lateral, vertical, not converging forward, opening 
about twice eye diameter. Pectoral fins absent. Dorsal-fm origin above gill opening. Median fins low, 
extending to 'A HL before pointed end of caudal fin. 

Head pores minute but apparent (Fig. 5); supraorbital pores 1 + 3, infraorbital pores 5 + 3, 5 pores 
along mandible, 3 pores overlying preopercle, 3 supratemporal pores. A single temporal and interor¬ 
bital pore. Two infraorbital pores between anterior and posterior nostrils. Approximately 171 lateral¬ 
line pores present, 7-10 (7 left, 10 right) above branchial basket, 83 before mid-anus, last pore about 
13 mm before tail tip. 

Teeth (Fig. 6) small, conical, numerous and densely packed. Anterior ethmovomerine teeth the 
largest, slightly retrorse, followed by 6-7 irregular rows of smaller teeth, followed by a row of 25 
smaller vomerine teeth. Teeth of maxillary and mandibular in patches, becoming smaller and more 
numerous posteriorly. Maxillary patch is approximately 150 teeth which become smaller and more 
densely packed posteriorly. Mandibular patch is approximately 250 teeth, becoming smaller and more 
densely packed posteriorly. (Teeth of the single specimen too difficult to accurately examine without 
dissection.) 

Gill arches (Fig. 3) mostly cartilaginous: basibranchials 1-4 cartilaginous; hypobranchials 1-4 
cartilaginous; ceratobranchial 1 cartilaginous, 2-4 ossified, 5 minute and cartilaginous; epibranchial 1 
cartilaginous, 2—4 ossified; inffapharyngobranchials 2-3 ossified; upper and lower pharyngeal tooth 
plates with 5 nearly regular rows of small, conical, densely packed and slightly retrorse teeth, the upper 
pharyngeal tooth plate with 5 longitudinal rows of ca. 15 teeth, plates closely sutured but not fused; 
the lower pharyngeal tooth plate with 20-25 conical teeth in 5 irregular longitudinal rows. Teeth com¬ 
parable in size and appearance to those of jaws. 

Body coloration in ethyl alcohol (coloration of fresh specimen shown in Figs. 1-2) uniform yel¬ 
lowish-tan, overlain with two rows of distinctive uniformly brown spots equal to or slightly smaller 
than orbit. Upper row with 70-75 round spots arranged slightly closer than then interspaces, meeting 
but not crossing at dorsal midline. Lower row has smaller and more numerous spots centered along lat¬ 
eral line that become enlarged in mid-trunk region. Snout spots much smaller, increasing in size behind 
orbit. Median fins pale. Cheeks, chin, ventral surface and peritoneum overlain with fine brown speck¬ 
ling. Anal opening within a pale spot. Throat and palate pale. Lateral-line pores within pale spots. Tail 
tip pale like ground color. 

size .— Known only from the holotype, a 407 mm TL female with maturing ova. 

n , v mo, ov.— From the Latin Camelopardalis (giraffe), in reference to its coloration, to be treat¬ 
ed as a noun in apposition. 

D i S T r .B D x io n .— Known only from the holotype, captured by trawl at 150 m depth in the East 
China Sea. 

remarks .— The holotype of the new species is a female and becoming sexually mature. The lat- 


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eral-line pores above the branchial basket are abnormally distributed for an ophichthid; (Fig. 5); they 
appear to be in pairs rather than equally spaced. The remainder of the lateral-line pores along the flank 
appear to be normally spaced. 

The new species would not be mistaken for any known ophichthid, on the basis of its elongate 
body, extended lower jaw, minute dentition, its anterior nostril location, distinctive coloration, and its 
lack of a pectoral fm. 

We look forward to the capture of additional specimens of this remarkable snake eel. 

Acknowledgments 

We wish to thank: Mysi Hoang (CAS) for staining and clearing the gill arches of the holotype; 
Erika Garcia (CAS) for photographing the gill arches; Katherine Piatek for assistance with the fig¬ 
ures; the staff of the California Academy of Sciences (CAS) and M. Kawazu (Kyushu University) 
and G. Shinohara (NSMT) for advice and assistance with specimens; and Tomio Iwamoto (CAS), 
David Smith (USNM), and Yusuke Hibino (Mie University) for reading a draft of this manuscript. 

References 

Bohlke, E.B. 1982. Vertebral formulae of type specimens of eels (Pisces: Anguillifomies). Proceedings of the 
Academy of Natural Sciences of Philadelphia 134:31-49. 

Bohlke, E.B. 1989. Methods and Tenninology. Pages 1-7 in Fishes of the Western North Atlantic , Part Nine, 
Vol. One: Orders Anguilliformes and Saceopharyngiformes. Sears Foundation for Marine Research, Yale 
University, New Haven, Connecticut, USA. 

Dingerkus, G., and L.D. Uhler. 1977. Enzyme clearing of alcian blue stained whole small vertebrates for demon¬ 
stration of carthage. Stain Technology 52:229 232. 

McCosker, J.E. 1977. The osteology, classification, and relationships of the eel family Ophichthidae. Pro¬ 
ceedings of the California Academy of Sciences, ser. 4, 41(1): 1-123. 

McCosker, J.E., E.B. Bohlke, and J.E. Bohlke. 1989. Family Ophichthidae. Pages 254-412 in Fishes of the 
Western North Atlantic, Part Nine, Vol. One: Orders Anguilliformes and Saceopharyngiformes . Sears 
Foundation for Marine Research, Yale University, New Haven, Conneticut, USA. 

McCosker, J.E., W. Chen, and H. Chen. 2009. Comments on the snake eel genus Xyrias (Anguillifomies: 

Ophichthidae) with the description of a new species. Zootaxa 2289:61-67. 

Nelson, G.J. 1966. Gill arches of teleostean fishes of the order Anguilliformes. Pacific Science 20(4):391-408. 


PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES 


Series 4, Volume 63, No. 10, pp. 329-331, 1 fig. 


October 14, 2016 


A New Synonymy in the Genus Pison Jurine, 1808 
(Hymenoptera: Crabronidae) 


Wojciech J. Pulawski 

Department of Entomology, California Academy of Sciences, 55 Music Concourse Drive, 

Golden Gate Park, California 94118, USA; Email: wpulawski@calacademy.org 

Pison montanum Cameron, 1908 is newly synonymized with Pison atrum (Spinola, 

1808). The combined range of Pison atrum extends from the Mediterranean Basin 
into South Africa. 

My recent studies on the genus Pison have revealed a previously unnoticed synonymy that is 
discussed below. 


Pison atrum (Spinola) 

Alyson ater Spinola, 1808:253, sex not indicated. Lectotype: S, Italy: Liguria: Marassi near Genova (Torino), 
designated by de Beaumont, 1952:42. - As Pison ater (correctly; atrum): Shuckard, 1838:75 (new combi¬ 
nation). 

Pison jurinei Spinola, 1808:256, sex not indicated (as Jurini, incorrect original capitalization and termina¬ 
tion). Lectotype: <5, Italy: Liguria: no specific locality (Torino), designated by de Beaumont, 1952:42. Syn¬ 
onymized with Pison atrum by Shuckard, 1838:75, synonymy confirmed by de Beaumont, 1952:42 (as 
new synonym). 

Tachybulus niger Latreille, 1809:75, 9. Syntypes: France: Brive and Italy: Genova (destroyed). Syn¬ 

onymized with Pison atrum by Shuckard, 1838:75. 

Pison montanum Cameron, 1908:289, (as montanus , incorrect original termination). Holotype by mono- 

typy: S, Tanzania: Mount Kilimanjaro: Kibonoto (Naturhistoriska Riksmuseet Stockholm), examined. 

New synonym. 

The African species Pison montanum was included by Turner (1916), Arnold (1924), and 
Leclercq (1965) in their keys to Afrotropical Pison , but apparently none of them examined the 
holotype. Of these authors, only Leclercq noticed the species close similarity to P. atrum , although 
he did not synonymize the two names. 

Pison montanum , as traditionally interpreted, and also P allonymum Schulz, another Afrotrop¬ 
ical species, closely resemble P. atrum (the type species of the genus) in having the following com¬ 
bination of characters: presence of three submarginal cells, second recurrent vein joining second 
intersubmarginal cell or nearly so, punctures of upper frons coarse (Fig. 1), propodeum without 
longitudinal carina separating side from dorsum and posterior surface, male sternum VIII broadly, 
deeply cmarginate apically. Also, in most specimens the basal declivity of tergum I is covered with 
erect setae, and the hindcoxal dorsum has the inner carina produced into a conspicuous tooth basal- 
ly, although the tooth is reduced in size or totally absent in many males. 

Pison allonymum Schulz and P. montanum (as traditionally interpreted) differ by the position 
of the hindocelli: in P. allonymum , the ocellocular distance is larger than the hindocellar diameter; 
in P montanum it is smaller than the midocellar diameter in the female and about equal to the mido- 
cellar diameter in the male. A hitherto unnoticed difference is the shape of the clypeal lamella of 

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the female, which is obtusely, roundly angulate in P. allonymum and acutely angulate in P. mon- 
tanum. Leclercq (1965) claimed that these species differ by the degree of punctation of gastral 
terga, but I cannot confirm his statement. 

The holotype of Pison montanum has lost its head, but its remaining characters clearly indi¬ 
cate that it is either P. allonymum or P. montanum as traditionally interpreted. The absence of the 
head precludes recognition based on external characters, also because the position of the hindocel- 
li was not mentioned in the original description. However, the setae of the gonocoxite of the holo¬ 
type are as long as in the specimens determined as P. montanum in various collections and slight¬ 
ly longer than in P. allonymum, demonstrating that the species interpretation by Turner (1916), 
Arnold (1924), and Leclercq (1965) was correct. 

In addition, Pison montanum is morphologically identical to P. atrum. These two names are 
therefore synonyms. 


Figure 1 . Pison atrum (Spinola): upper frons of female. 


PULAWSKI: NEW SYNONYMY IN THE GENUS PISON 


331 


Pison atrum is known from southern Europe north to southern Germany, Austria, Czech 
Republic, Slovakia, Romania, southern Russia, Caucasus, and also from Algeria, Canary Islands, 
Morocco, Cyprus, Turkey and Iran. Pison montcnium , described from Tanzania, was subsequently 
recorded from Ethiopia, South Africa, Zambia, and Zimbabwe. The combined geographic range 
extends from the Mediterranean Basin to South Africa. 

Acknowledgment 

1 sincerely thank Dr. Hege Vardal of the Swedish Museum of Natural History for sending the 
holotype of Pison montanum Cameron, and Robert L Zuparko and David Kavanaugh, both of the 
California Academy of Sciences, for reviewing the manuscript. 

References 

Arnold, G. 1924. The Sphegidae of South Africa. Part V. Annals of the Transvaal Museum 11:1-73, pis. 
I-II. 

Cameron, P. 1908. Hymenoptera, 7. Fossores, pp. 197-296 in Y. Sjostedt, ed., Wissenschaftliche Ergebnisse 
der schwedischen zoologischen Expedition nach deni Kilimanjaro, deni Meru und den umgebenden Mas- 
saisteppen Deutsch-Ostafrikas 1905-1906 , 2. Band, Abteilung 8-14. P. Palmquists Aktiebolag, Stock¬ 
holm 844 pp., 10 pis. 

de Beaumont, .1. 1952. Sphecidae palearctiques decrits par M. Spinola (Hym.). Bollettino del Museo di Zoolo- 
gia delVUniversita di Torino 3 (1951 1952):39—51. 

Latreille, PA. 1806-1809. Genera Crustaceorum et Insectorum secundum ordinem naturalem in familias 
disposita, iconibus exemplisque plurimis explicata. Amand Koenig, Parisiis et Argentorati [= Paris and 
Strasbourg]. Tomus primus, 18 + 302 pp., 16 pis. (1806); Tomus secundus, 280 pp. (1807); Tomus ter- 
tius, 258 pp. (1807); Tomus quartus et ultimus, 399 pp. (1809). 

Leclercq, J. 1965. Sphecidae (Hymenoptera Apocrita). Subfam. Trypoxyloninae in Exploration du Parc 
National de la Garamba - Mission H. de Saeger en collaboration avec P. Baert, G. Demoulin, I. Denisoff, 
J. Martin, M. Micha, A. Noirfalise, P. Sehoemaker, G. Troupin et J. Verschuren (1949-1952), Fasc. 46 
(5):67—153. 

Shuckard, W.E. 1838 (1837). Descriptions of new exotic aculeate Hymenoptera. The Transactions of the 
Entomological Society of London 2:68-82, pi. VIII. 

Spinola, M. 1806-1808. Insectorum Liguriae species novae aut rariores quas in agro Ligustico nuper detex- 
it, descripsit et iconibus illustravit Maximilianus Spinola, adjecto catalogo specierum auctoribus jam 
e nume rata rum, quae in eadem regione passim occurrunt. Yves Gravier, Genuae. Tom 1 L,S . xvii + 160 pp., 

2 pis, (21 Oct. 1806). Tom. 11. ii+262 pp., 5 pis. [pp. ii—82: 31 December 1807; pp. 
83-206: 17 Feb. 1808, pp. 207-262: 17 March 1808). 

Turner, R.E. 1916. Notes on the wasps of the genus Pison , and some allied genera. Proceedings of the Gen¬ 
eral Meetings for Scientific Business of the Zoological Society of London 1916:591-629. 


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Series 4, Volume 63, No. 11, pp. 333-340, 6 figs. 


October 14, 2016 


A Re-evaluation of the Generic Limits of Pison Jurine, and a 
New Species of the Genus Aulacophilinus Lomholdt 
(Hymenoptera: Crabronidae: Trypoxylini) 

Arnold S. Menke 

Ammophila Research Institute, 1429 Franklin St., Bisbee, Arizona 85603 

The status of subgenera in the genus Pison Jurine is discussed. Aulacophilinus 
Lomholdt and Entomopison Menke are restored to genus (revised status). The 
importance of the mandible in generic discretion is discussed. A new species of Aula- 
cophilinus from New Guinea, A, amhlygnathus , is described. Pison weiri Naumann, 

P. caliginosum Turner, P. mandibulatum Turner and P. pyrrhicum Naumann are 
transferred to the genus Aulacophilinus (all new status). An identification key to the 
six known species of Aulacophilinus is provided. 

The Cosmopolitan genus Pison has slightly less than 200 species and is one of the largest in 
the family Crabronidae. As currently interpreted, Pison contains a broad diversity of species mor¬ 
phology. The mandible in Pison is one of the more complexly variable features of the genus, and 
it is more diverse than indicated by Bohart and Menke (1976). The morphological diversity in 
Pisotl S.l. resulted in several authors describing generic taxa: Pisonoides Smith (1857); 
Krombeiniellum Richards (1962); and Entomopison Menke (1968). In my review of New World 
Pison (Menke, 1988), these genera were considered synonyms of Pison and species groups were 
established for the species diversity in the fauna of the Western Hemisphere. 

The purpose of this paper is to discuss the importance of the mandible as a generic character 
in Aulacophilinus, Pison, and Entomopison. My studies demonstrate some generic taxa considered 
as synonyms of Pisotl are really valid genera. Species of Aulacophilinus have a mandible which 
has a unique shape shared by both sexes. Thus 1 have reinstated Aulacophilinus as a genus (revised 
status). Species of Entomopison consistently have a large externoventral notch in both sexes, while 
species of Pison S.S. lack a notch or it is weakly formed. I now recognize the Neotropical Ento¬ 
mopison as a genus (revised status). The removal of Aulacophilinus and Entomopison from Pison 
leaves the latter an assemblage that is morphologically diverse. As such it is probably paraphylet- 
ic. Further study may suggest breaking up Pison S.S. into 2 or 3 additional genera. Krombeiniellum 
is one taxon that might be elevated to genus based on the densely setose eyes, 

Lomholdt (1980) described a new genus, Aulacophilinus, from the Solomon Islands in the 
Western Pacific, that he regarded as a close relative of the New World genus Aulacophilus Smith 
because of its peliolate abdomen. Indeed, the abdomen of Aulacophilus and Aulacophilinus is very 
similar, but Lomholt’s genus lacks the many parallel pleural carinae that characterize Aulacophilus. 
I have studied Lomholdt’s type material and find that Aulacophilinus is a close relative of Pison. 
The genus differs from Pison in the form of the mandible. In both sexes the apex has a rather broad 
and distinctive cutting edge (Figs. 3, 8). Furthermore, the outer surface of the mandible is rather 
uniformly and densely covered with short setae in Aulacophilinus. This mandible is unique and not 


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found in any species of Pison. I have discovered additional undescribed species of Alllacophilinus 
in the Western Pacific with the same type of mandible, but the abdomen is not petiolate. Thus Allla- 
cophilinus contains species that are more like Pison in their general facies. The distinctive 
mandible defines Alllacophilinus. The petiolate abdomen of Alllacophilinus rennellensis is strikng 
(Fig. 1) but this condition is known in some Pison S.S.\ pistillllin Mcnke, 1988 and woji Menke, 
1988, both from New Guinea. There are other petiolate orclavatc species but the examples just list¬ 
ed are the most extreme. The genus Pisonoides was described for obliteration Smith (1857), a peti¬ 
olate species known from India and Indonesia, but abdominal structure is too variable to be used 
as a generic character. Other petiolate species like Pisonoides obliteration are the Australian Pison 
icariodes Turner, 1908, and P. difficile Turner, 1908. Antropov (1999) synonymized Aulacophili- 
nus with Pison , a genus with which it is more closely allied. But Lomholdt’s genus is here resur¬ 
rected from synonymy. 

Naumann (1990) described and keyed four Australian species of Pison, two of which were 
new, that have an apically truncate mandible as in Alllacophilinus. He called the assemblage the 
caliginosum group, and included caliginosuin Turner, mandibulatum Turner, pyrrhicum Naumann, 
and weiri Naumann. I have discovered more species with this type of mandible in New Guinea, one 
of which is described below. All of these species belong in the genus Aulacophilinus and are new 
combinations. 

Sources of material 

American Entomological Institute, Logan, Utah (David Wahl) 

Bishop Museum. Honolulu, Hawaii (Gordon Nishida) 

The Natural History Museum, London, England (Colin Vardy)(BMNH) 

Zoological Museum, Copenhagen, Denmark (Ole Lomboldt) 

Terminology and procedures 

I follow Bohart and Menke (1976) and Harris (1979) for terms. Scanning electron photographs 
were made by me at the Smithsonian SEM facility back in the 1980s. 

Genus Aulacophilinus Lomholdt, revised status 

Aulacophilinus Lomholdt, 1980. The Natural History of Rennell Island, British Solomon Islands 8:27. Mono- 
typic. 

Aulacophilinus , Antropov, 1999. Zoologischeskiy Zhurnal 78:562. Synonymized Aulacophilinus with Pison. 

When Lomholdt described his new genus, he compared it to the Neotropical genus Aula- 
cophilus Turner because of similar abdominal elongation, the presence of only two submarginal 
cells, and the absence of a carina at the lop of the propodeal side. These are superficial resem¬ 
blances, however, and in my opinion Aulacophilinus is more closely allied with Pison. In fact 
Antropov (1999) realized the true affinities of Lomholdt’s genus and synonymized it with Pison. 
The peculiar elongate abdom inal petiole of rennellensis (Fig. 1) is approached or paralleled by sev¬ 
eral species of Pison (woji Menke, pistillum Menke, icarioides Turner, difficile Turner). The broad 
mandible common to the various species discussed here is an apomorphy that in my opinion ele¬ 
vates Aulacophilinus to genus. The mandible in this genus is acuminate apically, but the inner (or 
culling) edge is broadly expanded in an angular fashion near its apical one-third (Figs. 3, 8). In 
addition, the margin of the cutting edge is narrowly polished. This smooth rim extends along the 
truncation and around the sharp mandibular apex (Figs. 3, 7). The ventral (posterior) edge of the 
mandible is fringed with very short, dense setae. Both sexes share this unique type of mandible, 


MENKE: NOTES ON WASP GENERA PISON, ENTOMOPISON, AULACOPHILINUS 


335 


Figures 1-4, Aulacophilinus rennellensis, male features. 1, lateral profile of abdomen. 2, front view of head. 3 and 4, 
mandible, clypeus, and labrurn 


and I agree with Naumann (1990) that this is an apomorphy for Aulacophilinus. Another feature 
common to species of Aulacophilinus is a narrow labrum (Figs. 4, 8) and it may prove to be an 
additional apomorphy. 

The species of Aulacophilinus are rather diverse morphologically. Two species, A. caliginosus 
and A. weiri, have two submarginal cells, the rest three; A. weiri has an omaulus on the mesopleu- 
ron and a transverse carina on the pronotal collar; two species have a crenulate ridge at the top of 
the propodeal side (A. caliginosus , A. mandibulatUS ); and among the included species the occipital 
carina varies from a complete circle to interrupted at the midventral line of the head. 


Aulacophilinus amblygnathus Menke, sp. nov. 

Types.— Hoiotype male: New Guinea: Wau, October 1969, P. Shanahan (American Entomo¬ 
logical Institute). Paratypes: one female with same data as hoiotype (American Entomological 
Institute); one female, Wau, 1200 m, Nov. 1, 1965, P. Shanahan (Bishop Museum). 

Description: Hoiotype male.— Black, sh iny except frons and antenna dull, flagellomeres 
VII-XI pale beneath; clypeus and lower frons with short, appressed silver setae that obscure sculp¬ 
ture; head and thorax with long, erect pale setae except setae in ocellar triangle shorter and black; 
gaster with short, decumbent, pale setae; wing veins dark brown, membrane slightly infuscate. 


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Figures 5-8, Aulacophilinus amblygnathus features. 5 and 6, male genitalia, lateral view, and % view respectively. 7, 
female clypeus and mandibles. 8, male clypeus, labrurn and mandibles. 


Upper interocular distance 0.56X lower interocular distance; ocellocular distance 0.22X hin- 
docellus diameter; frons with large, shallow punctures that are 1 to 2 diameters apart, interspaces 
minutely roughened, dull; flagellum without tyli or other adornments, lengths of flagellomeres I- 
III equal, each slightly more than twice as long as wide, VI-X only slightly longer than wide; 
clypeus with broad, obtusely triangular median lobe that ends in small tooth (Fig. 8), edge of lobe 
thickened laterad (ventral view); labrum narrowly quadrangular, projecting beyond clypeal edge 
(Fig. 8); mandible broadening to an obliquely, arcuately truncate apex (Fig. 8); occipital carina 
essentially complete but becoming very low at midventral line, narrowly separated from hypos- 
tomal carina. Pronotum with anteromedial round pit that is about 0.5X hindocellus diameter, sev¬ 
eral irregular transverse rugae behind pit; collar not carinate but with obtuse elevation at middle; 
scutum punctate, punctures densest anteriorly (0.5 diameters apart), 1 to 2 diameters apart on disk, 
interspaces imbricate (Harris, 1979); scutellum similarly punctate and sculptured; mesopleuron 
coarsely punctate, punctures 0.5-2 diameters apart, interspaces smooth except minutely roughened 


MENKE: NOTES ON WASP GENERA P1SON, ENTOMOPISON, AULACOPHIL1NUS 


337 


on venter, hypoepimeral area horizontally rugosopuncatate; metapleural flange narrowly lamelli- 
form; prop odeum m ostly smooth, punctate, most sparsely soon side (1-1.5 diameters apart), punc¬ 
tures of dorsum finer than those on scutum; base of dorsum with short, strong ridges, dorsum with 
series of short, transverse rugae along midline; propodeal side not delimited dorsad by carina but 
there is a vague linear series of short transverse rugae between petiole socket and spiracle. Gaster 
more finely punctate than thorax, interspaces smooth, punctures coarsest on segment I, resembling 
pinpricks on last few segments, sterna unspecialized, VIII concavely truncate apically. Male geni¬ 
talia (Figs. 5, 6). Hindtarsomere I swollen beneath subapically in lateral profile, distal third flat¬ 
tened ventrally, this area densely covered with very short white setae, distal two thirds of II cov¬ 
ered with similar setal patch ventrally, III-IV with plantulae. Forewing with three submarginal 
cells, recurrent vein I ending on submarginal cell I but nearly interstitial, recurrent vein II ending 
on submarginal cell III, separated from II by about an ocellus diameter, forewing media diverging 
beyond cu-a. Length 6.5 mm. 

Pciratypes : female (two).— Color as in male except antenna completely black, and appressed 
silver setae of clypeus sparser, not obscuring sculpture. 

Similar to male except upper interocular distance 0.47-0.48X lower interocular distance; ocel- 
locular distance 0.07-0.10X hindocellus diameter; flagellomeres VII-IX nearly 2X as long as wide; 
clypeal lobe more angular (Fig. 7); cutting edge of mandible with small indentation basad of trun¬ 
cation (Fig. 7); propodeal side not delimited dorsad in any way; hindlarsomeres I-II unmodified; 
recurrent vein I ending on submarginal cell I about ocellus diameter from II; Length 8 mm. 

Discussion.— Aulacophilinus cimblygnathus is the only member of Aulacophilinus from New 
Guinea with three submarginal cells in the forewing. Others with three submarginal cells known to 
me are mctndibulatllS and pyrrhiCUS, both of which are found in Australia. 

Etymology.— Amblygnathus, a noun, is based on the Greek words amblys (= blunt, trun¬ 
cate) and gnathos (= jaw), a reference to the peculiar mandible. 

Key to Species of Aulacophilinus 


la. Forewing with two submarginal cells.2 

lb. Forewing with three submarginal cells.4 


2a. Gastral segment I in the form of a slender petiole with tergum swollen only at apex (Fig. 1); 

propodeal side not delimited above by carina; Solomon Islands (Rennell I.). 

. rennellensis Lomholdt 

2b. Gaster sessile, segment I not forming a petiole; propodeal side sometimes delimited above by 
carina; Australia, Norfolk I ..... 3 

3a. Mesopleuron with omaulus; face, scutum and mesopleuron areolate rugulose; pronotum with 
transverse crenulate carina; northern Australia. weiri (Naumann) 

3b. Mesopleuron without omaulus; face, scutum and mesopleuron punctate; pronotum without 
transverse carina; widespread in Australia, Norfolk I. caliginosilS (Turner) 

4a. Body entirely black; New Guinea.... . amblygnathus sp. nov. 

4b. B ody not entirely black, either mandibles or legs and abdomen extensively amber colored; Aus¬ 
tralia ....... 5 

5a. Mandible amber colored, abdomen and legs black; ocellocular distance 0.75 or more times hin¬ 
docellus diameter; frons densely punctate, interspaces dull; propodeal side delimited dorsad by 
carina that may be irregular and crenulate; anterior veinlet of third submarginal cell about one 
third length of posterior veinlet; southwestern Australia. mandiblllatUS (Turner) 


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5b. Mandible black, abdomen and legs largely amber colored; ocellocular distance less than half 
diameter of hindocellus; frons coarsely punctate-areolate, interspaces shiny; propodeal side 
without carina dorsad; anterior veinlet of third submarginal cell half length of posterior vein- 
let; northern and eastern Australia. pyrrhicus (Naumann) 

Notes on the types of A. rennellensis 

I studied the holotype female and paratype male in 1989. The mandible of the female has a 
polished edge from apex along lower side as in other species of the genus, but the cutting edge 
seems to lack this. It is dull (worn?). The male, on the other hand, has the usual polished cutting 
edge (Figs. 3, 4). The male antenna has polished, elevated tyli on flagellomeres I-IV (not II-V as 
stated by Lomholdt who apparently regarded the pedicel as flagellomere I). The male clypeal edge 
is quite thick. Seen in ventral view it is about an ocellus diameter wide. The male has a tiny labrum 
just like the female (Fig. 4). It is about as wide as an ocellus diameter. In the female, the pronotum 
has a round anterodorsal pit that is smaller than ocellus. The pit is within a broad flat but not high¬ 
ly polished area bordering the anterior margin. This flat area is suggestive of the lamella of the pilo- 
sum group of Etltomopison , but it is not a lamella, nor is it polished. It is setose. Its hind margin is 
slightly elevated over a length that is about two thirds width of collar. The occipital carina is a com¬ 
plete circle that is well separated from hypostomal carina, the two separated by about an ocellus 
diameter. Inner carina of hindcoxa is diagonal and widely separated from apical U-notch for 
trochanter, outer carina absent. The hindcoxa of amblygncithlis is intermediate between rennellen¬ 
sis and mandibulatus. The hindbasitarsis of the female is ordinary. The male abdomen is shown in 
Fig. 1. 


Notes on type of A. mandibulatus (Turner), 1916 

I studied the lectotype of Turner’s species many years ago (BMNH). The following notes are 
based on this specimen as well as other material. 

The amber mandible (and clypeal apex in female) is immediately diagnostic. The propodeal 
side has a well formed carina at the lop in the female, but in the male it is somewhat irregular and 
more crenulale. The ocellocular distance is broader than in any of the other species in the mandibu¬ 
latus group (i.e., amblygncithlis and the other New Guinea species), being almost as broad as the 
ocellar diameter in the female, and broader than the ocellocular distance in the male. The frons is 
strongly swollen just above the sockets, somewhat wedgelike, forming two humps when viewed 
from below (pyrrhicus is weakly this way). The New Guinea species are merely rounded off in this 
area. The frons is closely punctate, punctures less than diameter apart, some almost contiguous, and 
the interspaces are dull. The scutum is similarly punctate, but more uniformly nearly contiguous 
and the interspaces are somewhat shiny. The mesopleural punctures are also dense, but slightly 
larger and the interspaces are even shinier. The propodeal dorsum has a median carina that is met 
by many transverse carinae that are mostly perpendicular and which extend over the middle third 
of the surface. These carinae fade into dense striatopunctation laterad. The propodeal side is shiny 
and densely punctate, their size about as on scutum (smaller than mesopleural punctures). The 
abdominal terga are shiny and densely punctate, the punctures being smaller than anywhere else. 
The hindcoxa has a long inner carina that is essentially parallel with long axis of segment, and nar¬ 
rowly separated from apical U-shaped em argination. The outer carina is strong on apical half. The 
hindbasitarsis is not straight like Cimblygnathus. It has a slight curve from base to apex in lateral 
profile (the hind face when seen in lateral profile has a concave curvature). The male hindbasitar¬ 
sis is nearly straight. The male antenna lacks tyli or other adornments. 


MENKE: NOTES ON WASP GENERA P1SON, ENTOMOPISON, AULACOPHIL1NUS 


339 


Genus Entomopison Menke, revised status 

Entomopison Menke, 1968. Los Angeles County Museum Contributions in Science (1 35):5. Type species 
Pison pilosum Smith, 1 873, original designation. 

Entomopison is restricted to the New World tropics and contains the following 11 species: alini 
Antropov, I 996, aureofaciale Strand, 1910, convexifrons Taschenberg, 1870, cooperi Menke, 1988, 
gnythos Menke, 1988, longicome Menke, 1988, Oaxaca Menke, 1988, pilosum F. Smith, 1873, 
sphaerophallus Menke, 1988, vincenti Menke, 1988, and wasbcilieri Menke, 1988.1 (Menke 1988) 
segregated them into two species groups: the convexifrons group and the pilosum group. Pison alini 
Antropov (1996), belongs in the COtl vexifrons group. Elevation of Entomopison to genus makes the 
contained species all new combinations. 

Genus Pison Jurine 

Pison Jurine in Spinola, 1808:255. Type species Pison jurini Spinola 1808, monotypic (properly jlirinei Spin- 
ola, = Alyson citer Spinola, 1808). 

Pisonoides Smith, 1857. Type species Pison obliteration F. Smithl859, monotypic. 

Krombeiniellum Richards, 1962:118. New name for Paraceramius Radoszkowski, 1 887, nec Paraceramius 
Saussure, 1854. Type species: Paraceramius koreensis Radoszkowski, 1887 (junior synonym of agile 
(F. Smith), 1 869). 

For complete list of generic synonyms see Bohart and Menke (1976). Pulawski’s online cata¬ 
log of Crabronidae lists 198 species of Pison. However, a few of these are now in Aulacophilinus, 
11 are now in Entomopison, and many others await descripton. Pison likely has more than 200 
species worldwide. 


Acknowledgments 

Michael Ohl critically reviewed the manuscript and offered helpful suggestions. Wojciech 

Pulawski helped with literature and translated Antropov’s 1999 Russian paper for me. An anony¬ 
mous reviewer also provided additonal helpful recommendations. 

Literature cited 

Antropov, A ,V. 1996, A new' species of the digger wasps of the gens [sic] Pison (Hymenoptera, Sphecidae) 
from the New World. Zologischeskiy Zhurnal 75:629-631. 

Antropov, A.V. 1999, Digger wasps of the genus AulacophilllS (Hymenoptera, Sphecidae, Trypoxylini). Zool- 
ogischeskiy Zhurnal 78:561-572. 

Bohart, R.M., and A.S. Menke. 1976. Sphecid Wasps of the World. University of California Press, Berkeley, 
vi + 695 pp. 

Harris, R. 1979. A glossary of surface sculpturing. Occasional Papers in Entomology, State of California, 
Dept, of Food and Agriculture 28:1-31. 

Lomholdt, Ole. 1980. The Sphecidae (Hymenoptera) of the Rennell and Bellona Islands. The Natural Histo¬ 
ry of the Rennell Island , British Solomon Islands 8:27-32. 

Menke, A.S. 1988. Pison in the New World: a Revison (Hymenoptera: Sphecidae: Trypoxylini). Contributions 
of the American Entomological Institute 24(3): 1 — 171. 

Naumann, ED. 1990. Description of the caliginosum species group of the genus Pison Jurine. Journal of the 
Australian Entomological Society 29:233-245. 

Richards, O.W. 1962. A Re visional Study of the Masarid Wasps (Hymenoptera, Vespoidea). British Museum 
(Natural History), London, vii + 294 pp. 

Smith, F. 1857. Catalogue of the hymenopterous insects collected at Sarawak, Borneo, Mount Ophir, Malac¬ 
ca; and at Singapore, b A. R. Wallace. Journal and Proceedings of the Linnean Society 2:42-130. 


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Turner, R.E. 1908. Notes on the Australian fossorial wasps of the family Sphegidae, with descriptions of new 
species. Proceedings of the General Meetings for Scientific Business of the Zoological Society of London 
1908:457-535. 

Turner, R.E. 1916. Notes on the wasps of the genus Pison, and some allied genera. Proceedings of the Zoo¬ 
logical Society of London 1916:591-629. 


PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES 


Series 4, Volume 63, No. 12, pp. 341^155, 48 figs. 


October 14, 2016 


Inventory of the Carabid Beetle Fauna of the Gaoligong 
Mountains, Western Yunnan Province, China: Species of the 
Tribe Trechini (Coleoptera: Caraboidea), with Descriptions of 
Four New Genera, One New Subgenus and 19 New Species. 

Thierry Deuve l , David H. Kavanaugh 2 *, and Hongbin Liang 3 

1 Institut de Systematique, Evolution, Biodiversite, 1SYEB UMR 7205 MNHN, CNRS, UPMC, EPHE, 
Museum National d’Histoire Naturelle, Sorbonne-Universites, 57 rue Cuvier, CP50, F-75231 Paris cedex 
05, France. 2 Department of Entomology, California Academy of Sciences, 55 Music Concourse Drive, 
Golden Gate Park, San Francisco, CA 94118, U.S.A. 3 Key Laboratory? of Zoological Systematics, 
Institute of Zoology, Chinese Academy of Sciences, Beijing 100101, China 
* Corresponding author. David H. Kavanaugh (dkavanaugh@calacademy.org) 


Our study of 525 specimens of trechine carabids collected during a ten-year biodi¬ 
versity inventory project in the Gaoligong Shan region of western Yunnan Province, 
China, recognized a total of 29 different species representing nine different genera. 
Four of these genera are described as new: Miliutotrechus Deuve & Kavanaugh 
(Type species: StevetlsillS minutUS Ueno), Ttecliepaphiopsis Deuve & Kavanaugh 
(Type species: T. UllipOVOSa sp. nov.), EpaphiotvecllUS Deuve & Kavanaugh (Type 
species E. fortipesoides sp. nov.), and Tvechepaphiama Deuve & Kavanaugh (Type 
species T. gaoligOIlg sp . n o v.). GdOligOllgtrechllS D c u v e & Kavanaugh is described as 
a new subgenus of QueiniiectrechllS Deuve (Type species: O. (G.) balli sp. nov.). In 
addition, 19 of the 29 species are described as new: PerileptllS piisilloides Deuve & 
Liang (type locality: China, Yunnan, Tengchong County, Hehua Township, 5.4 km S 
of Hehua at Deng man village along Daying Jiang,N24.92346°/E 98.38612°, 1105 m), 
AgonotrechllS fllgongensis Deuve and Liang (type locality: China, Yunnan, Fugong 
County, Shiyueiiang Township, 0 to 2 km W of Shibali on Shibali Road, 2300-2530 
m ), A. xiaoheishan Deuve and Kavanaugh (type locality: China, Yunnan, Longling 
County, Longjiang Township, Xiaoheishan Forest Reserve, 2067 m), Queintiec- 
trechus (S. str.) griswoldi Deuve & Kavanaugh (type locality: China, Yunnan, Gong- 
shan County, Cikai Township, 0.1 km SE of Heipu Yakou, in valley below tunnel, 
N27.77437°/E 098.4 4793°, 3270 m ), Q. (S. Str.) gOtlgshanicUS Deuve and Liang (type 
locality: China, Yunnan, Gongshan County, 11 in g zh on g lu o Township, SW slope of 
Kawakarpu Shan, on slope NE of Chukuai Lake, N 27 .98206 °/E 98.48027°, 3950 m ), 
Q. ( Gaoligongtrechus) balli Deuve and Kavanaugh (type locality: China, Yunnan, 
Gongshan County, Qiqi/Dulong divide area, N 27.69655°/E 98.45389°, 3300-3680 m), 
Trechus shiyueiiang Deuve & Kavanaugh (type locality: China, Yunnan, Fugong 
County, Shiyueiiang Townhip, 10-11 km W ofShibalion Shibali Road, 3200-3280 m), 
T. mingguangensis Deuve and Liang (type locality: China, Yunnan, Tengchong Coun- 
ty, Mingguang Township, Eighth Boundary Post Pass, N 2 5.8098 4°/E 98.620 84 °, 2287 
m ), T. qiqiensis Deuve & Kavanaugh (type locality: China, Yunnan, Gongshan Coun¬ 
ty, Qiqi Trail at No. 12 Camp, N 2 7.7 1 503 °/E 98.5 0 244 °, 2775 m ), T. pseudo qiqiensis 
Deuve and Liang (type locality: China, Yunnan, Fugong County, Shiyueiiang Town¬ 
ship, 11.5 km above Shibali on Yaping Road, N 27 .2067 6 °/E 98 .7 1 763°, 3290 m ), T. 
luzhangensis Deuve & Liang (type locality: China, Yunnan, Lush u i County, Luzhang 


341 


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Township, Pianma Road at Feag.vue Yakou [Pass], N25.9 7228° /E 98.6 8 33 6°, 3150 m ), 

T. gongshanensis D euv e & Liang (type locality: C h in a, Yu n na n, G on gsh an County, 
Dongshaofang area, N 27.69655"/E98.45389% 3300-3600 m ), T. shibalicUS D eu ve & 
Kavanaugh (type locality: China, Yunnan, Fugong County, Shiyueliang Township, 

8,5 km above Shibali on Shibali Road, North Fork of Yamu He, 

N27.1 8416°/E 98.72026% 3100 m ), Trecliepaplliopsis uniporosa D euv e & Liang (type 
locality: China, Yunnan, Lushui County, Pianma Township, 9.3 km ESE of Pianma, 

N 25.99363°/E 98.6665 1 % 2460-2470 m ), T. linisetulosa Deuve & Kavanaugh (type 
locality: China, Yunnan, Fugong County, Shiyueliang Township, 0.2 km W of 
Shibali, N27.16650°/E 09 8.77936°, 2537 m ), T. ItlOHOchueta Deuve & Kavanaugh (type 
locality: China, Yunnan, Gongshan County, Q iq i Trail at No. 12 Bridge Camp area, 

16.3 air km W of Gongshan, N27.71503°/E 98.50244°, 2775 m .), T. lUlipilosa Deuve & 

Liang (type locality: China, Yunnan, Fugong County, Shiyueliang Town ship, 8.5 km 
above Shibali on Shibali Road, North Fork of Yamu He, N 2 7.184 1 6°/E 98.72026°, 

3100 m), Epaphiotrechus fortipesoides Deuve & Kavanaugh (type locality: China, 

Yunnan, Tengchong County, Houqiao Township, 8.5 airkni NNE of Houqiao at 
Gaoshidong, N25.39858°/ E98.30533°, 2580 m ), and Trechepaphiama gdOligOllg 
Deuve & Kavanaugh (type locality: China, Yunnan, Gaoligong Shan, Lushui Coun¬ 
ty, 9.3 km ESE of Pianma, N 2 5 .9 9 3 6 3 °/E 9 8.6 6 6 5 1 °, 2460-2470 ni .), Lectotypcs arc- 
designated for Perileptus imaicus jeannei and P. pusUlus jeannei. Agonotrechiotes 

Deuve is proposed as a junior synonym of EoCtliddS Jeannei, and AgOTlOtV£Chiot£S 
longiantennatus Deuve as a junior synonym of EoCIliddS frugllis Ueno. VV c present 
keys for id en tifica t in n of adults o f a 11 species in the study area, first to genus and then 
to species for each genus, as well as nom enclatural data, diagnoses, illustrations of 
dorsal habitus and male genitalia, and information about geographical, altitudinal 
and habitat distributions w ithin the study area and overall geographical distribution 
for each species. Geographical and altitudinal distributions of the species within the 
study area are compared, and broader geographical range patterns are character¬ 
ized, and syntopic among species analyzed. A possible role of the Gaoligong Shan 
region as one source area for the present-day fauna of the Himalaya and southern 
edge of the Q in gh a i-X iza n g (Tibetan) Plateau is also discussed. 

Keywords: Coleoptera, Caraboidea, Trechini, China, Yunnan, Gaoligong Shan, taxono¬ 
my, new genera, new species, distribution, biodiversity hotspot 

The Gaoligong Shan (Gaoligong Mountains) of extreme western Yunnan Province, China, 
form the westernmost range of the Hengduan Mountains system of southeastern Xizang 
Autonomous Region (Tibet), northern and western Yunnan, and western Sichuan (Fig. 1). They 
extend north to south for more than 600 km, and, in the central part of the range, their crest forms 
the border between China and Myanmar. They also separate and form parts of the watersheds of 
two of Southeast Asia’s major rivers, the Irrawaddy and the Salween (known in China as the 
Nujiang). Elevations within the region range from a low of about 650 m in the south to more than 
5000 m in the north. Chaplin (2006) reviewed the physical geography of the region. Because of its 
geographic isolation and rugged topography, much of this area has remained less disturbed than 
most other parts of China; and previous biological exploration of the area over the past 150 years 
has revealed exceptionally high species richness, based almost exclusively on records for verte¬ 
brates (e,g., Stattersfield et al. 1998) and vascular plants (Li et al. 2000). Because of these traits, 
two large nature reserves have been established in the area, and the region has been included in the 
Three Parallel Rivers of Yunnan World Heritage Site (UNESCO 2003). 

In late 1997, the California Academy of Sciences was invited to participate in a joint project 
with the Kunming Institutes of Botany and Zoology of the Chinese Academy of Sciences to con- 


DEUVE ET AL.: C ARAB ID BEETLE FAUNA OF THE GAOLIGONG MOUNTAINS 


343 


Figure 1. Map of Asia with study region outlined; scale line = m 500 km. Modified from Wikimedia Commons, World 
Atlas of the World, at URL: <http://upload.wikimedia. 0 rg/Mnkipedia/commons/ 8 / 8 f/Whole_world_-Jand and_ 
oceans J2000.jpg> [last accessed 28 September 2016] 


duct a biodiversity inventory of the Gaoligong Mountains. Scientists from several additional insti¬ 
tutions, including the Institute of Zoology, Beijing, and Royal Botanical Garden (Edinburgh) joined 
in the collaboration. Principal target groups for the inventory included bryophytes and vascular 
plants, all vertebrate groups, and arachnids, myriapods, and insects, especially the Neuropteroidea, 
Mecoptera, and Coleoptera (the Carabidae in particular). Multidisciplinary and multi-institutional 
teams carried out biotic sampling through more than 25 separate expeditions during the period 
1998 to 2007. More than 100 reports on the project have been published to date, including partial 
results for bryophytes (e.g., Long 2006, Shevock 2005), plants (e.g., Fritsch et al. 2008, Zhou et al. 
2006), birds (Dumbacher et al. 2011), amphibians (e.g., Liu et al. 2000), fishes (e.g., Chen et al. 
2005), spiders (e.g., Miller et al. 2009, Wang et al. 2010), and carabid beetles (Kavanaugh and 
Liang 2004 and 2006; Kavanaugh and Long 1999; Liang and Imura 2003; Liang and Kavanaugh 
2006 and 2007; Liu et al. 2010 and 2011). 

Prior to the start of the project the carabid beetle fauna of the region was very poorly known. 
The fauna for the entire Hengduan region included only about 50 species (Yu 1992), and most of 
these were widespread species from low elevation areas. The region in general and the higher ele¬ 
vations in particular were virtually unexplored with respect to the carabid fauna. As a result of our 
work on this project to date, we now recognize more than 525 species occurring in the Gaoligong 
Shan, with additional species undoubtedly represented among materials for groups not yet fully 
studied. For several of the groups currently under study, (e.g., Leistus (Nebriini), Broscosoma 
(Broscini), Amerizus {Tiruka) (Bembidiini) and Aristochroa (Pterostichini), species diversity is 
much higher in this area than is known anywhere else that these taxa occur. 


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This report, on the tribe Trechini, represents the second of an intended series of treatments on 
the carabid beetle fauna of the Gaoligong Shan region, each dealing with one or more tribes or gen¬ 
era represented in the fauna. The first report (Kavanaugh et al. 2014) described the zabrine 
(Caraboidea: Zabrini) fauna of the region. Subsequent reports will appear as taxonomic work on 
each group is completed and not in any particular taxonomic or phylogenetic order. 

Trechini is a megadiverse taxon, worldwide in distribution, including more than 3,300 
described species and subspecies distributed among more than 250 genera and subgenera (Lorenz 
2005), and additional new species and genera continue to be described each year, especially from 
the eastern Palearctic Region. Trechines also occupy a diversity of habitats, from lowland marsh¬ 
es and stream edges to alpine tundra and fellfields; and they are especially diverse in caves, with 
members of many genera and species morphologically highly specialized for troglobitic life (see a 
list of the Chinese cavernicolous trechine genera in Tian et al. (2016). 

As is the case with most other terrestrial anthropod groups, the trechine fauna of the study area 
has not been well documented previously. Most of our current knowledge of the Southeast Asian 
regional fauna is from the works of Belousov and Kabak (2003, 2014a, 2014b, 2016), Deuve 
(1988, 1992a, 1992b, 1995, 2004,2005, 2013a); Jeannel (1923, 1928, 1935, 1954), Schmidt (2009) 
and Ueno (1953, 1962,1972a, 1972b, 1973, 1977, 1981, 1995, 1996a, 1996b, 1997. 1999a, 1999b), 
with additional contributions by Andrewes (1936), Casale and Magrini (2009), Deuve et al. (2015), 
Donabauer (2010), Putzeys (1870), and Ueno and Yin (1993). To date, only five trechine species 
have been recorded from the Gaoligong Shan region: Agonotrechus yunnanus Ueno (1999a), 
Stevensius mintttus Ueno (1997), Trechus asetosus Ueno (1997), Trechus unisetiger Ueno (1997), 
and Trechus unisetosus Deuve (2004). Based on our study of the material collected for the project 
and additional specimens from the region housed in other collections, we recognize a total of 29 
trechine species found to occur in the study area. These species represent nine different genera, four 
of which are described here as new; and 19 of the 29 species are also described as new. 

We present here a key for identification of adults to genus and separate keys for identification 
to species for each genus. Treatments for each species include nomenclatural data, diagnoses, illus¬ 
trations of dorsal habitus and male genitalia, and information about geographical and habitat dis¬ 
tributions within the study area and overall geographical distribution for each species. We also dis¬ 
cuss geographical distributions of the species with respect to the seven core areas and to each other, 
as well as broader geographical range patterns and the altimdinal ranges of the species. 

Materials and Methods 

The natural physiographic limits of the study area for the project are as shown in Fig. 2 and 
include areas in eastern Myanmar and southern Xizang Autonomous Region (Tibet); but we had 
permission to survey only those parts in Yunnan Province. Specialists for all taxonomic groups 
concentrated their efforts on seven core areas within the project region (Fig. 3), selected to facili¬ 
tate comparisons of possible north to south and east to west spatial differences within the regional 
biota, as well as recognition of areas of local endemism. Other areas were sampled as time and 
opportunity permitted. The entomological team made a total of 13 expeditions to the Gaoligong 
region. Our sampling sites within the region are shown in Fig. 4. Habitats included in the study area 
range from subtropical lowland rainforest to the margin of glaciers and snowfields. In all, more 
than 35,000 carabid specimens were collected during the project by using a variety of collecting 
methods, including hand collecting both day and night, beating vegetation, sifting litter with sub¬ 
sequent extraction by hand or by mini-Winkler units, and Malaise flight traps and pitfall traps. All 
specimens were sorted to morphospecies (i.e., presumptive species units based on features of exter- 


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Figures 2-4. Fig. 2. Map showing natural extent of study area, colored in green (however, sampling was permitted 
only in those portions in Yunnan Province . Fig. 3. Map showing location of core sampling areas. Fig. 4. Map showing loca¬ 
tions of all entomological sampling sites. 


nal structure and male and female genitalic traits) and detailed systematic studies of taxonomic 
groups are ongoing. 

The present study is based on the examination of 525 specimens of trechine species from the 
Gaoligong Shan region. Specimens acquired during the project have been divided among and are 
deposited in the collections of our home institutions. Codens used throughout this report for col¬ 
lections in which specimens, including primary types, are deposited are as follows: 

BMNH British Museum (Natural History), London, United Kingdom 
CAS California Academy of Sciences, San Francisco, U.S.A. 

IOZ National Zoological Museum of China, Institute of Zoology, Beijing, China 

MNHN Museum National d’Histoire Naturelle, Paris, France 
NSMT National Science Museum (Natural History), Tokyo, Japan 

SCAU South China Agricultural University, Guangzhou, China 

Measurements.— The following measurements were recorded: body length (BL), measured 
longitudinally from the apex of the mandibles to the apex of the elytra; pronotal width (PW), meas- 


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ured transversely at the widest point on the pronotum; and pronotal length (PL), measured longi¬ 
tudinally between the anterior and posterior pronotal margins along the midline. The means of 
these measures were combined in the ratio, PW/PL, as an indicator of pronotal shape for each 
species. 

dissections of male genitalia.— Urite IX (the “ring sclerite”) and acdeagus were extract¬ 
ed as a unit manually by using a sharp point to slit the intersegmental membrane between segments 
VIII and IX. The dissected assemblage was then placed in a dilute solution of KOH at room tem¬ 
perature for 24 hours. After rinsing in water, the dissections were dehydrated using 95% EtOH, 
then urite IX was separated from the acdeagus by severing the connecting membranes. The parts 
were then mounted in a Euparal preparation between two small coverslips over a hole in a small 
card and pinned beneath the specimen. 

Illustrations.— Digital images of whole specimens and particular structures were taken 
using a Leica imaging system including an M165C dissecting microscope, DFC550 video camera, 
and two KL1500 LCD light sources. Stacked images were captured and combined into single mon¬ 
tage images using the Leica Application Suite V4.2.0. Plates of images were created using Adobe 
Photoshop CS5. Distribution maps for each species were generated from geographical coordinate 
data maintained in a Biota Version 3.0 database (Colwell 2012) using the ArcMap program in 
ArcGIS for Desktop Version 10.2 software from Esri. 

Geographical Coordinate Data .— All geographical coordinate data are presented in dec¬ 
imal degree format, with the first entry degrees North and the second degrees South, separated by 
“/”. Exceptions to this format include verbatim label data only. 

Treatment seor new species.— For all new species, label data for all specimens of the type 
series are quoted verbatim (between quotation marks), with data for multiple labels separated by 
“/” and any editorial additions included between brackets (“[-]”). 

Taxonomy 

Adult specimens of trechine species represented in the Gaoligong Shan region can be distin¬ 
guished using the keys provided in this paper. Each key is provided only for distinguishing mem¬ 
bers of taxa (different genera or species) represented in this region and may not be appropriate for 
more general use. 

Key for Identification ofAdults of Trechine Genera of the Gaoligong Shan Region 

1. Dorsal surface covered with fairly long pubescence; eyes also pubescent. 

. Perileptus Schaum 

1 ’ Dorsal surface glabrous except for isolated fixed setae typical for trechines; eyes also 

glabrous.2 

2(1’) Mentum and submentum fused or at least partially fused.3 

2’ Mentum and submentum not fused, separated by a distinct suture.5 

3(2) Body length (BL) less than 3 mm, the protibiae not furrowed ... Minutotrechus gen. nov. 

3’ BL more than 4 mm, the protibiae longitudinally furrowed.4 

4(3’) Fully winged, elytra with discal striae distinctly impressed. Agonotrechus Jeannel 

4’ Apterous, elytra with discal striae effaced. Queinnectrechus Deuve 

5(2’) Form slender and flattened, the pronotum relatively small; antennae long and slender, 

extended nearly to the apical one-fourth of the elytra. Eocnides Jeannel 

5’ Form more compact and convex, the pronotum average size; antennae shorter, extended 
only to the basal one-fourth of the elytra or less.6 


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6(5’) Right mandible with the anterior point of the retinaculum free and displaced distally to fonn 
a separate tooth; elytra without or with up to two discal setae, elytral surface with or with¬ 
out iridescence.7 

Right mandible tridentate, but premolar tooth fused with the retinaculum to fonn a trifid 
molar with the anterior point not displaced distally; elytra with two or three discal setae; 
elytral surface not iridescent. Treclms Clairville 

Tempora pubescent; size small, BL less than 3.8 mm; pronotum with basal angles obtuse and 

rounded; elytra without (in T. asetosa) or with only a single discal seta near stria 3. 

. Trechepaphiopsis gen. nov. 

Tempora glabrous; size larger, BL more than 4.0 mm; pronotum with basal angles sharp and 
rectangular; elytra with either two discal setae or none.8 

Elytra with two discal setae near stria 3, elytral disc slightly flattened along the length of the 

median suture. Epaphiotrechus gen. nov. 

Elytra without discal seta, elytral disc evenly convex. Trechepaphiama gen. nov. 

Genus PerileptllS Schaum, 1860 

Perileptus Schaum, 1860:663. 

type species. — Carabus areolatus Creutzer, 1799. 

diagnosis. — Adults of this genus (Figs. 5a, 6a) can be recognized by the following combi¬ 
nation of character states: size small to medium (BL = 2.0 to 3.5 mm), fully winged, dorsal surface 
covered with more or less long pubescence; eyes large, convex, pubescent; frons flat, frontal fur¬ 
rows wide and deep, attenuated posteriorly; terminal palpomeres slender, attenuated apically; men- 
tum free, not fused with submentum; submentum with 10 or 12 setae; antennal scape distinctly 
pubescent; pronotum narrowed basally, with base broadly projected posteriorly, median longitudi¬ 
nal furrow deeply and sharply defined; elytra elongate and flattened, recurrent stria indistinct; legs 
short, protibiae without longitudinal furrows. 

Geographical distribution. — Perileptus is a moderately diverse genus with about 50 
described species arrayed in four subgenera (Lorenz 2005). It is represented in the Palearctic, Ori¬ 
ental, Afrotropical, Australian and Neotropical Regions. The study area is within the previously 
known range of this genus. 


6 ’ 

7(6) 

T 

8(7’) 


Key for Identification of Adults of PerileptllS Species of the Gaoligong Shan Region 

1. Body size larger (BL = 2.6 to 2.8 mm); body color reddish, apex of the elytra darker; prono¬ 
tum distinctly convex, with sparse setiferous punctures, basal angles markedly projected lat¬ 
erally; elytra smooth, shiny, microsculture effaced, discal striae coarsely punctuate. 

... Perileptus imaicus Jeannel 

Body size smaller (BL = 2.3 mm), body color pale yellowish brown, elytra concolorous; 
pronotum less convex, with dense setiferous punctures, basal angles less projected laterally; 
elytra dull, irregular isodiametric microsculpture evident, discal striae distinctly but finely 
punctuate. Perileptuspusilloides sp. nov. 

Perileptus imaicUS Jeannel, 1923 

(Figs. 5, 35a, 45-48) 

Perileptus imaicus Jeannel 1923. Lectotype, here designated, a male, in BMNH, labeled: “Lectotype” [red 
label];/ “W. Almora, Kumoan U.P., India H.G.C.’V P. imaicus Jeannel det.’V “H.E. Andrewes Coll., B.M. 


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1945-97.”/ “Perileptus imaicus Jeannel, lectotype, Deuve & Kavanaugh des. 2016”. Type locality: India, 
Uttar Pradesh, Kumoan, West Almora. Paralectotypes (a total of 8), labeled same as lectotype, in BMNH 
and MNHN. 

diagnosis. — Adults of this species (Fig. 5a) can be distinguished from those of all other 
species in the region by the following combination of character states: size small (BL = 2.6 to 2.8 
mm), fully-winged; body color reddish with apical part of elytra darker; microsculpture of elytra 
effaced; dorsal pubescence sparse and long; pronotum slightly transverse (ratio PW/PL = 1.32), 
distinctly convex, with lateral margins markedly rounded in anterior half, straightened just anteri¬ 
or to basal angles, the latter acute and sharp and laterally projected, median basal area coarsely 
punctate; elytra shiny, moderately flattened, with striae coarsely punctate. 

Comments. — Perileptus denticollis Jeannel (1923), known from the Dali region of Yunnan 
Province, is probably conspecific with P imaicus because there appear to be no morphological fea¬ 
tures to distinguish members of these two nominal species. Members of Perileptus davidsoni 
Deuve (1989), described from the Kathmandu area in Nepal, also are externally similar in form and 
structure to those of P imaicus, but the apex of the median lobe of the aedeagus of males of 
P davidsoni is more broadly rounded (more narrowly rounded in P. imaicus males (Fig. 5b)). 

h abitat distribution. — Members of this species have been found in daytime on sandy flats, 
in gravel and under stones along the open, unshaded banks of small to large streams running 
through agricultural areas and other disturbed areas at elevations ranging from 680 to 2030 m (Fig. 
35a). Most specimens were driven from their hiding places by splashing the banks with water from 
the stream. At the collecting site in Longyang County (see below), members of this species were 
found syntopic with members of Perileptus pusilloides sp. nov. 

G eographical distribution w ithin the G AonGONG Shan . — Fig. 5c. We examined a total 
of 84 specimens (25 males and 59 females) from the following localities: Gongshan County: 
Bingzhongluo Township (Xiao Shangla He at Shuangla Village, N27.975147E98.65502°, 1550 m, 
9 October 2002, Stop # DHK-2002-043, D.H. Kavanaugh, H.B. Liang, & W.D. Ba collectors 
[2 males and 30 females; CAS, lOZ, MNHN]). Longyang County: Bawan Township (Kunhong He 
at Bingmen, N25.090657E98.83721°, 680 m, 22 May 2005, Stop# 2005-019, D.H. Kavanaugh & 
H.B. Liang collectors [12 males and 14 females; in CAS, IOZ)], 1 June 2005, Stop# 2005-039, 
D.H. Kavanaugh & H.B. Liang collectors [11 males and 11 females; CAS, IOZ]). Tengchong 
County: Jietou Township (0.75 km N of Dahetou Ligganjiao on Longtang He, N25.746227 
E98.69612°, 2030 m, 18 May 2006, Stop # DHK-2006-029, D.H. Kavanaugh, R.L. Brett, & H.B. 
Liang collectors [3 females; in CAS, IOZ]; stream 0.7 km N of Jietou, N25.431287E98.64773°, 
1564 m, 22 May 2006, Stop # DHK-2006-036A, D.H. Kavanaugh & R.L. Brett collectors 
[1 female; CAS]). These localities are at low to middle elevations in Core Areas 2, 6 and 7. 

Overall geographical distribution. — Fig. 45. This species currently is known from the 
Himalayan region of India from Himachal Pradesh to Darjeeling and Sikkim. It has also been 
reported from Yunnan under the name P denticollis Jeannel (1923) (see Ueno 1996b) 

Perileptus pusilloides Deuve and Liang sp. nov. 

(Figs. 6, 46-48) 

Type material.— Holotype, a male, in IOZ, labeled: “CASENT 10392597 “CHINA, Yun¬ 
nan, Tengchong County, Hehua Township, 5.4 km S of Hehua at Dengman village along Daying 
Jiang, N24.923467E98.38612°,”/ “1105 m, 2 June 2006, Stop # DHK-2006-053, D.H. Kavanaugh, 
R.L. Brett & D.Z. Dong collectors”/ “HOLOTYPE Perileptus pusilloides Deuve & Liang, sp. nov. 
designated 2016” [red label]. Para types (a total of 4): 2 males and 1 female (in CAS, IOZ, MNHN) 


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Figure 5. Perileptus imaicus Jeannel; a. Dorsal habitus (CASENT1036287). b. Median lobe of aedeagus of male 
(CASENT10363020), left lateral aspect, c. Map of locality records (red circles) for P. imaicus in the Gaoligong Shan region. 
Scale lines a, b = 0.5 mm, c = 100 km. 


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labeled same as holotype except first label “CASENT 1039257” and “CASENT 1039260” and 
“CASENT 1039258”, respectively; 1 female (in IOZ) labeled: “CASENT 1036294”/ “CHINA, 
Yunnan, Longyang County, Bawan Township, Kunhong He at Bingmen, N25.09065°/ 
E98.83721 0 ,”/ “680 m, 1 June 2005, Stop# 2005-039, D.H. Kavanaugh & H.B. Liang collectors”. 
All paratypes also bear the following label: “PARATYPE Perileptus pusilloides Deuve & Liang, 
sp. nov. designated 2016” [yellow label]. 

type locality.— China, Yunnan, Tengchong County, Hehua Township, 5.4 km S of Hehua 
at Dengman village along Daying Jiang, N24.92346°/E98.38612°, 1105 m. 

Etymology. — The species epithet, pusilloides , is a combination of the species epithet, 
pusillus, and the Greek suffix, -eiSt)g (transliterated into Latin as - oides ), meaning resembling, in 
reference to the similarity of members of this species to those of T. pusillus Jeannel (1923). 

diagnosis. — Adults of this species (Fig. 6a) can be distinguished from those of all other 
species in the region by the following combination of character states: size very small (BL = 2.3 
mm), fully-winged; body color light yellowish brown, elytra concolorous throughout; microsculp¬ 
ture of elytra irregularly isodiametric and moderately impressed; dorsal pubescence dense and 
rather short; pronotum slightly transverse (ratio PW/PL = 1.32) only slightly convex, with lateral 
margins moderately rounded in anterior half, straightened just anterior to basal angles, the latter 
acute and sharp but small, median basal area mgulose; elytra alutaceous, distinctly flattened, stri¬ 
ae finely punctate. 

Description. — Size very small, BL = 2.3 mm. Color of dorsum pale yellowish brown, head 
and pronotum slightly pale reddish in some specimens, appendages pale yellow, antennomeres 3 to 
11 slightly darker yellowish brown; dorsal surface covered with dense but short pubescence. 

Head. Relatively large, with eyes large and convex. Tempora very short. Frons flattened, near¬ 
ly smooth, shiny but finely punctate; frontal furrows arcuate and deeply impressed to posterior 
margin of tempora; two pairs of supraorbital setae present and distinctly longer than setae of pubes¬ 
cence. Labrum broad with apical margin distinctly emarginate or concave. Mandibles small, slen¬ 
der. Mentum deeply concave, with median tooth broad and truncate. Submentum with a transverse 
row of ten setae anteriorly. Gula broad. Antennae of moderate length, antennomeres 2 and 3 of 
equal length. 

Pronotum. Slightly transverse (ratio PW/PL = 1.32) and flattened, widest at anterior one- 
fourth, with lateral margins markedly rounded anteriorly, less so posteriorly, briefly and deeply sin¬ 
uate just anterior to basal angles, the latter acute and sharp but small; basal margin projected pos¬ 
teriorly as a short, truncate lobe medial to lateral sinuations. Disc moderately punctate, flattened 
medially and slightly convex laterally; median basal area rugulose, only faintly defined anteriorly; 
median longitudinal impression distinct throughout, narrow anteriorly, widened toward base. Lat¬ 
eral margination slender throughout, slightly reflexed dorsally. One lateral seta (at anterior one- 
fifth) and one basolateral seta (on basal angle) present on each side. 

Elytra. Alutaceous, moderately elongate, with humeri very distinct, rectangular but broadly 
rounded; elytral disc flattened, only moderately convex laterally; lateral margination slender and 
slightly reflexed dorsally; basal margin terminated medially at the origin of stria 5; striae 1 to 4 or 
5 dinstinctly impressed and moderately punctate, striae 5 or 6 to 8 effaced. Recurrent stria absent. 
Umbilicate setal series with setae of humeral group equidistance from each other, those of the 
median group inserted posterior to middle. 

Legs. Short. Male protarsomeres 1 and 2 dilated and apicomedially toothed. 

Venter. Ventral surface of head and thoracic pleurae, prosternum, metastemum, metepisterna 
and abdominal ventrites punctate. Pubescence denser on abdominal ventrites than on thoracic ven¬ 
ter. 


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Figure 6. Perileptus pusilloides sp. nov.; a. Dorsal habitus (CASENT1039260). b. Median lobe of aedeagus of male 
(CASENT1039259), left lateral aspect, c. Map of locality records (red circles) for P. pusilloides in the Gaoligong Shan 
region. Scale lines a, b = 0.5 mm, c = 100 km. 


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Male aedeagus. Median lobe (Fig. 6b) with apex narrowly lobate, apically rounded. 

Comments.— Based on features of form and structure, this species appears to be closely relat¬ 
ed to P pusillus, described from northern Vietnam. Its members can be distinguished from those of 
the latter in having their elytra wider, less shiny, more alutaceous, and with more finely punctate 
striae, and the pronotum relatively larger, with a less smooth surface, and with the greatest prono- 
tal width more anterior. The type series of P. pusillus originally consisted of ten syntypcs, seven of 
which are in the Jeannel Collection in MNHN, each with a “Type” label. Among these, six speci¬ 
mens match Jeannel’s description of P. pusillus veiy closely. The seventh specimen, a male, instead 
may be a member of our new species, P pusilloides. Consequently, in order to properly establish 
the identity to P. pusillus, we here designate a lectotype as follows: a male, in MNHN, labeled: 
“Hoa Binh, Tonkin”/ “Perileptus pusillus Jeannel, lectotype, design. 2016, ex coll. R. Jeannel, in 
coll. MNHN, Paris”. The remaining six MNHN syntypes are all paralectotypes of P. pusillus, 
including the lone possible male of P. pusilloides, which we are not including in the type series of 
the latter species. 

had itat distribution .— Members of this species have been found in daytime on sandy flats, 
in gravel and under stones along the open, unshaded banks of small to large streams running 
through agricultural areas with subtropical crops at elevations ranging from 680 to 1105 m. Most 
specimens were driven from their hiding places by splashing the banks with water from the stream. 
At the collecting site in Longyang County (see below), members of this species were found syn- 
topic with members of Perileptus imaicus. 

Geographical distribution within the Gaoligong Shan .— Fig. 6c. We examined a total 
of 5 specimens (3 males and 2 females) from low elevations on both western and eastern slopes of 
the southern part of the Gaoligong Shan in Tengchong and Longyang Counties, (see Type materi¬ 
al above for exact collection data), which are in Core Areas 6 and 7, respectively. 

Overall geographical distribution. — This species currently is known only from the 
southern part of the Gaoligong Shan region in western Yunnan Province, China. 

Genus AgOllOtrechllS Jeannel, 1923 

Agonotrechus Jeannel, 1923:428. 

Paragonotrechus Ueno, 1981:2. 

Bhutcinotrechus Ueno, 1977:188. 

type species. — Trechus birmanicus Bates, 1892. 

diagnosis. — Adults of this genus (Figs. 7a-9a) can be recognized by the following combi¬ 
nation of character states: size medium to large for a trechine (BL = 4.5 to 7.0 mm) members of 
most species with full hindwings and large, convex eyes, but those of a few species apterous and 
with eyes reduced in size and/or convexity, some even microphthalmous or nearly anophthalmous; 
frons flattened, depressed; dorsolateral margin of mandibular scope with a row of small setae 
aligned as a sparse comb in some members (Belousov & Kabak 2003); right mandible (Fig. 16a) 
bidentate, the premolar tooth fused with the retinaculum, from which the anterior tooth is distinct¬ 
ly projected and in a forward position and the posterior tooth is absent or flush; left mandible with 
only a short, subconical, more or less trifid ridge; clypcus with six setae in most members; labium 
with anterior margin not or only slightly concave; mentum fused with submentum, incompletely in 
some members; submentum with six setae in most members, but some with eight setae; pronotum 
little narrowed basally, with lateral explanation distinctly broadened basally in many members; ely¬ 
tra with discal striae varied, Rom complete to effaced, punctate in many members, parascutellar 
striole very long in many members; anterior discal seta present in all members and inserted at the 


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basal one-fifth or one-sixth of elytra, middle discal seta present or absent, preapical seta present at 
or near stria 2 in most members, absent from a very few members; legs slender, protibiae furrowed, 
the basal two protarsomeres dilated and toothed; abdominal ventrites glabrous except for usual 
paramedial setae; aedeagus with a spoon-shaped copulatory piece. 

Taxonomic notes.— Members of the first known Agonotrechus species had only the anteri¬ 
or discal seta of the elytron inserted at the basal one-fifth or one-sixth. Consequently, this feature 
was used by Jeannel (1923) to define his new genus. Members of Par agonotrechus Ueno (1981) 
are very similar and differ only in having a more slender body form and the parascutellar striole 
very long. Those of Bhutanotrechus Ueno (1977), described from Bhutan, were distinguished by 
having the elytral discal striae effaced and a second elytral discal seta (the middle seta) present. The 
presence of the middle seta is a plesiotypic feature in Trechini, but species with members having 
two discal setae sometimes have been grouped together or assigned to genus Bhutanotrechus 
(Deuve 1992b, 1995). Genus Agonotrechus is really a homogenous group, despite the differences 
among its members in impression of the elytral discal striae, length of the parascutellar stiole, num¬ 
ber of elytral discal setae and size of the eyes, differences in the last feature being associated with 
differential flight capability. There appears to be no reason to maintain Paragonotrechus and 
Bhutanotrechus as distinct genera. 

The bidentate dentition of the right mandible of Agonotrechus , with the premolar tooth com¬ 
pletely fused with retinaculum, distinguishes this group from members of the Stevensius Complex, 
with which Jeannel (1923, 1928) had grouped it. 

Geographical distribution. — This genus, which at present includes 16 species, is known 
from the southeastern part of the Palearctic Region and northern edge of the Oriental Region, from 
Nepal eastward to southern Gansu Province in the north and Myanmar, Vietnam, and Hubei and 
Shaanxi Provinces; and one species has been described from Japan. The study area is within the 
previously known range of this genus. 

Key for Identification of Adults of AgOllOtrecllUS Species of the Gao li gong Shan Region 

1. Eyes less projected, less convex and only about twice as long as the tempora; pronotum only 
slightly transverse, lateral explanation narrow anteriorly; lateral elytral discal striae deep and 

markedly punctate, elytral intervals convex.A. xiaoheishan sp. nov. 

Eyes large and projected, more convex and more than twice as long as the tempora.2 

2. Size smaller, BL less than 5.5 mm. Pronotum smaller, lateral explanation narrow anteriorly; 

lateral elytral discal striae deep and elytral intervals only slightly convex. A. yunnanus Ueno 
Size larger, BL = 5.8 to 6.5 mm. Pronotum larger, lateral explanation wider anteriorly; lateral 
elytral discal striae more faintly impressed and elytral intervals nearly flat.3 

3. Elytral silhouette subovoid, rather short; pronotum narrower, with lateral explanation narrow, 
slightly widened basally; elytral recurrent stria abruptly terminated, not connected to apex of 

discal stria 5. A. fugongensis sp. nov. 

Elytral silhouette more subquadrate; pronotum markedly transverse, with lateral explanation 
wide anteriorly, also widened and slightly explanate basally; elytral recurrent stria continuous 
anteriorly with discal striae 5. A. wuyipeng Deuve 

Agonotrechus fugongensis Deuve and Liang, sp. nov. 

(Figs. 7, 35b, 36a, 46-48) 


Type material.— Holotype, a male, in IOZ, labeled: “CASENT 1019979”/ “CHINA, Yun- 


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nan, Fugong County, Lishadi Township, 0 to 2 km E of Shibali on Shibali Road, N27.16536°/ 
E098.78003 0 to N27.16100 0 /”/ “E098.79370 0 , 2300-2530 m, 18 August 2005, Stop# DHK-2005- 
096, D. Z. Dong collector”/ “HOLOTYPE Agonotrechns fugongensis Deuve & Liang, sp. nov. des¬ 
ignated 2016” [red label]. Paratypes (only 1): a female (in CAS) labeled: “CASENT 1006974”/ 
“CHINA, Yunnan, Gaoligong Shan, Nujiang Prefecture, Nujiang State Nature Reserve, Qigi He, 
9.9 airkm W of Gongshan/7 “N27.71542 0 / E98.56529 0 , 2000m, 9-14 July 2000, Stop#00-22A, 
D.H. Kavanaugh, C.E. Griswold, Liang H.-B., D. Ubick, & Dong D.-Z. collectors”/ “IMAGE” 
[pale green label]/ “PARATYPE Agonotrechus fugongensis Deuve & Liang, sp. nov. designated 
2016” [yellow label]. 

type locality.— China, Yunnan, Fugong County, Shiyueliang Township, 0 to 2 km W of 
Shibali on Shibali Road, 2300-2530 m. 

derivation of species name.— The species epithet, fugongensis, is derived from the name 
of the county (Xian) in which the holotype was collected, Fugong, and the Latin suffix, -ensis, 
denoting place. 

diagnosis. — Adults of this species (Fig. 7a) can be distinguished from those of all other 
species in the region by the following combination of character states: size large (BL = 5.8 to 6.0 
mm), fully-winged, dorsum dark piceous to reddish brown, shiny, pronotum and elytra slightly iri¬ 
descent; eyes convex, moderately projected; clypeus with four setae; mentum and submentum 
fused, submentum with six setae; pronotum small but transverse, ratio PW/PL = 1.30, median basal 
area smooth, framed by two deep, oblique furrows, lateral explanation rather narrow, broadened 
only posteriorly, lateral margins with a short sinuation anterior to the sharp, rectangular or slightly 
acute basal angles; elytra with all discal striae evident and finely punctate, striae 1 to 4 moderate¬ 
ly impressed, striae 5 to 8 more faintly impressed, parascutellar striole rather long, recurrent stria 
deeply impressed but abruptly terminated anteriorly, not connected with stria 5, intervals slightly 
convex, two discal seta (anterior and middle setae) present in stria 3, the anterior seta inserted with¬ 
in basal one-sixth of elytra, the middle seta inserted near elytral middle, preapical seta also pres¬ 
ent; median lobe of aedeagus of male (Fig. 7b) long and slender, endophallus with copulatory piece 
spoon-shaped. 

description. — Size large, BL = 5.8 to 6.0 mm. Color of dorsum piceous to reddish brown, 
shiny, pronotum and elytra slightly iridescent, appendages paler, palpi yellowish tan. 

Head. Moderate in size, slightly elongate, eyes large, convex, moderately projected, their 
diameter more than two times length of tempora. Tempora short, only slightly convex and glabrous, 
joined to neck region at ca. 135° angle. Frons more or less flattened, with two pairs of supraorbital 
setae, frontal furrows distinct, impressed posterior to or beyond posterior supraorbital setae. 
Clypeus with four setae. Labrum with six setae, apical margin slightly concave. Mandibles slen¬ 
der; right mandible bidentate, the anterior tooth far forward and long, the basal tooth reduced, 
obtuse and blunt; left mandible with a very slender trifid process, formed from fusion of the reti¬ 
naculum with premolar tooth. Palpi with apical palpomeres fusiform. Mentum and submentum 
fused. Mentum bifossulate, divided into three sectors, a median and two lateral parts, separated by 
two deep, longitudinal furrows; medial tooth simple, subtriangular with blunt apex, half as long as 
lateral lobes. Submentum with six setae subapically, gula broad. Genae with a single ventral seta 
on each side. Antennae of moderate length, with only four antennomeres extended beyond the 
pronotal base; antennonmeres 3 and 4 virtually the same length and each longer than antennomere 
2. 

Pronotum. Rather small, moderately narrow posteriorly, ratio PW/PL = 1.30, widest slightly 
anterior to middle, lateral margins with short sinuation just anterior to sharp, rectangular or slight¬ 
ly acute basal angles; pronotal disc convex, median longitudinal impression very fine and shallow, 


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Figure 7. Agonotrechus fugongensis sp. nov.; a. Dorsal habitus (CASENT1019979). b. Median lobe of aedeagus of 
male (CASENT1019979), left lateral aspect, c. Map of locality records (red circles) for A. fugongensis in the Gaoligong 
Shan region. Scale lines a, b = 0.5 mm, c = 100 km. 


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extended posteriorly to near basal margin, but not as close anteriorly to apical margin; basal foveae 
small vaguely delimited; median basal area smooth, delimited laterally by short but deep and 
oblique furrows; basal margin nearly straight, slightly sinuate. Lateral explanation moderately nar¬ 
row and slightly reflexed, widened posteriorly but not flattened. Midlateral pair of setae inserted 
anterior to middle and basolateral pair inserted at hind angles. 

Elytra. Moderately wide, elytral silhouette subovoid, humeri distinct but rounded, disc convex. 
All discal striae evident and finely punctate, striae 1 to 4 moderately impressed, striae 5 to 8 more 
faintly impressed, partially effaced, parascutellar stride rather long, recurrent stria deeply 
impressed but abruptly terminated anteriorly, not connected with discal stria 5, intervals slightly 
convex. Parascutellar setiferous pore present at base at common origin of discal striae 1 and 2. Two 
discal seta (anterior and middle setae) present in stria 3, the anterior seta inserted within basal one- 
sixth of elytra, the middle seta inserted near elytral middle. Preapical seta also present, inserted on 
interval 2 near stria 2, closer to sutural elytral margin than to apex. Umbilicate setal series with 
setae of humeral group equidistant for each other and those of median group both inserted posteri¬ 
or to middle of elytra. 

Legs. Slender, moderately long. Protibiae furrowed, without anteroapical pubescence. Male 
protarsi with tarsomeres 1 and 2 dilated and apicomedially toothed. 

Abdomen. Ventrites each with a pair of paramedial setae, ventrite VII of males with a single 
pair of paramedial apical setae, of females with two pairs. 

Male aedeagus. Median lobe (Fig. 7b) long and slender, endophallus with a spoon-shaped scle- 

rite. 

Comments. — Males of this species are most similar to those of A. wuyipeng in features of the 
aedeagus; but they are smaller, their pronota distinctly narrower, more slender, and with much nar¬ 
rower lateral margination, and the recurrent stria does not connect anteriorly with stria 5 as it does 
in A. wuyipeng members. 

h abitat distribution .— The holotype specimen of this species was found under a stone on 
moist substrate along a roadcut through an agricultural area formed in a large clearcut in what had 
been mixed broadleaf evergreen and conifer forest at an elevation somewhere between 2300 and 
2530 m (Fig. 36a). The paratype specimen was collected under stones on the shaded bank of the 
Qiqi River just above the Forestry station at Qiqi at an elevation of 2000 m (Fig. 35b). Members 
of no other Agonotrechus or other trechine species have been found syntopic with those of 
A. fugongensis. 

Geographical distribution within the Gaoligong Shan .— Fig. 7c. We examined a total 
of 2 specimens (1 male and 1 female) from Fugong and Gongshan Counties, respectively (see Type 
material above for exact collection data). 

Specimens of this species were collected only in the northern half of the study area (Core 
Areas 2 and 3) and only on the eastern side of the mountain range. This distribution pattern may 
be an artifact of inadequate sampling on the western slope of the mountain range in the north, much 
of which is in Myanmar. The geographical range of this species overlaps that of A. wuyipeng , but 
members of the latter species appear to occupy slightly higher elevations in the same general areas. 

Overall geographical distribution. — This species currently is known only from the 
northern half of the Gaoligong Shan in western Yunnan Province, China. 

Agonotrechus wuyipeng Deuve, 1992 

(Figs. 8, 36b, 45-48) 

Agonotrechus wuyipeng Deuve, 1992b: 172. Holotype, a male, in IOZ. Type locality: China, Sichuan, Wo long, 

Wuyipeng, 2500 m. 


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diagnosis. — Adults of this species (Fig. 8a) can be distinguished from those of all other 
species in the region by the following combination of character states: size large (BL = 6.5 to 6.7 
mm), fully-winged, dorsum piceous to reddish brown and distinctly iridescent; head slender, eyes 
markedly projected; clypeus with four or six setae; mentum and submentum incompletely fused, 
submentum with six or eight setae; mandibles with four to six small setae along the dorsolateral 
margin of the scrobe; pronotum transverse (ratio PW/PL = 1.30), with lateral explanation broader, 
especially basally, basal angles rectangular and sharp with a small apical tooth projected laterally; 
elytra broad, with all discal striae evident and finely punctate, striae 1 to 5 moderately impressed, 
striae 6 to 8 more faintly impressed, parascutellar striole rather long, recurrent stria continuous 
anteriorly with stria 5, intervals only faintly convex, two discal setae (anterior and middle setae) 
present, the anterior seta inserted within basal one-sixth of elytra in stria 3, the middle seta near 
elytral middle in stria 3; median lobe of aedeagus of male (Fig. 8b) long and slender, with apex 
short, recurved dorsally and bluntly pointed, endophallus with a spoon-shaped sclerite. 

Comments. — The polymorphism we observed in the number of setae on both the clypeus 
(four or six) and submentum (six or eight) among specimens from the Gaoligong Shan populations 
was surprising. Belousov & Kabak (2003) described Agonotrechus dubius, based on a single 
female from Gansu Province, and noted similarities with A, wuyipeng. We have examined a male 
specimen from Shaanxi Province (Ningshan County, Huoditang Township, 1549 m, collected by 
Matt Brantley on 9 July 2005) which shares features described for the holotype of A. dubius , 
including six setae on the submentum, which Belousov & Kabak (2003) contrasted with the eight 
setae reported for A. wuyipeng members (Deuve 1992b). Together with other similarities, the poly¬ 
morphism in this feature found among Gaoligong Shan specimens suggests that these two species 
may be better treated as conspecific, with A. dubius as a distinct subspecies, members of which 
have smaller size, relatively wider pronota, and more coarsely punctate elytral discal striae but are 
otherwise similar to members of the nominate form. 

habitat distribution. — Specimens of this species were collected in daytime from under 
stones in shaded roadside and trailside areas with scattered grasses at elevations ranging from 2687 
to 2770 m (Fig. 36b). Members of no other Agonotrechus or other trechine species have been found 
syntopic with those of A. wuyipeng. 

Geogr aphical distribution w ithin the G aoligong Shan .— Fig. 8c. We examined a total 
of 10 specimens (7 males and 3 females) from the following localities: Fugong County: 
Shiyueliang Township (1 km above Shibali on Yaping Road, 2687 m, 1 May 2004, H.B. Liang col¬ 
lector [1 female; MNHN], Gongshan County: Qiqi Trail at No 12 Bridge, N27.71500°/ 
E98.50222°, 2770 m, 2 May 2002, H.B. Liang & W.D. Ba collectors [7 males and 2 females; CAS, 
IOZ, MNHN], 

Specimens of this species were collected only in the northern half of the study area (Core 
Areas 2 and 3) and only on the eastern side of the mountain range. This distribution pattern may 
be an artifact of inadequate sampling on the western slope of the mountain range in the north, much 
of which is in Myanmar. The geographical range of this species overlaps that of A. fugongensis, 
but members of the latter species appear to occupy slightly lower elevations in the same general 
areas. 

Overall geographical distribution. — Fig. 45. This species currently is known only from 
the type locality in northcentral Sichuan and the northern half of the Gaoligong Shan in western 
Yunnan, but it probably occurs in the intervening region as well. 


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Figure 8. Agonotrechus wuyipeng Deuve; a. Dorsal habitus (CASENT1010854). b. Median lobe of aedeagus of male 
(CASENT1010854), left lateral aspect, c. Map of locality records (red circles) for A. wuyipeng in the Gaoligong Shan 
region. Scale lines a, b = 0.5 mm, c = 100 km. 


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Agonotrechus xiaoheishan Deuve and Kavanaugh, sp. nov. 

(Figs. 9, 16a, 44, 46-48) 

Type material.— Holotype, a male, in IOZ, labeled: “CASENT 1036866”/ “CHINA, Yun¬ 
nan, Longling County, Longjiang Township, Xiaoheishan Forest Reserve, N24.83671 0 / 
E098.76185°,”/ “2067 m, 28 May 2005, Stop# HBL-05-19, H.B. Liang, H.M. Yan & K.J. Gao col¬ 
lectors”/ “HOLOTYPE Agonotrechus xiaoheishan Deuve & Kavanaugh, sp. nov. designated 2016” 
[red label]. Paratypes (a total of 13): 2 males and 7 females (in CAS, IOZ, MNHN) labeled same 
as holotype except first label “CASENT 1036874”, “CASENT 1036875”, “CASENT 1036867”, 
“CASENT 1036868”, “CASENT 1036869”, “CASENT 1036870”, “CASENT 1036871”, 
“CASENT 1036872” and “CASENT 1036873”, respectively; 1 male and 1 female (in CAS) 
labeled “CASENT 1036863” and “CASENT 1036864”, respectively/ “CHINA, Yunnan, Longling 
County, Longjiang Township, Xiaoheishan Forest Reserve, Guchengshan, 2020 m, N24.82888°/ 
E098.76001°,”/ “28 May 2005, Stop# 2005-033B, D.H. Kavanaugh, H.B. Liang, D.Z. Dong & J.L. 
Yang collectors”; 2 females (in CAS) labeled “CASENT 1031915” and “CASENT 1031916”, 
respectively/ “CHINA, Yunnan, Longling County, Longjiang Township, Xiaoheishan Forest 
Reserve, Guchengshan, 2020 m, N24.82888 0 / E098.76001 0 ,”/ “28 May 2005, Stop# 2005-033C, 
D.H. Kavanaugh, C.E. Griswold, H.B. Liang, D.Z. Dong, H.M. Yan & K.J. Guo collectors”. All 
paratypes also bear the following label: “PARATYPE Agonotrechus xiaoheishan Deuve & 
Kavanaugh, sp. nov. designated 2016” [yellow label]. 

t ype locality .— China, Yunnan, Longling County, Longjiang Township, Xiaoheishan For¬ 
est Reserve, N24.83671°/E098.76185°, 2067 m. 

derivation of species name. — The species epithet, xiaoheishan , is a noun in apposition, 
derived from the name of the area in which the holotype was collected. 

diagnosis. — Adults of this species (Fig. 9a) can be distinguished from those of all other 
species in the region by the following combination of character states: size medium (BL = 5.3 to 
5.7 mm), fully-winged, dorsum piceous to reddish brown and distinctly iridescent; head slender, 
eyes only moderately projected; clypeus with four setae; mentum and submentum fused, submen- 
tum with six setae; mandibles with two or three small setae along the dorsolateral margin of the 
scrobe; pronotum small and narrow (ratio PW/PL = 1.21), basal angles subrectangular, basal flat¬ 
tened area restricted; midlateral seta inserted at anterior one-fourth, basolateral seta at hind angle; 
elytra with all discal striae evident, deeply impressed and punctate, however striae 1 to 4 effaced 
or nearly so near base, parascutellar striole rather long, recurrent stria continuous anteriorly with 
stria 5, intervals convex, only a single discal seta (anterior seta) present, inserted at basal one- 
eighth of elytra in stria 3; median lobe of aedeagus of male (Fig. 9b) short, with apex bent ventrally 
and bluntly pointed, endophallus with a spoon-shaped sclerite. 

Description.— Size medium, BL = 5.3 to 5.7 mm. Color of dorsum piceous, shiny, irides¬ 
cent, femora concolorous, tibiae, tarsi, antennae and mandibles paler reddish tan, palpi yellowish 
tan. 

Head. Relatively slender, eyes only moderately projected, but nonetheless convex and with 
diameter twice length of tempora. Tempora not or only slightly convex, joined to neck region at ca. 
120° angle. Frons slightly flattened, with two pairs of supraorbital setae, frontal furrows linear, dis¬ 
tinct between the eyes, but effaced posterior to insertion of second supraorbital seta. Clypeus with 
four setae. Labrurn slightly widened apically, apical margin slightly concave. Mandibles sharp, the 
right mandible (Fig. 16a) bidentate with the anterior tooth spaced well forward of the posterior 
tooth and the left mandible with only a small subtriangular process, also with two or three small 
setae along the dorsolateral margin of the scrobe. Mentum and submentum fused. Mentum with 
medial tooth rather broad, with the apex either obtuse or truncate, less than one-half as long as lat- 


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Figure 9. Agonotrechus xiaoheishan sp. nov.; a. Dorsal habitus (CASENT1036866). b. Median lobe of aedeagus of 
male (CASENT1036866), left lateral aspect, c. Map of locality records (red circles) for A. xiaoheishan in the Gaoligong 
Shan region. Scale lines a, b = 0.5 mm, c = 100 km. 


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eral lobes. Submentum with six setae subapically, gula broad. Genae with a single seta ventrally on 
each side. Antennae long and slender, extended to (in females) or slightly beyond (in males) the 
middle of the elytra, with five or six antennomeres beyond the pronotal base; antennomneres 3 and 
4 virtually the same length and antennomere 2 slightly shorter. 

Pronotum. Relatively small, not or only slightly narrowed basally, only slightly transverse, 
ratio PW/PL = 1.21, widest at anterior one-third; lateral margins rectilinear in basal half, not sinu¬ 
ate except for a very slight and short inflexion just anterior to the basal angle, which is sub rectan¬ 
gular and sharp; pronotal disc smooth, glabrous, moderately convex; median longitudinal impres¬ 
sion very fine; basal foveae deep, basal flattened area rather small, delimited laterally by short but 
deep and oblique furrows. Lateral explanation slender and moderately reflexed in anterior two- 
thirds, then progressively broader in basal one-third without. Midlateral pair of setae inserted at 
anterior one-fifth and basolateral pair inserted at hind angles. 

Elytra. Convex, large and broad, especially in relation to pronotum, elytral silhouette ovoid, 
about equally narrowed apically and basally, humeri evident but rounded. All discal striae evident, 
regular, deeply impressed and punctate, however striae 1 to 4 effaced or nearly so near base, paras- 
cutellar striole rather long, recurrent stria continuous anteriorly with discal stria 5, intervals con¬ 
vex, only a single discal seta (anterior seta) present, inserted at basal one-eighth near stria 3. Paras- 
cutellar setiferous pore present at base at common origin of discal striae 1 and 2. Anterior discal 
seta present, inserted at basal one-eighth of elytra in stria 3, which is effaced anterior to that point. 
Middle discal seta absent. Preapical seta present on interval 3 near stria 2 opposite the anterior edge 
of the subapical sinuation. Umbilicate setal series with setae of humeral group equidistant for each 
other and those of median group both inserted posterior to middle of elytra. 

Legs. Slender but only moderately long. Protibiae furrowed. Male protarsi with tarsomeres 1 
and 2 dilated and apicomedially toothed. 

Male aedeagus. Median lobe (Fig. 9b) short, with apex bent ventrally and bluntly pointed, 
endophallus with a spoon-shaped sclerite. 

Comments, — Members of this new species are morphologically similar to those of 
Agonotrechus binnanicus Bates (1892), described from Kachin State in Myanmar, and 
Agonotrechus tenuicollis Ueno (1986), described from eastern Nepal. However, they can be dis¬ 
tinguished from members of both of these other species by the following features: pronotum with 
anterior margin straight, not concave, anterior angles only faintly projected and more broadly 
rounded, and lateral borders more slender anteriorly and at middle; and apex of the median lobe of 
the male aedeagus narrower and more curved. 

habitat distribution,— Members of this species have been found in and under rotting logs 
in dark, closed-canopy broadleaf evergreen forest (Fig. 44) at elevations ranging from 2020 to 2067 
m. Although members of no other Agonotrechus species have been found syntopic with those of A. 
xiaoheishan, specimens of Trechus indicus were collected in the same samples. 

Geocraphicai, distribution within the Gaougong Shan. — Fig. 9c. We examined a total 
of 14 specimens (4 males and 10 females), all from Xiaoheishan Forest Reserve in the southern 
part of the Gaoligong Shan (see Type material above for exact collection data). 

This species was recorded only from near the top of the western slope in the southern part of 
the study area (Core Area 6). Its known geographical range does not overlap with that of any other 
Agonotrechus species, although A. yunnanus has been recorded from the adjacent Core Area 7 on 
the eastern slope of the Gaoligong Shan, 13.3 km to the north. 

Overall geographical distribution .— This species currently is known only from the type 
area in the southern part of the Gaoligong Shan, in western Yunnan Province, China. 


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Agonotrechus yunnanus Ueno, 1999 
(Figs. 10, 46-48) 

Agonotrechus yunnanus Ueno, 1999a:215. Holo- 

type, a male, in NSMT. Type locality: China, 

Yunnan, Gaoligong Shan, Longyang County, 

24.957 98.75°, 2200-2500 m. 

Notes on type material.— We have not 
had an opportunity to study the unique holo- 
type of this species, so features noted below 
are taken from Ueno’s orginal description. 

Diagnosis. — Adults of this species (see 
Ueno 1999a, Fig. 1) can be distinguished from 
those of all other species in the region by the 
following combination of character states: size 
slightly small for the genus (BL = 5.4 mm); 
eyes large and projected; tempora short; prono- 
tum rather small and only slightly transverse 
(ratio PW/PL = 1.28), lateral explanation slen¬ 
der anteriorly, slightly widened basally, basal 
angles subrectangular and sharp; elytra with 
intervals only slightly convex, discal striae 
deeply impressed and punctate, two discal 
setae (anterior and middle) present near stria 3; 
median lobe of aedeagus of male (see Ueno 
1999a, Figs. 2-3) with apex short and broadly 
rounded. 

Habitat distribution.— Although no 
precise habitat information accompanied the 
unique holotype specimen, Ueno (1999a:219) 
suggested it was probably collected near the 
pass across the crest of the Gaoligong Shan on 
the route from “Bawan to Shang’ying, which 
lies near the lower edge of the Rhododendron 
zone.” According to Ueno (1999a), specimens 
of Trechus indicus Putzeys (which he recorded 
as Trechus macrops Jeannel) and Epa- 
phiotrechus fortipes (Ueno) comb. nov. were 
also collected in the same area. 

Geographical distribution within 
the Gaoligong Shan.— Fig. 10 . Loilgycing 
County: N24.957E98.75 0 , 2200-2500 m, 8-16 
May, O. Semala collector [1 male; NSMT]. No 
specimens of this species were collected dur¬ 
ing this study. 

The type locality for this species is on the 
eastern slope of the southern part of the 


Figure 10. Agonotrechus yunnanus Ueno; Map of 
locality record (red circle) in the Gaoligong Shan region. 
Scale line =100 km. 


DEUVE ET AL.: CARABID BEETLE FAUNA OF THE GAOLIGONG MOUNTAINS 


363 


Gaoligong Shan in the northern part of Core Area 7. This area is not within the geographical range 
of any other Agonotrechus species, although A. xiaoheishan has been recorded from the adjacent 
Core Area 6 on the western slope of the Gaoligong Shan, 13.3 km to the south. 

o verall geographical distribution .— This species currently is known only from the type 
locality in the southern part of the Gaoligong Shan, in western Yunnan Province, China. 

Genus Minutotrechus Deuve and Kavanaugh, gen. nov. 
type species. — Stevensius minutus Ueno, 1997. 

Derivation of genus group name.— The genus group name (masculine) is a combination 
of the Latin adjective, minutus , meaning very small, and the generic name, Trechus, in reference to 
the small size of members of this genus. 

d ia g n o sis .— Adults of this genus (Fig. 11 a) can be recognized by the following combination 
of character states: size small (BL = 2.7 to 2.9 mm), apterous, body color brown to black; head 
large with small but protruding eyes, their diameter shorter than length of tempora; mandibles 
short, obtusely bidentate, mentum and submentum at least partial fused, labial suture partially per¬ 
ceptible, mentum with medial tooth truncate; pronotum small, cordate, narrow (ratio PW/PL = 
1.25), very convex, globulose, glabrous, basal angles small and subrectangular, slightly obtuse, 
with basal margin broadly lobate, basal area convex, both midlateral and basolateral setae present; 
elytra ovoid and markedly convex, with discal striae 2 to 8 striae effaced, stria 1 deeply impressed 
and punctate, both anterior and middle discal setae present, preapical seta absent, lateral groove 
abruptly terminated anteriorly at humerus; legs short, protibiae without longitudinal furrows. 

Comments. — This new genus is known from only six female specimens that were originally 
assigned by Ueno (1997) to genus Stevens ius Jeannel (1923) of the eastern Himalayan region. 
However, they can be distinguished from members of that genus by their smaller head size, protib¬ 
iae without longitudinal furrows, pronotum more cordate and with basal angles smaller and basal 
margin broadly lobate and elytra with the lateral groove abruptly terminated anteriorly at humerus. 
Minutotrechus appears to be more closely related to Hubeitrechus Deuve (2005), but its members 
can be distinguished from those of the latter in having the mentum and submentum at least partial 
fused, the mandibular teeth short and obtuse, the pronotum with the median basal area more con¬ 
vex and basal margin broadly lobate and without margination, and lateral groove abruptly termi¬ 
nated at the humerus. Because no male specimens of Minutotrechus have been collected to date, 
we do not know if male protarsomeres 1 and 2 are elongate as in males of Hubeitrechus or broad 
as in Stevens ius males. Members of this new genus can also be compared with those of Uenoites 
Belousov and Kabak (2016), from which they differ in having the right mandible obtusely bifid 
(tridentate in Uenoites members), the protibiae without longitudinal furrows (longitudinal furrows 
present in Uenoites members), the mentum and submentum at least partial fused (not fused in 
Uenoites members), elytra with only two discal setae (three or more discal setae present in 
Uenoites members), the preapical seta absent (present in Uenoites members) and the lateral groove 
abruptly terminated at humerus (gradually narrowed anterior to humerus in Uenoites members). 

Geographical distribution. — This genus currently is known only from the type species, 
which is known only from the southern part of the Gaoligong Shan region of western Yunnan 
Province, China. 

Minutotrechus minutus (Ueno, 1997 ) 

(Figs. 11,46M8) 

Stevensius minutus Ueno, 1997:182. Holotype, a female, in NMST. Type locality: China, Yunnan, Gaoligong 
Shan, Tengchong County, Dabei, 2430 m. 

Minutotrechus minutus (Ueno) Nf.w Combination. 


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Figure 11. Minutotrechus minutus (Ueno); a. Dorsal habitus (paratype). scale line = 1.0 mm; b. Map of locality 
records (red circle) for M. minutus in the Gaoligong Shan region. Scale lines a = 0.5mm, b = 100 km. 


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Notes on type material. — We have not had an opportunity to study the holotype of this 
species, but we have examined a paratype female deposited in IOZ. Features noted below are based 
on our examination of that paratype and Ueno’s orginal description. 

d ia g n o sis .— Adults of this species (Fig. 1 la), the only known species in this new genus, can 
be distinguished from those of all other species in the region by the combination of character states 
noted in the generic diagnosis. 

habitat distribution. — According to Ueno (1997) specimens of the type series were col¬ 
lected at an elevation of 2430 m in a dense Rhododendron forest by sifting moist leaf litter accu¬ 
mulations on the ground. He also noted that many specimens of “Trechus asetosus Ueno” (1997) 
were also collected in the same litter samples. 

Geographical distribution within the Gaoligong Shan. — Fig. lib. This Species is 
known only from the six female specimens of the type series collected at the type locality, in Teng- 
chong County, high on the western slope of the southern part of the Gaoligong Shan in Core 
Area 6. 

Overall geographical distribution. — This species currently is known only from the type 
locality in the southern part of the Gaoligong Shan, in western Yunnan Province, China. 

Genus Queinnectrechus Deuve, 1992 

Queinnectrechus Deuve, 1992b:354. 

type species. — Queinnectrechus excentricus Deuve, 1992a 

diagnosis. — Adults of this genus (Figs. 12-14) can be recognized by the following combi¬ 
nation of character states: size moderate (BL = 3.5 to 4.8 mm), apterous, body dark, reddish brown 
to piceous, surface micaceous; head with small eyes, mentum and submentum fused; right 
mandible tridentate, the premolar tooth distinct but joined with the retinaculum (see Deuve 1992b, 
Fig. 23); pronotum cordiform and markedly convex, lateral margination effaced posteriorly, with 
two setae (midlateral and basolateral) present on each side; elytra markedly convex, inflated, 
slightly tear-shaped, humeri effaced, elytral discal striae absent or vestigial, with two (in most 
members) or three (in a few members) discal setae present, aligned on interval 3 near stria 3, 
preapical seta absent from most members, in some of these members inserted forward in a subdis- 
cal position, in very few placed in typical trechine position nearer elytral apex and next to stria 2; 
protibiae furrowed; abdominal ventrites IV to VI glabrous except for a single pair of paramedial 
setae; endophallus of male aedeagus with two sclerites. 

Comments. — Two new genera closely related to Queinnectrechus have recently been 
described. Members of Dactylotrechus Belousov and Kabak (2003) are distinguished by the super¬ 
numerary setae present on the external margins of the pronotum and on more lateral areas of the 
elytral disc. Members of Puertrechus Belousov and Kabak (2014a) are distinguished by the pres¬ 
ence of a single discal setae subbasally on interval 5. Members of both taxa have a preapical seta 
inserted near stria 2, a plesiomorphic feature among trechines. Taxonomic limits and phylogenetic 
relationships among the “genera” Stevensius, Kozlovites Jeannel (1935), Queinnectrechus, Dacty¬ 
lotrechus, Puertrechus, Sinotrechiama Ueno (2000), Uenoite s Belousov and Kabak (2016) and 
Minutotrechus (describe above) are still poorly resolved. This is why we describe below a new 
taxon, Gaoligongtrechus, provisionally with the rank of subgenus. This assignment can be changed 
as needed in the future. 

Geographical distribution. — This genus, which currently include 13 described species 
(Belousov and Kabak 2003, Casalle and Magrini 2009, Deuve 1992a, and Ueno 1998a and 1998b) 
is currently known from the Min Shan region of northern Sichuan Province southwest to western 


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Yunnan Province. The Gaoligong Shan forms the southwestern limit of the known distributional 
range of the genus. 


Key for Identification of Subgenera of Queinnectrechus in the Gaoligong Shan Region 

1. Pronotum with basal angles prolonged posteriorly as slender digitiform processes (Figs. 12a, 

13 a).Subgenus Queinnectrechus Deuve 

1’ Pronotum with basal angles simple (Fig. 14a). . . Subgenus Gaoligongtrechus subgen. nov. 

Subgenus Queinnectrechus Deuve, 1992 

Queinnectrechus Deuve, 1992a:3 54. 

diagnosis.— Adults of this subgenus (Figs. 12a, 13a) can be recognized by the following 
combination of character states: size moderate (BL = 3.5 to 4.8 mm), apterous, body dark, reddish 
brown to piceous, surface micaceous; pronotum with basal angles prolonged posteriorly as slender 
digitiform processes, with two setae (midlateral and basolateral) present on each side; elytra 
markedly convex, inflated, slightly tear-shaped, humeri effaced, elytral discal striae effaced, with 
two (in most members) or three (in a few members) discal setae present, aligned on interval 3 near 
stria 3, preapical seta absent from most members. 

Geographical d is trib u tio n .— Same as for genus (see above). 

Key for Identification of Adults of Subgenus Queinnectrechus species 

of the Gaoligong Shan Region 

I. Size larger (BL = 4.3 to 4.8 mm), elytra (Fig. 12a) more elongate; median lobe of male aedea- 

gus broadest at mid-shaft, with apex long and recurved. Q. griswoldi sp. nov. 

Size smaller (BL = 3.5 to 3.8 mm), elytra (Fig. 13a) shorter, ovoid; median lobe of male aedea- 
gus slender, with apex short and not at all curved. Q. gongshanicus sp. nov. 

QlieinnectrechllS (S. Str.) griswoldi Deuve and Kavanaugh, sp. nov. 

(Figs. 12, 37b, 38a, 38b, 39b, 46-48) 

Type material.— Holotype, a male, in IOZ, labeled: “CASENT 1026334”/ “CHINA, Yun¬ 
nan, Gongshan County, Cikai Township, 0.1 km SE of Heipu Yakou in valley below tunnel, 
N27.76978°/ E98.44681 0 ,”/ “3720 m, 13 August 2006, Stop #DHK-2006-073 D.H. Kavanaugh & 

J. A. Miller collectors”/ “HOLOTYPE Queinnectrechus (s. str.) griswoldi Deuve & Kavanaugh, sp. 
nov. designated 2016” [red label]. Paratypes (a total of 20): 5 males and 3 females (in CAS, IOZ, 
MNHN) labeled same as holotype except first label “CASENT 1026333”, “CASENT 1026335” to 
“CASENT 1026338” and “CASENT 1026330” to “CASENT 1026332”, respectively; 1 female (in 
IOZ) labeled “CASENT 1010344”/ “CHINA, Yunnan, Gongshan County, Cikai Township, 52.6 
lcm W of Gongshan on Dulong Valley Road, 3360-3380 m,”/ “N27.77032 0 / E098.44661 0 , 1-2 
October 2002, Stop #DHK-2002-034, D.H. Kavanaugh & P.E. Marek collectors”; 1 male and 3 
females (in CAS, IOZ) labeled “CASENT 1024862” and “CASENT 1024863” to “CASENT 
1024865”, respectively/ “CHINA, Yunnan, Gongshan County, Bingzhongluo Township, SW slope 
of Kawakarpu Shan, on slope NE of Chukuai Lake, 3950,”/”N27.98206°/ E098.48027 0 ,20 August 
2006, Stop #DHK-2006-086 Y. Liu, P. Hu, D.Z. Dong, & J. Wang collectors”; 3 males and 1 female 
(in CAS, IOZ, MNHN) labeled “CASENT 1026813” to “CASENT 1026815” and “CASENT 
1026815”, respectively/ “CHINA, Yunnan, Gongshan County, Bingzhongluo Township, SW slope 
of Kawakarpu Shan at Chukuai Lake, 3720 m,”/ “N27.98121 0 / E098.47580 0 , 18 August 2006 Stop 
#DHK-2006-079 J.A. Miller, D.Z. Dong, & Y. Liu collectors”; 1 female (in CAS) labeled 


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Figure 12. Queinnectrechus ( s. str .) griswoldi sp. nov.; a. Dorsal habitus (CASENT1026334). b. Median lobe of 
aedeagus of male (CASENT1026334), left lateral aspect, c. Map of locality records (red circles) for Q. griswoldi in the 
Gaoligong Shan region. Scale lines a, b = 0.5 mm, c = 100 km. 


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“CASENT 1025813”/ “CHINA, Yunnan, Gongshan County, Bingzhongluo Township, SW slope of 
Kawakarpu Shan 0.3 km SW of Chukuai Lake at campsite,”/, “N27.976867 E098.47799 0 , 3750 m, 
18 August 2006 Stop #DHK-2006-078 D.H. Kavanaugh collector”; 2 females (in IOZ) labeled 
“CASENT 1025836” and “CASENT 1025837”, respectively / “CHINA, Yunnan, Gongshan Coun¬ 
ty, Bingzhongluo Township, SW slope of Kawakarpu Shan, 0.3 km SW of Chukuai Lake at camp¬ 
site,”/ “N27.976S67 E098.44779 0 , 3750 m, 19 August 2006, Stop #DHK-2006-082 Y. Liu collec¬ 
tor”. All paratypes also bear the following label: “PARATYPE Queinnectrechus (, s. str.) griswoldi 
Deuve & Kavanaugh, sp. nov. designated 2016” [yellow label]. 

Type locality.— China, Yunnan, Gongshan County, Cikai Township, 0.1 km SE of Heipu 
Yakou, in valley below tunnel, N27.774377 E098.44793 0 , 3270 m. 

derivation of species name. — The species epithet, griswoldi , is the Latinized form (in the 
genitive case) of the surname of Charles E. Griswold, now Curator Emeritus and former Schlinger 
Chair of Arachnology at the California Academy of Sciences, who participated in several of the 
expeditions to the study area and helped collect many carabid specimens for this project. We are 
pleased to name this elegant species in his honor. 

diagnosis. — Adults of this species (Fig. 12a) can be distinguished from those of all other 
species in the region by the following combination of character states: size medium (BL = 4.3 to 
4.8 mm), body dark reddish brown, very shiny; eyes small and convex, tempora slightly convex; 
pronotum markedly narrowed posteriorly, markedly convex, globulose, narrowly cordate, disc 
smooth, basal angles with distict digitorm projections, each side with a single midlateral and baso- 
lateral seta; elytra markedly convex, tear-shaped, discal striae effaced, with three discal setae in a 
row along the presumed location of stria 3, preapical seta absent; median lobe of male aedeagus 
(Fig. 12b) large, elongate, broadest at mid-shaft, with apex long and recurved dorsally, endophal- 
lus with internal sclerites acuminate apically. 

description.— Size medium, BL = 4.3 to 4.8 mm. Color of body, antennae and legs dark 
reddish brown, palpi paler, yellowish tan, body surface very shiny, micaceous, smooth and 
glabrous. 

Head. Moderate in size, eyes small and convex, their convexity greater than and their diame¬ 
ter about as long as tempora, the latter only slightly convex and glabrous. Frons with frontal fur¬ 
rows deep, rounded, and not interrupted posteriorly; two suporaorbital setae present, the anterior 
inserted opposite midpoint of eye, the posterior inserted in postocular groove. Clypeus with four 
setae. Labrum with six setae, anterior margin distinctly concave. Right mandible tridentate. Left 
mandible with a small, minutely tridentate process. Mentum and submentum fused. Mentum with 
medial tooth wide, bifid or truncate, one half the length of the lateral lobes. Submentum with six 
setae anteriorly. Gula wide. Genae with a single seta ventrally on each side. Antennae of moderate 
length, extended posteriorly almost to or slightly beyond basal one-fourth of elytra, with 3.5 anten- 
nomeres in females and 4.5 antennomeres in males extended beyond basal pronotal margin, anten- 
nomeres slightly broadened, antennomere 3 slightly longer than antennomeres 2 or 4. 

Pronotum. Shape narrowly cordifonn, only slightly transverse (ratio PW/PL = 1.1), widest at 
the anterior one-fourth, markedly narrowed posteriorly, lateral margins distinctly sinuate anterior 
to basal angles, the latter extended posteriorly as slender, pointed, digitiform processes. Disc 
markedly convex, globulose, smooth and glabrous, with median longitudinal impression very faint¬ 
ly impressed or effaced, basal fovea small and smooth; median basal area smooth; basal margin 
slightly convex and rounded. Lateral border of pronotum slender, distinctly defined only in anteri¬ 
or one-third to one-half, effaced in posterior one-half to two -thirds. Single midlateral setae on 
each side inserted at anterior one-third; single basolateral seta on each side, inserted on basal angle 
at base of digitform process. 


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Elytra. Elytral silhouette slightly tear-shaped, with humeri effaced, disc markedly convex, 
smooth and glabrous. All discal striae effaced except for faintly impressed parascutellar striole and 
a short, faintly impressed recurrent stria. Basal setiferous pore present. Three discal setae present, 
aligned along presumed track of stria 3. Preapicale seta absent, one or two apicoangular setules 
present. Umbilicate setal series with setae of humeral group equidistance from each other, with the 
first slightly more medially inserted than the others, setae of median group inserted distinctly pos¬ 
terior to middle. 

Legs. Moderately long but slender. Protibiae with longitudinal furrow, sparely pubescent api- 
cally on anterior surface. Male protarsomeres 1 and 2 dilated and apicomedially toothed. 

Abdomen. Abdominal ventrites glabrous, except for a single paramedial seta on each side, and 
ventrite VII of males apically with one pair of paramedial setae, females with two pairs. 

Male aedeagus. Median lobe (Fig. 12b) large, elongate, broadest at mid-shaft, with apex long 
and recurved dorsally; endophallus with internal sclerites acuminate apically. 

Comments. — Based on similarities in the form of the aedeagus of males, this species is close¬ 
ly related to Queinnectrechus jiuhecola Deuve & Kavanaugh (Deuve et al. 2015), described from 
Lijiang County, northwestern Yunnan. However, members of this new species are distinguished 
from those of the latter by their much smaller size (BL = 5.3 to 5.8 mm in Q. jiuhecola members), 
more elongate body form, more convex elytra, discal striae fully effaced (at least three medial stri¬ 
ae evident in Q. jiuhecola members) and, most significantly, preapical seta absent (present near 
stria 2 subapically in Q. jiuhecola members). 

In a recent paper, Belousov and Kabak (2016) established a new genus, Uenoites, in which 
they placed 0. jiuhecola , as well as three other species previously included in Kozlovites or Deu- 
veotrechus Ueno (1995), based on the presence of a preapical seta, more convex tempora, and the 
median lobe of the male aedeagus not markedly hooked apically. However, all these features are 
plesiomorphic among trechines, which is problematic for demonstrating phylogenetic affinity of 
the included taxa. Clearly, phylogenetic relationships among these groups of species remain unre¬ 
solved. As noted above, the genitalia of 0. griswoldi males are very similar to those of O. jiuheco¬ 
la males, wherease those of the new species described below, Queinnectrechus gongshanicus sp. 
nov., are of a very different form, although both of the new species described here are members of 
genus Queinnectrechus . 

h abit at distribution. — Members of this species have been found in a variety of microhab¬ 
itats in the alpine zone near the crest of the Gaoligong Shan and the eastern slope, at elevations 
ranging from 3270 to 3950 m. Specimens were collected under stones in moist meadows and on 
tundra slopes and ridges with sparse to thick herbaceous vegetation (Figs. 37b, 38a, 38b, 39b), at 
the edges of small streams and seeps from talus slopes, and at the edges of Rhododendron thickets 
up to 1.5 m tall. Members of this species have been found syntopic with specimens of Queinnec¬ 
trechus ( Gaoligongtrechus ) lialli sp. nov., Queinnectrechus (s. str.) gongshanicus sp. nov., Trechus 
gongshanensis sp. nov., Trechus qiqiensis sp. nov. and Trechepaphiopsis monochaeta sp. nov. 

Geographical distribution within the Gaoligong Shan. — Fig. 12c. We examined a 
total of 21 specimens (10 males and 11 females), all from the northern part of the Gaoligong Shan, 
in Bingzhongluo and Cikai Townships, Gongshan County (see Type material above for exact col¬ 
lection data). These localities are all on the crest or eastern slope of the Gaoligong Shan in Core 
Area 2. 

Overall geographical distribution.— This species currently is known only from the 
northern part of the Gaoligong Shan, in western Yunnan Province, China. 


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Queinnectrechus {s. str.) gongshanicus Deuve and Liang, sp. nov. 

(Figs. 13, 39a, 46-48) 

Type material.— Holotype, a male, in IOZ, labeled: “CASENT 1024868”/ “CHINA, Yun¬ 
nan, Gongshan County, Bingzhongluo Township, SW slope of Kawakarpu Shan, on slope NE of 
Chukuai Lake, 3950 m,”/ “N27.98206/ E098.48027 0 , 20 August 2006, Stop #DHK-2006-086 
Y. Liu, P. Hu, D.Z. Dong, & J. Wang collectors”/ “HOLOTYPE Queinnectrechus (s, str .) gong¬ 
shanicus Deuve & Liang, sp. nov. designated 2016” [red label], Paratypes (a total of 10): 2 females 
(in CAS, IOZ) labeled same as holotype except first label “CASENT 1024866” and “CASENT 
1024867”, respectively; 1 male and 1 female (in IOZ, MNHN) labeled: “CASENT 1025160” and 
“CASENT 1025161”, respectively/ “CHINA, Yunnan, Gongshan County, Bingzhongluo Town¬ 
ship, SW slope of Kawakarpu Shan, 0.75 km NW of Chukuai Lake,”/ “N27.986317 E098.47069 0 , 
21 August 2006, Stop #DHK-2006-095 Y. Liu, P. Hu, & J. Wang collectors”; 1 male and 3 females 
(in CAS, IOZ) labeled “CASENT 1025935” and “CASENT 1025936” to “CASENT 1025938”, 
respectively/ “CHINA, Yunnan, Gongshan County, Bingzhongluo Township, SW slope of 
Kawakarpu Shan, 0.3 km SW of Chukuai Lake at campsite, “N27.976867 E098.44779 0 ”/ “3750 
m, 19-22 August 2006, Stop #DHK-2006-095C D.H. Kavanaugh, J.A. Miller, D.Z. Dong, Y. Liu, 
P. Hu, & J. Wang collectors”; 2 males (in CAS, MNHN) labeled “CASENT 1026203” and 
“CASENT 1026204”, respectively/ “CHINA, Yunnan, Gongshan County, Bingzhongluo Town¬ 
ship, S W slope of Kawakarpu Shan, 0.3 km NNE of Chukuai Lake, “N27.983937 E098.47491°”/ 
“3745 m, 19 August 2006, Stop #DHK-2006-081 D.H. Kavanaugh, J.A. Miller, & D.Z. Dong col¬ 
lectors”. All paratypes also bear the following label: “PARATYPE Queinnectrechus (s. str .) gong¬ 
shanicus Deuve & Liang, sp. nov. designated 2016” [yellow label]. 

Type locality.— China, Yunnan, Gongshan County, Bingzhongluo Township, SW slope of 
Kawakarpu Shan, on slope NE of Chukuai Lake, N27.98206/E098.48027 0 , 3950 m. 

d er i vat io n o f spe c ie s name .— The species epithet, gongshanicus , is derived from the name 
of the county (Xian) in which the holotype was collected, Gongshan, and the Latin adjectival suf¬ 
fix, -icus, meaning belonging to or pertaining to. 

diagnosis.— Adults of this species (Fig. 13a) can be distinguished from those of all other 
species in the region by the following combination of character states: same features as members 
of 0. griswoldi except, size smaller (BL = 3.5 to 3.8 mm), body form short, with elytra, in partic¬ 
ular, shorter and more oval, less elongate; digitiform projections of pronotal basal angles slightly 
more divergent laterally; elytral recurrent stria slightly deeper impressed, median lobe of male 
aedeagus (Fig. 13b) markedly different, with shaft thin and straighter, abruptly bent basally, apex 
short and rectangular, endophallus with a single apically acuminate internal sclerite. 

description.— Size smaller, BL = 3.5-3.8 mm. Body color dark, reddish-piceous, antennae 
and legs reddish tan, palpi and tarsi paler, yellowish tan. 

Head. Moderate in size; eyes small and convex, their convexity greater than and their diame¬ 
ter about as long as tempora, the latter moderately convex and glabrous. Frons not flattened; frontal 
furrows deep, rounded, slightly attenuated posterior to the eyes. Clypeus with four setae. Labram 
with six setae, anterior margin distinctly concave. Mentum and submentum fused. Mentum with 
medial tooth short, wide, bifid, less than one-half the length of the lateral lobes. Submentum with 
six setae anteriorly. Gula wide. Genae with a single seta ventrally on each side. Antennae slightly 
shorter, with only two antennomeres extended posteriorly beyond basal pronotal margin, anten- 
nomeres slightly broadened, antennomere 3 slightly longer than antennomeres 2 or 4. 

Pronotum. Cordate (ratio PW/PL = 1.15), markedly narrowed posteriorly, greatest width near 
anterior one-third, lateral margins rounded anteriorly, then straightened posteriorly just anterior to 
basal angles, the latter prolonged as slender, slightly divergent digitiform processes. Disc marked- 


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Yunnan 


MYANMAR 


Yengcftaeg 


Yingpng 


Figure 13. Queinnectrechus ( s. str .) gongshanicus sp. nov.; a. Dorsal habitus (CASENT1024868). b. Median lobe of 
aedeagus of male (CASENT1024868), left lateral aspect, c. Map of locality records (red circles) for Q. gongshanicus in the 
Gaoligong Shan region. Scale lines a, b = 0.5 mm, c = 100 km. 


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ly convex, smooth and glabrous, median longitudinal impression only superficially impressed; 
basal foveae distinct but small and round; median basal area smooth and markedly transverse, 
delimited anteriorly by a faint transverse impression; basal margin convex and rounded. Lateral 
border of pronotum rudimentary, extremely slender, distinctly defined only in anterior one-third to 
one-half, effaced in posterior one-half to two -thirds. Single midlateral setae on each side inserted 
at anterior one-third; single basolateral seta on each side, inserted on basal angle at base of digit- 
form process. 

Elytra. Short, ovoid, very slightly tear-shaped, narrower anteriorly than posteriorly, humeri 
effaced. Disc markedly convex and smooth. All discal striae effaced, except for a faintly impressed 
parascutellar stride and a short and shallow recurrent stria. Basal setiferous pore present. Three 
discal setae present, aligned along presumed track of stria 3. Preapicale seta absent, two or three 
apicoangular setules present. Umbilicate setal series with setae of humeral group equidistance from 
each other, with the first slightly more medially inserted than the others, setae of median group 
inserted distinctly posterior to middle. 

Legs. Slightly short but slender. Protibiae with longitudinal furrow, glabrous or sparely pubes¬ 
cent apically on anterior surface. Male protarsomeres 1 and 2 dilated and apicomedially toothed. 

Abdomen. Abdominal ventrites glabrous, except for a single paramedial seta on each side, and 
ventrite VII of males apically with one pair of paramedial setae, of females with two pairs. 

Male aedeagus. Median lobe (Fig. 13b) with shaft with shaft thin and straight, abruptly bent 
basally, apex short and rectangular, endophallus with a single long, apically acuminate internal 
sclerite. 

habitat distribution .— Members of this species have been found in a variety of microhab¬ 
itats in the alpine zone on the southwest slope of Kawakarpu Shan, at elevations ranging 3745 to 
3950 m. Specimens were collected from under stones in moist meadows, on tundra slopes and 
ridges with sparse to thick herbaceous vegetation, at the edges of small streams and seeps from 
talus slopes They were also collected in pitfall traps placed at the edges of Rhododendron thickets 
up to two meters tall. Members of this species have been found syntopic with specimens of Quein- 
nectrechus ( Gaoligongtrechus ) balli sp. nov., Queinnectrechus ( 5 , str.) giiswoldi and Trechus gong- 
shanensis sp. nov. 

Geographical distribution within the Gaoligong Shan.— Fig. 13c. We examined a 
total of 11 specimens (4 males and 7 females), all from the southwest slope of Kawakarpu Shan in 
the northern part of the Gaoligong Shan, in Bingzhongluo Township, Gongshan County (see Type 
material above for exact collection data). These localities are all in Core Area 2. 

Overall geographical distribution.— This species currently is known only from the 
northern part of the Gaoligong Shan, in western Yunnan Province, China. 

Subgenus GaoligOllgtrechllS Deuve and Kavanaugh, subgen. nov. 

Type species. — Queinnectrechus ( Gaoligongtrechus ) balli sp. nov. 

derivation of genus group name.— The genus group name (masculine) is a combination 
of Gaoligong , the name of the mountain range where this taxon was discovered, and the genus 
name Trechus. 

diagnosis.— Adults of this subgenus (Fig. 14a) can be recognized by the following combi¬ 
nation of character states: size moderate (BL = 4.3 to 4.8 mm), apterous, body dark and very shiny, 
micaceous; head slightly elongate, eye small, their diameter less than length of tempora; right 
mandibles tridentate; mentum and submentum fused; pronotum narrowed posteriorly, narrowly 
cordate, pronotal disc markedly convex and glabrous, basal angles acute but without digitiform 
extensions, basal margin roundly convex and fitted to concave elytral base, laterally with two or 


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three anteromedial setae and a single basal seta on each side; clytral silhouette tear-shaped, widest 
distinctly posterior to middle, humeri effaced, disc very convex and smooth, without distinct dis- 
cal striae, except recurrent stria short but distinctly impressed, with four to six discal setae aligned 
near presumed location of stria 3, preapical seta present; abdominal ventrite IV to VI each with a 
pair of parainedial setae; male aedeagus with median lobe rather slender, bent basally about 90° to 
shaft, endophallus with a pair of slender sclerites tapered to points on both ends. 

Geographical distribution.- — At present this subgenus includes the single species 
described below, which is known only from the northern part of the Gaoligong Shan in western 
Yunnan Province, China. 

Queinnectrechus ( Gaoligongtrechus ) balli Deuve and Kavanaugh, sp. nov. 

(Figs. 14, 37b, 39b, 40a, 46-48) 

Type material.— Holotype, a male, in IOZ, labeled: “CASENT 10019357 “CHINA, Yun¬ 
nan Province, Gaoligong Shan, Nujiang Prefecture, Nujiang State Nature Reserve, Dulong/Gong- 
shan Yakou [= Qiqi/Dulong divide] area, 21 airkm W of Gongshan,7 “N27.696557 E98.45389 0 , 
3300-3680m, 16-17 July 2000, Stop#00-24C, D.H. Kavanaugh, C.E. Griswold, Liang H.-B., D. 
Ubick, & Dong D.-Z. collectors”/ “HOLOTYPE Queinnectrechus ( Gaoligongtrechus ) balli Deuve 
& Kavanaugh, sp. nov. designated 2016” [red label]. Paratypes (a total of 32): 2 males and 12 
females (in CAS, IOZ, MNHN) labeled same as holotype except first label “CASENT 1001933” 
to “CASENT 1001934” and “CASENT 1001936” to “CASENT 1001947”, respectively; 1 female 
(in CAS) labeled “CASENT 10103437 “CHINA, Yunnan, Gongshan County, Cikai Township, 
52.6 km W of Gongshan on Dulong Valley Road, 3360-3380 m,7 “N27.770327 E098.44661 0 , 1- 
2 October 2002, Stop #DHK-2002-034, D.H. Kavanaugh & P.E. Marek collectors”; 1 female (in 
CAS) labeled “CASENT 10243757 “CHINA, Yunnan, Gongshan County, Cikai Township, south¬ 
east slope of Heipu Yakou, 3365 m, N27.770327 E098.44674 0 ,”/ “11 August 2006, Stop #DHK- 
2006-069A, D.H. Kavanaugh, J.A. Miller, D.Z. Dong, & Y. Liu collectors”; 3 males and 2 females 
(in IOZ, MNHN) labeled “CASENT 1025840” to “CASENT 1025842” and “CASENT 1025838” 
to “CASENT 1025839”, respectively/ “CHINA, Yunnan, Gongshan County, Bingzhongluo Town¬ 
ship, SW slope of Kawakarpu Shan, 0.3 Ion SW of Chukuai Lake at campsite,”/ “N27.976867 
E098.44779°, 3750 m, 19 August 2006, Stop #DHK-2006-082 Y. Liu collector”; 1 female (in CAS) 
labeled “CASENT 1026202”/ “CHINA, Yunnan, Gongshan County, Bingzhongluo Township, SW 
slope of Kawakarpu Shan, 0.3 km NNE of Chukuai Lake, N27.983937 E098.47491°,”/ “3745 m, 
19 August 2006, Stop #DHK-2006-081 D.H. Kavanaugh, J.A. Miller, & D.Z. Dong collectors”; 5 
males and 2 females (in CAS, IOZ) labeled ““CASENT 1026323” to “CASENT 1026327” and 
“CASENT 1026328” to “CASENT 1026329”, respectively/ “CHINA, Yunnan, Gongshan County, 
Cikai Township, 0.1 km SE of Heipu Yakou in valley below tunnel, N27.769787 E98.44681 0 ,”/ 
“3720 m, 13 August 2006, Stop #DHK-2006-073 D.H. Kavanaugh & J.A. Miller collectors”; 1 
male (in CAS) labeled “CASENT 10267077 “CHINA, Yunnan, Gongshan County, Dulongjiang 
Township, NW slope of Heipu Yakou, 3350 m, N27.774377 E09S.44793°,7 “13 August 2006, 
Stop #DHK-2006-075 D.H. Kavanaugh & J.A. Miller collectors”; 2 females (in IOZ) labeled 
“CASENT 1026817” and “CASENT 1026818”, respectively/ “CHINA, Yunnan, Gongshan Coun¬ 
ty, Bingzhongluo Township, SW slope of Kawakarpu Shan at Chukuai Lake, 3720 m,7 
“N27.981217 E098.47580 0 , 18 August 2006 Stop #DHK-2006-079 J.A. Miller, D.Z. Dong, & Y. 
Liu collectors”. All paratypes also bear the following label: “PARATYPE Queinnectrechus 
(Gaoligongtrechus) balli Deuve & Kavanaugh, sp. nov. designated 2016” [yellow label]. 

Type locality.— China, Yunnan, Gaoligong Shan, Gongshan County, Qiqi/Dulong divide 
area, N27.69655°/E98.45389°, 3300-3680 m. 


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Figure 14. Queinnectrechus ( Gaoligongtrechus ) balli sp. nov.; a. Dorsal habitus (CASENT1001935). b. Median lobe 
of aedeagus of male (CASENT1001935), left lateral aspect, c. Map of locality records (red circles) for Q. balli in the 
Gaoligong Shan region. Scale lines a, b = 0.5 mm, c = 100 km. 


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Derivation of species name.— The species epithet, bctlli, is the Latinized form (in the gen¬ 
itive case) of the surname of George Eugene Ball, Professor Emeritus at the University of Alberta, 
Edmonton, Alberta, Canada — our mentor, dear friend, and one of the world’s most accomplished 
and inspirational systematists. We are pleased to name this extraordinary species in his honor. 

diagnosis. — Adults of this species (Fig. 14a) can be distinguished from those of all other tre- 
chine species in the region by the combination of character states noted in the diagnosis for this 
genus. 

Description. — Size moderate, BL = 4.3 to 4.8 mm. Color of dorsum dark, forebody dark 
reddish brown, elytra piceous to black, antennae and legs reddish brown, palpi slightly paler, red¬ 
dish tan; surface smooth and markedly shiny. 

Head. Slightly elongate, with eyes small but convex, their diameter less than length of tempo- 
ra. Tempora not or only slightly convex, glabrous, and joined to neck region at a markedly obtuse 
(ca. 150°) angle. Frons with deep frontal furrow that delimit three (two lateral and medial) very 
convex areas, however furrows abruptly interrupted or less impressed posteriorly; two pairs of 
supraorbital seta present, the anterior pair inserted near middle of eyes, the posterior pair inserted 
dorsally on the tempora near the postocular furrow. Clypeus with four setae. Labrum with six setae, 
anterior margin distinctly concave. Right mandible tridentate, with the middle tooth reduced, 
obtuse, left mandible with premolar tooth fused with retinaculum to form a small caniniform 
process with a sharp tip. Mentum and submentum fused. Mentum with medial tooth broad, trun¬ 
cate, about half as long as lateral lobes, the latter apically pointed. Submentum with six setae ante¬ 
riorly. Genac with a single ventral seta on each side. Antennae pubescent from apical half of scape 
distally, extended posteriorly to basal one-third of elytra, with four antennomeres posterior to the 
pronotal base, antennomere 3 slightly longer than antennomere 4. 

Pronotum. Narrowly cordate, not transverse, about as long as wide (PW/PL = 1.0), markedly 
narrowed posteriorly, the lateral margins distinctly sinuate anterior to basal angles, the latter acute, 
projected, but without digitiform extensions. Disc markedly convex, globose, glabrous; median 
longitudinal impression faintly impressed; basal foveae formed as small, deep, circular pits, medi¬ 
an basal area smooth, faintly delimited, basal margin dilated medially as a broad, round projection. 
Lateral margination narrow, evident only in anterior one-third, effaced in posterior two-thirds. Two 
or three anteromedial setae and a single basal seta present on each side, the latter inserted slightly 
but distinctly anterior to basal angle. 

Elytra. Elytral silhouette tear-shaped, narrower anteriorly than posteriorly, widest distinctly 
posterior to middle, humeri effaced, basal part of lateral explanation not visible from above. Disc 
markedly convex and smooth, with basal part abruptly and truncate, concave, fitted to convex base 
of pronotum, without distinct discal striae, except recurrent stria short but evidently impressed. 
Parascutellar setiferous pore present. Four to six discal setae present and aligned near presumed 
location of stria 3. Preapical seta present, inserted slightly more medially than the row of discal 
setae. Umbilicate setal series with setae of humeral group aggregated with distance between first 
and second setae less than between second and third and third and fourth, setae of median group 
distinctly posterior to middle. 

Legs. Moderately long but slender; protibiae with longitudinal furrow, sparely pubescent api¬ 
cally on anterior surface. Male protarsomeres 1 and 2 dilated and apicomedially toothed. 

Abdomen. Abdominal ventrites glabrous, except for a single paramedial seta on each side and 
ventrite VII of females with two setae on each side. 

Male aedeagus. Median lobe (Fig. 14b) slender, bent basally about 90° to shaft, basal bulb with 
a large sagittal aileron, apex short and slightly narrowed, blunt; endophallus with a pair of slender 
sclerites tapered to points on both ends. 


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Comments. — Members of this new genus exhibit practically the same chaetotaxic pattern as 
those of genus Kozlovites Jeannel (1935), with a row of four to six discal setae apparently aligned 
on interval 3 in or near stria 3 and a preapical seta apparently inserted on interval 3 near stria 2 but 
in a position forward of the level of the anterior tip of the recurrent stria. However, 
Gaoligongtrechus members are distinguished from those of Kozlovites (and of Uenoites Belousov 
and Kabak 2016) by the extreme convexity of both the pronotum and elytra, which arc of similar 
fonn to that seen in members of Queinnectrechus (s. str. ), and by the posterior projection of the 
pronotal basal area and margin and its fit with the modified elytral base. Additional distinguishing 
features include the lateral marginations, each of which bears one or two anteromedial setae and is 
effaced in the posterior two-thirds, the effaced elytral discal striae, the medial tooth of the mentum 
truncate rather than bifid, the abdominal ventrites each with only a single pair of paramcdial setae, 
except for the female ventrite VII which has two pairs of subapical paramedial setae, as is typical 
among Trechini. In addition, Belousov and Kabak (2016) considered the apex of the male median 
lobe formed as a large apical hook as a synapomorphy for Kozlovites species. The apex of the medi¬ 
an lobe of O. (G. ) balli males (Fig. 14b) has no trace of a hook; and the slender shaft and abrupt¬ 
ly bent (at a 90° angle) basal region are unlike that seen in males of any described Kozlovites or 
Uenoites species. 

Members of subgenus Gaoligongtrechus differ from those of subgenus Queinnectrechus 
Deuve and Dactylotrechus Belousov and Kabak, (2003), in having the basal pronotal angles sim¬ 
ple, without the digitiform extensions seen in members of these two genera. Like Dactylotrechus 
members, those of our new subgenus have anterolateral setae in the pronotal margins and a preapi¬ 
cal seta apparently inserted near stia 2; however, this seta is inserted farther forward, in a subdis- 
cal position, in Q. (G.) balli members. 

Habitat distribution. — Members of this species have been found under stones on moist, 
organic substrate in alpine meadows, slopes and ridges with low, dense to sparse herbaceous veg¬ 
etation, at elevations ranging from 3300 to 3750 m (Fig. 37b, 39b, 40a). One specimen was col¬ 
lected at night, found walking on the barren slope of a roadcut at 3350 m elevation. Members of 
this species have been found together (syntopic) repeatedly with specimens of Queinnectrechus 
gongshanicus, Queinnectrechus griswoldi , Trechus gongshanensis sp. nov., and Trechus qiqiensis 
sp. nov. 

Geographical distribution within the Gaoligong Shan.— Fig. 14c. We examined a 
total of 33 specimens (12 males and 21 females), all from the northern part of the Gaoligong Shan, 
in Bingzhongluo, Cikai and Dulongjiang Townships in Gongshan County (see Type material above 
for exact collection data). These localities are in Core Areas 1 and 2. 

Overall geographical distribution.— This species currently is known only from the 
northern part of the Gaoligong Shan, in western Yunnan Province, China. 


Genus EoCtlideS Jeannel, 1954 

Eocnides Jeannel, 1954:10. 

Agonofrechiotes Deuve 2010:17. new synonymy 

Type species. — Eocnides assamensis Jeannel, 1954 

diagnosis. — Adults of this genus (Fig. 15a) can be recognized by the following combination 
of character states: size large (BL = 4.8 to 5.2 mm); body form long and slender, fully-winged; eyes 
moderately large; frontal furrows deep; mentum and submentum fused or not; antennae long and 
slender; pronotum small, elytra long and flattened, discal striae partially effaced basally and later¬ 
ally, with two discal setae present inserted in stria 3, umbilicate setae aggregated into humeral and 


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median groups; legs slender; protibiae longitudinally furrowed; abdominal ventrites with long, 
very sparse pubescence medially in addition to two or more apical paramedial setae; median lobe 
of male aedeagus slender, the apex slightly recurved ventrally, endophallus with internal sclerite 
very long and tapered. 

Comments. — This genus was described by Jeannel (1954) based on a single female speci¬ 
men, which he considered as close to genus Cnides Motschulsky (1862), based on overall form and 
features of elytral sculpture. However, the discovery of a second species and two males allowed 
Ueno (1989) to find that the median lobe of the male aedeagus was formed as a closed tube, not at 
all open on its ventral face. He concluded that it should be considered as a taxon in subtribe Trechi- 
na and placed near the “ Trechoblemus ” and “ Trechus ” Complexes. We here recognize for the first 
time Agonotrechiotes Deuve (2010) as a junior synonym of Eocnides Jeannel (1954). 

Geographical distribution .— This genus currently is known from only two species with a 
combined disjunct distribution including Assam (northeastern India) and Sichuan Province, China. 
The discovery of members of this genus in the study area in western Yunnan Province partially fills 
a gap in the distribution of the genus. 

Eocnides fragilis Ueno, 1989 
(Figs. 15,45-48) 

Eocnides fragilis Ueno, 1989:14. Holotype, a male, in NMST. Type locality: China, Sichuan, Nanping 

County, Jiuzhaigou, Xiajijie Hai, ca. 2600 m. 

Agonotrechiotes longiantennatus Deuve 2010:17. new Synonymy 

diagnosis. — Adults of this species (Fig. 15a) can be distinguished from those of all other 
species in the region by the following combination of character states: size large (BL = 4.8 to 5.2 
mm), fully-winged; body color pale, brownish-tan; head with eyes only moderate in size; frontal 
furrows attenuated; anterior pair of supraorbital setae not foveate; pronotum slightly transverse 
(ratio PW/PL= 1.48), median longitudinal impression distinctly impressed, especially basally, least 
so at middle; basal foveae prolonged anteriorly parallel to lateral margin in basal one-half; elytra 
with discal stria 2 partially effaced near base, striae 5 and 6 faintly but distinctly impressed; medi¬ 
an lobe of male aedeagus (Fig. 15b) slender, apex slightly recurved ventrally. 

Habitat distribution . — Members of this species were found by splashing gravel bars at the 
edges of a small, lowland stream adjacent to disturbed, agricultural areas, at elevations ranging 
from 1610 to 1630 m. This microhabitat is similar to that described by Ueno (1999b) for specimens 
collected at Munigou He, 2670 m, in Songpan County, northern Sichuan. No other trechines were 
collected at these sites. 

Geographical distribution within the Gaoligong Shan. — Fig. 15c. We examined a 
total of 2 specimens (1 male and 1 female) from the following localities: Gongshan County: 
Bingzhongluo Township (Niwaluo He, just below Nu Jiang Road, 1630 m, N28.05140 0 / 
E98.59319 0 , 8 October 2002, D.H. Kavanaugh, PE. Marek and D.Z. Dong collectors [1 male; 
CAS]; Bingzhongluo Township (Yimaluo He, just below Nu Jiang Road, 1610 m, N28.02499°/ 
E98.62564°, 8 October 2002, D.H. Kavanaugh, PE. Marek and H.B. Liang collectors [1 female; 
IOZ]. Both of these localities are in Core Area 2. 

Overall geographical distribution . — Fig. 45. This species has been recorded previous¬ 
ly from five localities in northern Sichuan Province, in Jiuzhaigou and Songpan Counties (Ueno 
1989, 1999b) and in Luding County (Deuve 2010). We report here the following new record for 
Sichuan: Wenchuan County : Qionglai Shan, Sanjiang Township (Sanjiang Nature Reserve, river 
at gate to reserve, 1300 m, N30.94697°/E103.31217°, 9 September 2007, D.H. Kavanaugh collec- 


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Figure 15. Eocnides fragilis Ueno; a. Dorsal habitus (CASENT1029485). b. Median lobe of aedeagus of male 
(CASENT1029485), left lateral aspect, c. Map of locality records (red circles) for E. fragilis in the Gaoligong Shan region. 
Scale lines a, b = 0.5 mm, c = 100 km. 


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379 


tor [1 female; CAS]. This record extends the range of E. fragilis in Sichuan. The discovery of 
E. fragilis in the northern Gaoligong Shan region, in western Yunnan Province, extends its known 
range an additional 260 km SW. 


Genus TrechllS Clairville, 1806 

Trechus Clairville, 1806:22. 

type species. — Carabus rubens, Clairville, 1806 [nee Fabricius, 1801] (= Carabus quadris- 
triatus Schrank, 1781), designated by Blanchard (in Audouin et al. 1841, plate 25). See also com¬ 
ments on type species by Bousquet (2012: 505). 

diagnosis. — Adults of this genus (Fig. 17-25) can be recognized by the following combina¬ 
tion of character states: size very small to small for family (BL = 2.5 to 7.0 mm), fully-winged or 
apterous, eyes large and projected or reduced, with some members eyeless; body color varied, from 
pale yellowish-tan to black; body form varied, compact and convex in most members, more slen¬ 
der and depressed in some members; labram with anterior margin concave; right mandible biden- 
tate or tridentate, but with the premolar fused with the retinaculum; submentum free, not fused with 
mentum, with six setae anteriorly in most members; pronotum with disc glabrous, two pairs of 
setae lateral present, one each side near middle and near basal angle; elytra with discal striae dis¬ 
tinctly impressed and complete or more or less effaced, recurrent stria distinct, discal setae only on 
interval 3, near or in stria 3 in most members, two setae present in most members, but with more, 
one or none present in a few members; preapical seta present near discal stria 2 or absent, in a few 
members present and inserted more anteriorly in a discal position on interval 3 near stria 2 or 3; 
umbilicate setae of the humeral group equally spaced; protibiae longitudinally furrowed or not. 

Comments. — As can be inferred from the above diagnosis, the large genus Trechus is 
markedly heterogeneous and probably polyphyletic. This hypothesis of polyphyletism has been 
corroborated by early results of analyzes of nucleic acid sequences data (Faille et al. 2010, 2013a). 

Among morphological features commonly used in comparative systematic studies of genus 
Trechus, two deserve special attention because they have allowed us to distinguish three new gen¬ 
era, described below, among species occuring in the Gaoligong Shan Mountains. These features 
include (1) the presence and position or the absence of a preapical elytral seta, and (2) the denti¬ 
tion of the mandibles. 

1) Presence and position of preapical seta: The primary elytral discal setae of carabid beetles 
are located on the odd intervals: 3, 5, 7 and 9 (Jeannel 1941). Jeannel named the setae of interval 
9 the “ serie ombiliquee ”, the umbilicate series. 

In members of subtribe Trechina, the umbilicate series includes a group of four consecutive 
humeral setae, a middle group of two consecutive setae, and two more isolated posterior setae; dis¬ 
cal setae are absent from interval 7; they occur on interval 5 in some members (eg., in Trechiama 
Jeannel (1927) and Epaphiopsis Ueno (1953)). In most members, there are three setae on interval 
3, inserted subbasally, near the middle and preapically, respectively. Although this can be consid¬ 
ered the basic number for members of the sub tribe, but this number is varied, more or less. 

In members of genus Trechus, there are typically three discal setae in interval 3, the first two 
inserted in or against stria 3, the third in prepical position inserted against stria 2. In some species 
(eg., Trechus perissus Andrewes (1936), described from Sikkim (see Ueno 1972a), or Trechus 
setitemporalis Deuve (2005), described from southern Xizang Autonomous Region), an additional 
discal seta is present, inserted next to stria 3, between the middle and preapical setae. In a few 
species, it is the preapical seta itself, typically inserted preapically against stria 2 that is advanced 
anteriorly to a discal position on the 3rd interval and inserted either in the center of the interval or 


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even against stria 3. In the latter case, it is difficult to know if it is actually the preapical seta shift¬ 
ed forward or a supernumerary discal seta with the preapical seta absent. Forward displacement of 
the preapical seta has been observed in other Trechina as well, such as in members of Epaphiopsis 
subgenus Pseudepaphius Ueno (1962), occuring in subtropical China. 

Among the representatives of the genus Trechus occuring in the Gaoligong Shan, we have 
identified a particular group of species that we call the “ Trechus qiqiensis Group”, members of 
which share this forward displacement of the preapical seta. We suggest that this is a synapomor- 
phy for this group. Moreover, with the exception of T. shiyueliang sp. nov., members of this group 
share another synapomorphy: the recurrent stria is continuous anteriorly with stria 7, which is 
unusual in genus Trechus , although previously observed in Trechus yasudai Ueno (1973) from east¬ 
ern Nepal. Based on these two unusual features, this group of species appears to represent a natu¬ 
ral, monophyletic group. 

Members of the Trechus qiqiensis group may be related to some Himalayan species, such as 
Trechus himalayanus Ueno (1972b), in which the preapical seta is displaced forward (Deuve 1988) 
but the recurrent stria is not in line anteriorly with stria 7. Forward displacement of the preapical 
seta has long been known to occur also in members of genus Epaphius Samouelle (1819). 

2) Dentition of the mandibles'. Jeannel (1926, 1941) separated the Trechini into two groups: (1) 
the “Tridentati”, with a premolar tooth on the mandibles, which grouped what he called the “more 
primitive” lineages; and (2) the “Bidentati’, “without a premolar tooth”, which corresponded to the 
Trechina, including genus Trechus. This fundamental dichotomy was accepted by most authors, 
who, as seemed appropriate, described the mandibles as either “tridentate” or “bidentate”. Howev¬ 
er, this was actually a source of confusion because assigned to the Bidentati were some members 
with a cleft retinaculum that appeared trifid. The mandibles of those Bidentati with a trifid reti¬ 
naculum were thus often called “tridentate” in species descriptions. However, Jeannel defined the 
distinction between the Bidentati of Tridentati precisely, with the criterion being the presence or 
absence of the premolar tooth. 

The mandibles are naturally asymmetrical in order to allow meshing of the teeth when closed. 
It is the right mandible which serves as the benchmark because it best shows the components: 
molar, premolar and retinacular blade [For a good understanding of this classic nomenclature, see 
Acom and Ball (1991)]. 

Study of cave Trechina of China recently has revealed that, within the genus Guizhaphaenops 
Vigna Taglianti (1997), members of some species had the right mandible tridentate while in those 
of other species it was bidentate. The explanation given was that the premolar tooth was merged 
with the retinaculum to form a trifid (“tridentate”) process, and that in the species with the “biden¬ 
tate” mandible, the median point of this process had been lost subsequently, resulting in a biden¬ 
tate process, formed in reality of the merged premolar tooth and the retinaculum retaining only its 
anterior tooth (Deuve and Queinnec 2014). It appears that what was described by Jeannel as dis¬ 
appearance of the premolar tooth was actually the result of a merger of the latter with the retinac¬ 
ulum. 

The combination of the unifid premolar with the bifid retinaculum to form a trifid process is 
evident in genus Queinnectrechus, for which the right mandible has been described and illustrated 
by Deuve (1992a, 1992b), then by Belousov and Kabak (2003), and in which the two teeth, pre¬ 
molar and retinacular, are not yet fully merged. 

Among Trechus members, the right mandible has been considered as of the “bidentate” type, 
but with both bidentate and tridentate retinacula represented (Jeannel 1941). In fact, in both cases 
the premolar tooth is present but fused with the retinaculum. This is the case among all the true 


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381 


Trechus we have examined from the Gaoligong Shan, which all show a trifid state (Fig. 16b-d). 
However, in members of a very specific and homogeneous group of species in the study area, the 
right mandible has dentition of another type: the premolar tooth is incompletely fused with retina- 
cle (a relatively symplesiomorphic condition) and the anterior tooth of the retinaculum is located 
forward (a synapomorphy), a greater distance from the posterior tooth (Fig. 16f). Members of this 
group of species also present two additional remarkable features. First, there is only one or no dis- 
cal setae on elytral interval 3; and second, the tempora are sparsely pubescent, whereas they are 
glabrous in all other “ Trechus ” of the Gaoligong Shan region. Members of some species in Tibet, 
such as Trechus setitemporalis Deuve (2005), or those of the “ Trechus dacatraianus Group” 
(Schmidt 2009), also have pubescent tempora. However, it is the combination of the three states 
mentioned above, especially the mandibular structure, which leads us to exclude these species from 
Trechus and consider them as representing a distinct genus within the Epaphiopsis Complex of 
genera. Descriptions of this genus and the included species are provided below. A related species, 
but one in which members have the tempora glabrous and the premolar and retinaculum of the right 
mandible (Fig. 16h) more fully fused, is also excluded from Trechus and described as a separate 
genus. Finally, we also exclude two additional species from the southern part of the Gaoligong 
Shan from Trechus. In members of these species, the right mandible (Fig. 16g) is very similar to a 
bidentate type, but the premolar tooth is not fused with the retinaculum. We consider these also as 
representing a new and distinct genus described below. 

Geographical distribution.— Genus Trechus is a megadiverse taxon with more than 900 
described species and subspecies arrayed in eight subgenera (Lorenz 2005); and about 95% of 
these species-group taxa are currently classified in the nominate subgenus. The genus is predomi¬ 
nately Holarctic in distribution and widespread in that Region (Jeannel 1927, Casale & Laneyrie 
1982), with a few species also recorded from the Afrotropical and Oriental Regions. Several 
species from subtropical or tropical parts of Southeast Asia have been described in genus Trechus. 
Examples of such species include Trechus thai Deuve (1995) from Thailand, Trechus myanmaren- 
sis Deuve (2005) and Trechus natmataungensis Donabauer (2010) from Myanmar and Trechus 
vietnamicus Ueno (1995) from Vietnam. However, the genus is not known from tropical or sub¬ 
tropical parts of China, where it appears to be replaced by species representing the Epaphiopsis 
Complex of genera. 


Key for Identification ofAdults of TrechllS Species of the Gaoligong Shan Region 

1. Size larger (BL = 4.2 to 5.0 mm), fully-winged; elytra elongate, recurrent stria terminated 
abruptly on interval 6; preapical seta absent or vestigial and inserted near elytral apex, also 

without a third discal seta in apical one-fouth of elytra. T. indicus Putzeys 

Size varied, but generally smaller (BL = 3.3 to 4.7 mm), elytra short, recurrent stria continu¬ 
ous anteriorly with stria 5 or 7, preapical seta present and inserted near stria 2 in a subdiscal 
position or as third discal seta near stria 3 in apical one-fourth of elytra ( Trechus qiqiensis 
Group).2 

Trechus qiqiensis Group: 

2. Elytra with recurrent stria in line anteriorly with stria 5, preapical seta inserted near stria 2; size 
moderately large for group (BL = 4.0 to 4.2 mm) ( Fugong County). . T. shiyueliang sp. nov. 
Elytra with recurrent stria in line anteriorly with stria 7, preapical seta inserted in interval 


3, either near stria 2 or stria 3; size varied.3 

3. Size larger, BL at least 4.0 mm.4 

Size smaller, BL less than 4.0 mm.6 


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4. Size larger (BL = 4.5-4.7 mm); pronotum less transverse (ratio PW/PL = 1.40), with anterior 
transverse impression indistinct; elytra broad, disc slightly flattened ( Tengchong County). .. 

. T. mingguangensis sp. nov. 

Size smaller (BL = 4.0 to 4.2 mm); pronotum more transverse (ratio PW/PL = 1.44 to 1.46), 
with anterior transverse impression distinct; elytra ovoid, disc convex, not at all flattened . 5 

5. Elytra with discal striae very distinctly punctate and slight attenuated toward elytral apex; 

pronotum with basal foveae deep, distinct ( Gongshan County) . T. qiqiensis sp. nov. 

Elytra with discal striae only faintly punctate and not attenuated toward elytral apex; prono¬ 
tum with basal foveae shallow, less well defined ( Fugong County) . 

... T. pseudoqiqiensis sp. nov. 

6. Size larger (BL = 3.7-3.8 mm), elytra with lateral discal striae more shallowly impressed but 

still clearly evident ( Lushui County) . T. luz.hangensis sp. nov. 

Size smaller (BL = 3.3-3.5 mm); elytra with discal striae 6 and 7 effaced, faintly or not at all 
evident.7 

7. Tempora only slightly convex ( Gongshan County) (Fig. 23a) . ... T. gongshanensis sp. nov. 

Tempora markedly convex ( Fugong County’) (Fig. 24a). T. shibalicus sp. nov. 

Trechus ( Trechus ) indicus Putzeys, 1870 

(Figs. 16b, 17,41a, 44, 45-48) 

Trechus indicus Putzeys, 1870:175. Holotype, lost (see Jeannel 1923, footnote p. 416). Type locality: eastern 
India. 

Trechus ( s. sir.) indicus Putzeys: Jeannel, 1927:157, 158 (in part) 

Trechus (.S', str.) mucrops Jeannel, 1927:157, 160 (in part). Type locality: China, Yunnan. Synonymized by 
Jeannel (1935: 275). 

Trechus macrops Jeannel: Andrewes, 1935 63, 67. 

Trechus indicus Putzeys: Jeannel, 1935:275. 

Trechus indicus Putzeys: Deuve, 1988:80. 

Trechus (s. str.) indicus Putzeys: Ueno 1977:181. 

Trechus (s. str) macrops Jeannel: Ueno & Yin, 1993:354. These authors considered T. macrops as a distinct 
species. 

diagnosis.— Adults of this species (Fig. 17a) can be distinguished from those of all other 
species in the region by the following combination of character states: size large for genus (BL = 
4.2 to 5.0 mm); fully-winged; eyes markedly large, tempora very short, joined to neck region at 
nearly a right angle; frontal furrows angulatc at midlength; right mandible as in Fig. 16b; mentum 
and submentum not fused; mentum with medial tooth simple, triangular; antennae with anten- 
nomeres 3 and 4 subequal in length, antennomere 2 slightly shorter; pronotum transverse (ratio 
PW/PL = 1.43), with lateral margins not sinuate posteriorly, straight just anterior to basal angles, 
the latter small and rectangular, lateral explanation very wide basally, basal margin slightly convex 
medally; elytra oblong, lateral discal striae effaced, only the medial four or five striae distinctly 
impressed, recurrent stria markedly impressed; preapical seta absent or vestigial, two discal setae 
present, inserted in stria 3 at anterior one-fifth and at middle, respectively; median lobe of male 
aedeagus (Fig. 17b) not arcuate, with apex slender, ventrally hooked, endophallus with two elon¬ 
gate, projecting sclerites, one apically tapered, the other apically lobate. 

Comments.— Ueno (1977:182) correctly pointed out that this widely distributed species, with 
members ftilly-winged, is morphologically varied across its range. In particular, the preapical seta 


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383 


tends to be absent from members of eastern populations. The many specimens we examined from 
localities in the study area confirm his observation. Among them, the prepical seta is absent from 
most specimens. If present, the seta is very small or vestigial, and only present unilaterally in most 
such cases. This led Ueno (1999a:215), while reporting the presence of this species in the Gaoli- 
gong Shan for the first time, to consider the eastern populations, including those in the study area, 
as a separate species, T. macrops Jeannel. 

h a B i t a t distribution — Members of this species have been found in a variety of microhab¬ 
itats at elevations ranging from 1230 to 2486 m. They have been collected in daytime from under 
stones on the shaded and open banks of small to large streams, along roadcuts, and in closed 
canopy forest. At night, they have been found active on open sandy beaches and floodplain flats of 
larger streams (Fig. 41a) and on the ground along roadcuts. In almost all of the localities where 
specimens of T indicus were founded, they were the only trechine collected. However, they were 
found syntopic with specimens of Agon otrechus xiaoheishan at Xiaoheishan Forest Reserve in the 
southern part of the study area. Ueno (1999a) reported that specimens of this species (which he 
recorded as Trechus macrops Jeannel) were found in the same area as specimens of Agonotrechus 
yunnanus and Epaphiotrechus fortipes (Ueno) comb. nov. 

Geographical distribution within the Gaoligong Shan. — Fig. 17c. We examined a 
total of 86 specimens (45 males and 41 females) from the following localities: Fugong County-. 
Lumadeng Township (Lumadeng, 1230 m, N27.02606°/E98.86269°, 23 April 2004, D.H. 
Kavanaugh collector [1 female; CAS]). Gongshan County-. Dulongjiang Township (Bapo, 1412 m, 
N27.73902°/E98.34975°, 3 November 2004, H.B. Liang collector [1 male; IOZ]; Bapo area, 
Dulong Jiang at Mulangdang, 1355 m, N27.75256°/E98.34745°, 4 November 2004, H.B. Liang 
collector [1 male; IOZ]; 0.6 km N of Dizhengdang village on Dulong Jiang, 1880 m, N28.084427°/ 
E98.32652°, 29-30 October 2004, D.H. Kavanaugh, D.Z. Dong & G. Tang collectors [9 male and 
8 females; CAS, IOZ]; Dulong Jiang at Elideng village, 1640 m, N28.000287°/E98.32145°, 
3 November 2004, D.H. Kavanaugh, D.Z. Dong & G. Tang collectors [13 male and 8 females; 
CAS, IOZ]; Dulong Jiang at Xianjiudang village, 1580 m, N27.940927E98.33340°, 4 November 
2004, D.H. Kavanaugh, M.A. Dixon, D.Z. Dong & G. Tang collectors [1 male and 7 females; CAS, 
IOZ]; 0.2 km S of confluence of Dulong Jiang and Muke Wang, 1450 m, N27.841257E98.33979°, 
7 November 2004, D.H. Kavanaugh, V.F. Lee & D.Z. Dong collectors [1 female; CAS]; 0.5 km N 
of Kongdang, 1500 m, N27.881117E98.34063°, 25 October 2004, D.H. Kavanaugh, H.B. Liang, 
D.Z. Dong & G. Tang collectors [1 female; CAS]; Moqie Wang at Gongshan-Dulong Road Km 91, 
1550 m, N27.899347E98.34999°, 6 November 2004, D.H. Kavanaugh & H.B. Liang collectors [2 
males and 4 female; CAS, IOZ]). Longling County-. Longjiang Township (small stream 1.2 km 
SSE of Km 23.5 on Route 23.5, 2020 m, N24.28887E98.76001 0 , 25 May 2005, D.H. Kavanaugh, 
H.B. Liang & D.Z. Dong collectors [1 male; CAS]; Xiaoheishan Forest Reserve, 2067 m, 
N24.83671°/E98.76185°, 28 May 2005, H.B. Liang, K.J. Guo & H.M. Yan collectors [3 males; 
CAS, IOZ]); Zhen’an Township (Bangbie village, 1540 m, N24.81306°/E98.83306°, 30 October 
2003, H.B. Liang & X.C. Shi collectors [1 male; IOZ]). Longyang County: Bawan Township 
(Baoshan-Tenchong Road Km 24 at Nankang Yakou, 2130 m, N24.825837E98.77222°, 26 Octo¬ 
ber 2003, H.B. Liang & X.C. Shi collectors [1 female; IOZ]; Bawan-Tengchong Road at Km 40- 
41, 2404 m, N24.93750°/E98.75083°, 12 October 2003, H.B. Liang & X.C. Shi collectors [1 male; 
IOZ]; Bawan-Tengchong Road at Km 41, 2486 m, N24.93750°/E98.75083°, 11 October 2003, 
H.B. Liang collector [1; female; IOZ]; Bawan-Tengchong Road at Km 42-46, 2290 m, 
N24.95361 °/E98.74222°, 14 October 2003, H.B. Liang & X.C. Shi collectors [1 female; IOZ]; 
Luoshuidong area at Sancha He, 2300 m, N24.94833°/E98.75667°, 26-31 October 1998, D.H. 
Kavanaugh & C.E. Griswold collectors [1 female; CAS]). Tengchong County: Jietou Township 


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Figure 16. Trechine right mandibles, ventral aspect, a. Agonotrechus xiaoheishan sp. nov. b. Trechus indicus Putzeys 
with dentition enlarged, c. Trechus gongshanensis sp. nov. d. Trechus shibalicus sp. nov. e. Pseudepaphius gonggaicus 
Deuve with dentition enlarged (China, Sichuan Province, Moxi Township, NE slope of Gongga Shan), f. Trechepaphiopsis 
uniporosa sp. nov. with dentition enlarged, g. Epaphiotrechus fortipesoides sp. nov. with dentition enlarged, h. Trechepa- 
phiama gaoligong sp. nov. Scale lines = 0.5 mm. 


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Figure 17. Trechus indicus Putzeys; a. Dorsal habitus (CASENT1016115). b. Median lobe of aedeagus of male 
(CASENT1015874), left lateral aspect, c. Map of locality records (red circles) for T. indicus in the Gaobgong Shan region. 
Scale lines a, b = 0.5 mm, c = 100 km. 


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(Zhoujia-po village, 1740 m, N25.33222°/E98.67611°, 24 October 2003, D.Z. Dong collector 
[1 male; CAS]); Qushi Township (Longchuan Jiang at Xiaojiangqiao, 1445 nm, N25.23944 0 / 
E98.63722°, 21 October 2003, H.B. Liang & X.C. Shi collectors [1 male; IOZ]); Shangying Town¬ 
ship (Bawan-Tengchong Road Km 46-51, 2220 m, N24.95722°/E98.73667°, 17 October 2003, 
H.B. Liang & X.C. Shi collectors [1 male; IOZ]); Bawan-Tengchong Road Km 48-51 at Dahaop- 
ing Forest Station, 2014 m, N24.97556°/E98.73000°, 18 October 2003, H.B. Liang collector 
[6 males and 6 females; CAS, IOZ, MNHN]; Bawan-Tengchong Road Km 65, beside Longchuan- 
jiang, 1335 m, N24.04167°/E98.67306°, 19 October 2003, H.B. Liang & X.C. Shi collectors 
[1 male and 1 female; CAS IOZ]; Bawan-Tengchong Road Km 65 at Longwenqiao, 1285 m, 
N25.02396°/E98.67675°, 20 October 2003, H.B. Liang & X.C. Shi collectors [1 male; IOZ]); 
Wuhe Township (Xiaoheishan Forest Station, 2025 m, N24.82889°/E98.76000°, 29 October 2003, 
H.B. Liang collector [1 male and 1 female; CAS, IOZ]). 

Members of this species have been collected in both northern and southern parts of the study 
area and on both eastern and western slopes of the Gaoligongshan. We have recorded this species 
from Core Areas 1, 3, 6, and 7. The absence of records from Core Areas 2, 4 and 5 is likely the 
result of inadequate sampling. 

Overall geographical distribution.— Fig. 45. This species is widely distributed along 
the southern edge of the Himalayan Mountains and Qinghai-Xizang (Tibetan) Plateau from eastern 
Afghanistan to Sichuan Province, China. Attainment and maintenance of this broad geographical 
range is no doubt facilitated by the flight capability of members of this species. 


Trechus ( Trechus ) shiyueliang Deuve and Kavanaugh,sp. nov. 

(Figs. 18, 40b, 42b, 46-48) 

Type material.— Holotype, a male, in IOZ, labeled: “CASENT 1021004”/ “CHINA, Yun¬ 
nan, Fugong County, Lishadi Township, 10-11 km W of Shibali on Shibali Road, 3200-3280 m, 
NN27.199807E98.71375° to N27.20654°/E98.71772°,7 8 August 2005, Stop#DHK-2005-068 
D.H. Kavanaugh, H.B. Liang, P. Paquin, & D.Z. Dong collectors”/ “IMAGE” [green label]/ 
“HOLOTYPE Trechus shiyueliang Deuve & Kavanaugh, sp. nov. designated 2016” [red label]. 
Paratypes (a total of 8): 1 female (in IOZ) labeled same as holotype, except first label: “CASENT 
1021005”; 1 male and 1 female (in IOZ) labeled: “CASENT 1018374” and “CASENT 1018373”, 
respectively/ “CHINA, Yunnan, Fugong County, Lishadi Township, 10 km W of Shibali on Shibali 
Road, 3221 m,7 , N27.20055°/E98.71399°, 16 August 2005, Stop #PP-3805 P. Paquin collector”; 
1 female (in CAS) labeled: “CASENT 1023759”/ “CHINA, Yunnan, Fugong County, Lishadi 
Township, 8.5 km above Shibali on Shibali Road, North Fork of Yamu He, 
N27.18416°/E98.72026°,7 “3100 in, 7 May 2004 Stop #DHK-2004-038AD.H. Kavanaugh, C.E. 
Griswold, Liang H.-B., & Zhu B.-X. collectors”; 1 male (in IOZ) labeled: “CASENT 1020019”/ 
“CHINA, Yunnan, Fugong County, Lumadeng Township, ridge S of Shibali Yakou, 
N27.20802°/E98.69644°,”/ “3740 m, 12 August 2005, Stop #DHK-2005-092 D. H. Kavanaugh 
collectors”; 2 males and 1 female (in CAS) labeled: “CASENT 1021323” to “CASENT 1021324” 
and “CASENT 1021325”, respectively/ “CHINA, Yunnan, Fugong County, Lumadeng Township, 
Lao Shibali Yakou, 3270 m, N27.06429°/E98.75123°, 13 August 2005,”/ “Stop# DHK-2005-079, 
D.H. Kavanaugh, H.B. Liang, D.Z. Dong, & G. Tang collectors”. All paratypes also bear the fol¬ 
lowing label: “PARATYPE Trechus shiyueliang Deuve & Kavanaugh, sp. nov. designated 2016” 
[yellow label]. 

type locality.— China, Yunnan, Fugong County, Shiyueliang Township, 10-11 km W of 
Shibali on Shibali Road, 3200-3280 m. 

derivation of species name.— The species epithet, shiyueliang , a noun in apposition, is 


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387 


derived from the name of the township in which the type locality is located, Shiyueliang. This is 
the current name for the former Lishadi township. 

diagnosis. — Adults of this species (Fig. 18a) can be distinguished from those of all other 
species in the region by the following combination of character states: size relatively large (BL = 
4.0 to 4.2 mm), apterous; body color reddish-brown; head slightly elongate; tempora glabrous; 
pronotum moderately transverse (ratio PW/PL = 1.42); elytra with striae not or faintly punctate, 
recurrent stria continuous anteriorly with stria 5, two discal setae present and inserted next to stria 
3, preapical seta present and inserted next to stria 2 in forward position near apical one-fourth of 
elytra; median lobe of male aedeagus with apex thin and faintly reflexed dorsally in lateral view 
(Fig. 18b), endophallus with only a single, small thin elongate sclerite. 

Description .— Size relatively large, BL = 4.0 to 4.2 mm. Body color reddish-brown, shiny, 
appendages slightly paler. Body surface smooth, head capsule finely alutaceous. 

Head. Moderate in size, slightly elongate and thick; eyes small and moderately convex, their 
diameter about 1.5 times as long as tempora, the latter short, moderately convex and glabrous. 
Frons slightly flattened; frontal furrows deep, rounded, slightly attenuated posterior to the eyes. 
Two pairs of supraorbital setae present, the anterior pair inserted in foveae. Clypeus with four setae. 
Labrum with six setae, anterior margin distinctly concave. Mandibles short; right mandible triden- 
tate with middle tooth closer to basal tooth (premolar) than to distal tooth. Mentum and submen- 
tum not fully fused but nearly so, suture between them only faintly impressed. Mentum with medi¬ 
al tooth apically truncate, less than one-half the length of the lateral lobes. Submentum with six 
setae anteriorly, swollen anteriorly. Gula wide. Genae with a single seta ventrally on each side. 
Antennae short, with less than two antennomeres extended posteriorly beyond basal pronotal mar¬ 
gin; antennomeres 3 and 4 slightly longer than antennomere 2. 

Pronotum. Moderately transverse (ratio PW/PL = 1.42), narrowed posteriorly, greatest width 
near anterior one-third; lateral margins markedly rounded, straightened only just anterior to small, 
rectangular and sharp basal angles. Disc convex, smooth and glabrous, median longitudinal 
impression deeply impressed, interrupted anteriorly a short distance from anterior margin; basal 
foveae distinct; median basal area with several faint, irregular, punctiform depressions, delimited 
laterally by oblique furrows. Lateral borders of pronotum moderately slender, regular, slightly 
reflexed, lateral grooves deeply impressed. Single midlateral setae on each side inserted at anteri¬ 
or one-third; single basolateral seta on each side, inserted slightly anterior to apex of basal angle. 

Elytra. Ovoid, humeri distinct but rounded. Disc convex and smooth. Striae not or only faint¬ 
ly punctate, striae 1 to 5 deeply impressed, striae 6 to 8 very faintly impressed but evident. Paras- 
cutellar striole present. Recurrent stria continuous anteriorly with stria 5. Basal setiferous pore 
present at common origin of striae 1 and 2. Two discal setae present and inserted next to stria 3, 
one at anterior one-fourth and one near middle of elytra. Preapical seta present and inserted on 
interval 3 next to stria 2 in forward position near apical one-fourth of elytra. 

Umbilicate setal series with setae of humeral group equidistance from each other and setae of 
median group inserted slightly posterior to middle. 

Legs. Short, protibiae with longitudinal furrow. Male protarsomeres 1 and 2 dilated and api- 
comedially toothed. 

Abdomen. Abdominal ventrites glabrous, except for a single paramedial seta on each side, and 
ventrite VII of males apically with one pair of paramedial setae, of females with two pairs. 

Male aedeagus. Median lobe (Fig. 18b) moderately broad basally with sagittal aileron present 
and moderate in size; shaft gradually narrowed toward apex; apical lamella short, rectilinear, 
extremely thin and slightly recurved dorsally in lateral view; endophallus with a single thin and 
elongate sclerite. 


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Figure 18. Trechus shiyueliang sp. nov.; a. Dorsal habitus (CASENT1021004). b. Median lobe of aedeagus of male 
(CASENT 1021004), left lateral aspect, c. Map of locality record s(red circles) for T. shiyueliang in the Gaoligong Shan 
region. Scale lines a, b = 0.5 mm, c = 100 km. 


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habitat distribution Members of this species have been found in a variety of microhab¬ 
itats at elevations ranging from 3100 to 3740 m. They have been collected in daytime from under 
stones in thickets of bamboo and Primus , in rocky open areas cleared by snow avalanches (Fig. 
40b), in meadows adjacent to bamboo and Rhododendron thickets (Fig. 42b) and along roadcuts 
through such thickets. They have also been collected at night, found walking on the substrate along 
roadcuts through areas of bamboo with an overstory of scattered A hies trees. At higher elevations, 
they were the only trechine encountered in this area; but they were found syntopic with specimens 
of Trechus shibalicus sp. nov., Trechepaphiopsis unipilosa sp. nov. and Trechepaphiopsis unisetu- 
losa sp. nov. at the upper limit of the altitudinal range of this last mentioned species (3100 m). 

Geographical distribution within the Gaoligong Shan.— Fig. 18c. We examined a 
total of 9 specimens (5 males and 4 females), all from Fugong County on the eastern slope and crest 
of the northcentral part of the Gaoligong Shan (see Type material above for exact collection data). 
These localities are all in Core Area 3. 

Overall geographical distribution.— This species currently is known only from the 
northcentral part of the Gaoligong Shan, in western Yunnan Province, China. 

Trechus ( Trechus ) mingguangensis Deuve and Liang, sp. nov. 

(Figs. 19, 41b, 46-48) 

Type material.— Holotype, a male, in IOZ, labeled: “CASENT 10386927 “CHINA, Yun¬ 
nan, Tengchong County, Mingguang Township, Eighth Boundary Post Pass, N25.80984°/ 
E98.62084°, 2287 m, 23 May 2006,”/ “Stop # DHK-2006-037A, D.H. Kavanaugh, R.L. Brett, X.P. 
Wang & D.Z.Dong collectors”/ “HOLOTYPE Trechus mingguangensis Deuve & Liang, sp. nov. 
designated 2016” [red label]. Paratypes (only 1): a female (in CAS) labeled: “CASENT 1038840”/ 
“CHINA, Yunnan, Tengchong County, Mingguang Township, small stream on SW-facing slope 
below 7.9 airkm N of Zizhi village, 2200 m,7 “N25.80314YE98.62117°, 2200 m, 23 May 2006,7 
“Stop # DHK-2006-038A, D.H. Kavanaugh & D.Z. Dong collectors”/ “PARATYPE Trechus ming¬ 
guangensis Deuve & Liang, sp. nov. designated 2016” [yellow label]. 

Type locality.— China, Yunnan, Tengchong County, Mingguang Township, Eighth Bound¬ 
ary Post Pass, N25.803147E98.62117°, 2287 m. 

derivation of species name.— The species epithet, mingguangensis, is derived from the 
name of the township in which the type locality is located, Mingguang, and the Latin suffix, -ensis, 
denoting place. 

diagnosis. — Adults of this species (Fig. 19a) can be distinguished from those of all other 
species in the region by the following combination of character states: size large (BL = 4.5 to 4.8 
mm), apterous; body color reddish-brown; tempora glabrous, short and convex; pronotum trans¬ 
verse (ratio PW/PL = 1.40), basal angles acute and sharp; elytra very slightly elongate, striae fine¬ 
ly punctate, striae 1 to 4 deeply impressed, stria 5 more faintly impressed, striae 6 to 8 effaced or 
nearly so, recurrent stria terminated anteriorly at a slight convexity in line with the presumed loca¬ 
tion of sUia 7 apically; two normal discal setae present, inserted next to stria 3, preapical seta pres¬ 
ent, inserted in a discal position next to stria 3 at apical one-fourth of elytra; median lobe of male 
aedeagus with a large sagittal aileron, apical lamella long and rectilinear, nearly straight, apex thin, 
endophallus with copulatory piece poorly defined (Fig. 19b). 

description.— Size large, BL = 4.5 to 4.8 mm. Body color dark reddish-brown, shiny, 
appendages paler, yellowish- or reddish-tan, palpi pale yellow. Body surface smooth, only head 
capsule faintly alutaceous. 

Head. Moderate in size; eyes slightly reduced, moderately convex, but their diameter less than 
twice as long as tempora, the latter short, moderately convex and glabrous. Frons moderately flat- 


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Figure 19. Trechus mingguangensis sp. nov.; a. Dorsal habitus (CASENT1038692). b. Median lobe of aedeagus of 
male (CASENT1038692), left lateral aspect, c. Map of locality records (red circles) for T. mingguangensis in the Gaoligong 
Shan region. Scale lines a, b = 0.5 mm, c = 100 km. 


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tened; frontal furrows deep, curved, prolonged and not attenuated posterior to the eyes. Two pairs 
of supraorbital setae present, the anterior pair inserted in foveae. Clypeus with four setae. Labrum 
with six setae, anterior margin distinctly concave. Mentum and submentum not fused but suture 
between them only faintly impressed. Mentum with medial tooth broad and apically truncate, about 
one-half the length of the lateral lobes. Submentum with six setae anteriorly. Gula wide. Genae 
with a single seta ventrally on each side. Antennae short, not quite extended to the basal one-fourth 
of elytra, male with about 2.5 to 3 antennomeres extended posteriorly beyond basal pronotal mar¬ 
gin; antemiomeres 2 and 3 about equal in length, antennomere 4 slightly shorter. 

Pronotum. Moderately transverse (ratio PW/PL = 1.40), narrowed posteriorly, greatest width 
anterior to middle; lateral margins widely rounded, straightened only just anterior to basal angles, 
the latter small, but projected, acute and sharp. Disc convex, smooth and glabrous, median longi¬ 
tudinal impression sharply impressed, not quite extended to anterior margin; basal foveae distinct, 
subcircular; median basal area with several faint, longitudinal rugulae, delimited laterally by 
oblique furrows. Basal margin slightly bisinuate, slightly reflexed and oblique laterally, slightly 
projected medially. Lateral borders of pronotum moderately slender, regular, slightly reflexed, lat¬ 
eral grooves deeply impressed. Single midlateral setae on each side inserted slightly anterior to 
middle; single basolateral seta on each side, inserted at basal angle. 

Elytra. Ovoid, only very slightly elongate, more narrowed basally than apically, humeri dis¬ 
tinct but rounded. Disc convex and smooth. Striae very finely punctate, striae 1 to 4 deeply 
impressed, stria 5 more faintly impressed, striae 6 to 8 effaced or nearly so. Parascutellar striole 
present. Recurrent stria terminated anteriorly by a slight convexity at the presumed location of stria 
7 apically. Basal setiferous pore present at common origin of striae 1 and 2. Two discal setae pres¬ 
ent and inserted next to stria 3, one at anterior one-fourth and one near middle of elytra. Preapical 
seta present and inserted in a discal position on interval 3, next to stria 3 in forward position at api¬ 
cal one-fourth of elytra. Umbilicate setal series with setae of humeral group equidistance from each 
other and setae of median group inserted slightly posterior to middle. 

Legs. Relatively short, protibiae with longitudinal furrow. Male protarsomeres 1 and 2 dilated 
and apicomedially toothed. 

Abdomen. Abdominal ventrites glabrous, except for a single paramedial seta on each side, and 
ventrite VII of males apically with one pair of paramedial setae, of females with four pairs. 

Male aedeagus. Median lobe (Fig. 19b) only slightly enlarged basally but with a large sagittal 
aileron present; shaft markedly narrowed at middle and broader subapically; apical lamella long, 
rectilinear, extremely thin in lateral view; endophallus voluminous but without well-defined scle- 
rites. 

habitat distribution. — One specimen of this species was found under stones and debris 
along a roadcut (Fig. 41b) through slightly disturbed primary forest of Tsuga , with small, scattered 
small bamboo thickets; and the other specimen was found under stones along a small stream cross¬ 
ing the road in the same general area. No other trechines were collected in either of these sites, 
which range in elevation from 2200 to 2287 m. 

Geographical distribution within the Gaoligong Shan.— Fig. 19c. We examined a 
total of 2 specimens, both from Mingguang Township in Tengchong County on the western slope 
of the southern part of the Gaoligong Shan (see Type material above for exact collection data). Both 
of these localities are in Core Area 6. 

Overall geographical distribution.— This species currently is known only from the 
western slope of the southern part of the Gaoligong Shan, in western Yunnan Province, China. 


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Trechus ( Trechus) qiqiensis Deuve and Kavanaugh, sp. nov. 

(Figs. 20, 37b, 42a, 46-48) 

Type material.— Holotype, a male, in IOZ, labeled: “CASENT 1007384”/ “CHINA, Yun¬ 
nan Province, Gaoligong Shan, Nujiang Prefecture, Nujiang State Nature Reserve, No. 12 Bridge 
Camp area, 16.3 airkm W of Gongshan”/ “N27.71503°/E98.50244°, 2775 m, 15-19 July 2000, 
Stop#00-23A, D.H. Kavanaugh, C.E. Griswold, Liang H.-B., D. Ubick, & Dong D.-Z. collectors”/ 
“IMAGE” [green label]/ “HOLOTYPE Trechus qiqiensis Deuve & Kavanaugh, sp. nov. designat¬ 
ed 2016” [red label]. Paratypes (at total of 11): 2 males and 4 females (in CAS, IOZ, MNHN) 
labeled: same as holotype, except first label “CASENT 1007380” to “CASENT 1007381” and 
“CASENT 1007382” to “CASENT 1007383” and “CASENT 1007385” to “CASENT 1007386”, 
respectively; 1 male and 2 females (in CAS, IOZ) labeled: “CASENT 1010345” and “CASENT 
1010346” to “CASENT 1010347”/ “CHINA, Yunnan, Gongshan County, Cikai Township, 52.6 km 
W of Gongshan on Dulong Valley Road, 3360-3380 m,7 “N27.77032°/E98.44661°, 1-2 October 
2002, Stop #DHK-2002-034, D.H. Kavanaugh & P.A. Marck collectors”; 1 male (in CAS) labeled: 
“CASENT 1015626”/ “CHINA, Yunnan, Gongshan County, Cikai Township, 48 km W of Gong¬ 
shan on Dulong Valley Road, 3330 m,”/ “N27.78075°/E98.47000°, 13 November 2004, Stop # 
DHK-2004-086, D.H. Kavanaugh collector”; 1 male (in IOZ) labeled: “CASENT 1025832”/ 
“CHINA, Yunnan, Gongshan County, Bingzhongluo Township, SW slope of Kawakarpu Shan, 0.3 
km SW of Chukuai Lake at campsite,”/ “N27.976867E98.47799°, 3750 m, 19 August 2006, Stop 
#DHK-2006-082, Y. Liu collector”. All paratypes also bear the following label: “PARATYPE 
Trechus qiqiensis Deuve & Kavanaugh, sp. nov. designated 2016” [yellow label]. 

type locality. — China, Yunnan, Gongshan County, Qiqi Trail at No. 12 Camp, 
N27.71503°/E98.50244°, 2775 m. 

derivation oe species name .— The species epithet, qiqiensis, is derived from the name of 
the ancient trail, Qiqi Trail, passing through the type locality, and the Latin suffix, -ensis, denoting 
place. 

diagnosis. — Adults of this species (Fig. 20a) can be distinguished from those of all other 
species in the region by the following combination of character states: size relatively large (BL = 
4.0 to 4.2 mm), apterous; body color reddish-brown; eyes slightly convex; tempora glabrous, short 
and convex; pronotum transverse (ratio PW/PL = 1.44), basal angles rectangular and sharp; elytra 
with striae 1 to 4 deeply impressed on disc but attenuated apically, remaining striae successively 
less distinct, the outermost effaced or nearly so; recurrent stria terminated anteriorly by a slight 
convexity in line with stria 7; two discal setae present, inserted on interval 3 next to stria 3; preapi- 
cal seta present and inserted in a discal position on interval 3, next to either stria 2 or 3 in forward 
position near apical one-fourth of elytra; median lobe of male aedeagus large and robust, subapi- 
cally bent ventrally, apical lamella sinuate and reflexed dorsally (Fig. 20b), endophallus with a 
more heavily sclerotized voluminous lobe or scaly fold. 

Description. — Size relatively large, BL = 4.0 to 4.2 mm. Body color reddish-brown, shiny, 
appendages concolorous but paler, palpi pale yellow. Body surface smooth, only head capsule 
faintly alutaceous. 

Head. Moderate in size; eyes only slightly projected, slightly more convex, their diameter less 
than twice as long as tempora, the latter short, convex and glabrous. Frons slightly flattened; frontal 
furrows deep, curved, continuous but attenuated posterior to the eyes. Two pairs of supraorbital 
setae present, the anterior pair inserted in foveae. Clypeus with four setae. Labrum with six setae, 
anterior margin distinctly concave. Mandibles short; right mandible distinctly tridentate with mid¬ 
dle tooth closer to basal tooth (premolar) than to distal tooth, left mandible with a small slightly 
bifid process. Mentum and submentum not fused but suture between them only faintly impressed. 


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CHIN 


Yunnan 


MYANMAR 


ftrgjang 


Uanghe 


Long ling 


Figure 20. Trechus qiqiensis sp. nov.; a. Dorsal habitus (CASENT1007384). b. Median lobe of aedeagus of male 
(CASENT1007384), left lateral aspect, c. Map of locality records (red circles) for T. qiqiensis in the Gaoligong Shan region. 
Scale lines a, b = 0.5 mm, c = 100 km. 


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Mentum with medial tooth broad and apically truncate, about one-half the length of the lateral 
lobes. Submentum with six setae anteriorly. Gula wide. Genae with a single seta ventrally on each 
side. Antennae short, extended posteriorly to basal one-sixth of elytra, with about 2 or 3 anten- 
nomeres extended posteriorly beyond basal pronotal margin; antennomeres 2 and 3 about equal in 
length, antennomere 4 slightly shorter. 

Pronotum. Transverse (ratio PW/PL = 1.44), narrowed posteriorly, greatest width near anteri¬ 
or one-fourth; lateral margins markedly rounded, with a short sinuation just anterior to basal angles, 
the latter small, rectangular and sharp. Disc convex, smooth and glabrous, median longitudinal 
impression finely impressed, but continuous between anterior and posterior margins; basal foveae 
distinct, circular; median basal area faintly rugulose, delimited laterally by short, oblique furrows. 
Basal margin nearly rectilinear, slightly projected medially. Lateral borders of pronotum moder¬ 
ately slender, regular, narrowly and regularly reflexed, lateral grooves narrow but distinctly 
impressed. Single midlateral setae on each side inserted slightly anterior to middle; single basolat- 
eral seta on each side, inserted at basal angle. 

Elytra. Ovoid, humeri distinct but rounded. Disc convex and smooth. Striae finely punctate, 
intervals faintly convex; striae 1 to 4 deeply impressed but attenuated apically, stria 5 more faint¬ 
ly impressed, striae 6 to 8 effaced or nearly so. Parascutellar striole present. Recurrent stria termi¬ 
nated anteriorly by a slight convexity at the presumed location of stria 7 apically. Basal setiferous 
pore present at common origin of striae 1 and 2. Two discal setae present and inserted next to stria 
3, one at anterior one-fourth and one near middle of elytra. Preapical seta present and inserted in a 
discal position on interval 3, next to either stria 2 or 3 in forward position at apical one-fourth of 
elytra in most specimens; in a few specimens two preapical setae present, one inserted farther for¬ 
ward next to stria 3, as a third discal seta, the second inserted less far forward and next to stria 2. 
Umbilicate setal series with setae of humeral group equidistance from each other and setae of medi¬ 
an group inserted slightly posterior to middle. 

Legs. Medium proportions, slightly short, protibiae with longitudinal furrow. Male protar- 
someres 1 and 2 dilated and apicomedially toothed. 

Abdomen. Abdominal ventrites glabrous, except for a single paramedial seta on each side, and 
ventrite VII of males apically with one pair of paramedial setae, of females with two pairs. 

Male aedeagus. Median lobe (Fig. 20b) large and robust, with a moderate-sized sagittal 
aileron; shaft subapically bent ventrally; apical lamella sinuate and reflexed dorsally, bluntly point¬ 
ed apically; endophallus with a more heavily sclerotized voluminous lobe or scaly fold. 

habitat distribution. — Members of this species have been found at elevations ranging 
from 2775 to 3750 m in a variety of habitats. At the lowest elevation (2775 m), specimens were 
collected by sifting meager leaf litter and mosses on the forest floor and on rotting logs (Fig. 42a). 
This area had abundant conifers {Abies, Thuja, and Picea species) that formed a partly open 
canopy. Litter and mosses were moist and substrate underneath was composed of crumbling 
granitic sand. Specimens of Trechepaphiopsis monochaeta sp. nov. were also collected in the same 
litter samples at this site. At higher elevations (above 3300 m), specimens were found under stones 
in meadows, talus slopes, in low Rhododendron thickets, and on heath-covered tundra slopes (Fig. 
37b). In such areas, T. cjicjiensis specimens were found syntopic with those of Queinnectrechus 
( Gaoligongtrechus ) balli and Queinnectrechus ( s . str.) griswoldi, and at highest elevation (3750 m) 
also with members of Trechus gongshanensis sp. nov. 

Geographical distribution within the Gaoligong Shan.— Fig. 20c. We examined a 
total of 12 specimens (5 males and 7 females), all from Bingzhiongluo and Cikai Townships in 
Gongshan County on the crest and eastern slope of the northern part of the Gaoligong Shan (see 
Type material above for exact collection data). These localities are all in Core Area 2. 


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Overall geographical distribution .— This species currently is known only from Gong- 
shan County on the crest and eastern slope of the northern part of the Gaoligong Shan, in western 
Yunnan Province, China. 

Trechus ( Trechus) pseudoqiqiensis Deuve and Liang, sp.nov. 

(Figs. 21, 42b, 46-48) 

Type material. — Holotype, a female, in IOZ, labeled: “CASENT 1014205”/ “CHINA, Yun¬ 
nan, Fugong County, Lishadi Township, 11.5 km above Shibali on Yaping Road, 
N27.20676°/E98.71763°,”/ “3290 m, 8 May 2004 Stop #DHK-2004-040 D.H. Kavanaugh, C.E. 
Griswold, Liang H.-B., Li X.-Y., & Zhu B.-X. collectors”/ “HOLOTYPE Trechus pseudoqiqiensis 
Deuve & Liang, sp. nov. designated 2016” [red label]. 

Type locality. — China, Yunnan, Fugong County, Shiyueliang Township, 11.5 km above 
Shibali on Yaping Road, N27.20676°/E98.71763°, 3290 m. 

Derivation of species name. — The species epithet, pseudoqiqiensis, is a combination of the 
Greek prefix, y/cvSijg (translated into Latin as pseudo-) meaning false, and the species epithet, 
qiqiensis, in reference to the similarity of the unique holotype female of this species to members 
of T. qiqiensis . 

diagnosis. — Adults of this species (Fig. 21a) can be distinguished from those of all other 
species in the region by the following combination of character states: similar to T. qiqiensis in size 
(BL = 4.0 mm) and most features, except body color paler, more reddish; elytral striae more faint¬ 
ly punctate, almost impunctate toward elytral apex, more deeply impressed on center of disc and 
not attenuated near apex, recurrent stria less abruptly interrupted by a convexity and nearly con¬ 
tinuous anteriorly with stria 7; preapical seta present and inserted on interval 3 next to stria 2 near 
apical one-fourth of elytra. 

Description .— Size relatively large (BL = 4.0 mm). Body color bright brownish red, shiny, 
appendages yellowish-tan, palpi pale yellow. 

Head. Moderate in size; eyes slightly convex, their diameter about 1.5 times as long as tem- 
pora, the latter short, convex and glabrous. Frons not flattened; frontal furrows deep, rounded, not 
or only slightly attenuated posterior to the eyes. Two pahs of supraorbital setae present, the anteri¬ 
or pair inserted in foveae. Clypeus with four setae. Labrum with six setae, anterior margin dis¬ 
tinctly concave. Mentum and submentum not fused. Mentum distinctly concave at middle, with 
medial tooth broad and apex slightly bifid, about one-half the length of the lateral lobes. Submen¬ 
tum with six setae anteriorly. Gula wide. Genae with a single seta ventrally on each side. Antennae 
short, with about 2 antennomeres extended posteriorly beyond basal pronotal margin; anten- 
nomeres 2 and 4 about equal in length, antennomere 3 slightly longer. 

Pronotum. Transverse (ratio PW/PL = 1.45), slightly narrowed posteriorly, greatest width near 
anterior one-third; lateral margins broadly rounded, not sinuate subbasally, straightened only 
immediately anterior to basal angles, the latter small and rectangular. Disc convex, smooth and 
glabrous, median longitudinal impression finely impressed, interrupted slightly posterior to anteri¬ 
or margin but extended to basal margin; basal foveae distinct, but shallower and slightly convex at 
center; median basal area mainly smooth, but with several small foveae anteriorly near the poste¬ 
rior transverse impression. Basal margin rounded and slightly projected medially. Lateral borders 
of pronotum slender, narrowly reflexed, lateral grooves distinctly impressed. Single midlateral 
setae on each side inserted near anterior one-third; single basolateral seta on each side, inserted at 
basal angle. 

Elytra. Ovoid and only slightly elongate, humeri not very broad but rounded. Disc convex and 
smooth. Striae very finely punctate, nearly smooth in apical half, intervals moderately convex; 


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Figure 21. Trechus pseudoqiqiensis sp. nov.; a. Dorsal habitus (CASENT1014204). b. Map of locality record (red cir¬ 
cle) for T. pseudoqiqiensis in the Gaoligong Shan region. Scale lines a = 0.5mm, b = 100 km. 


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striae 1 to 4 deeply impressed and not attenuated apically, stria 5 to 8 successively more faintly 
impressed by still evident. Parascutellar striole present, short. Recurrent stria less abruptly inter¬ 
rupted by a convexity and nearly continuous anteriorly with stria 7. Basal setiferous pore present 
at common origin of striae 1 and 2. Two discal setae present and inserted next to stria 3, one at ante¬ 
rior one-fourth and one near middle of elytra. Preapical seta present and inserted in a discal posi¬ 
tion on interval 3 next to stria 2 in forward position at apical one-fourth of elytra. Umbilicate setal 
series with setae of humeral group equidistance from each other and setae of median group insert¬ 
ed slightly posterior to middle. 

Legs. Slightly short, protibiae with only a faint longitudinal furrow. 

Abdomen. Abdominal ventrites glabrous, except for a single paramedial seta on each side, and 
ventrite VII of female apically with two pairs of paramedial setae. 

h a b itat distribution.— The unique holotype female of this species was found under a stone 
in an open area cleared of forest but surrounded by bamboo thickets and scattered Abies, Rhodo¬ 
dendron and Primus species at an elevation of 3290 m (Fig. 42b). One specimen of Trechus shibali- 
cus sp. nov. also was found at the same site. 

Geographical distribution within the Gaoligong Shan.— Fig. 21b. This Species is 
known from a single locality in Fugong County, on the eastern slope of the northcentral part of the 
Gaoligong Shan (see Type material above for exact collection data). This locality is in Core 
Area 3. 

Overall geographical distribution .— This species currently is known only from the type 
locality on the eastern slope of the northcentral part of the Gaoligong Shan, in western Yunnan 
Province, China. 

Trechus ( Trechus) luzhangensis Deuve and Liang, sp. nov. 

(Figs. 22, 43a, 46-48) 

Type material.— Holotype, a male, in IOZ, labeled: “CASENT 1017595”/ “CHINA, Yun¬ 
nan, Lushui County, Luzhang Township, Piana Road at Fengxue Yakou, N25.97228°/E98.68336°, 
3150 m, 11 May 2005,”/ “Stop# 2005-007, D.H. Kavanaugh, H.B. Liang, C.E. Griswold, D.Z. 
Dong & K.J. Guo collectors”/ “HOLOTYPE Trechus luzhangensis Deuve & Liang, sp. nov. des¬ 
ignated 2016” [red label]. Paratypes (only 1): a female (in CAS) labeled same as holotype, except 
first label “CASENT 1017596” and last label “PARATYPE Trechus luzhangensis Deuve & Liang, 
sp. nov. designated 2016” [yellow label]. 

type locality.— China, Yunnan, Lushui County, Luzhang Township, Pianma Road at 
Fengxue Yakou [Pass], N25.97228°/E98.68336°, 3150 m. 

e t y m o l o g y . — The species epithet, luzhangensis, is derived from the name of the township, 
Luzhang, in which the type locality is found, and the Latin suffix, -ensis, denoting place. 

diagnosis.— Adults of this species (Fig. 22a) can be distinguished from those of all other 
species in the region by the following combination of character states: size medium (BL = 3.7 to 
3.8 mm), apterous; body color dark piceous, shiny, elytra with interval 1, lateral margins and api- 
comedial area reddish; tempora glabrous, convex, half as long as diameter of eyes; pronotum trans¬ 
verse (ratio PW/PL = 1.46), basal angles rectangular and sharp; elytra ovoid, convex, striae finely 
impressed, striae 1 to 3 deeply impressed on disc, not attenuated apically, remaining striae succes¬ 
sively less distinct, the outermost very faint but still evident; recurrent stria terminated anteriorly 
at distinct convexity of interval 7; two discal setae present, inserted on interval 3 next to stria 3; 
preapical seta present and inserted in a discal position on interval 3, near stria 2 in forward posi¬ 
tion near apical one-fourth of elytra; median lobe of male aedcagus (Fig. 22b) with moderate-sized 


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Figure 22. Trechus luzhangensis sp. nov.; a. Dorsal habitus (CASENT1017595). b. Median lobe of aedeagus of male 
(CASENT1017595), left lateral aspect, c. Map of locality records (red circles) for T. luzhangensis in the Gaoligong Shan 
region. Scale lines a, b = 0.5 mm, c = 100 km. 


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sagittal aileron, apical lamella short and thick, with apex blunt, endophallus with a sclerotized line 
and a scaly area. 

description.— Size medium, BL = 3.7 to 3.8mm. Body color dark piceous, shiny, elytra 
with interval 1, lateral margins and apicomedial area more or less reddish, appendages paler, yel¬ 
lowish-orange, palpi pale yellow. Body surface smooth, head capsule faintly alutaceous. 

Head. Moderate in size; eyes only slightly projected but convex, their diameter twice as long 
as tempora, the latter short, moderately convex and glabrous. Frons not flattened; frontal furrows 
deep, rounded, prolonged and not attenuated posterior to the eyes. Two pairs of supraorbital setae 
present, the anterior pair inserted in foveae. Clypeus with four setae. Labrum with six setae, ante¬ 
rior margin distinctly concave. Mandibles short; right mandible distinctly tridentate with middle 
tooth closer to basal tooth (premolar) than to distal tooth. Men turn and submentum not fused but 
nearly so, suture between them only faintly impressed. Mentum with medial tooth apically trun¬ 
cate, less than half the length of the lateral lobes. Submentum with six setae anteriorly, swollen 
anteriorly. Gula wide. Genae with a single seta ventrally on each side. Antennae rather short, with 
about 2 antennomeres extended posteriorly beyond basal pronotal margin; antennomeres 2 and 3 
about equal in length, antennomere 4 slightly shorter. 

Pronotum. Transverse (ratio PW/PL = 1.46), moderately narrowed posteriorly, greatest width 
anterior to middle; lateral margins widely rounded, straightened only just anterior to basal angles, 
the latter very small, but projected, acute and sharp. Disc convex, smooth and glabrous, median 
longitudinal impression slender but sharply impressed, extended anteriorly to near anterior margin; 
basal foveae distinct, subcircular; median basal area faintly, longitudinally rugulose, delimited lat¬ 
erally by obliquely curved furrows. Lateral borders of pronotum moderately slender, narrowly 
reflexed, lateral grooves deeply impressed. Single midlateral setae on each side inserted near ante¬ 
rior one-third; single basolateral seta on each side, inserted at basal angle. 

Elytra. Ovoid, only slightly more narrowed basally than apically, humeri distinct but rounded. 
Disc convex and smooth. Striae finely impressed, not or only faintly punctate; striae 1 to 3 deeply 
impressed on disc, not attenuated apically, remaining striae successively less distinct, the outermost 
very faint but still evident. Parascutellar striole present. Recurrent stria terminated anteriorly at dis¬ 
tinct convexity of interval 7. Basal setiferous pore present at common origin of striae 1 and 2. Two 
discal setae present and inserted next to stria 3, one at anterior one-fourth and one near middle of 
elytra. Preapical seta present and inserted in a prediscal position on interval 3, closer to stria 2 than 
3 in forward position at apical one-fourth of elytra. Umbilicate setae of humeral group equidistance 
from each other and setae of median group inserted slightly posterior to middle. 

Legs. Relatively short, protibiae with longitudinal furrow. Male protarsomeres 1 and 2 dilated 
and apicomedially toothed. 

Abdomen. Abdominal ventrites glabrous, except for a single paramedial seta on each side, and 
ventrite VII of males apically with one pair of paramedial setae, of females with two pairs. 

Male aedeagus. Median lobe (Fig. 22b) only moderately enlarged basally but with a medium¬ 
sized sagittal aileron present; shaft narrow subbasally, progressively thicker toward subapical por¬ 
tion; apical lamella short and thick, with apex blunt, endophallus with a sclerotized line and a scaly 
area, similar to that seen in males of T. shiyueliang sp. nov. (Fig. 18b). 

Habitat distribution. — Members of this species have been found under stones on slopes 
above (Fig. 43a) and below the road and both side of the pass at an elevation of 3150 m. The habi¬ 
tat in this area includes broken scrub vegetation of two to four meter high bamboo and Rhododen¬ 
dron thickets on a thin layer of organic substrate, as well as open areas with stones on granitic sand 
substrate, and small seeps. Both specimens were found at the edges of thickets on organic substrate. 
No other trechines were found at this site. 


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Geographical distribution within the Gaoligong Shan . Fig. 22c. We examined a total 
of 2 specimens (1 male and 1 female), both from the crest of the southcentral part of the Gaoligong 
Shan in Lushui County (see Type material above for exact collection data). This locality straddles 
Core Areas 4 and 5. 

Overall geographical distribution. This species currently is known only from the type 
locality on the crest of the southcentral part of the Gaoligong Shan, in western Yunnan Province, 
China. 

Trechus (Treehus) gongshanensis Deuve and Liang, sp. nov. 

(Figs. 16c, 23, 38a, 39a, 39b, 40a, 46-48) 

Type material.— Holotype, a male, in IOZ, labeled: “CASENT 10019297 “CHINA, Yun¬ 
nan Province, Gaoligong Shan, Nujiang Prefecture, Nujiang State Nature Reserve, Dulong/Gong- 
shan Yakou area, 21 airkm W of Gongshan,”/ “N27.696557 E98.45389 0 , 3300-3680 m, 16-17 July 
2000, Stop#00-24C, D.H. Kavanaugh, C.E. Griswold, Liang H.-B., D. Ubick, & Dong D.-Z. col¬ 
lectors”/ “HOLOTYPE Trechus gongshanensis Deuve & Liang, sp. nov. designated 2016” [red 
label]. Paratypes (a total of 51): 1 male and 4 females (in CAS, IOZ) labeled: same as holotype, 
except first label “CASENT 1001928” and “CASENT 1001930” to “CASENT 1001932” and 
“CASENT 1008148”, respectively; 8 males and 4 females (in CAS, IOZ) labeled: “CASENT 
1010348” to “CASENT 1010355” and “CASENT 1010356” to “CASENT 1010359”, respective¬ 
ly/ “CHINA, Yunnan, Gongshan County, Cikai Township, 52.6 km W of Gongshan on Dulong Val¬ 
ley Road, 3360-3380 m,7 “N27.770327 E98.44661 0 , 3360-3380 m, 1-2 October 2002, D.H. 
Kavanaugh & P.E. Marek collectors”; 1 female (in IOZ) labeled: “CASENT 10248697 “CHINA, 
Yunnan, Gongshan County, Bingzhongluo Township, SW slope of Kawakarpu Shan, on slope NE 
of Chukuai Lake, 3950 m,7 “N27.982067 E98.48027 0 , 20 August 2006 Stop #DHK-2006-086 Y. 
Liu, P. Hu, D.Z. Dong & J. Wang collectors”; 1 male (in CAS) labeled: “CASENT 1025105”/ 
“CHINA, Yunnan, Gongshan County, Bingzhongluo Township, SW slope of Kawakarpu Shan, 0.4 
km NW of Chukuai Lake,”/ “N27.982317 E98.47069 0 , 3808 m, 21 August 2006 Stop #DHK- 
2006-094 D.Z. Dong collector”/ 2 males and 2 females (in CAS, IOZ) labeled: “CASENT 
1025164” to “CASENT 1025165” and “CASENT 1025162” to “CASENT 1025163”, respective¬ 
ly/ “CHINA, Yunnan, Gongshan County, Bingzhongluo Township, S W slope of Kawakarpu Shan, 
0.75 km NW of Chukuai Lake,”/ “N27.986317 E98.47069 0 , 3820 m, 21 August 2006 Stop #DHK- 
2006-095 Y. Liu, P. Hu, & J. Wang collectors”; 1 male and 2 females (in CAS, IOZ) labeled: 
“CASENT 1025833” and “CASENT 1025834” to “CASENT 1025835”, respectively/ “CHINA, 
Yunnan, Gongshan County, Bingzhongluo Township, SW slope of Kawakarpu Shan, 0.3 km SW 
of Chukuai Lake at campsite,”/ “N27.986317 E98.47069 0 , 3820 m, 21 August 2006 Stop #DHK- 
2006-082 Y. Liu collector”; 1 female (in IOZ) labeled: “CASENT 1025920”/ “CHINA, Yunnan, 
Gongshan County, Bingzhongluo Township, SW slope of Kawakarpu Shan at Chukuai Lake, 3720 
m,7 “N27.981217 E98.47580 0 , 19 August 2006 Stop #DHK-2006-095B D.H. Kavanaugh, J.A. 
Miller, D.Z. Dong, Y. Liu, P. Hu, & J. Wang collectors”; 1 female (in CAS) labeled: “CASENT 
1025939”/ “CHINA, Yunnan, Gongshan County, Bingzhongluo Township, SW slope of 
Kawakarpu Shan, 0.3 km SW of Chukuai Lake at campsite, N27.976867 E98.47799 0 ,”/ “3750 m, 
19 August 2006 Stop #DHK-2006-095C D.H. Kavanaugh, J.A. Miller, D.Z. Dong, Y. Liu, P. Hu, 
& J. Wang collectors”; 4 males and 2 females (in CAS, IOZ) labeled: “CASENT 1026196” to 
“CASENT 1026199” and “CASENT 1026200” to “CASENT 1026201”, respectively/ / “CHINA, 
Yunnan, Gongshan County, Bingzhongluo Township, SW slope of Kawakarpu Shan, 0.3 km NNE 
of Chukuai Lake, N27.983937 E98.47491 0 ,”/ “3745 m, 19 August 2006 Stop #DHK-2006-081 


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Figure 23. Trechus gongshanensis sp. nov.; a. Dorsal habitus (CASENT1001929). b. Median lobe of aedeagus of male 
(CASENT1001929, left lateral aspect, c. Map of locality record s(red circles) for T. gongshanensis in the Gaoligong Shan 
region. Scale lines a, b = 0.5 mm, c = 100 km. 


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D.H. Kavanaugh, J.A. Miller, & D.Z. Dong collectors”; 3 males (in CAS, IOZ) labeled: “CASENT 
1026320” to “CASENT 1026322”, respectively/ “CHINA, Yunnan, Gongshan County, Cikai 
Township, 0.1 km SE of Heipu Yakou in valley below tunnel, N27.76978°/ E98.44681 0 ,”/ “3720 
m, 13 August 2006, Stop #DHK-2006-073 D.H. Kavanaugh & J.A. Miller collectors”; 1 male (in 
CAS) labeled: “CASENT 1026366”/ “CHINA, Yunnan, Gongshan County, Bingzhongluo Town¬ 
ship, SW slope of Kawakarpu Shan at Chukuai Lake, 3720 m,”/ “N27.98121 0 / E98.47580 0 , 19 
August 2006 Stop #DHK-2006-080 D.H. Kavanaugh & J.A. Miller collectors”; 8 males and 5 
females (in CAS, IOZ, MNHN) labeled: “CASENT 10263824” to “CASENT 10263831” and 
“CASENT 10263819” to “CASENT 10263823”, respectively/ “CHINA, Yunnan, Gongshan Coun¬ 
ty, Bingzhongluo Township, SW slope of Kawakarpu Shan at Chukuai Lake, 3720 m,”/ 
“N27.98121 0 / E98.47580°, 18 August 2006 Stop #DHK-2006-079 D.H. Kavanaugh, J.A. Miller, 
D.Z. Dong, & Y. Liu collectors”. All paratypes also bear the following label: “PARATYPE Trechus 
gongshanensis Deuve & Liang, sp. nov. designated 2016” [yellow label]. 

type locality.— China, Yunnan, Gongshan County, Dongshaofang area, 
N27.69655°/E95.45389°, 3300-3600 m. 

derivation of species name.— The species epithet, gongshanensis, is derived from the 
name of the county, Gongshan, in which all specimens of the type series were collected, and the 
Latin suffix, -ensis, denoting place. 

diagnosis. — Adults of this species (Fig. 23a) can be distinguished from those of all other 
species in the region by the following combination of character states: size small (BL = 3.3 to 3.5 
mm), apterous; body color dark piceous, elytra slightly reddish near sutural and lateral margins; 
tempora glabrous and only slightly convex; pronotum transverse (ratio PW/PL=1.45), basal 
angles acute and sharp; elytra convex, discal striae 1 to 3 or 4 finely impressed, striae 6 to 8 effaced, 
recurrent stria joined anteriorly with apex of stria 7, two discal setae present on interval 3 next to 
stria 3, preapical seta present and inserted in a discal, forward position near apical one-fourth of 
elytra on interval 3, near stria 2 in most individuals, nearer to stria 3 in a few; aedeagus of male 
with base robust, apical lamella elongate with apex blunt, endophallus with a slightly sclerotized 
scaly area. 

description. — Size small, BL = 3.3 to 3.5 mm. Body color dark piceous and shiny, elytra 
slightly reddish near sutural and lateral margins, appendages paler reddish yellow, palpi paler yel¬ 
low. Body surface smooth, head capsule faintly alutaceous. 

Head. Moderate in size, eyes moderately convex, their diameter only slightly longer than 
length of tempora, the latter glabrous and only slightly convex. Frontal furrows deep, rounded, 
slightly attenuated posteriorly but prolonged posterior to the eyes. Clypeus with four setae. Labrum 
with six setae, apical margin distinctly concave. Right mandible as in Fig. 16c. Mentum and sub- 
mentum not fused but nearly so, suture between them very fine and only faintly impressed. Men¬ 
tum with medial area concave, median tooth short, apically bifid, less than half as long as lateral 
lobes. Submentum with six setae anteriorly, swollen anteriorly. Gula wide. Genae with a single seta 
ventrally on each side. Antennae short, extended posteriorly to basal one-fifth of elytra, with about 
2 antennomeres extended posteriorly beyond basal pronotal margin; antennomere 3 slightly longer 
than antennomere 2 or 4. 

Pronotum. Transverse (ratio PW/PL = 1.45), widest at anterior one-third, narrowed posterior¬ 
ly, lateral margins rounded, then abruptly sinuate just anterior to basal angles, the latter acute, sharp 
and slightly projected laterally. Disc convex, smooth, median longitudinal impression finely 
impressed but prolonged to the basal margin, slightly widened and deepened in the median basal 
area, the latter also with several faint longitudinal rugulae and delimited anteriorly by an arcuate 
posterior transverse impression that is partially effaced medially. Basal foveae distinct but smooth. 


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Basal margin nearly rectilinear. Lateral margination slender throughout and narrowly reflexed, lat¬ 
eral grooves distinctly impressed. Single midlateral setae on each side inserted near anterior one- 
third; single basolateral seta on each side, inserted slightly anterior to apex of basal angle. 

Elytra. Ovoid, humeri distinct but rounded. Disc convex, with striae 1 to 3 or 4 finely 
impressed, the more lateral striae successively more faintly impressed, striae 7 and 8 effaced, stri¬ 
ae slightly and irregularly punctate. Parascutellar striole rudimentary. Recurrent stria distinct, mod¬ 
erately deep, terminated at the posterior end of stria 7. Basal setiferous pore present at common ori¬ 
gin of striae 1 and 2. Two discal setae present and inserted next to stria 3, one at anterior one-fourth 
and one near middle of elytra. Preapical seta present and inserted in a prediscal position on inter¬ 
val 3, closer to stria 2 than 3 in most specimens (closer to stria 3 in a few) in forward position at 
apical one-fourth of elytra. 

Legs. Relatively short, protibiae with longitudinal furrow. Male protarsomeres 1 and 2 dilated 
and apicomedially toothed. 

Abdomen. Abdominal ventrites glabrous, except for a single paramedial seta on each side, and 
ventrite VII of males apically with one pair of paramedial setae, of females with two pairs. 

Male aedeagus. (Fig. 23b). Median lobe with basal bulb robust and with a thick sagittal 
aileron; apical lamella elongate, slightly tapered, with apex blunt. Endophallus nearly unarmed, 
with only a faintly sclerotized scaly area. 

habitat distribution .— Members of this species have been found in a variety of microhab¬ 
itats in the alpine zone at elevations ranging from 3360 to 3950 m. Specimens were collected from 
under stones in moist meadows, on tundra slopes and ridges with sparse to thick herbaceous veg¬ 
etation (Fig. 38a, 39b, 40a) and at the edges of small streams and seeps from talus slopes (Fig. 39a). 
They were also collected in pitfall traps placed at the edges of Rhododendron thickets up to two 
meters tall. Members of this species have been found syntopic with specimens of Queinnectrechus 
( Gaoligongtrechus ) halli, Queinnectrechus (s. str) griswoldi, Queinnectrechus (s. str.) gongshard¬ 
ens, and Trechus qiqiensis at one or more sites. 

Geographical distribution within the Gaoligong Shan.— Fig. 23c. We examined a 
total of 52 specimens (22 males and 30 females), all from Bingzhiongluo and Cikai Townships in 
Gongshan County on the crest and eastern slope of the northern part of the Gaoligong Shan (see 
Type material above for exact collection data). These localities are all in Core Area 2. 

o veraee geographical distribution .— This species currently is known only from Gong¬ 
shan County in the northern part of the Gaoligong Shan, in western Yunnan Province, China. 

Trechus ( Trechus ) shibalicus Deuve and Kavanaugh, sp. nov. 

(Figs. 24, 16d, 40b, 42b, 46-48) 

Type material.— Holotype, a male, in IOZ, labeled: “CASENT 1017532”/ “CHINA, Yun¬ 
nan, Fugong County, Lishadi Township, 8.5 km above Shibali on Shibali Road, north bank of North 
Fork of Yamu He, N27.18416 0 / E98.72026 0 ,”/ “3100 m, 8 August 2005 Stop #DHK-2005-067A 
D.H. Kavanaugh, H.B. Liang, D.Z. Dong, & J.F. Zhang collectors”/ “HOLOTYPE Trechus shibali¬ 
cus Deuve & Kavanaugh, sp. nov. designated 2016” [red label]. Paratypes (a total of 43): 1 male 
and 3 females (in IOZ, CAS) labeled: same as holotype, except first label “CASENT 1017531” and 
“CASENT 1017533” to “CASENT 1017535”, respectively; 1 male (in CAS) labeled: “CASENT 
1014204”/ “CHINA, Yunnan, Fugong County, Lishadi Township, 11.5 km above Shibali on Yap- 
ing Road, N27.20676 0 / E98.71763 0 ,”/ “3290 m, 8 May 2004 Stop #DHK-2004-040 D.H. 
Kavanaugh, C.E. Griswold, Liang H.-B., Li X.-Y., & Zhu B.-X. collectors”; 1 male (in IOZ) 
labeled: “CASENT 1018365”/ “CHINA, Yunnan, Fugong County, Lishadi Township, headwaters 
of North Fork Yamu He just E of Shibali Yakou, 3450 m,”/ “N27.21034°/ E98.70141°, 7 August 


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2005, Stop# LHB-05-52, H.B. Liang & J.F. Zhang collectors”; 1 male (in IOZ) labeled: “CASENT 
10186057 “CHINA, Yunnan, Fugong County, Lishadi Township, headwaters of North Fork Yamu 
He just E of Shibali Yakou, 3450 m,”/ “N27.21034 0 / E98.70141 0 , 12 August 2005, Stop# LHB-05- 
54, H.B. Liang & J.F. Zhang collectors”; 1 male (in CAS) labeled: “CASENT 1018844”/ “CHINA, 
Yunnan, Fugong County, Lishadi Township, Shibali area, 2535 m, N27.165367 E98.78003°, 4-17 
August 2005,”/ “Stop# DHK-2005-059 D.H. Kavanaugh, H.B. Liang, P. Paquin, & D.Z. Dong col¬ 
lectors”; 1 female (in CAS) labeled: “CASENT 1020015”/ “CHINA, Yunnan, Fugong County, 
Lumadeng Township, second cirque S of Shibali Yakou, 3675 m,”/ “N27.202447 E98.69526°, 17 
August 2005, Stop# DHK-2005-093, D.H, Kavanaugh collector”; 2 males and 2 females (in CAS, 
IOZ) labeled: “CASENT 1020903” to “CASENT 1020904” and “CASENT 1020905” to 
“CASENT 1020906”, respectively/ “CHINA, Yunnan, Fugong County, Lishadi Township, Shibali 
Yakou, 3612 m, N27.21231 0 / E98.69575 0 , 7 August 2005,”/ “Stop# DHK-2005-066, D.H. 
Kavanaugh, H.B. Liang, P. Paquin, & D.Z. Dong collectors”; 1 female (in CAS) labeled: 
“CASENT 1021290”/ “CHINA, Yunnan, Fugong County, Lishadi Township, 0.5 km NE of Shibali 
Yakou, N27.21447°/ E98.70064 0 ,7 “3460 m, 12 August 2005, Stop# DHK-2005-077, D.H. 
Kavanaugh, P. Paquin, & D.Z. Dong collectors”; 6 males and 5 females (in CAS, IOZ) labeled: 
“CASENT 1022286” to “CASENT 1022291” and “CASENT 1022292” to “CASENT 1022296”, 
respectively/ “CHINA, Yunnan, Fugong County, Lumadeng Township, 8.5 km above Shibali on 
Shibali Road, North Fork of Yamu He,”/ “N27.18416°/ E98.72026 0 ,”/ “3100 m, 5 May 2004, Stop 
#LHB-04-023 Lian H.-B., Li X.-Y., & Zhu B.-Q. collectors”; 1 female (in CAS) labeled: 
“CASENT 1023606”/ “CHINA, Yunnan, Fugong County, Lishadi Township, 11.5 km above 
Shibali on Shibali Road, N27.206767 E98.771763 0 ,”/ “3290 m, 6 May 2004 Stop #DHK-2004- 
036 D.H. Kavanaugh, C.E. Griswold, Liang H.-B., & Zhu B.-X. collectors”; 9 males and 4 females 
(in CAS, IOZ, MNHN) labeled: “CASENT 1023743” to “CASENT 1023745” and “CASENT 
1023747” to “CASENT 1023752” and “CASENT 1023753” to “CASENT 1023756”, respective¬ 
ly/ “CHINA, Yunnan, Fugong County, Lishadi Township, 8.5 km above Shibali on Shibali Road, 
North Fork of Yamu He, N27.184167 E98.72026 0 ,”/ “3100 m, 7 May 2004 Stop #DHK-2004- 
038A D.H. Kavanaugh, C.E. Griswold, Liang H.-B., & Zhu B.-X. collectors”; 1 male and 3 females 
(in CAS, IOZ) labeled: “CASENT 1023775” and “CASENT 1023776” to “CASENT 1023778”, 
respectively/ “CHINA, Yunnan, Fugong County, Lumadeng Township, 8.5 km above Shibali on 
Shibali Road, North Fork of Yamu He,”/ “ N27.183267 E98.72002 0 , 3100 m, 7 May 2004 Stop 
#DHK-2004-038B D.H. Kavanaugh collector”. All paratypes also bear the following label: 
“PARATYPE Trechus shibalicus Deuve & Kavanaugh, sp. nov. designated 2016” [yellow label]. 

type locality. — China, Yunnan, Fugong County, Shiyucliang Township, 8,5 km above 
Shibali on Shibali Road, North Fork of Yamu He, N27.184167E98.72026 0 , 3100 m. 

derivation of species name.— The species epithet, shibalicus , is derived from the name of 
settlement, Shibali, at and near which specimens of the type series were collected, and the Latin 
adjectival suffix, -icus, meaning belonging to or pertaining to. 

diagnosis.— Adults of this species (Fig. 24a) can be distinguished from those of all other 
species in the region by the following combination of character states: size small (BL = 3.3 to 3.5 
mm), apterous; body color reddish brown; tempora glabrous, short and distinctly convex; prono- 
tum transverse (ratio PW/PL = 1.44), basal angles rectangular or slightly acute and sharp; elytra 
with striae finely punctate, lateral striae more or less effaced, recurrent stria abruptly interrupted at 
slight convexity at posterior end of stria 7, two typical discal setae present, preapical seta present 
and inserted in a discal, forward position near apical one-fourth of elytra on interval 3 next to stria 
2; male aedeagus with sagittal aileron present but reduced, apical lamella slender with blunt apex, 
endophallus with a distinct scaly area. 


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CHIN 


Yunnan 


Tefiqchonfl 


Yinfl^rg 


Ltfi?i$#W 


Longing 


Figure 24. Trechus shibalicus sp. nov.; a. Dorsal habitus (CASENT1017532). b. Median lobe of aedeagus of male 
(CASENT1017532), left lateral aspect, c. Map of locality records (red circles) for T. shibalicus in the Gaoligong Shan 
region. Scale lines a, b = 0.5 mm, c = 100 km. 


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Description. — Size small, BL = 3.3 to 3.5 mm. Body color dark brown and shiny, elytra 
slightly reddish near sutural and lateral margins, appendages paler reddish yellow, palpi paler yel¬ 
low. Body surface smooth. 

Head. Short and thick, eyes slightly convex, their diameter slightly greater than length of tem- 
pora, the latter glabrous and distinctly convex. Frons not flattened; frontal furrows deep, broadly 
rounded, not attenuated posteriorly. Two pairs of supraorbital setae present, the anterior pair insert¬ 
ed in foveae. Clypeus with four setae. Labrum with six setae, anterior margin distinctly concave. 
Right mandible as in Fig. 16d. Mentum and submentum not fused. Mentum with median tooth bifid 
apically, half as long as lateral lobes. Submentum with six setae anteriorly. Gula wide. Genae with 
a single seta ventrally on each side. Antennae rather short, with about 1.5 to 2 antennomeres 
extended posteriorly beyond basal pronotal margin; antennomeres 2 and 3 about equal hi length, 
antcnnomere 4 slightly shorter. 

Pronotum transverse (ratio PW/PL = 1.44), greatest width slightly anterior to middle, slightly 
narrowed posteriorly, lateral margins rounded, abruptly sinuate just anterior to basal angles, the lat¬ 
ter rectangular or slightly acute and sharp. Disc convex; median longitudinal impression distinctly 
impressed, slightly widened and deepened in the median basal area, the latter also with several lon¬ 
gitudinal rugulae and delimited laterally by short, oblique furrows. Basal foveae distinct, small and 
rounded. Lateral margination narrow and slightly widened basally, lateral border slightly reflexed, 
lateral grooves distinctly impressed. Single midlateral setae on each side inserted near anterior one- 
third; single basolateral seta on each side, inserted at basal angle. 

Elytra. Ovoid, humeri distinct but rounded. Disc convex; striae finely punctate, striae 1 to 4 
distinctly impressed, the more lateral striae more or less effaced, stria 6 barely perceptible, stria 7 
and 8 indistinct. Scutellar striole short but deeply impressed. Recurrent stria distinct, abruptly inter¬ 
rupted at slight convexity at posterior end of stria 7. Basal setiferous pore present at common ori¬ 
gin of striae 1 and 2. Two discal setae present and inserted next to stria 3, one at anterior one-fourth 
and one near middle of elytra. Preapical seta present and inserted in a prediscal position on inter¬ 
val 3 next to stria 2 in forward position more than twice as far from apex as from sutural margin). 

Legs. Short, protibiae with longitudinal furrow. Male protarsomeres 1 and 2 dilated and api- 
comedially toothed. 

Abdomen. Abdominal ventrites glabrous, except for a single paramedial seta on each side, and 
ventrite VII of males apically with one pair of paramedial setae, of females with two pairs. 

Male aedeagus. Median lobe (Fig. 24b) with base moderate in size, sagittal aileron present but 
reduced, shaft moderately thick subbasally, apical lamella narrowed, moderately elongate, with 
blunt apex; endophallus only faintly sclerotized but with a distinct scaly area. 

habitat distribution.— Members of this species have been found in a variety of microhab¬ 
itats over a broad elevational range, from 2535 to 3675 m. They have been collected in daytime 
from under stones in thickets of bamboo (various elevations) (Fig. 42b), in subalpine forests of 
scattered Abies and Rhodondendron mixed with bamboo thickets (at 3290 m), in rocky open areas 
cleared by snow avalanches but shaded by 3 meter high herbaceous cover (3100 m) (Fig. 40b), in 
meadows adjacent to bamboo and Rhodondendron thickets (3400 m) and along roadcuts through 
such thickets, and on talus and vegtetated slopes in a glacial cirque (3675 m). At the lowest record¬ 
ed elevation (2535 m), one specimen was collected under a stone along a roadcut on moist, shad¬ 
ed ground. At both highest and lowest elevations, they were the only trechine encountered; but they 
were found syntopic with specimens of Trechus shiyueliang, Trechus pseudoqiqiensis, Trechepa- 
phiopsis unisetulosa sp. nov., and Trechepaphiopsis unipilosa sp. nov. at one or more mid-eleva¬ 
tion (3100 to 3290 m) sites. 

Geographical distribution within the Gaoligong Shan.— Fig. 24c. We examined a 


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407 


total of 44 specimens (23 males and 21 females), all from Fugong County on the eastern slope and 
crest of the northcentral part of the Gaoligong Shan (see Type material above for exact collection 
data). These localities are all in Core Area 3. 

Overall geographical distribution .— This species currently is known only from Fugong 
County in the northcentral part of the Gaoligong Shan, in western Yunnan Province, China. 

Genus Tvechepaphiopsis D e u v e and Kavanaugh, gen. nov. 

type species. — Trechepaphiopsis uniporosa , sp. nov. 

derivation of genus group name .— The genus group name (feminine) is a combination of 
two other trechine generic names, Trechus and Epaphius, plus the Greek suffix, o\|/iq (translated 
into Latin as - opsis) meaning having the aspect of, here referring to a similarity with Epaphius 
members. 

diagnosis. — Adults of this genus (Figs. 25-31) can be recognized by the following combi¬ 
nation of character states: size small to moderate (BL = 2.8 to 3.7 mm), apterous; body color light 
to dark brown, reddish-brown, or piccous-brown, most members with dorsum slightly iridescent, 
dorsal surface glabrous except for isolated fixed setae typical for trechines, eyes also glabrous; 
head short, with eyes small; tempora convex, swollen in some members, sparsely pubescent; right 
mandible (Fig. 16e) with premolar tooth not fused with retinaculum and anterior point of the reti¬ 
naculum free and displaced distally to form a separate tooth; mentum and submentum not fused; 
mentum with median tooth apical truncate or bifid; submentum with six setae; pronotum trans¬ 
verse, disc glabrous, basal foveac only slightly distinct, median basal area short and very trans¬ 
verse, delimited laterally by short, obliquely curved furrows, basal margin slightly projected pos¬ 
teriorly in most members, basal angles small, obtuse, rounded; elytra distinctly convex, with stri¬ 
ae finely impressed, more or less punctate, lateral striae attenuated or effaced, recurrent stria ter¬ 
minated anteriorly with a bend or hook on interval 5 or 6, with a single discal seta on interval 3 
next to stria three or without discal setae; preapical seta present, inserted next to stria 2; median 
lobe of male aedeagus of varied form, but endophallus membraneous, with spiny or scaly areas in 
some members, more or less sclerotized but without distinct sclerites. 

Comments. — In subtropical China, the “Epaphiopsis Complex” of genera is represented 
mainly by Pseudepaphius Ueno (1962), members of which are distinguished from true Epaphiop¬ 
sis members by their smooth pronotum and the presence of a single discal seta on interval 5 next 
to stria 5 (Deuve 1995). Members of our new genus, Trechepaphiopsis , are easily distinguished by 
their elytra chaetotaxy. Genus Junnanotrechus Ueno and Yin (1993) also belongs in this generic 
complex and is probably closely related to Pseudepaphius , based on both morphological (Deuve 
2013a) and molecular (A. Faille, unpublished) data. The illustration of the mandibles of Junnano¬ 
trechus elegantulus Belousov and Kabak (2014b, Fig. 1) confirms this phylogenetic affinity. The 
right mandible presents the same morphological features shared with Pseudepaphius (Fig. 16e) and 
Trechepaphiopsis members (Fig. 16f), namely the unfused premolar and the anterior tip of the reti¬ 
naculum distinctly displaced anteriorly to form a separate tooth. 

Geographical distribution. — This genus currently is known only from the Gaoligong 
Shan region of western Yunnan Province, China, where it is represented by the seven species treat¬ 
ed here. Three were previously described, all in genus Trechus , and four are described here as new. 
Each species apparently occupies only a narrow geographical range within the Gaoligong Shan, but 
their combined known ranges cover all but the northernmost part of the study area. 

As is reflected in the key to species presented below, only male members of most of these 
species can be reliably distinguished, and that only by extraction and examination of their genital- 
ic structure. To date, only two of these species have been recorded as sympatric and syntopic. 


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namely T, unisetosa (Deuve) and T. uniporosa sp. nov. (in Core Area 4). Two species, T. unisetu- 
losa sp. nov. and T. unipilosa sp. nov. occur in the same general area but apparently have different, 
non-overlapping altitudinal ranges. So, at least for the present, locality data may aid in tentative 
identifications of females and undissected males, except for the first two species mentioned above. 


Key for Identification of Adults of Trechepaphiopsis Species of the Gaoligong Shan Region 

1. Elytra without discal setae. T. asetosa (Ueno) 

Elytra with a single discal seta, at or near mid-elytral length, next to stria 3.2 

2. Size larger, BL = 3.5 to 3.7 mm; specimen from southeastern part of Gaoligong Shan region 

( Baoshan County) . T. unisetigera (Ueno) 

Size smaller, BL = 2.8 to 3.5 mm; specimen from more northerly part of the Gaoligon Shan 
region.3 

3. Median lobe of male aedeagus (Fig. 27b) with a sinuous fonn, apical lamella very thin in lat¬ 
eral view, trilobed in apical view (Fig. 27c), cndophallus formed as a long sleeve densely cov¬ 
ered with long, fine spines; specimen from the southcentral part of the Gaoligong Shan region 

( Lushui County ). T. unisetosa (Deuve) 

Median lobe of male aedeagus (Figs. 28b-31b) with a more simple form, apical lamella thick¬ 
er in lateral view, monolobate in apical view (Fig. 28c), cndophallus with scaly areas but with¬ 
out long spines.4 

4. Median lobe of male aedeagus (Fig. 28b) with apical lamella progressively narrowed in later¬ 
al view, apex narrow, bluntly rounded and slightly recurved dorsally {Lushui County •). 

. T. uniporosa sp. nov. 

Median lobe of male aedeagus (Figs. 29b-31b) with apex thicker and more broadly rounded 
in lateral view.5 

5. Size larger, BL = 3.3 to 3.5 mm; median lobe of male aedeagus (Fig. 29b) with shaft thicker 
subbasally, then moderately narrowed toward apex, apex broadly rounded, slightly lobate 

(Fugong County) . T. unisetulosa sp. nov. 

Size smaller, BL = 2.7 to 3.1 mm; median lobe of male aedeagus (Figs. 30b-31b) with shaft 
of more consistent thickness throughout its length, apex short and rounded, not dilated. ... 6 

6. Median lobe of male aedeagus (Fig. 30b) with scaly spoon-shaped area of endophallus only 

slightly narrowed toward apex (Gongshan County) . T. monochaeta sp. nov. 

Median lobe of male aedeagus (Fig. 31b) with scaly spoon-shaped area of endophallus 
markedly narrowed toward apex (Fugong County) . T. unipilosa sp. nov. 

Trechepaphiopsis asetosa (Ueno), 1997 
(Figs. 25, 46-48) 

Trechus (s. sir.) asetosus Ueno, 1997: 185. Holotype, a male, in NMST. Type locality: China, Yunnan, 
Gaoligong Shan, Tengchong County, Dabei, 2430 m. 

Trechepaphiopsis asetosa (Ueno), New Co m b in at 10 n 

Notes on type material.— We have not had an opportunity to study the holotype or any 
other specimens of this species. Features noted below are based on Ueno’s (1997) original descrip¬ 
tion and illustrations. 

diagnosis.— Adults of this species (see Ueno 1997, Fig. 2) can be distinguished from those 
of all other species in the region by the following combination of character states: size small to 
medium (BL= 3.0 to 3.5 mm), apterous; body color dark brown to piceous, dorsum slightly irides- 


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409 


cent; eyes very small, flattened, not projected 
laterally beyond tempora ; tempora convex, 
slightly inflated; pronotum large and trans¬ 
verse (ratio PW/PL=1.32 to 1.49), widest 
near anterior two-fifths, lateral margins slight¬ 
ly more curved in apical half than in basal half, 
basal angles obtuse and rounded; elytra ovoid, 
relative long and narrow, convex, striae 1 and 
2 distinct, nearly complete and finely punctate, 
striae 3 and 4 faintly evident, stria 5 to 7 
effaced, striae 8 evident but interrupted in api¬ 
cal half; recurrent stria terminated anteriorly in 
presumed location of interval 6; discal setae 
absent; preapical seta present, inserted next 
stria 2, inserted closer to sutural than to apical 
margin; median lobe of male aedeagus (see 
Ueno 1997, Figs. 3 and 4) with apex short and 
narrow, faintly recurved dorsally, endophallus 
without sclerites, but covered with scales. 

Habitat distribution.— According to 
Ueno (1997), members of this species were 
collected by sifting moist leaf litter accumula¬ 
tions on the ground in the “Rhododendron 
zone” at elevations ranging from 2340 to 2440 
m. At one of three sites where specimens of 
T. asetosa were found (at 2340 m), specimens 
of Minutotrechus minutus were also collected 
in the same litter samples. 

Geographical distribution within 
the Gaoligong Shan.— Fig. 25. We exam¬ 
ined a single paratype specimen from the type 
locality, the only site from which the species 
has been recorded, in eastern Tengchong 
County in the southern part of the Gaoligong 
Shan region. This site is in Core Area 6. 

Overall geographical distribu¬ 
tion.— This species currently is known only 
from the type locality, in eastern Tengchong 
County in the southern part of the Gaoligong 
Shan, in western Yunnan Province, China. 

Trechepaphiopsis unisetigera (Ueno), 1997 
(Figs. 26, 46-48) 

Trechus (s. str .) unisetiger Ueno, 1997: 190. Holo- 

type, a female, in NMST. Type locality: China, 

Yunnan, Gaoligong Shan, Longyang County, 

Hongxinshu, 2700 m. 


habitus (paratype). b. Map of locality record (red circle) for T. 
asetosa in the Gaoligong Shan region. Scale line =100 km. 


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Trechepaphiopsis unisetigera (Ueno), new Combi¬ 
nation 

Notes on type material.— We have not 
had an opportunity to study the holotype or any 
other specimens of this species. Features noted 
below are based on Ueno’s (1997) original 
description. He did not provide any illustra¬ 
tions for this species. 

diagnosis. — Adults of this species can 
be distinguished from those of all other species 
in the region by the following combination of 
character states: size medium (BL = 3.3 to 3.7 
mm), apterous; body color dark reddish brown, 
dorsum slightly iridescent; eyes very small; 
tempora convex; pronotum transverse (ratio 
PW/PL = 1.34 to 1.40), widest slightly anteri¬ 
or to middle, lateral margins evenly curved 
throughout,with basal angles obtuse and 
rounded; elytra broad, relatively short, convex, 
stria 1 to 3 distinct, punctate, striae 4 to 7 faint¬ 
ly impressed but evident in basal two-thirds, 
striae 8 evident in apical half; recurrent stria 
terminated anteriorly in presumed location of 
interval 6; one discal setae present near apical 
one-third and inserted next to stria 3; preapical 
seta present, inserted next stria 2, inserted 
slightly closer to sutural than to apical margin. 

Habitat distribution.— According to 
Ueno (1997) the two specimens of the type 
series of this species were “sorted out from soil 
samples taken in a forest of Lithocarpus vari- 
olosus ” at an elevation of 2700 m. No other 
trechines have been found syntopic with mem¬ 
bers of this species. 

Geographical distribution within 
the Gaoligong Shan.— Fig. 26. This 
species is known only from the type locality in 
Longyang County in the southern part of the 
Gaoligong Shan region. This locality is in Core 
Area 7. 

Overall geographical distribu¬ 
tion.— This species currently is known only 
from Longyang County in the southern part of 
the Gaoligong Shan region, western Yunnan 
Province, China. 


Figure 26. Trechepaphiopsis unisetigera (Ueno); Map 
of locality record (red circle) for T. unisetigera in the 
Gaoligong Shan region. Scale line =100 km. 


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Trechepaphiopsis unisetosa (Deuve), 2004 (sp. 2) 

(Figs. 27, 37a, 43b, 46-48) 

Trechus unisetosus Deuve, 2004: 220. Holotype, a female, in SCAU. Type locality: China, Yunnan, Gaoligong 

Shan, Lushui County, Fengxue Yakou [Pass], 2600-2700 m. 

Trechepaphiopsis unisetosa (Deuve), New Combination 

diagnosis. — Adults of this species (Fig. 27a) can be distinguished from those of all other 
species in the region by the following combination of character states: size medium (BL = 3.1 to 
3.5 mm), apterous; body color light brown, dorsum slightly iridescent; eyes small but convex; tem- 
pora very convex, sparsely pubescent; pronotum transverse (ratio PW/PL=1.35), with basal 
angles obtuse and rounded; elytra convex, with median 2 or 3 striae deeply impressed, lateral stri¬ 
ae faintly impressed to effaced; recurrent stria terminated anteriorly in presumed location of inter¬ 
val 6; only one discal setiferous pore present, inserted at anterior one-third next to stria 3; preapi- 
cal seta present, inserted next to stria 2; median lobe of male aedeagus (Fig. 27b) highly distinc¬ 
tive, with ventral margin of shaft sinuous and apex short and thin in lateral view, with broad apical 
projects in ventral view; endophallus without sclerites, but with a long tubular sleeve covered with 
long, slender spines. 

Commen t s.— Among males of all the species of Trechepaphiopsis, those of T. unisetosa have 
an endophlallus most like that typical for members of the Epaphiopsis complex of genera, espe¬ 
cially of certain species of Epaphius described from China (Jeannel 1962, Deuve 1992b). These 
also have a long tubular sleeve covered with long spines, symmetrically arranged in ventral view 
(Fig. 27c) 

habitat distribution . — Members of this species have been found in mixed broadleaf ever¬ 
green/ deciduous forest at elevations ranging from 2460 to 2470 m and collected using both pitfall 
baps and sifting of leaf litter from the forest floor (Fig. 37a). Specimens of Trechepaphiosis uni- 
porosa sp. nov and Trechepaphiama goaiigong sp. nov were collected in the same sifted leaf litter 
samples and are therefore syntopic with T. unisetosa at this site. Additional specimens of T. unise¬ 
tosa were collected in pitfall traps set in Rhododendron and bamboo thickets on the east flank of 
the summit ridge just south of the Fengxue Yakou at 3150 m (Fig. 43b), a site slightly above the 
type locality for the species. 

Geographical distribution within the Gaoligong Shan. — Fig. 27d. We examined a 
total of 25 specimens (10 males and 16 females) from the following localities: Lushui County: 
Luzhang Township (100 m S of Fengxue Yakou on east side of pass, 3150 m, 
N25.97195°/E97.68381°, 11-21 May 2005, D.H. Kavanaugh, C.E. Griswold & K.J. Guo collectors 
[1 male and 1 female; CAS, IOZ]); Pianma Township (9.3 km ENE of Pianma along road to Lushui 
at Changyan He, 2460-2470 m, N25.99363°/E97.66651°, 15-18 October 1998, D.H. Kavanaugh, 
C.E. Griswold, C. Ferraris & C.L. Long collectors [7 males, 12 females; CAS, IOZ, MNHN], 12- 
21 May 2005, D.H. Kavanaugh, C.E. Griswold & K.J. Guo collectors [2 male and 2 females; CAS, 

I0Z] ). 

Members of this species have been collected only in the southcentral part of the study area, on 
both western and eastern slopes of the Gaoligongshan, which are in Core Areas 4 and 5, respec¬ 
tively. 

Overall geographical distribution . — This species currently is known only from Lushui 
County in the southcentral part of the Gaoligong Shan region, western Yunnan Province, China. 


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Figure 27. Trechepaphiopsis unisetosa (Deuve); a. Dorsal habitus (CASENT1024491). b-c. Median lobe of aedeagus 
of male (CASENT1001903); b. left lateral aspect; c. ventral aspect, d. Map of locality records (red circles) for T. unisetosa 
in the Gaoligong Shan region. Scale lines a-c = 0.5 mm, d = 100 km. 


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Trechepaphiopsis uniporosa Deuve and Liang, sp.nov. 

(Figs. 16f, 28, 37a, 46-48) 

Type material.— Holotype, a male, in IOZ, labeled: “CASENT 1001923”/ “CHINA, Yun¬ 
nan Province, Gaoligongshan Mountains, Nujiang Prefecture, 9 [actually 9.3] km ESE of Pianma, 
25°59.6’N/ 98°37.6’E.7 “2450 [actually 2460-2470] m, 15-18 October 1998, Stop #98-118D D.H. 
Kavanaugh, C.E. Griswold, C. Ferraris & C.-L. Long collectors”/ “IMAGE” [green label]/ 
“HOLOTYPE Trechepaphiopsis uniporosa Deuve & Liang, sp. nov. designated 2016” [red label]. 
Paratypes (a total of 5): 3 males and 2 females (in CAS, IOZ, MNHN) labeled: same as holotype, 
except first label “CASENT 1001920” to “CASENT 1001922” and “CASENT 1001924” to 
“CASENT 1001925”, respectively. All paratypes also bear the following label: “PARATYPE 
Trechepaphiopsis uniporosa Deuve & Liang, sp. nov. designated 2016” [yellow label]. 

Type locality. — China, Yunnan, Gaoligong Shan, Lushui County, Pianma Township, 9.3 
km ESE of Piamna, 25.99363°/ 98.66651°, 2460-2470 m. 

Derivation of species name. — The species epithet, uniporosa , is an adjective derived from 
the Latin words, unus, meaning one, and porus , meaning pore or hole. The name refers to the sin¬ 
gle discal setiferous pore found on the elytra of members of this species. 

diagnosis. — Adults of this species (Fig. 28a) can be distinguished from those of all other 
species in the region by the following combination of character states: size medium (BL = 3.3 to 
3.5 mm), apterous; body color light brown, slightly iridescent; eyes small; tempora distinctly con¬ 
vex, sparsely pubescent; pronotum transverse (ratio PW/PL = 1.34), with basal angles obtuse and 
rounded; elytra convex, with stria 1 deeply impressed and finely punctate, other striae more faint¬ 
ly impressed and lateral striae effaced; recurrent stria terminated anteriorly in presumed location of 
interval 6; only one discal setiferous pore present, inserted at anterior one-third next to stria 3; 
preapical seta present, inserted next to stria 3; median lobe of male aedeagus (Fig. 28b) with apex 
slender but bluntly rounded and slightly recurved dorsally, endophallus with a faintly sclerotized 
scaly area. 

d e script ion . — Size medium BL = 3.3 to 3.5 mm. Body color light brown, appendages con- 
colorous, except palpi paler, dorsum shiny, slightly iridescent. 

Head. Moderate in size, short; eyes small, not projected, their diameter about equal to length 
of tempora but their convexity less than that of tempora, the latter markedly convex, inflated and 
sparsely pubescent. Frontal furrows thin and linear, deeply impressed, arcuate, continuous posteri¬ 
orly to hind margins of tempora. Two pairs of supraorbital setae present, the anterior pair inserted 
in foveae. Clypeus with four setae. Labrum with six setae, anterior margin distinctly concave or 
emarginate. Mandibles short and slender, right mandible as in Fig. 16f. Mentum and submentum 
not fused. Mentum with medial tooth apically truncate, about one-half the length of the lateral 
lobes. Submentum with six setae anteriorly. Antennae rather short, not quite extended posteriorly 
to basal one-fourth of the elytra, only 2.5 antennomeres extended posteriorly beyond basal prono- 
tal margin; antennomeres 2 and 3 about equal in length, antennomere 4 slightly shorter. 

Pronotum. Distinctly transverse (ratio PW/PL = 1.34), with greatest width near anterior one- 
third, only slightly narrowed posteriorly; lateral margins rounded, slightly more so anteriorly, 
slightly straighten just anterior to basal angles, the latter obtuse and rounded. Disc smooth and con¬ 
vex, median longitudinal impression finely impressed, but continuous between anterior and poste¬ 
rior margins; basal foveae shallow, faintly impressed; median basal area reduced, faintly delimit¬ 
ed, slightly punctate in some specimens. Basal margin nearly rectilinear. Lateral pronotal borders 
moderately slender, regular, narrowly and regularly reflexed, lateral grooves narrow but distinctly 
impressed. Single midlateral setae on each side inserted near anterior one-third; single basolateral 
seta on each side, inserted at basal angle. 


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Figure 28. Trechepaphiopsis uniporosa sp. nov.; a. Dorsal habitus (CASENT1001923). b-c. Median lobe of aedeagus 
of male (CASENT1001923); b. left lateral aspect; c. ventral aspect. d..Map of locality records (red circles) for T. uniporosa 
in the Gaoligong Shan region. Scale lines a-c = 0.5 mm, d = 100 km. 


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Elytra. Ovoid, only slightly more narrowed anteriorly than posteriorly, humeri distinct but 
rounded. Disc convex, striae faintly impressed and finely punctate, the medial three or four striae 
distinctly impressed, the more lateral striae more or less effaced, but perceptible if only by the pres¬ 
ence of fine punctures seen as rows of brown dots visible through the integument. Parascutellar 
striole present. Recurrent stria abruptly terminated anteriorly in presumed location of interval 6. 
Parascutellar setiferous pore present at base at common origin of discal striae 1 and 2. Only one 
discal setiferous pore present, inserted slightly anterior to middle next to stria 3. Preapical seta pres¬ 
ent on interval 3 near stria 3, closer to elytra apical margin that to sutural margin. 

Legs. Short, protibiae with longitudinal furrow. Male protarsomeres 1 and 2 dilated and api- 
comedially toothed. 

Abdomen. Abdominal ventrites glabrous, except for a single paramedial seta on each side, and 
ventrite VII of males apically with one pair of paramedial setae, of females with two pairs. 

Male aedeagus. Median lobe (Fig. 28b) with sagittal aileron very small, shaft moderately 
broad basally, expanded near mid-length, then gradually narrowed apically to a thin, bluntly round¬ 
ed apex, the latter slightly recurved dorsally; endophallus with a small scaly sclerotized area (Fig. 
28c). 

habitat distribution .— Members of this species have been found in mixed broadleaf ever¬ 
green/ deciduous forest at elevations ranging from 2460 to 2470 in and collected by sifting leaf lit¬ 
ter on the forest floor (Fig. 37a). Specimens of Trechepaphiosis unisetosa and Trechepaphiama 
gaoligong sp. nov were collected in the same sifted leaf litter samples and are therefore syntopic 
with T. uniporosa at this site. 

Geographical distribution within the Gaoligong Shan.— Fig. 28d. We examined a 
total of 6 specimens (4 males and 2 females) from the type locality on the western slope of the 
southcentral part of the Gaoligong Shan in Lushui County (see Type material above for exact col¬ 
lection data). This locality is in Core Area 4. 

Overall geographical distribution.— This species currently is known only from west¬ 
ern Lushui County in the southcentral part of the Gaoligong Shan region, western Yunnan 
Province, China. 

Trechepuphiopsis Ullisetulosa Deuve and Kavanaugh, sp. nov. 

(Figs. 29, 40b, 46—48) 

Type material.— Holotype, a male, in IOZ, labeled: “CASENT 1018407”/ “CHINA, Yun¬ 
nan Province, Fugong County, Lishadi Township, 0.2 km W of Shibali, 2357 m, 
N27.16650°/E098.77936°”/ “18 August 2005, in leaf litter. Stop# PP-4405, P. Paquin”/ “IMAGE” 
[green label]/ “HOLOTYPE Trechepuphiopsis unisetulosa Deuve & Kavanaugh, sp. nov. desig¬ 
nated 2016” [red label]. Paratypes (a total of 26): 1 female (in IOZ) labeled: same as holotype, 
except first label “CASENT 1018408”; 1 female (in CAS) labeled: “CASENT 1014246”/ 
“CHINA, Yunnan Province, Fugong County, Lishadi Township, 0.4 km SE of Shibali along North 
Fork of Yamu He, N27.163377E098.78208°, 2475 m, 8-11 May 2004”/ “from mini-Winkler 
extraction of leaf litter siftate, Stop #CGY37, C. E. Griswold, D. H. Kavanaugh, & Yan H.-M. col¬ 
lectors”; 1 male (in CAS) labeled: “CASENT 1019222”/ “CHINA, Yunnan Province, Fugong 
County, Lishadi Township, Shibali Road at Shibali, N27.16786°/E098.77741°,7 “ 2560 m, 3 May 
2004 Stop #DHK-2004-024, D.H. Kavanaugh & C. E. Griswold, collectors”; 1 female (in IOZ) 
labeled: “CASENT 1020952”/ “CHINA, Yunnan Province, Fugong County, Lishadi Township, 8.5 
km above Shibali on Shibali Road, north bank of North Fork of Yamu He, 
N27.1184167E098.72026°”/ “3100 m, 9 August 2005, Stop DHK-2005-067A D.H. Kavanaugh, 
H.B. Liang, D.Z. Dong, & P Paquin collectors”; 1 male and 1 female (in CAS) labeled: “CASENT 


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1021953” and “CASENT 1021952”, res/ “CHINA, Yunnan Province, Fugong County, Lishadi 
Township, 0.3 km above Shibali on Shibali Road, N27.ll6791°/E098.77655°”/ “2563 m, in pitfall 
trap, 3-11 May 2004, Stop #CGY21, C. E. Griswold, D. H. Kavanaugh, Liang H.-B., & Yan H.-M. 
collectors”; 10 males and 10 females (in CAS, 10Z, MNHN) labeled: “CASENT 1023285” to 
“CASENT 1023294” and “CASENT 1023295” to “CASENT 1023304”, respectively/ “CHINA, 
Yunnan Province, Fugong County, Lishadi Township, 0.3 km above Shibali on Shibali Road, 
N27.1166361 °/E098.77667°”/ “2563 m, in pitfall trap, 4 May 2004, Stop #DHK-2004-026, D. H. 
Kavanaugh & C. E. Griswold collectors”. All paratypes also bear the following label: “PARATYPE 
Trechepaphiopsis unisetulosa Deuve & Kavanaugh, sp. nov. designated 2016” [yellow label]. 

type locality.— China, Yunnan, Fugong County, Shiyueliang Township, 0.2 km W of 
Shibali, N27.16650°/E098.77936°, 2537 m, 

d e r i vat ion o e s pe c ie s name .— The species epithet, unisetulosa, is an adjective derived from 
the Latin words, units, meaning one, and setulosus, meaning bearing bristles or setae. The name 
refers to the single discal seta found on the elytra of members of this species. 

diagnosis.— Adults of this species (Fig. 29a) can be distinguished from those of all other 
species in the region by the following combination of character states: size small to medium (BL 
= 3.3 to 3.5 mm), apterous; body color dark reddish brown, dorsum shiny, very slightly iridescent; 
eyes small, only slightly projected laterally; tempora convex and sparsely pubescent; pronotum 
transverse (ratio PW/PL = 1.40), with basal angles obtuse and rounded; elytra convex, with medi¬ 
al three or four stria distinct, finely punctate, more lateral striae more or less effaced; recurrent stria 
terminated anteriorly in presumed location of interval 6; only one discal setiferous pore present, 
inserted near anterior one-third next to stria 3; preapical seta present, inserted next to stria 2; medi¬ 
an lobe of male aedeagus (Fig. 29b) with apex lobated and broadly rounded; endophallus with a 
large, scaly sclerorized area. 

Description. — Size small to medium, BL = 3.3 to 3.5 mm. Body color dark reddish brown, 
interval 1 of elytra slightly paler, reddish, appendages yellowish brown, palpi paler yellow; dorsum 
smooth, pronotum and elytra shiny, very slightly iridescent, head slightly alutaceous from more 
deeply impressed isodiametric microsculpture. 

Head. Rather broad; eyes small, only slightly projected laterally, their diameter about equal to 
length of tempora, the latter convex and sparsely pubescent. Frons convex, with frontal furrows 
evenly rounded and deeply impressed, continuous to posterior margins of tempora. Two pairs of 
supraorbital setae present, the anterior pair inserted in foveae. Clypeus with four setae. Labrum 
with six setae, anterior margin distinctly emarginate. Mentum and submentum not fused. Mentum 
with medial tooth broad and apically bifid, less than half the length of the lateral lobes. Submen¬ 
tum with six setae anteriorly. Gula broad. Genae with a single ventral seta one each side. Antennae 
short, with only about 1.5 antennomeres extended posteriorly beyond basal pronotal margin; anten- 
nomeres 2 and 4 about equal in length, antennomere 3 slightly longer. 

Pronotum. Transverse (ratio PW/PL = 1.40), with greatest width near anterior one-third, mod¬ 
erately narrowed posteriorly; lateral margins rounded, slightly more so anteriorly, slightly straight¬ 
en, but not at all sinuate, just anterior to basal angles, the latter bluntly obtuse. Disc convex, medi¬ 
an longitudinal impression finely impressed, continuous from posterior side of apical median 
swelling to posterior margin; basal foveae shallow, faintly impressed; median basal area well 
defined but short and transverse. Lateral borders slender and finely relexed dorsally, lateral grooves 
deeply impressed. Single midlateral setae on each side inserted near anterior one-third; single baso- 
lateral seta on each side, inserted at basal angle. 


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Figure 29. Trechepaphiopsis unisetulosa sp. nov.; a. Dorsal habitus (CASENT1018407). b. Median lobe of aedeagus 
of male (CASENT1001903), left lateral aspect, c. Map of locality records (red circles) for T. unisetulosa in the Gaoligong 
Shan region. Scale lines a, b = 0.5 mm, c = 100 km. 


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Elytra. Ovoid, not or only slightly elongate, about equally narrowed anteriorly and posterior¬ 
ly, humeri distinct but rounded. Disc convex, striae faintly impressed and finely punctate, the medi¬ 
al three or four striae distinctly impressed, the more lateral striae more or less effaced, bearly per¬ 
ceptible. Parascutellar stride present, longer than average for genus. Recurrent stria abruptly ter¬ 
minated anteriorly in presumed location of posterior end of stria 5. Parascutellar setiferous pore 
present at base at common origin of discal striae 1 and 2. Only one discal setiferous pore present, 
inserted at anterior two-fifth of elytral length next to stria 3. Prcapical seta present on interval 3 
next to stria 2 and about equidistant from apical and sutural elytral margins. Umbilicate setal series 
with setae of humeral group equidistance from each other and setae of median group inserted 
slightly posterior to middle. 

Legs. Short, protibiae with longitudinal furrow. Male protarsomeres 1 and 2 dilated and api- 
comedially toothed. 

Abdomen. Abdominal ventrites glabrous, except for a single paramedial seta on each side, and 
ventrite VII of males apically with one pair of paramedial setae, of females with two pairs. 

Male aedeagus. Median lobe (Fig. 29b) with sagittal aileron moderate in size, shaft broad 
basally, then gradually narrowed apically to a broadly rounded apex; endophallus with a large, 
scaly sclerorized area with two parts or folds. 

h abitat distribution,— Members of this species have been collected in pitfall traps set in 
slightly disturbed broadleaf forest with large trees, dense understory of ferns and other herbs, and 
a deep leaf litter layer, and also by sifting leaf litter in this same habitat. One specimen was beat¬ 
en from roadside vegetation with suspended leaf and twig debris in that vegetation in the same area. 
One specimen was collected at a higher elevation (3100 m) from under stones in a rocky open areas 
cleared by snow avalanches but shaded by three meter high herbaceous cover (Fig. 40b). Speci¬ 
mens of Trechus shiyueliang, T. shibalicus and Trechepaphiopsis umpilosa sp. nov.were found syn- 
topic with the specimen of T. unisetulosa at this last site. 

Geographical distribution within the Gaoligong Shan.— Fig. 29c. We examined a 
total of 27 specimens (13 males and 14 females) from sites on the eastern slope of the northcentral 
part of the Gaoligong Shan in Fugong County (see Type material above for exact collection data). 
All of these sites are in Core Area 3. 

Overall geographical distribution .— This species currently is known only from Fugong 
County in the northcentral part of the Gaoligong Shan region, western Yunnan Province, China. 

Trechepaphiopsis monochaeta Deuve and Kavanaugh, sp. nov. 

(Figs. 30, 38b, 42a, 46-48) 

type material.— Holotype, a male, in IOZ, labeled: labeled: “CASENT 1007387”/ 
“CHINA, Yunnan Province, Gaoligong Shan, Nujiang Prefecture, Nujiang State Nature Reserve, 
No. 12 Bridge Camp area, 16.3 airkmW of Gongshan ”/ “N27.71503°/E98.50244°, 2775 m, 15-19 
July 2000, Stop#00-23A, D.H. Kavanaugh, C.E. Griswold, Liang H.-B., D. Ubick, & Dong D.-Z. 
collectors”/ “IMAGE” [green label]/ “HOLOTYPE Trechepaphiopsis monochaeta Deuve & 
Kavanaugh, sp. nov. designated 2016” [red label]. Paratypes (at total of 56): 14 males and 17 
females (in CAS, IOZ, MNHN) labeled: same as holotype, except first label “CASENT 1007388” 
to “CASENT 1007400” and “CASENT 1021923” and “CASENT 1007401” to “CASENT 
1007416” and “CASENT 1021924”, respectively; 10 males and 14 females (in CAS, IOZ) labeled: 
“CASENT 1006508” to “CASENT 1006517” and “CASENT 1006518” to “CASENT 1006530” 
and “CASENT 1021922”, respectively/ “CHINA, Yunnan Province, Gaoligong Shan, Nujiang Pre¬ 
fecture, Gongshan County, Dazhu He drainage, 13.5 airkm SW of Gongshan, 2830m”/ 
“N27.62947°/E98.62010°, 30 June- 5 July 2000, Stop#00-17I, D.H. Kavanaugh, C.E. Griswold, 


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Liang H.-B., D. Ubick, & Dong D.-Z. collectors”; 1 female (in CAS) labeled: “CASENT 
1025812”/ “CHINA, Yunnan, Gongshan County, Bingzhongluo Township, SW slope of 
Kawakarpu Shan 0.3 km SW of Chukuai Lake at campsite,”/, “N27.97686 0 / E098.47799 0 , 3750 m, 
18 August 2006 Stop #DHK-2006-078 D.H. Kavanaugh collector”. All paratypes also bear the fol¬ 
lowing label: “PARATYPE Trechepaphiopsis monochaeta Deuve & Kavanaugh, sp. nov. desig¬ 
nated 2016” [yellow label]. 

Type locality .— China, Yunnan, Gaoligong Shan, Gongshan County, Qiqi Trail at No. 12 
Bridge Camp area, 16.3 airkm W of Gongshan, N27.71503°/E98.50244°, 2775 m, 

derivation of species name.— The species epithet, monochaeta, is an adjective derived 
from the Greek words, povog (transliterated into Latin as mono), meaning one or single, and yarnx 
(transliterated into Latin as chcieta), meaning hah or bristle. The name refers to the single discal 
seta found on the elytra of members of this species. 

Diagnosis. — Adults of this species (Fig. 30a) can be distinguished from those of all other 
species in the region by the following combination of character states: size small (BL = 2.8 to 3.0 
mm), apterous; body color reddish brown, dorsum shiny, very slightly iridescent; eyes small but 
convex; tempora convex and sparsely pubescent; pronotum transverse (PW/PL = 1.38), with basal 
angles obtuse and rounded; elytra convex, medial four or five striae distinctly impressed, more lat¬ 
eral striae more or less effaced; recurrent stria terminated anteriorly in presumed location of inter¬ 
val 6; only one discal setiferous pore present, inserted anterior to and next to stria 3; preapical seta 
present, inserted next to stria 2; median lobe of male aedeagus (Fig. 30b) with apex moderately 
broad and apically rounded, slightly deflected ventrally; endophallus with an elongate and scaly 
sclerotized area not narrowed apically. 

Description. — Size small, BL = 2.8 to 3.0 mm. Body color reddish brown, appendages 
slightly paler, yellowish brown, palpi paler yellow; dorsum shiny, slightly iridescent. 

Head. Medium in size; eyes small but moderately convex, their diameter about equal to length 
of tempora, the latter distinctly convex and sparsely pubescent. Frons convex, with frontal furrows 
slightly angulate and deeply impressed, continuous to posterior margins of tempora, not or slight¬ 
ly attenuated posterior to margins of eyes. Two pairs of supraorbital setae present, the anterior pair 
inserted in foveae. Clypeus with four setae. Labrum with six setae, anterior margin distinctly emar- 
ginate or concave. Mentum and submentum not fused. Mcntum with medial tooth broad and api¬ 
cally truncate or faintly bifid, about half the length of the lateral lobes. Submentum with six setae 
anteriorly. Gula broad. Genae with a single ventral seta one each side. Antennae short, with only 
about 1.5 antennomeres extended posteriorly beyond basal pronotal margin; antennomeres 2 and 3 
about equal in length, antennomere 4 slightly shorter. 

Pronotum. Transverse (ratio PW/PL = 1.38), with greatest width slightly anterior to middle; 
lateral margins rounded, slightly more so anteriorly, not or only slightly straighten posteriorly just 
anterior to basal angles, the latter obtuse and bluntly or rounded. Disc convex and smooth, median 
longitudinal impression finely impressed, continuous from middle of apical median swelling to 
posterior margin; basal foveae very small and shallow, faintly impressed; median basal area well 
defined but short and transverse, smooth but with several small longitudinal foveae. Lateral bor¬ 
ders slender and finely relexed dorsally, lateral grooves deeply impressed. Single midlateral setae 
on each side inserted near anterior one-third; single basolateral seta on each side, inserted at basal 
angle. 

Elytra. Ovoid, only slightly narrowed anteriorly and posteriorly, humeri distinct but rounded. 
Disc convex, striae finely impressed and not or only faintly punctate, the medial three or four stri¬ 
ae clearly impressed, the more lateral striae more or less effaced, but still evident. Parascutellar stri¬ 
de present and distinct. Recurrent stria deeply impressed, its anterior recurved end terminated in 


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Figure 30. Trechepaphiopsis monochaeta sp. nov.; a. Dorsal habitus (CASENT1007411). b. Median lobe of aedeagus 
of male (CASENT1007411), left lateral aspect, c. Map of locality records (red circles) for T. monochaeta in the Gaoligong 
Shan region. Scale lines a, b = 0.5 mm, c = 100 km. 


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the posterior end of stria 5. Only one discal setiferous pore present, inserted slightly anterior to 
middle next to stria 3. Preapical seta present, inserted on interval 3 next to stria 2 and about equi¬ 
distant from apical and sutural elytra! margin. 

Legs. Short, protibiae with longitudinal furrow. Male protarsomeres 1 and 2 dilated and api- 
comedially toothed. 

Abdomen. Abdominal ventrites glabrous, except for a single paramedial seta on each side, and 
ventrite VII of males apically with one pair of paramedial setae, of females with two pairs. 

Male aedeagus. Median lobe (Fig. 30b) with sagittal aileron very small, shaft moderately 
broad basally, then gradually narrowed, with apex moderately broad and apically rounded, slight¬ 
ly deflected ventrally; endophallus with an elongate and scaly sclerotized area not narrowed api¬ 
cally. 

Comments.— This species appears to be very closely related (perhaps sister species) to T. 
unipilosa sp. nov., but its male members can be distinguished from those of the latter by having the 
base of the median lobe of the male aedeagus broader (narrower in T. unipilosa members, see Fig. 
3 lb) and the scaly sclerotized area of the endophallus of more equal width throughout and not nar¬ 
rowed distally (as it is in T. unipilosa males). 

habitat distribution.— Members of this species have been collected by sifting leaf litter 
and mosses from the floor of closed canopy forest with an understory of ferns and large-leafed (ca. 
60 cm long leaves) Rhododendron sp. at elevations ranging from 2775 to 2830 m (Fig. 42a). At the 
lowest elevation, specimens of Trechus qiqiensis were collected in the same litter samples. One 
specimen was also found under stones or wood chips in a disturbed open meadow area surround¬ 
ed by one meter high Rhododendron thickets at an elevation of 3750 m (Fig. 38b); and specimens 
of Queinnectrechus griswoldi were found in this same area 

Geographical distribution within the Gaoligong Shan.— Fig. 30c. We examined a 
total of 57 specimens (23 males and 34 females) from sites on the crest or eastern slope of the 
northern part of the Gaoligong Shan in Gongshan County (see Type material above for exact col¬ 
lection data). All of these sites are in Core Area 2. 

o verall geographical distribution .— This species currently is known only from Gong¬ 
shan County in the northern part of the Gaoligong Shan region, western Yunnan Province, China. 

Trechepaphiopsis unipilosa Deuve and Liang, sp. nov. 

(Figs. 31,40b, 46M8) 

Type material.— Holotype, a male, in IOZ, labeled: “CASENT 1023757”/ “CHINA, Yun¬ 
nan, Fugong County, Lishadi Township, 8.5 km above Shibali on Shibali Road, North Fork of 
Yamu He, N27.18416°/E98.72026°,”/ “3100 m, 7 May 2004 Stop #DHK-2004-038A D.H. 
Kavanaugh, C.E. Griswold, Liang H.-B., & Zhu B.-X. collectors”/ “HOLOTYPE Trechepaphiop¬ 
sis unipilosa Deuve & Liang, sp. nov. designated 2016” [red label]. Paratypes (at total of 17): 2 
females (in CAS, IOZ) labeled: same as holotype, except first label “CASENT 1023746” and 
“CASENT 1023758”, respectively; 1 male and 2 females (in CAS, IOZ) labeled: “CASENT 
1017488” and “CASENT 10174896” to “CASENT 1017490”, respectively/ “CHINA, Yunnan, 
Fugong County, Lumadeng Township, 8.5 km above Shibali on Shibali Road, south bank of North 
Fork of Yamu He, N27.18326 0 / E98.72002 0 ”/ 3100 m, 8 August 2005 Stop #DHK-2005-067B 
D.H. Kavanaugh, H.B. Liang, D.Z. Dong, & J.F. Zhang collectors”; 7 males and 1 female (in CAS, 
IOZ, MNHN) labeled: “CASENT 1018375” to “CASENT 1018381” and “CASENT 1018382”, 
respectively/ “CHINA, Yunnan, Fugong County, Lishadi Township, 10 km W of Shibali on Shibali 
Road, 3250 m,”/ “N27.20055°/E98 J1399°, 16 August 2005 Stop #PP-3805 P. Paquin collector”; 
1 male (in CAS) labeled: “CASENT 1018392”/ “CHINA, Yunnan, Fugong County, Lishadi Town- 


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ship, 10.5 km W of Shibali on Shibali Road, 3221 m ”/ “N27.20192°/E98.71329°, 17 August 2005 
Stop #PP-4105 P. Paquin collector”; 3 males (in CAS, IOZ) labeled: “CASENT 1023607”, 
“CASENT 1023608” and “CASENT 1023820”, respectively/ “CHINA, Yunnan, Fugong County, 
Lishadi Township, 11.5 km above Shibali on Shibali Road, N27.20676°/E98.71763°,”/ “3290 m, 6 
May 2004 Stop #DHK-2004-036 D.H. (Cavanaugh, C.E. Griswold, Liang H.-B., & Zhu B.-X. col¬ 
lectors”. All paratypes also bear the following label: “PARATYPE Trechepaphiopsis unipilosa 
Deuve & Liang, sp. nov. designated 2016” [yellow label]. 

Type locality. — China, Yunnan, Fugong County, Shiyueliang Township, 8.5 km above 
Shibali on Shibali Road, North Fork ofYamu He, N27.184167E98.72026°, 3100 m. 

derivation of species name. — The species epithet, unipilosa , is an adjective derived from 
the Latin word, unus, meaning one, and the Greek word, izIXoq (transliterated into Latin as pilus ), 
meaning hair. The name refers to the single discal seta found on the elytra of members of this 
species. 

diagnosis. — Adults of this species (Fig. 31a) can be distinguished from those of all other 
species in the region by the following combination of character states: size small (BL = 2.7 to 3.1 
mm), apterous; body color reddish brown, dorsum shiny, pronotum and elytra slightly iridescent; 
eyes small but convex; tempora convex and sparsely pubescent; pronotum transverse (ratio PW/PL 
= 1.42), with basal angles obtuse and rounded; elytra convex, medial striae (1 to 5 or 6) distinct, 
finely punctate, more lateral striae more or less effaced; recurrent stria terminated anteriorly on 
interval 6; only one discal setiferous pore present, inserted at anterior one-third next to stria 3; 
preapical seta present, inserted next to stria 2; median lobe of male aedeagus (Fig. 31b) with apex 
slender and rounded, endophallus with a faintly sclerotized scaly area distinctly narrowed distally. 

Description .— Size small, BL = 2.7 to 3.1 mm. Body color reddish brown, antennmae con- 
colorous, legs slightly paler, yellowish brown, palpi paler yellow; dorsum shiny, pronotum and ely¬ 
tra slightly iridescent, head slightly alutaceous from more deeply impressed, irregularly isodia- 
metric microsculpture. 

Head. Broad; eyes small but convex, their diameter about equal to length of tempora, the lat¬ 
ter short, distinctly convex and sparsely pubescent Frons convex, frontal furrows sharply impressed 
and rounded, not attenuated posterior to eyes. Two pairs of supraorbital setae present, the anterior 
pair inserted in foveae. Clypeus with four setae. Labrum with six setae, anterior margin distinctly 
emarginate. Mentum and submentum not fused. Mentum with medial tooth apically bifid, about 
half the length of the lateral lobes. Submentum with six setae anteriorly. Gula broad. Genae with a 
single ventral seta one each side. Antennae short, with only about two antennomeres extended pos¬ 
teriorly beyond basal pronotal margin; antennomeres 2 and 4 about equal in length, antennomere 3 
slightly longer. 

Pronotum. Transverse (ratio PW/PL = 1.42), with greatest width at anterior one-third; lateral 
margins rounded, slightly more so anteriorly, not or only slightly straighten posteriorly just anteri¬ 
or to basal angles, the latter obtuse and blunted, Disc convex and smooth, median longitudinal 
impression finely impressed, continuous from middle of apical median swelling to posterior mar¬ 
gin; basal foveae small and shallow, faintly impressed; median basal area well defined but short 
and transverse, smooth or with a few small longitudinal foveae. Lateral borders slender and finely 
relexed dorsally, lateral grooves deeply impressed. Single midlateral setae on each side inserted 
near anterior one-third; single basolateral seta on each side, inserted at basal angle. 

Elytra. Ovoid only slightly elongate, not or only very slightly more narrowed anteriorly than 
posteriorly, humeri distinct but rounded. Disc convex, striae finely punctate, only medial striae 1 
to 3 distinctly impressed, more lateral striae more or less effaced, but still evident in most speci¬ 
mens. Parascutellar striole present, distinct but short. Recurrent stria deeply impressed, its anterior 


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Figure 31. Trechepaphiopsis imipilosa sp. nov.; a. Dorsal habitus (CASENT1023757). b. Median lobe of aedeagus of 
male (CASENT1023757), left lateral aspect, c. Map of locality records (red circles) for T. unipilosa in the Gaoligong Shan 
region. Scale lines a, b = 0.5 mm, c = 100 km. 


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end abruptly terminated in the posterior end of stria 5. Parascutcllar setiferous pore present at base 
at common origin of discal striae 1 and 2. Only one discal setiferous pore present, inserted at ante¬ 
rior two-fifth of elytral length next to stria 3. Only one discal setiferous pore present, inserted 
slightly anterior to middle next to stria 3. Preapical seta present, inserted on interval 3 next to stria 
2 and about equidistant from apical and sutural elytral margin. Umbilicate setal series with setae of 
humeral group equidistance from each other and setae of median group inserted slightly posterior 
to middle. 

Legs. Short, protibiae with longitudinal furrow. Male protarsomeres 1 and 2 dilated and api- 
comedially toothed. 

Abdomen. Abdominal ventrites glabrous, except for a single paramedial seta on each side, and 
ventrite VII of males apically with one pair of paramedial setae, of females with two pairs. 

Male aedcagus. Median lobe (Fig. 31b) with sagittal aileron very small, shaft moderately 
broad basally, then gradually narrowed apically to a narrowly rounded apex; endophallus with a 
faintly sclerotized scaly area distinctly elongate and nan-owed distally. 

Comments. — This species appears to be very closely related (perhaps sister species) to 
I monochaetus, but its male members can be distinguished from those of the latter by having the 
base of the median lobe of the male aedeagus narrower (broader in T. monochaeta members, see 
Fig. 30b) and the scaly sclerotized area of the endophallus distinctly narrowed distally (of more 
equal width throughout in T. monochaeta males). 

Habitat distribution. — Members of this species have been collected by sifting leaf litter 
from forests of scattered, large Abies sp. trees with a dense understory of bamboo or Rhododen¬ 
dron spp. at elevations ranging from 3221 to 3290 m. Specimens of Trechus shiyueliang and T. 
shibalicus were collected in one or more of the same litter samples. Specimens of T. unipilosa were 
also collected at 3100 m elevation under stones at the open edges of a small stream draining a 
north-facing glacial cirque with a large snowfield in its basin (Fig. 40b). A specimen of Trechepa- 
phiopsis unisetulosa sp. nov. was also collected in this area, at the upper limit of the altitudinal 
range of this last mentioned species. 

Geographical distribution within the Gaoligong Shan.— Fig. 31c. We examined a 
total of 18 specimens (13 males and 5 females) from sites on the eastern slope of the northcentral 
part of the Gaoligong Shan in Fugong County (see Type material above for exact collection data). 
All of these sites are in Core Area 3. 

Overall geographical distribution .— This species currently is known only from Fugong 
County in the northcentral part of the Gaoligong Shan region, western Yunnan Province, China. 

Genus EpapIliotrechllS Deuve and Kavanaugh, gen. nov. 

type species .— Epaphiotreckusfortipesoides sp. nov. 

Derivation of genus group name. — The genus group name (masculine) is a combination of 
two other trechine generic names, Epaphius and Trechus. 

diagnosis. — Adults of this genus (Fig. 33a) can be recognized by the following combination 
of character states: size large (BL = 4.5 to 4.7 mm), apterous; body color dark piceous with elytral 
interval 1 and lateral areas of pronotum and elytra more or less reddish; dorsum shiny, distinctly 
iridescent, dorsal surface glabrous except for isolated fixed setae typical for trechines, eyes also 
glabrous; right mandible (Fig. 16g) with premolar not fused with retinaculum but closely associat¬ 
ed with the latter [possibly representative of an intennediate state in the evolution of the “biden- 
tate” mandibular type (see discussion above for genus Trechus)), anterior tip of retinaculum 
enlarged as a distinct tooth and displaced anteriorly (but not quite as far as in members of Trechep- 
aphiopsis species); pronotum with basal angles small and rectangular; elytra elongate, oblong. 


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slightly flattened along the median suture area, striae finely impressed, crenulate or finely punc¬ 
tate, lateral striae partly effaced but striae 6 and 7 still evident; two discal setae present on interval 
3 next to stria 3, inserted near the anterior one-sixth and near mid-elytral length, respectively; 
preapical seta present, inserted next to stria 2. 

Comments.— When Ueno (1999) described Trechus (s. str.) fortipes, he noted that it was a 
“strange species similar to certain Epaphiopsis ”, with the distinct premolar on the right mandible. 
However, characters of the male aedeagus, particularly the presence of a distinct cndophallic scle- 
rite, led him to consider this species as a basal member of genus Trechus in which the plesiomor- 
phic right mandibular dentition was retained. 

Geographical distribution. This genus currently is known from only two species, both 
found only in the southern part of the Gaoligong Shan region of western Yunnan Province, China. 

Key for Identification ofAdults of Epaphiotrechus Species of the Gaoligong Shan Region 

1. Median tooth of mentum small and simple, apex not bifid; elytra with stria 5 distinctly deep¬ 
ened at the base, recurrent stria short, terminated anteriorly at a slight convexity at apex of stria 

5; preapical seta inserted equidistant from both apical and sutural margins. 

. E. fortipes (Ueno) 

Median tooth of mentum long and wide, apex bifid; elytra with stria 5 not or only faintly deep¬ 
ened at the base, recurrent stria long, terminated anteriorly at a slight convexity at apex of stria 
6; preapical seta inserted closer to sutural than to apical margin . .. E. fortipesoides sp. nov. 

Epaphiotrechusfortipes (Ueno), 1999 
(Fig. 32, 46-48) 

Trechus (.S’, str.) fortipes Ueno, 1999a: 219. 

Epaphiotrechus fortipes ( Ueno), n e w Combination 

Type material. — Holotype, a male, in NSMT. Type locality: China, Yunnan, Gaoligong 
Shan, Baoshan County, N24°577 E98°45', 2200-2500 m 

notes on type material.— We have not had an opportunity to study the holotype or any 
other specimens of this species. Features noted below are based on Ueno’s (1999a) original 
description and illustrations. 

diagnosis.— Adults of this species (see Ueno 199a, Fig. 4) can be distinguished from those 
of all other species in the region by the following combination of character states: 

Size large (BL = 4.5 mm), apterous; female holotype with unusually broad protarsomeres; 
body color dark piceous, slightly paler reddish on elytral interval 1; right mandible tridentate; men¬ 
tum with median tooth short and simple; pronotum slightly narrowed (ratio PW/PL = 1.26), with 
basal angles small and rectangular; elytra elongate, slightly flattened medially, striae finely 
impressed, medial three or four striae distinctly crenulate (due to strial punctures), more lateral stri¬ 
ae slightly effaced, but striae 6 and 7 evident, stria 5 deepened at its base; parascutellar striole pres¬ 
ent, slightly elongate; recurrent stria short, terminated near apex of stria 5; two discal setiferous 
pores present, inserted at anterior one-sixth and slightly posterior to middle, respectively, on inter¬ 
val 3 near stria 3; preapical seta present, inserted on interval 3 next to stria 2 and about equidistant 
from apical and sutural elytral margins; median lobe of male aedeagus (see Ueno 1999a, Figs. 5 
and 6) with apex, expanded, securiform in lateral view; endophallus with distinct, elongate-trian¬ 
gular sclerite. 

habitat distribution.— No members of this species were found during field collecting for 
this project, and no habitat information accompanied the type material. Ueno (1999a) suggested 
that the type locality was located “near the lower edge of the Rhododendron zone. He visited the 


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CHIN 


XizSng 


Yunnan 


MYANMAR" 

wi [y 


J'Bagchepg 


Lotn*iiq 


y-naMno 


area and collected very briefly in “ Rhododen¬ 
dron and bamboo thickets” there without suc¬ 
cess. We concur with him that such thickets 
represent the most likely habitat for members 
of this species. Ueno (1999a) reported that 
specimens of this species (which he described 
as Trechus fortipes ) were found in the same 
area as specimens of Agonotrechus yunnanus 
and Trechus indicus (which he recorded as 
Trechus macrops Jeannel). 

Geographical distribution within 
the Gaoligong Shan.— Fig. 32. This 
species is known only from a single male spec¬ 
imen from the type locality in Baoshan Coun¬ 
ty in the southern part of the Gaoligong Shan 
region. This locality is in Core Area 7. 

Overall geographical distribu¬ 
tion.— This species currently is known only 
from Baoshan County in the southern part of 
the Gaoligong Shan region, western Yunnan 
Province, China. 

Epaphiotrechus fortipesoides Deuve and 

Kavanaugh, sp. nov. 

(Figs. 16g, 33, 46-48) 

type material .— Holotype, a female, in 
IOZ, labeled: “CASENT 1039089”/ “CHINA, 
Yunnan, Tengchong County, Houqiao Town¬ 
ship, 8.5 airkm NNE of Houqiao at Gaoshi- 
dong, 2580 m, N25.39858°/E98.30533°,”/ “27 
May 2006, Stop # DHK-2006-043, D.H. 
Kavanaugh, R.L. Brett, & D.Z. Dong collec¬ 
tors”/ “IMAGE” [green label]/ “HOLOTYPE 
Epaphiotrechus fortipesoides Deuve and 
Kavanaugh, sp. nov. designated 2016” [red 
label]. 

type locality.— China, Yunnan, Teng¬ 
chong County, Houqiao Township, 8.5 airkm 
NNE of Houqiao at Gaoshidong, N25.39858°/ 
E98.30533°, 2580 m. 

Derivation of species name.— The 
species epithet , fortipesoides, is a combination 
of the species epithet, fortipes , and the Greek 
suffix, -sidtjg (transliterated into Latin as - 
oides ), meaning resembling, in reference to the 
similarity of the unique holotype female of this 
species to members of T. fortipes. 


Figure 32. Epaphiotrechus fortipes (Ueno); Map of 
locality record (red circle) in the Gaoligong Shan region. 
Scale line =100 kin. 


DEUVE ET AL.: CARABID BEETLE FAUNA OF THE GAOLIGONG MOUNTAINS 


427 


diagnosis. — Adults of this species (Fig. 33a) can be distinguished from those of all other 
species in the region by the following combination of character states: size large (BL = 4.7 mm), 
apterous; body color piceous, sutural and lateral elytral margins slightly paler, reddish; tempora 
glabrous; median tooth of mentum apically truncate, almost bifurcate; antennae long; pronotum 
slightly narrowed (ratio PW/PL=1.28), basal angles small, sharp, rectangular; elytra oblong, 
slightly flattened along median suture, striae finely impressed, finely punctate, striae 1 to 2 or 3 
more deeply impressed, more lateral striae more or less effaced but still evident, stria 5 not more 
deeply impressed at its base; recurrent stria long, deeply impressed, abruptly terminated on inter¬ 
val 7; two discal setiferous pores present, inserted at anterior one-sixth and slightly posterior to 
middle, respectively, on interval 3 near stria 3; preapical seta present, inserted 1.5 times as far from 
apical elytral margin as from sutural margin. 

description. — Size large, BL = 4.7 mm. Body color dark piceous, head, basal and lateral 
pronotal margins, and lateral and sutural margins of elytra more or less reddish, appendages paler 
reddish brown, palpi yellowish brown, dorsum smooth, shiny with pronotum and elytra moderate¬ 
ly iridescent, head faintly alutaceous due to moderately impressed isodiametric microsculpture. 

Head. Broad, slightly elongate. Eyes slightly convex, their diameter slightly less than twice 
length of tempora, the latter short but distinctly convex. Frons convex, with frontal furrows sharply 
impressed, not attenuated posterior to eyes. Two pairs of supraorbital setae present, the anterior pair 
inserted in slight foveae. Clypeus with four setae. Labrum with six setae, anterior margin distinct¬ 
ly concave. Right mandible (Fig. 16g) with premolar not fused with retinaculum but closely asso¬ 
ciated with the latter, anterior tip of retinaculum enlarged as a distinct tooth and displaced anteri¬ 
orly. Mentum and submentum not fused. Mentum with medial tooth apically truncate, about one- 
half the length of the lateral lobes. Submentum with six setae anteriorly. Genae with a single ven¬ 
tral seta one each side. Antennae moderate in length, with about 2.5 antennomeres extended pos¬ 
teriorly beyond basal pronotal margin; antennomeres 3 and 4 about equal in length, antennomere 2 
slightly shorter. 

Pronotum. Slightly narrowed (ratio PW/PL = 1.28), with greatest width anterior to middle, 
only slightly narrowed posteriorly; lateral margins rounded, straightened for a short distance pos¬ 
teriorly, then slightly sinuate just anterior to basal angles, the latter small, sharp and rectangular; 
basal margin with median region slightly and convexly projected posteriorly. Disc convex, with 
median longitudinal impression distinct, markedly deepened in median basal area, not extended 
anteriorly to apical margin; basal foveae moderate in size and depth; median basal area delimited 
laterally by a pair of short, oblique furrows, smooth except for a pair of paramedial longitudinal 
foveae. Single midlateral setae on each side inserted anterior to middle; single basolateral seta on 
each side, inserted at basal angle. 

Elytra. Elongate ovoid, slightly narrowed anteriorly, humeri rounded and only slightly evident. 
Disc moderately convex, slightly flattened in sutural area; striae finely impressed, finely punctate, 
striae 1 to 2 or 3 more deeply impressed, more lateral striae more or less effaced but still evident, 
stria 5 not more deeply impressed at its base. Parascutellar striole present, deeply impressed and 
long. Recurrent stria long, deeply impressed, abruptly terminated on interval 7 slightly anterior to 
the insertion point of preapical seta. Two discal setiferous pores present, inserted at anterior one- 
sixth and near middle, respectively, on interval 3 near stria 3. Preapical seta present, inserted 1.5 
times as far from apical elytral margin as from sutural margin on interval 3 next to stria 2. 

Legs. Short; protibiae with longitudinal furrow. 

Abdomen. Abdominal ventrites glabrous, except for a single paramedial seta on each side, and 
ventrite VII of female apically with two pairs of paramedial setae. 

Comments. — This species appears to be closely related to E. fortipes, but its members can be 


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Figure 33. Epaphiotrechus fortipesoides sp. nov.; a. Dorsal habitus (CASENT1039089). b. Map of locality record (red 
circle) for E. fortipesoides in the Gaoligong Shan region. Scale lines a = 0.5 mm, b = 100 km. 


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429 


distinguished from those of the latter by the truncate, almost bifurcate, median tooth of the men- 
tum (simple in E. fortipes members), elytral stria 5 not more deeply impressed basally (distinctly 
more deeply impressed in E. fortipes members), recurrent stria longer (shorter in E. fortipes mem¬ 
bers) and terminated on interval 7 (terminated on interval 5 in E. fortipes members), and the preapi- 
cal seta inserted 1.5 times as far from apical elytral margin as from sutural margin (inserted about 
equidistant from apical and sutural margins in E. fortipes members). 

h abitat distribution. — The unique holotype female of this species was found on the 
ground under a stone or log in a managed conifer forest along a roadcut at an elevation of 2580 m. 
No other trechines were found in this area. 

Geographical distribution within the Gaoligong Shan.— Fig. 33b. This species is 
known only from a single female specimen from the type locality in western Tengchong County in 
the southern part of the Gaoligong Shan region. This locality is in Core Area 6. 

overall geographical distribution. — This species currently is known only from Teng¬ 
chong County in the southern part of the Gaoligong Shan region, western Yunnan Province, China. 

Genus TrechepctphicutKl Deuve and Kavanaugh, gen. nov. 

type species. — Trechepaphiama gaoligong sp. nov. 

derivation of genus group name. — The genus group name (masculine) is a combination 
of two other trechine genus group names, Trechus and Epaphiama. 

diagnosis. — Adults of this genus (Fig. 34a) can be recognized by the following combination 
of character states: size large (BL = 4.2 mm), apterous; body color reddish brown, dorsum shiny, 
slightly iridescent, dorsal surface glabrous except for isolated fixed setae typical for trechines, eyes 
also glabrous, small, only slightly convex; tempora glabrous; right mandible (Fig. 16h) with pre¬ 
molar tooth fused with retinaculum, anterior tip of retinaculum enlarged to form a distinct tooth 
and displaced distally, posterior tip of retinaculum reduced, nearly flattened; mention and submen- 
tum not fused; mentum with median tooth simple with apex rounded; submentum [probably] with 
six setae [only five setae visible in unique female holotype of type species]; pronotum slightly con¬ 
vex, disc glabrous, basal foveae only slightly distinct, median basal area delimited laterally by 
oblique furrows, basal margin slightly curved medially, slightly sinuate and oblique laterally; ely¬ 
tra broadly rounded, more narrowed apically than basally, humeri broad, disc convex with striae 
finely impressed, slightly punctate, lateral striae attenuated, recurrent stria deeply impressed, its 
anterior end straight, almost continuous anteriorly with the apex of stria 5; no discal setae present, 
preapical seta present, inserted next to stria 2. 

Comments. — This genus, known from only a single female specimen, belongs to the “Epa- 
phiopsis Complex” of genera as evidenced by the dentition of the mandibles. Its members can be 
distinguished from those of other genera of this lineage by the absence of elytral discal setae, which 
is exceptional among trechines, and by their markedly convex, dorsally inflated elytra, imparting 
an appearance similar to that of Epaphiama Jeannel (1962) members — short and convex. In con¬ 
trast, Trechepaphiopsis members are similar in habitus to small Epaphiopsis members while those 
of Epaphiotrechus are more similar in habitus to large Trechus, members. Abetter understanding of 
phylogenetic affinities of this new genus must await discovery of a male and study of its genital 
structure and/or comparative molecular study. 

Geographical distribution.— This genus currently is known only from the type species, 
which is known only from the southcentral part of the Gaoligong Shan region of western Yunnan 
Province, China. 


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Trechepaphiama gcioligong Deuve and Kavanaugh, sp. nov. 

(Figs. 16h, 34, 37a, 46-48) 

t V p E M A T E RIA LHolotype, a female, in IOZ, labeled: “CASENT 1001926”/ “CHINA, Yun¬ 
nan Province, Gaoligongshan Mountains, Nujiang Prefecture, 9 [actually 9.3] km ESE of Pianma, 
25°59.6’N/ 98°37.6 , E.”/ “2450 [actually 2460-2470] m, 15-18 October 1998, Stop #98-118D D.H. 
Kavanaugh, C.E. Griswold, C. Ferraris & C.-L. Long collectors”/ “IMAGE” [green label]/ 
“HOLOTYPE Trechepaphiama gaoligong Deuve & Kavanaugh, sp. nov. designated 2016” [red 
label]. 

Type locality. — China, Yunnan, Gaoligong Shan, Lushui County, 9.3 km ESE of Pianma, 
N25.99363°/ E98.66651 0 , 2460-2470 m. 

derivation of species name. — The species epithet, gaoligong , is a noun in apposition, 
derived from the name of the mountain range, the Gaoligong Shan, in which the holotype was col¬ 
lected. 

diagnosis. — Adults of this species (Fig. 34a) can be distinguished from those of all other 
species in the region by the following combination of character states: size large (BL = 4.2 mm), 
apterous; body color reddish brown, dorsum shiny; eyes reduced, their diameter about equal to 
length of tempora, the latter distinctly convex and glabrous; pronotum moderately transverse (ratio 
PW/PL=1.26), basal angles subrectangular; elytra broad, markedly convex, slightly inflated, 
humeri broadly rounded, striae 1 to 5 deeply impressed and slightly punctate, more lateral striae 
more or less effaced, intervals 1 to 4 slightly convex, more lateral intervals flat; recurrent stria ante¬ 
riorly nearly continuous with posterior end of stria 5; discal setiferous pores absent; preapical seta 
present, inserted on interval 3 next to stria 2. 

description .— Size large, BL 4.2 mm. Body color reddish brown, appendages coneolorous, 
except palpi yellowish brown, dorsum smooth, shiny, pronotum and elytra very faintly iridescent, 
head faintly alutaceous due to moderately impressed isodiametric microsculpture 

Head. Moderate in size, slightly broad; eyes small, only slightly projected laterally, their diam¬ 
eter about equal to length of tempora, the latter short, convex and glabrous. Frons only slightly con¬ 
vex, with frontal furrows deeply impressed but less so posterior to hind margins of eyes. Two pairs 
of supraorbital setae present, the anterior pair inserted in distinct foveae. Clypcus with four setae. 
Labrum with six setae, anterior margin moderately concave. Right mandible (Fig. 16h) with pre¬ 
molar tooth fused with retinaculum, anterior tip of retinaculum enlarged to form a distinct tooth 
and displaced distally, posterior tip of retinaculum reduced, nearly flattened. Left mandible with 
only a relatively small, feebly trifid tooth. Mentum and submentum not fused. Mentum with medi¬ 
al tooth broad, apically rounded, about one-half the length of the lateral lobes. Submentum with 
four setae anteriorly. Gula broad. Genae with a single ventral seta one each side. Antennae rather 
short, only about two antennomeres extended posteriorly beyond basal pronotal margin; anten- 
nomeres 2 and 4 about equal in length, antennomere 3 slightly longer. 

Pronotum. Slightly narrow (ratio PW/PL = 1.26), with greatest width anterior to middle, only 
slightly narrowed posteriorly; lateral margins rounded, straightened for a short distance posterior¬ 
ly, then very slightly sinuate just anterior to basal angles, the latter blunt and subrectangular, very 
slightly and bluntly projected laterally; basal margin with median region slightly and convexly pro¬ 
jected posteriorly. Disc distinctly convex, with median longitudinal impression very finely 
impressed, superficial, effaced in the median anterior area but extended to basal margin; basal 
foveae distinct but shallow; median basal area small, delimited laterally by a pair of oblique fur¬ 
rows, smooth except for several short longitudinal depressions; basal margin slightly bisinuate, 
slightly convex medially; lateral borders slender, gradually widened posteriorly, faintly reflexed 


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431 


Yunnan 


MYANMAR 


Tcngchong 


LftflgpWig 


Uanghp 


Longing 


Figure 34. Trechepaphiama gaoligong sp. nov.; a. Dorsal habitus (CASENT1001926). b. Map of locality record (red 
circle) for T. gaoligong in the Gaoligong Shan region. Scale lines a = 0.5 mm, b = 100 km. 


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dorsally, lateral grooves widened near base. Single midlateral setae on each side inserted anterior 
to middle; single basolateral seta on each side, inserted at basal angle. 

Elytra. Broad, ovoid, slightly more narrowed posteriorly than anteriorly, markedly convex, 
slightly inflated; humeri broadly rounded; striae 1 to 5 deeply impressed and slightly punctate, 
more lateral striae more or less effaced, barely evident; intervals 1 to 4 slightly convex, more lat¬ 
eral intervals flat; basal margination terminated medially at origin of stria 5. Parascutellar striole 
present, short but distinct. Recurrent stria anteriorly nearly continuous with posterior end of stria 
5. Discal setiferous pores absent. Preapical seta present, inserted on interval 3 next to stria 2. 
Umbilicate setal series with setae of humeral group equidistance from each other and setae of medi¬ 
an group inserted slightly posterior to middle. 

Legs. Short; protibiae with longitudinal furrow. 

Abdomen. Abdominal ventrites glabrous, except for a single paramedial seta on each side, and 
ventrite VII of female apically with two pairs of paramedial setae. 

h abitat distribution .— The unique holotype female of this species was collected by sifting 
leaf litter taken on sandy substrate on a secondary floodplain covered by a broadleaf forest canopy 
at an elevation of 2460 m (Fig. 37a). Specimens of Trechepaphiopsis uniporosa and T unisetosa 
were collected in the same sifted litter samples at this site. 

Geographical distribution within the Gaoligong Shan.— Fig. 34b. This Species is 
known only from a single female specimen from the type locality in western Lushui County on the 
western slope of the southcentral part of the Gaoligong Shan region. This locality is in Core 
Area 4. 

Overall geographical distribution .— This species currently is known only from Lushui 
County in the southcentral part of the Gaoligong Shan region, western Yunnan Province, China. 

Discussion 

The Gaoligong Shan region is at the very heart of one of the world’s biodiversity hotspots 
(Myers et al. 2000), where faunal elements from the Palearctic and Oriental Regions meet. Adding 
to this diversity is a distinct regional Chinese element, probably of mixed Palearctic/Oriental ori¬ 
gin (Deuve 2013b), which either became isolated and evolved independently within the region or 
has been replaced elsewhere by present-day Palearctic and/or Oriental elements. Perhaps the most 
well-known representative of this element is the giant panda, Ailuropoda melanoleuca David 
(1869); but there are also several endemic carabid generic and subgeneric representatives as well 
(e.g., the nebriine genus Archcistes Jedlicika (1935) and the trechine genus Eocnides ). 

The trechine carabid fauna of the region is hyperdiverse, and the fauna of the Gaoligong Shan 
region itself is exceptionally species-rich. Of the 29 trechine species recorded from the latter, 
including the 19 species reported here as new, 25 are known from nowhere else. This pattern con¬ 
trasts dramatically with that found among the zabrines of the region (Kavanaugh et al. 2014), all 
13 of which have been recorded from outside the Gaoligong Shan region and none of which were 
new to science. In the following sections, we discuss broader geographical distribution patterns of 
the four trechine species known from outside the region and of the supraspecific taxa of which 
Gaoligong Shan trechines are members. We also examine geographical and altitudinal distribution 
patterns and patterns of syntopy (co-occurrence in the same habitat) among species in the region¬ 
al fauna. 

broad geographical distribution patterns. — The overall geographical ranges of the 
four trechine species known to occur both inside and outside of the Gaoligong Shan region are 
graphically approximated, superimposed on one another, in Fig. 45. Among the geographical 


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433 


ranges of these species, three general range patterns are apparent. The first pattern (1) is shown by 
one species, Perileptus imaicus , with a geographical range that includes only a narrow swath along 
the southern base of the Himalayan Range from Himachal Pradesh, India in the west to the 
Gaoligong Shan region, where it reaches its eastern distributional limit. Among the zabrines, 
Amara elegantula Tschitscherine shares this same pattern but in higher elevation habitats 
(Kavanaugh et al. 2014). The second (2) pattern is shown by two species, Agonotrechus wuyipeng 
and Eocni des fra gills , both of which have a geographical range that extends from central or north¬ 
ern Sichuan, respectively, southwest to the northern half of the Gaoligong Shan region, where they 
reach their western distributional limit. Finally, the fourth species, Trechus indicus, shows the third 
pattern (3), which is a combination of the first two. The range of T. indicus extends from eastern 
Afghanistan eastward along the southern edge of Himalayan range and the Qinghai-Xizang 
(Tibetan) Plateau to northcentral Sichuan, with its southern limit in the mountains of the Gaoligong 
Shan region. Several zabrine species, including Amara sikkimensis Andrewes, A. chalciope 
(Bates), A. dissimilis Tschitscherine, A, latithorax Baliani, and A. birmana Baliani share this same 
pattern (Kavanaugh et al. 2014, Fig. 27 ), although with varied eastern and western extents. Deuve 
(1997, 2013b) recognized what he called subzones within both the southern Palearctic and north¬ 
ern Oriental Regions based mainly on his biogeographic analysis of the Carabus fauna of China 
and adjacent areas. Our general ranges patterns (1) to (3) correspond well to the southern parts of 
his Sichuano-Tibetan subzone (“Sous-zone Sichuano-Tibetame”, subzone III in Deuve 2013b, Fig. 
13); and pattern (2) is similar to his Yunnan Plateau subregion (“Plateau du Yunnan”, subzone Ilia 
in the same figure). 

It is likely no coincidence that all four of the trechine species with known ranges extended 
beyond the Gaoligong Shan region have adults that are fully-winged. The ability to fly undoubt¬ 
edly supports the maintenance of larger occupied ranges as well as greater potential for dispersal 
to new areas. Within the trechine fauna of the study area, four additional species have fully-winged 
adults: Perileptus pusilloides, Agonotrechus Jugongensis , A. xiaoheishan , and A, yunnanus. 
Although all of these species are currently known only from the study area, the discoveiy of one 
or more of them in additional, adjacent areas is possible or even likely in the future. However, it 
would be unexpected to discover populations of any of the remaining 21 species, all of which have 
flightless adults, outside the Gaoligong Shan region. 

With such a large proportion of the trechine fauna of the region known from nowhere else (25 
of 29 known species [86%] and four of eight genera [50%, plus one additional subgenus] ), a look 
at the overall distributions of the genera or subgenera to which these trechines belong may provide 
a broader geographic context for understanding the composition of fauna. Perileptus is widely dis¬ 
tributed in tropical to temperate portions of all continents in the Eastern Hemisphere and in the 
Caribbean portion of the Neotropical Region, probably introduced into the latter from Africa with 
commercial trade. Agonotrechus is restricted to the southern portion of the Sichuano-Tibetan sub¬ 
zone and the Subtropical China subzone (“Sous-zone dc Chine subtropicale”, subzone V in Deuve 
2014b, Fig. 13) including Japan. Eocnides is restricted to the southcentral portion of the Sichuano- 
Tibetan subzone and Queinnectrechus to the Yunnan Plateau subregion of that subzone. Trechus, 
with its present taxonomic inclusiveness, is Holarctic in distribution, and species in the study area 
likely have their closest relatives to the north. Three of the remaining genera, Trechepaphiopsis, 
Epaphiotrechus, and Trechepaphiama, are members of the Epaphiopsis complex of genera, which 
mainly occupy the Subtropical China subzone. The phyletic affinity of Minutotrechus remains 
unclear based on morphological features. As noted above, it is likely related to either Uenoites, and 
therefore has a Sichuano-Tibetan affinity, or to Hubeitrechus, and hence has a Subtropical China 
affinity. 


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Kavanaugh et al. (2014) suggested that the Gaoligong Shan region may have been an area of 
differentiation, speciation and origin of montane elements from which, rather than to which, at least 
some of the species that now range more broadly subsequently spread. This hypothesis was based 
on meager evidence provided by the zabrine fauna of the Gaoligong Shan, but also on geologic evi¬ 
dence. The Hengduan Mountains date their origins to the late Mesozoic, whereas the uplift of the 
Himalayan Ranges and Qinghai-Xizang Plateau began later, in the early Cenozoic (Chaplin 2005). 
Hence the biota of the Gaoligong Shan region probably predates that of these other areas as well. 
The trechine fauna of the region supports this hypothesis even more strongly. The occurrence of 
four precinctive genera and one apparently precinctive subgenus of small, flightless beetles, two of 
which are represented by two and seven species, respectively, suggests differentiation and diversi¬ 
fication within the region. The occurrence of another nine likely precinctive species in other gen¬ 
era suggests speciation in situ. Finally, even the four species with ranges extended beyond the 
Gaoligong Shan region all have those ranges either centered on the region of anchored there. As 
with the zabrines, a better understanding of phylogenetic relationships among the Gaoligong Shan, 
Eurasian and Oriental trechine species and genera is required in order to test this hypothesis, and 
such an analysis has not yet been undertaken. 

Regional geographical and altitudinal distribution patterns. — Within the 

Gaoligong Shan study area, most of the trechine species represented are narrowly distributed, both 
geographically and altitudinally. This is not surprising given the high percentage (72%) of species 
with flightless adults and their preferences for moist, undisturbed habitats. Such areas are restrict¬ 
ed within the region mainly to remaining forested areas at low to middle elevations and alpine 
meadows, moist tundra, stable talus slopes, stream edges and bamboo and Rhododendron thickets 
at higher elevations. Many such areas in the region are separated from each other by deep valleys 
and, increasingly, by disturbance associated with agriculture and human habitation. While human 
disturbance is relatively recent, topographic diversity of the region has been developing since the 
Miocene (Chaplin 2005). 

The chart in Fig. 46 summarizes the recorded regional distributions of the species with respect 
to our project-designated Core Areas (see Fig. 3); and the recorded altitudinal ranges for each 
species are shown in Fig. 47. These charts clearly demonstrate the relatively narrow geographical 
and altitudinal ranges of most of the trechine species occurring in the region. This is especially 
apparent from comparisons with the ranges of zabrine species in the same area (see Kavanaugh et 
al. 2014, Figs. 28 and 29). Among the 13 zabrine species in the fauna, one is recorded from all 7 
Core Areas, one from 6 Core Areas (all except 5) and three from four or five Core areas; and most 
of these species are likely to occur in all seven Core Areas. In contrast, only one species, Trechus 
indicus , is recorded from as many as four Core Areas (1, 3, 6 and 7) and also has a relatively broad 
known altitudinal range (1230 to 2486 m). Given that it is recorded from the northernmost and 
southernmost core areas, as well as from both eastern and western slopes of the Gaoligong Shan, 
it is likely to be found in additional, if not all, core areas with further sampling. Only one additional 
species, Perileptus imaicus, is recorded from as many as three Core Areas (2, 6 and 7) and also has 
a broad known altitudinal range (from 680 to 2030 m). Like T. indicus, it is recorded from both 
northernmost and southernmost core areas and from both slopes of the mountain range, so it is like¬ 
ly to be more widespread in the region than is presently known, at low to middle elevations. Five 
additional species are recorded from as many as two core areas: Perileptuspusilloides (Core Areas 
6 and 7, known altitudinal range from 680 to 1105 m); Agonotrechus fugongensis (Core Areas 2 
and 3, known altitudinal range from 2300 to 2530 m); A. wuyipeng (core areas 2 and 3, known alti¬ 
tudinal range from 2687 to 2770 m); Queinnectrechus balli (Core Areas 1 and 2, known altitudinal 
range from 3300 to 3750 m); and Trechepaphiopsis unisetosci (Core Areas 4 and 5, known altitu- 


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435 


dinal range from 2460 to 3150 m). For each of these species, Core Areas known to be occupied are 
immediately adjacent. A sixth species, Trechus luzhangensis is recorded only from the pass divid¬ 
ing Core Areas 4 and 5 (at 3150 m) and so has been attributed to both Core Areas. Among these 
six species, only P. pusilloides, A. fugonensis and A. wuyipeng have frilly-winged adults and are 
likely to have ranges greater than are presently known. The other three species, 0. balli, T. unise- 
tosa, and T. luzhangensis , have flightless adults and are not likely to have ranges outside their 
presently known Core Areas. The remaining 21 species are presently known only from a single 
Core Area and have altitudinal ranges of varied breadth. Among these, only Agonotrechus xiaoheis- 
han. A, yunnanus, and Eocnidesfragilis have fully-winged adults and therefore are likely to be sig¬ 
nificantly more widespread within the region than presently known. 

A comparison of recorded diversity for trechine species among the seven Core Areas (Fig. 46) 
shows that all of them are occupied by at least two species, with highest diversity in Core Area 2 
(with 10 of the 29 species), second highest in Core Areas 3 and 6 (each with 8 species) and lowest 
recorded diversity in Core Areas 1 and 5, which are the Core Areas least extensively sampled and 
perhaps most heavily impacted by human disturbance, respectively. In contrast, the zabrines were 
found to be most diverse in Core Area 6, second most diverse in Core Area 2, and least diverse in 
Core Area 4 (Kavanaugh et al. 2014). At present, Core Areas 1 and 2 (West/East versants in the 
northernmost part of the region) are known to uniquely share only one species, Queinnectrechus 
balli, adults of which are flightless. Further sampling is likely to confirm the occurrence in Core 
Area 1 of additional species presently known only from Core Area 2, particularly those inhabiting 
high elevation habitats there. Core Areas 2 and 3 (areas adjacent North/South on the eastern slope 
of the Gaoligong Shan) share two species uniquely, Agonotrechus fugongensis and A. wuyipeng, 
both of which have fully-winged adults. Unfortunately, the China/Myanmar border forms the west¬ 
ern limit of Core Area 3 and also of our study area (see Material and Methods section above), so 
the fauna of the western versant of the Gaoligong Shan in the northcentral part of the range remains 
unknown. Core Areas 4 and 5 (West/East versants in the southcentral part of the region) uniquely 
share two species, Trechus luzhangensis and Trechepaphiopsis unisetosa, both with flightless 
adults. 

Five of the seven Core Areas are inhabited by at least one species recorded from no other Core 
Area. Six species are uniquely recorded from Core Area 2, the northernmost eastern versant of the 
region. All of these except Eocnides fragilis have flightless adults. Five species are uniquely 
recorded from Core Areas 3 and 6, respectively; and all of these except Agonotrechus xiaoheishan 
(in Core Area 6) have flightless adults. Three species are uniquely recorded from Core Area 7, the 
southernmost eastern versant of the region, and two from Core Area 4. Among these, all except 
Agonotrechus yunnanus have flightless adults. 

If we ignore the two most widespread species, Trechus indicus and Perileptus imaicus, it is 
clear that five different Core Areas or combinations thereof have distinctive trechine assemblages. 
(1) The northermost part of the Gaoligong Shan region (Core Areas 1 and 2 together) is the most 
diverse and distinctive region, with seven species (six of which have only flightless adults) unique 
to it. Only Agonotrechus fugongensis and A. wuyipeng, both with fully-winged adults, arc shared 
with any other area (adjacent Core Area 3). (2) Core Area 3, the north central part of the region, 
also has a distinctive trechine fauna, with five species (all with flightless members) unique to it 
(plus the two species shared with Core Area 2 as just noted). (3) The southcentral portion of the 
region (Core Areas 4 and 5 together) also has a distinct trechine assemblage of four species, all with 
flightless members, unique to it. Trechus indicus, Perileptus imaicus and P pusilloides, all with 
frilly-winged adults, are recorded from Core Areas 6 and 7, but otherwise the trechine assemblages 
of these areas are distinctive. (4) Four species are unique to Core Area 6, all except Agonotrechus 


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xiaoheishan with flightless adults; and (5) three species are uniquely recorded from Core Area 7, 
with only A. yunnanus having fully-winged adults. 

With respect to the distributions of genera within the Gaoligongshan region, a broader picture 
emerges. Both Perileptus and Agonotrechus range from Core Area 2 in the northeast to Core Areas 
6 and 7 in the south, with gaps in records from the middle part of region. Minutotrechus is record¬ 
ed only from Core Area 6 in the southwestern part of the region and Eocnides only from Core Area 
2 in the northernmost part. Queinnectrechus also appears restricted to Core Areas 1 and 2 in the 
northernmost part. Although Trechus indicus is widespread both within and beyond the region, the 
remaining species of Trechus , which we have informally labeled the “ qiqiensis group” appear 
restricted to the northern and central parts of the region (Core Areas 2, 3,4, and 5). Trechepaphiop- 
sis is widely distributed in the region, from northernmost to southernmost Core Areas but most 
diverse in the northcentral and southcentral areas (Core Areas 3 and 4). Epaphiotrechus is record¬ 
ed only from Core Areas 3 and 7, non-adjaccnt northcentral and southern parts of the region on the 
east slope of the Gaoligong Shan, and Trechepaphiama is known only from Core Area 4 on the 
western slope in the southcentral part of the mountain range. In general, genera that have affinities 
with faunas of the Palearctic Region (e.g., Trechus species) and with the Sichuano-Tibetan subzone 
in particular (e.g., Queinnectrechus and Eocnides ) mainly occupy northern and central parts of the 
region, whereas genera with affinities with the Oriental Region and with the Subtropical China sub¬ 
zone of the region in particular, mainly occupy the central and southern parts of the region. How¬ 
ever, the adaptations of individual species in each of these affinity classes to diverse local topo¬ 
graphic differences, which foster geographical and altitudinal isolation in the region, and differ¬ 
ences in the respective flight capabilities of their members, which support broader or narrower dis¬ 
tributional ranges, serve to partially obscure this overall pattern. 

With respect to the altitudinal distribution of the trechine fauna of the study area (Fig. 47), sev¬ 
eral points can be made. Highest diversity is concentrated in a broad zone between about 2250 m 
and 3750 m, with 27 of the 29 species occurring within this zone. Only two species, Perileptus 
imaicus and P pusilloides occur below 1000 m in the study area; and only two more, Eocnides 
fragilis and Trechus indicus , occur below 2000 m. As noted for each elsewhere above, all of these 
species have fully-winged adults. Eight species ( Queinnectrechus griswoldi , Q. gongshanicus , Q. 
halli , Trechus shiyueUang , T qiqiensis, T. gongshanensis, T. shibalicus , and Trechepaphiopsis 
monochaeta ) occur above 3500 m, but only O. gongshanicus appears to be restricted to that eleva¬ 
tion or above. All of these species have flightless adults. Compared with the distributions of 
zabrines in the region, the trechines occupy relatively restricted altitudinal ranges. The average dif¬ 
ference between highest and lowest elevations recorded for each of the 24 trechine species for 
which more than one elevational record is known is 500 m (range = 20 to 1350 m), whereas the 
average difference for zabrine species in the region is 1193 m (range = 225 to 2111 m), more than 
double that for trechine species (Kavanaugh et al. 2014). Also, six of 13 zabrine species occupy 
greater altitudinal ranges than any trechine species in the same area. 

In the preceding discussion, we have repeatedly noted relationships between the extent of geo¬ 
graphical and/or altitudinal ranges and flight capability. Why? As Schmidt et al. (2016) noted, 
“... considering the low dispersal ability of the tiny wingless beetles within their mountainous 
environment, veiy small distributional areas can be expected for each of the species and therefore 
the contemporary existence of a large number of closely related allopatric Trechus species.” This 
certainly describes very well the trechine fauna of the Gaoligong Shan region, and also explains 
how it differs so markedly from the zabrine fauna of the same region. 

Syntopy of spec ie s in the regional fa una .— Records of the co-occurrence of different tre¬ 
chine species at the same site and in the same habitat (i.e., syntopic) within the study area are sum- 


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437 


marized in Fig. 48. Syntopy appears to be relatively rare among trechines of the region, especially 
compared with the zabrines of the region. Four of 13 zabrine species have been recorded syntopic 
with eight other zabrine species, three other species syntopic with seven other species, and all 13 
species syntopic with at least one other species (Kavanaugh et al. 2014). In contrast, seven of the 
29 trechine species have not been found syntopic with any other species. Only one trechine species, 
Queinnectrechus griswoldi, has been found syntopic with five other species ( Q . balli, Q. gongshan- 
icus, Trecluis qiqiensis , T. gongshanensis, and Trechepaphiopsis monochaeta ), but at no single site 
with more than four of those species. Queinnectrechus balli and T. gongshanensis have been found 
syntopic with the same cadre of species except T. monochaeta , and T. qiqiensis has been found syn¬ 
topic with all of the above except Q. gongshanicus. All the above records of syntopy are from the 
northernmost part of the study area, in Core Area 2. Trechus shihalicus has been found syntopic 
with four other species ( Trechus shiyueliang, T. pseudoqiqiensis, Trechepaphiopsis unisetulosa and 
T. unipilosa ) in Core Area 3. Trechus shiyueliang has been found syntopic with the same cadre of 
species except T. pseudoqiqiensis in the same area. Trechus indicus has also been found syntopic 
with three other species (with Agonotrechus xiaoheishan in Core Area 6 and with A. yunnanus and 
Epaphiotrechus fortipes in Core Area 7). 

Given the low level of syntopy among the trechines of the area overall, it is perhaps surpris¬ 
ing to find it among congeneric species in several genera. The two Perileptus species in the fauna 
are syntopic in Core Areas 6 and 7. All three of the Queinnectrechus species are syntopic in Core 
Area 2. Among Trechus species in the region, T. qiqiensis and T. gongshanensis have been found 
syntopic in Core Area 2 and T. pseudoqiqiensis , T. shihalicus and T. shiyueliang in Core Area 3. 
Among the species of Trechepaphiopsis , only T. unisetulosa and T. unipilosa have been found syn¬ 
topic, also in Core Area 3. For several of these syntopic pairs of congeneric species we have noted 
evident differences in body size between the pair members. The ranges in size (BL) of Perileptus 
imaicus and P. pusilloides adults are 2.6 to 2.8 mm and 2.3 mm, respectively. Those of the two 
species of Queinnectrechus (s. str .), Q. griswoldi and Q. gongshanicus, are 4.3 to 4.8 mm and 3.5 
to 3.8 mm, respectively. Those of Trechus qiqiensis and T. gongshanensis are 4.0 to 4.2 mm and 
3.3 to 3.5 mm, respectively; and those of Trechus pseudoqiqiensis and T. shihalicus are 4.0 mm and 
3.3 to 3.5 mm, respectively (T. shiyueliang [BL = 4.0 to 4.2] appeal's to be more distantly related 
based on key morphological features). Finally, those of Trechepaphiopsis unisetulosa and T. unip¬ 
ilosa are 3.3 to 35 mm and 2.7 to 3.1 mm, respectively. Sokolov and Kavanaugh (2014) found sim¬ 
ilar size differences among closely related, small, flightless, litter-dwelling and syntopic anillines 
of the genus Geocharidius in Nuclear Centeral America. Just what role such size differences may 
play in facilitating syntopy is unclear, but perhaps they allow members of the different species to 
share slightly different microspaces and/or food (prey) in their shared habitat without or with 
reduced competition. 

Finally, we cannot resist commenting on the Agonotrechus fauna of the region. Four species 
occur there, all with fully-winged adults, and none of them has yet been recorded syntopic with any 
another, although A. fugongensis and A. wuyipeng have been collected in adjacent habitats in the 
southern part of Core Area 2. Unlike members of both Perileptus species, which are syntopic in the 
southern part of the region, those of all the Agonotrechus species apparently only occupy habitats 
above 2000 in in elevation (range = 2000 to 2770 m). Just how, when, or even if these beetles use 
their wings remains unknown. 


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Figure 35. Photographs of habitats for trechine species in the Gaoligong Shan region, a. Gongshan County, 
Bingzhongluo Township, Xiao Shangla He at Shuangla Village, elevation 1550 m; habitat in which specimens of Perilep- 
tus imaicus Jeannel were collected, b. Gongshan County, Qigi He, 9.9 airkm W of Cikai, elevation 2000 m; habitat in which 
specimens of Agonotrechus fugongensis sp. nov. were collected. Photos by David H. Kavanaugh. 


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Figure 36. Photographs of habitats for trechine species in the Gaoligong Shan region, a. Fugong County, Shiyueliang 
Township, 1 km E of Shibali on Shibali Road, elevation 2400 m; habitat in which specimens of Agonotrechus fugongensis 
sp. nov. were collected, b. Gongshan County, Cikai Township, Qiqi Trail at No. 12 Bridge Camp, elevation 2770 m; habi¬ 
tat in which specimens of Agonotrechus wuyipeng Deuve were collected. Photos by David H. Kavanaugh. 


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Figure 37. Photographs of habitats for trechine species in the Gaoligong Shan region, a. Lushui County, Pianma 
Township ,9.3 km ENE of Pianma along road to Lushui at Changyan He, elevation 2460 m; habitat in which specimens of 
Trechepaphiopsis unisetosa (Deuve), T. uniporosa sp. nov., and Trechepaphiama gaoligong sp. nov. were collected, b. 
Gongshan County, Cikai Township, slope S of of Heipu Yakou, elevation 3370 m; habitat in which specimens of Quein- 
nectrechus griswoldi sp. nov., Q. balli sp. nov., and Trechus qiqiensis sp. nov. were collected. Photos by David H. 
Kavanaugh. 


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Figure 38. Photographs of habitats for trechine species in the Gaoligong Shan region, a. Gongshan County, 
Bingzhongluo Township, SW slope of Kawakarpu Shan 0.3 km SW of Chukuai Lake, elevation 3750 m; habitat in which 
specimens of Queinnectrechus griswoldi sp. nov. and Trechus gongshanicus sp. nov. were collected, b. same area, but habi¬ 
tat in which specimens of Queinnectrechus griswoldi sp. nov. and Trechepaphiopsis monochaeta sp. nov. were collected. 
Photos by David H. Kavanaugh. 


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Figure 39. Photographs of habitats for trechine species in the Gaoligong Shan region, a. Gongshan County, 
Bingzhongluo Township, SW slope of Kawakarpu Shan, 0.3 km NNE of Chukuai Lake, elevation 3745 m; habitat in which 
specimens of Queinnectrechus gongshanicus sp. nov., Q. balli sp. nov. and Trechus gongshanensis sp. nov. were collected, 
b. Gongshan County, Bingzhongluo Township, SW slope of Kawakarpu Shan on slope NE of Chukuai Lake, elevation 3950 
m; habitat in which specimens of Queinnectrechus griswoldi sp. nov. and Trechus gongshanensis sp. nov. were collected. 
Photos by David H. Kavanaugh. 


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Figure 40. Photographs of habitats for trechine species in the Gaoligong Shan region, a. Gongshan County, 21 air km 
W of Cikai on east slope of Qiqi/Dulong divide, elevation 3600 m; habitat in which specimens of Queinnectrechus balli 
sp. nov. and Trechus gongshanensis sp. nov. were collected, b. Fugong County, Shiyueliang Township, 8.5 km above 
Shibali on Shibali Road, elevation 3100 m; habitat in which specimens of Trechus shiyueliang sp. nov., T. shibalicus sp. 
nov., Trechepaphiopsis unisetulosa sp. nov. and T. unipilosa sp. nov. were collected. Photos by David H. Kavanaugh. 


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Figure 41. Photographs of habitats for trechine species in the Gaoligong Shan region, a. Gongshan County, 
Dulongjiang Township, Dulong Jiang at Elideng village, elevation 1640 m; habitat in which specimens of Trechus indicus 
Putzeys were collected, b. Tengchong County, Mingguang Township, slope SW of Eighth Boundary Post Pass, elevation 
2887 m; habitat in which specimens of Trechus mingguangensis sp. nov. were collected. Photos by David H. Kavanaugh. 


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Figure 42. Photographs of habitats for trechine species in the Gaoligong Shan region, a. Gongshan County, Cikai 
Township, Qiqi Trail at No. 12 Bridge Camp area, elevation 2775 m; habitat in which specimens of Trechus qiqiensis sp. 
nov. and Trechepaphiopsis monochaeta sp. nov. were collected, b. Fugong County, Shiyueliang Township, 11.5 km above 
Shibali on Shibali Road, elevation 3290 m; habitat in which the holotype of Trechus pseudoqiqiensis sp. nov. was collect¬ 
ed. Photos by David H. Kavanaugh. 


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Figure 43. Photographs of habitats for trechine species in the Gaoligong Shan region, a. Lushui County, Luzhang 
Township, Piana Road 0.1 kin E of Fengxue Yakou, elevation 3150 m; habitat in which specimens of Trechus liizhangen- 
sis sp. nov. were collected, b. Lushui County, Luzhang Township, 100 m S of Fengxue Yakou on east side of pass, eleva¬ 
tion 3150 m; habitat in which specimens of Trechepaphiopsis unisetosa (Deuve) were collected. Photos by David H. 
Kavanaugh. 


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Figure 44. Photograph of habitat for trechine species in the Gaoligong Shan region. Longling County, Longjiang 
Township, Xiaoheishan Forest Reserve, elevation 2020 m; habitat in which specimens of Agonotrechus xiaoheishan sp. 
nov. and Trechus indicus Putzeys were collected. Photo by David H. Kavanaugh. 


Figure 45. Map showing approximate overall known geographical distributions of species occurring in the Gaoligong 
Shan region as well as outside the region. 1 = Peripleptus imaicus Jeannel. 2 = Agonotrechus wuyipeng Deuve. 3 = Eoc- 
nides fragilis Ueno. 4 = Trechus indicus Putzeys. Scale line = 500 km. 


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species 


Core Area 

2 3 4 5 6 7 


Periteptus imaicus 
Perileptus pusilloides 
Agonotrechus fugongensis 
Agonotrechus wuyipeng 
Agonotrechus xiaoheishan 
Agonotrechus yunnanus 
Minutotrechus minutus 
Queinnectrechus batti 
Queinnectrechus griswoldi 
Queinnectrechus gongshanicus 
Eocnides fragilis 
Trechus in die us 
Trechus shiyueliang 
Trechus mingguangensis 
Trechus qiqiensis 
Trechus pseudoqiqiensis 
Trechus iuzhangensis 
Trechus gongshanensis 
Trechus shibaiicus 
Trechepaphiopsis asetosa 
Trechepaphiopsis unisetigera 
Trechepaphiopsis unisetosa 
Trechepaphiopsis uniporosa 
Trechepaphiopsis unisetulosa 
Trechepaphiopsis monochaeta 
Trechepaphiopsis unipilosa 
Epaphiotrechus fortipes 
Epaphiotrechus fortipesoides 
Trechepaphiama gaoligong 


X 

X X 
X X 


X 


X X 
X X 


X 

X 

X 


X X 
X 


X 


X 


X 


X 


Figure 46. Chart showing the representation of trechine species in project-designated Core Areas (see Fig. 3) in the 
Gaoligong Shan region. 


DEUVE ET AL.: CARABID BEETLE FAUNA OF THE GAOLIGONG MOUNTAINS 


449 


Perileptus imaicus 
Perileptus pusilloides 
Agonotrechus fugongensis 
Agonotrechus wuyipeng 
Agonotrechus xiaoheishan 
Agonotrechus yunnanus 
Minutotrechus minutus 
Queinnectrechus balfi 
Queinnectrechus griswoldi 
Queinnectrechus gongshanicus 
Eocnides fragifis 
Trechus indie us 
Trechus shiyueliang 
Trechus mingguangensis 
Trechus qiqiensis 
Trechus pseudoqiqiensis 
Trechus luzhangensis 
Trechus gongshanensis 
Trechus shibaiicus 
Trechepaphiopsis asetosa 
Trechepaphiopsis unisetigera 
Trechepaphiopsis unisetosa 
Trechepaphiopsis uniporosa 
Trechepaphiopsis unisetulosa 
Trechepaphiopsis monochaeta 
Trechepaphiopsis unipilosa 
Epaphiotrechus fortipes 
Epaphiotrechus fortipesoides 
Trechepaphiama gaoiigong 


.. 


1000 2000 3000 4000 500 Q 

elevation (meters above sea level} 


Figure 47. Chart illustrating the altitudinal ranges of trechine species represented in the Gaoiigong Shan region. 


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species 


Perileptus imaicus 
Perileptus pusilloides 
Agonotrechus fugongensis 
Agonotrechus wuyipeng 
Agonotrechus xiaoheishan 
Agonotrechus yunnanus 
Minutotrechus minutus 
Queinnectrechus belli 
Queinnectrechus griswoldi 
Queinnectrechus gongshanicus 
Eocnides fragilis 
Trechus indicus 
Trechus shiyueliang 
Trechus mingguangensis 
Trechus qiqiensis 
Trechus pseudoqiqiensis 
Trechus luzhangensis 
Trechus gongshanensis 
Trechus shibalicus 
Trechepaphiopsis asetosa 
Trechepaphiopsis unisetigera 
Trechepaphiopsis unisetosa 
Trechepaphiopsis uniporosa 
Trechepaphiopsis unisetulosa 
Trechepaphiopsis monochaeta 
Trechepaphiopsis unipilosa 
Epaphiotrechus fortipes 
Epaphiotrechus fortipesoides 
Trechepaphiama gaoligong 


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Figure. 48. Chart illustrating the co-occurence (syntopy) of trechine species in samples from the same habitats and at 
the same sites in the Gaoligong Shan region. 


DEUVE ET AL.: CARABID BEETLE FAUNA OF THE GAOLIGONG MOUNTAINS 


451 


Acknowledgements 

We thank our home institutions, the Museum National d’Histoire Naturelle (Paris), the Cali¬ 
fornia Academy of Sciences (San Francisco) and the Institute of Zoology (Beijing), respectively, 
for ongoing support of our research programs. The Kunming Institutes of Botany and Zoology pro¬ 
vided organizational and logistical support for all our expeditions. The Yunnan Provincial Division 
of Forestry and Baoshan-Gaoligong and Nujiang Prefecture Forest Reserves granted access and 
collecting permits for our work in the reserves, and staffs of these institutions assisted with col¬ 
lecting and other efforts in these areas. Support for fieldwork for this project was provided through 
Grant No. DEB-0103795 from the National Science Foundation (Biotic Surveys and Inventories 
Program), Grant No. 305702 13 from the National Science Foundation of China, grants from the 
National Geographic Society and the John D. and Catherine T. MacArthur Foundation, and private 
donations to the China Natural History Project at the California Academy of Sciences. 

We owe special thanks to our colleagues who assisted with collecting trechine specimens used 
in this study. These included Marilyn A. Dickson, Dazhi Dong, Charles E. Griswold, Zichao Liu, 
Paul E. Marek, Xiaochun Shi, Guo Tang and Darrell Ubick. Habitus images were taken by Victor 
G. Smith (CAS, Entomology). 

We also wish to express our appreciation to Dr. Terry L. Erwin who reviewed the entire man¬ 
uscript and offered useful suggestions for its improvement. 


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Table of Contents 


Mary L. Butterwick and Thomas F. Daniel. Vegetation and Flora of the Southern Black 
Mountains of West-Central Arizona .269 

John E. McCosker and Makoto Okamoto. Chauligenion Camelopardalis , a new genus 
and species of deepwater snake eel (Anguilliformcs: Ophichthidac) from the East China 
Sea .321 

Wojciech Pulawski. A new synonymy in the genus Pison Jurine, 1808 (Hymcnoptera: 
Crabronidae).329 

Arnold S. Menke. A re-evaluation of the generic limits of Pison Jurine, and a new species 
of the genus Aulacophilinus Lomboldt (Hymcnoptera: Crabronidae: Trypoxylini) 
.333 

Thierry Deuve, David H. Kavanaugh, and Hongbin Liang. Inventory of the carabid 
beetle fauna of the Gaoligong Mountains, Western Yunan Province, China: Species of 
the Tribe Trechini (Coleoptera: Caraboidca), with descriptions of four new genera, one 
new subgenus and 19 new species.341