Corresponding author: David H. Kavanaugh (
Academic editor: A. Casale
A ten-year multidisciplinary, multi-national and multi-institutional biodiversity inventory project in the Gaoligong Shan region of western Yunnan Province, China generated more than 35,000 specimens of the beetle (Coleoptera) family Carabidae. In this report, first of a planned series, we focus on diversity in tribe Zabrini. Our study of just over 1300 specimens of zabrine carabids from the project, all in genus
Kavanaugh DH, Hieke F, Liang H, Dong D (2014) Inventory of the carabid beetle fauna of the Gaoligong Mountains, western Yunnan Province, China: species of the tribe Zabrini (Coleoptera, Carabidae). ZooKeys 407: 55–119. doi:
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 (
Map of Asia with study region outlined; scale line = 500 km. Modified from Wikimedia Commons, World Atlas of the World, at URL:
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 conduct a biodiversity inventory of the Gaoligong Mountains. Scientists from several additional institutions, 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
Prior to the start of the project the carabid beetle fauna of the region was very poorly known. The faunal for the entire Hengduan region included only about 50 species (
This report, on the tribe
As is the case with most other terrestrial anthropod groups, the
The natural physiographic limits of the study area for the project are as shown in
A total of 1,327 specimens representing zabrine species were collected during the project. All of these specimens have been divided among and are deposited in collections of our home institutions. Codens used throughout this report for collections in which specimens, including primary types, are deposited are as follows:
British Museum (Natural History), London, United Kingdom
California Academy of Sciences, San Francisco, U.S.A.
Collection of P. Meyer, Darmstadt, Germany
Collection of D. Wrase, Berlin, Germany
Deutsches Entomologisches Institut, Eberswalde, Germany
Forest Research Institute, Dehra Dun, India
National Zoological Museum of China, Institute of Zoology, Beijing, China
Institut Royal des Sciences Naturelles de Belgique, Brussels, Belgium
Museo Civico di Storia Naturale, Genoa, Italy
Muséum National d’Histoire Naturelle, Paris, France
Naturhistoriska Riksmuseet, Stockholm, Sweden
National Museum (Natural History), Prague, Czech Republic
National Zoological Collection, Zoological Survey of India, Calcutta, India
Zoological Institute Academy of Sciences, St.Petersburg, Russia
Zoologisches Forschungsmuseum “Alexander Koenig”, Bonn, Germany
Museum für Naturkunde an der Humboldt-Universität, Berlin, Germany
Zoological Museum, Moscow University, Moscow, Russia
The only measurements taken were of body length, measured from the anterior margin of the clypeus to the apex of the longer elytron. Digital images of dorsal habitus of a typical representative member of each morphospecies were taken using an Automontage imaging system from Synchroscopy with a JVC KY-F-75U digital camera and a Leica M420 dissecting microscope. The “CASENT” number associated with each image, as noted in figure captions, is a unique identifier that refers to the particular specimen photographed and its CAS database record. Distribution maps for each species were generated from geographical coordinate data maintained in a Biota Version 3.0 database (
Adult specimens of species represented in the Gaoligong Shan region can be distinguished using the following key.
1 | Medial protibial spurs trifid ( |
|
– | Medial protibial spurs simple ( |
2 |
2 | Last abdominal sternite of male with two pairs of setiferous punctures ( |
|
– | Last abdominal sternite of male with one pair of setiferous punctures ( |
3 |
3 | Body length more than 11 mm; tarsomere 5 of hind tarsi with five or six setal pairs ventrally ( |
|
– | Body length less; tarsomere 5 of hind tarsi with two (in a few specimens three) setal pairs ventrally ( |
4 |
4 | Elytron with parascutellar pore-puncture present; dorsal surface with metallic luster | 5 |
– | Elytra without parascutellar pore-puncture | 7 |
5 | Base of pronotum evenly convex from one side to the other, outer basal impressions absent or only very faintly suggested; body length 9–10 mm; sclerites of internal sac of median lobe of male aedeagus as in |
|
– | Base of pronotum slightly flattened at the sides, only the middle part evenly convex, outer basal impressions evident, either shallow and obliquely linear ( |
6 |
6 | Base of pronotum coarsely punctate; outer basal impressions shallow and obliquely linear ( |
|
– | Base of pronotum finely punctate; outer basal impressions deep and broadly foveate ( |
|
7 | Dorsal surface light-brown to brownish black, without metallic reflection, entire legs and antennae pale; body length 6.5–8.0 mm; male aedeagus with apical third of median lobe broader than middle third ( |
|
– | Dorsal surface darker, with or without distinct metallic reflection, at least femora dark (piceous to black) | 8 |
8 | Pronotum with lateral margins straight or faintly to distinctly sinuate just anterior to basal angles, rounded near middle, less rounded or nearly straight also in anterior one-third in most specimens, anterior angles distinctly and narrowly projected anteriorly beyond anterior margin; dorsal surface dark with distinct metallic blue-green reflection in most specimens, non-metallic black in a few specimens | 9 |
– | Pronotum with lateral margins more or less evenly rounded from apical to basal angle, anterior angles not or only faintly and broadly projected anteriorly beyond anterior margin; dorsal surface with or without metallic reflection | 10 |
9 | Pronotum ( |
|
– | Pronotum ( |
|
10. | Elytral microscuplture comprised of distinctly transverse meshes in both males and females (more transverse and less deeply impressed in males than in females); pronotum ( |
|
– | Elytral microscuplture comprised of distinctly isodiametric meshes in both males and females (more deeply impressed in females than in males); pronotum ( |
11 |
11 | Pronotum ( |
|
– | Pronotum ( |
12 |
12 | Dorsal surface with distinct metallic copper or bronze reflection, non-metallic black in very few specimens; pronotum ( |
|
– | Dorsal surface black or piceous, without or with only very faint metallic green reflection; pronotum with posterior angles obtusely angulate and slightly denticulate, basal impressions broadly and deeply foveate, outer basal impression distinctly delimited laterally by a broad convexity; elytral striae distinct throughout and deeply impressed; metatibiae of males with a brush-like patch of setae medially in the apical half; body length 7.7–8.7 mm |
Adults of this species (
Tibial and abdominal apices.
Tarsomere 5 of left hind tarsus, medial aspect.
Specimens of this species were collected from under stones in open roadside and waste areas (
Members of this species were collected in both the northern and southern parts of the study area (Core Areas 2, 6 and 7) but not in the central part. This gap in distribution is most likely an artifact of inadequate sampling and not a real disjunction.
Adults of this species (
Apical and subapical region of median lobe (dorsal aspect) and apex of right paramere (medial aspect) of aedeagus of males.
Specimens of this species were collected in daytime from under stones and other cover in open roadside areas with scattered grasses and shrubs (
Members of this species were collected from the northern to the southern parts of the study area (Core Areas 2, 3, 5 and 6), but they were found only on the eastern side of the mountain range in northern and central areas (Core Areas 2,3 and 5) and only on the western versant in the southern part (Core Area 6). This distribution pattern is most likely an artifact of inadequate sampling on the western slope of the mountain range, much of which is in Myanmar.
Adults of this species (
Specimens of this species were collected in daytime from under stones and other cover in open roadside areas (
Members of this species were collected from the northern to the southern parts of the study area (Core Areas 2, 3, 5 and 6), but they were found only on the eastern side of the mountain range in northern half of the study area (Core Areas 2 and 3), on both side in the central part (Core Areas 3 and 4) and only on the western versant in the southern part (Core Area 6). This distribution pattern is most likely an artifact of inadequate sampling on the western slope of the mountain range in the north, some of which is in Myanmar.
Adults of this species (
Specimens of this species were collected in daytime from under stones and other cover in open roadside areas (
Members of this species were collected in both the northern and southern parts of the study area (Core Areas 2, 5, 6 and 7), but not in the central part (Core Area 3). This gap in distribution is most likely an artifact of inadequate sampling and not a real disjunction.
Adults of this species (
Apical region of median lobe (dorsal aspect) and apex of right paramere (medial aspect) of aedeagus of males.
Specimens of this species were collected, often in great abundance, in daytime from under stones and other cover in open roadside areas (
Members of this species were collected only in the southern half of the study area (Core Areas 4, 5, 6 and 7), on both eastern and western slopes of the mountain range.
Adults of this species (
Specimens of this species were collected in daytime from under stones and other cover in open roadside areas (
Members of this species were collected from the northern to the southern parts of the study area (Core Areas 1, 2, 3, 5 and 6), but they were found only on the western side of the mountain range in the southern part (Core Area 6). This distribution pattern may be an artifact of inadequate sampling on the eastern slope of the mountain range in the south.
Adults of this species can be distinguished from those of all other species in the region by the following combination of character states: body length 8.5-9.0 mm; dorsal surface black, with very faint metallic blue or green metallic reflection in most individuals, more vivid (as in
Specimens of this species were collected in daytime from under stones and other cover in open roadside areas (
Members of this species were collected only in the northern half of the study area (Core Areas 1, 2 and 3), on both sides of the mountain range (Core Areas 1 and 2).
Adults of this species (
Specimens of this species were collected in daytime from under stones and other cover in open roadside areas (
This is the only species recorded from all seven Core Areas in the Gaoligong Shan region; but in the southern half of the study area, it is restricted to only the highest elevations, where members are found mainly on the passes over the crest of the mountain range and on the slopes just below the passes on both sides.
Adults of this species (
Specimens of this species were collected in daytime from under stones and other cover in open roadside areas (
This species was recorded from all but one Core Area (Core Area 5) in the Gaoligong Shan region. In the southern half of the study area, it is restricted to only the highest elevations, where members are found mainly on the passes over the crest of the mountain range.
Adults of this species (
Specimens of this species were collected in daytime from under stones and other cover in open roadside areas near the crest of the mountain range (
This species was recorded only from the northern part of the study area (Core Areas 1 and 2), where it is restricted to the highest elevations sampled, along the crest of the mountain range and both east and west slopes just below the crest.
Adults of this species (
Specimens of this species were collected in daytime from under stones on the open bank of a small stream at an elevation of 1740 m and at night on wet stones along a roadside at 1515 m elevation. The only other
This species was recorded only from the western slope of the southern part of the study area (Core Area 6).
Adults of this species (
Specimens of this species were collected only at night, active on the ground in marshy meadow margin areas (
Members of this species were collected at only two sites, one in the northeastern part (Core Area 2) and other in the southwestern part (Core Area 6) of the study area, but not in intervening areas. This gap in distribution is most likely an artifact of inadequate sampling and not a real disjunction.
Adults of this species (
Specimens of this species were collected, often in great abundance, in daytime from under stones and other cover in open roadside areas and waste areas (
Photographs of habitats for
This species was recorded from the southern two-thirds of the study area (Core Areas 2,4, 5, 6 and 7), on both eastern and western slopes of the mountain range.
Photographs of habitats for
Photographs of habitats for
Photographs of habitats for
Photographs of habitats for
Map showing approximate overall geographical distributions of species occurring in the Gaoligong Shan region (ranges that extend to the upper right corner of the map are truncated there but continue northeast to Japan and the Russian Far East); dashed lines connect apparently significant disjunct areas of the range of a species; scale line = 500 km.
Chart showing the representation of
Chart illustrating the altitudinal ranges of
Chart illustrating the co-occurence (syntopy) of
Although the Gaoligong Shan region is at the heart of one of the world’s biodiversity hotspots (
The second general geographical range pattern includes two species,
On first glance, the observed diversity of
The chart in
A comparison of recorded diversity for
With respect to the altitudinal distribution of the
In the future, it will be interesting to compare and contrast both the broad and regional geographical distributions and the altitudinal and ecological range patterns seen in other carabid groups represented in the area with those found among the
We thank our home institutions, the Museum für Naturkunde, Berlin, the California Academy of Sciences, San Francisco, Institute of Zoology, Beijing, and the Kunming Institute of Zoology, respectively, for ongoing support of our research programs. The Kunming Institutes of Botany and Zoology provided organization 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 collecting 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. 30570213 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 many colleagues who assisted with the collecting effort over the ten-year duration of the fieldwork for this project and who, collectively, contributed many important specimens for this study. These included Bruce Bartholomew, Roberta L. Brett, Thomas S. Briggs, Peter W. Fritsch, David Garfield, Charles E. Griswold, Vincent F. Lee, Dong Lin, Paul E. Marek, Jeremy A. Miller, Pierre Paquin, Norman D. Penny, Darrel Ubick and Xinping Wang (U.S.A. team members); Weidong Ba, Keji Guo, Qingbai Hou, Peng Hu, Chunhua Li, Cailian Li, Rong Li, Xueyan Li, Xingcai Liang, Ye Liu, Zichao Liu, Chunlin Long, S. Z. Ma, Xuejun Meng, Guangxu Peng, Xianjin Peng, Yuehua San, Hongliang Shi, Xiaochun Shi, Guo Tang, Jun Wang, Meng Xie, J. Xiong, Xiang Xu, Hengmei Yan, Jialian Yang, Jiayu Yang, Song Yang, Jinfeng Zhang and Benxiang Zhu (Chinese team members); and Marilyn A. Dickson, David G. Long and Philip Thomas (of the Royal Botanical Garden, Edinburgh). Habitus images were taken by Victor G. Smith (CAS, Entomology), and Stan Blum (CAS, Informatics) and Lauren Scheinberg (CAS, Herpetology) provided assistance with generation of distribution maps from digital GPS data.