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Cedric Collingwood and Harold Heatwole.
Ants from northwestern China (Hymenoptera, Formicidae).
Psyche 103:1-24, 2000.

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PSYCHE
Vol. 103 2000 NO. 1-2
ANTS FROM NORTHWESTERN CHINA
(HYMENOPTERA, FORMICIDAE)
Department of Zoology, North Carolina State University, Raleigh, NC 27695-7617, U.S.A.
An ecological survey of the ant fauna of the southern part of the Junggar Basin and adjacent mountains, Xinjiang, China, revealed 46 species of which 27 (59%) were new records for China. Most of the species are widespread and no endemics were found. A largely boreal fauna occupies the spruce forest zone at high elevations of the Tien- shan Mountains, giving way, lower down, in elm forest, to a mixed, but primarily mesic temperate fauna. Loess desert and degraded steppe at mid-elevations and in the foothills are overgrazed and have only a few species that elsewhere occur in temperate mesic and/or steppic habi- tats. The sandy deserts and poplar woodlands of the arid Junggar Basin have a fauna characteristic of deserts and steppes. The salt desert fauna has a strange mixture of a number of elements. The ant fauna of northwestern China is poorly known. Conse- quently, during an ecological study in the Junggar (Dzungar; Zhun- gel; Kzungar) Basin, Xinjiang Uygur Autonomous Region (Fig. I), in the summer of 1991, the junior author collected ants over a range of habitats at a variety of elevations. The present paper reports on that Present address: City Museum, Leeds LSI 3AA, UK Manuscript received 17 April 1995.




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T~ENSHAN MOUNTAINS
/ JUNGGAR BASIN )
Fig. 1. Map of Xinjiang Uygur Autonomous Region, western China, showing location of the principal study areas. 1, Loess Desert; 2, Sangong River; 3, Tien-Chi (Heavenly Lake). There were a number of habitats studied at Muosuowan Biologi- cal Station and Fukang Biological Station, and opportunistic collections were made at various sites in the vicinity of Shihezi and Urumqi. collection. A later publication, now in preparation, will treat the ecol- ogy of the region's ants.
In the following account, supplementary distributional data, unless accompanied by literature citations, are based on locality data from spec- imens examined by the senior author in various museum collections. Some of the voucher specimens from the present study were retained in the senior author's collection. The remainder were deposited at the following institutions: the Xinjiang Institute of Biol- ogy, Pedology and Desert Research, Ummqi, China; the Museum of Comparative Zoology at Harvard University; the Department of Ento- mology at North Carolina State University. Xinjang Province lies in the western part of the Gobi-Taklamakan Desert complex and contains two large, arid basins encircled by moun-



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20001 Collingwood & Heatwole 3
tains. The Junggar Basin, in which most of the present work was car- ried out, is the most northern of the two, and is separated from the southern Tarim Basin by the Tienshan (Tian Shan) Mountains. The characteristics of the area have been described by Walter et al. (1983), Chen (1988) and Zhang (1990) and were summarized in publicity brochures published by the Xinjiang Institute of Biology, Pedology and Desert Research (XIBPDR) and the Chinese Academy of Sciences. The following description relies heavily on those sources as well as on personal communications from Professor Zhang Li Yun of XIBPDR. The basin encompasses 50,000 km2 and contains the second largest sandy desert in China, the Kurbantonkut (Gurbantiiggiit) Desert. The floor of the basin lies below 500 m elevation. Proceeding toward the Tienshan Mountains and up the northern slopes, one passes through a series of progressively more mesic vegetation types. Histori- cally, there was a belt of poplar woodland (Populus euphractica diver- sifolia) fringing the desert (Zhang, personal communication), but human influence has reduced it to a few scattered remnants. Between 1,020 and 1,600 m elevation are steppes, now heavily overgrazed and degraded. Above 1,600 m are forests, first elm and then spruce, leading finally to the treeline and thence to perpetual snow. Alluvial fans and loess deposits spread out at the base of the mountains in some localities. In the desert there are local areas of halo- phytic vegetation on salty soils.
Where there is irrigation, natural vegetation gives way to cotton and a variety of vegetable crops. Moister conditions than those of the basin floor generally occur in valleys, on river banks and where groundwater from the mountains raises the watertable. The fringe of the desert near the mountains has higher humidities, greater rainfall, more subsurface water, greater run-off and a greater diversity and cover of vegetation, than occurs deeper in the basin. The wooded areas at lower elevations have been influenced heavily by humans. About 1958 approximately 50% of the natural forests of elms and poplars were destroyed and grass planted in an attempt to extend grazing lands (Zhang 1990).
The principal study sites were located in various desert and steppic habitats in the southern part of the Junggar Basin (Muosuowan and Fukang biological stations) and up the northern slopes of the Tienshan Mountains, through mesic forests to Tien-Chi (Tianchi) (Heavenly Lake) near treeline. In these areas, ants were collected by baited tran- sects, using sardines for bait, supplemented at some sites by pitfall traps, examination of the stomach contents of insectivorous vertebrates,



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and opportunistic collections. In order to obtain species with different activity rhythms, the transects were attended four times per day, morn- ing, mid-day, afternoon and night. In addition, incomplete, opportunis- tic collections were made in the environs of the cities of Shihezi and Ummqi (Ummchi; Wulumuchi) and in two desert sites on the southern side of the Tienshan Mountains. Localities from which specimens were collected are shown in Fig. 1. A brief description of the collecting sites follows.
1. Muosuowan Biological Station
Transects were used for a comprehensive collection in various habitats at the Muosuowan Biological Station, located approximately 200 km northwest of Urumqi near the terminus of a 100-km irrigation canal draining mnoff from the Tienshan Mountains into the desert. The station grounds lie at an elevation of 335-350 m above sea level. The region is characterized by stabilized dunes, partly stabilized dunes, and flat interdune plains. The average annual precipitation is about 115 mm. In the open, air temperature one meter above the ground reaches as high as 37OC in the summer. Wind is a common event. The vegeta- tion is of scattered perennial shrubs. In spring often there are ephemeral and ephemeroid plants, but these dry up by summer. The habitats studied were: (1) tops of sand dunes, (2) slopes of sand dunes, (3) natural interdune plains and (4) interdune plains on which plantations of shrubs had been established. 2. Vicinity of Shihezi
Small opportunistic collections were made in: (1) salt desert east of Shihezi, (b) subdesert 47 km north of Shihezi, (c) overgrazed Artemisia desert south of Shihezi on the foothills of the Tienshan Mountains at 1600 m elevation, and (d) a nearby sheep farm at 1650 m. 3. Urumqi City and Vicinity
Ants were collected opportunistically: (1) at various places in the city of Ummqi, (2) in micro-shrub desert near the Ummqi Reservoir, 10 km south of Urumqi at an elevation of 1,100 m, and (3) in a shrubby part of the Gobi Desert on the southern side of the Tienshan Mountains, 33 krn south-southeast of Ummqi.



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1. Fukang Biological Station
The Fukang Biological Station covers a large area (lat. 43' 50' to 44O 30' N; long. 87O 45' to 88' 05' E) west of the city of Fukang and 76 krn northeast of Ummqi. Mean annual precipitation is 164 mm, but annual evaporation is approximately 2,000 mm. Maximum tempera- tures in summer reach 42.6OC and winter minima descend to -41.6OC. Except for shrub plantations on interdune plains, the habitats occurring at the Muosuowan Biological Station also are represented at Fukang, as well as some additional ones. There are dune tops, dune slopes, interdune plains, salt desert, poplar woodland on sand, poplar woodland on clay, irrigated crops, and plantations of trees used for windbreaks. The habitat diversity results from the location being near the limit of runoff from the Tienshan Mountains and consequently there is a mixture of arid habitats toward the interior of the basin and more mesic ones toward the mountains.
Transects were employed in each of these habitats. NORTHERN SLOPES OF THE TIENSHAN MOUNTAINS A series of habitats was examined for ants, beginning in the foothills at the edge of the desert in the Junggar Basin and extending upward through progressively more humid altitudinal zones to near treeline at Tien-Chi. They are described below in order of increasing elevation. Baited transects were used at all elevations. 1. Loess Desert, elevation 800 m
Located 40 km northeast of Urumqi and 29 km south of the Fukang Biological Station are foothills of the Tienshan Mountains. These hills are covered by a mixture of glacial gravels and boulders, and clay deposits of loess origin. Precipitation averages about 200 mm annually. The vegetation is a mixture of small shrubs, herbs and grasses; the area is heavily overgrazed. Transects were established on both northern and southern slopes of these foothills.
2. Sangong (San-Gong) River, elevation 1250 m Two sites were studied at this location: (1) riverine woodland of elm trees (Ulmus pumila), now left as a park-like remnant in a village



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on the northern bank of the Sangong River and (2) a steppe of herbs, grasses and small shrubs desertified through overgrazing and soil com- paction.
The combination of higher elevation and proximity to the river resulted in moister conditions in the woodland than the mean annual precipitation of 250 mrn would suggest. Furthermore, winter tempera- tures are milder here than either in the basin, into which cold air drains, or further up-slope where altitudinal effects become evident. The degraded steppe, being farther from the river was drier. 3. Tien-Chi, elevation 1900 m
The lake Tien-Chi was formed by glaciers and reaches depths of 160 m. It is surrounded by spruce forest (Picea schrenkiana var. tian- shanica). There are also poplar trees (Populus tremula) and shrubs (Rosa) in the area. Mean annual precipitation is over 500 rnrn. Genus Myrmica Latreille 1804
Myrmica angulinodis Ruzsky 1905
Locality: Tien-Chi 26.viii.9 1.
This species was taken at only one site: prepared upland pasture
fringing spruce forest. In common with other Myrmica species, M. angulinodis is a general scavenger but may also attend extrafloral nectaries and other sugar sources including aphid honeydew. This species was described from Siberia and has been recorded from Mon- golia (Pisarski 1969). Wu and Wang (1995) list eight species of Myr- mica from China, including M. angulinodis; they indicate its distribution as Inner Mongolia, Japan, Korea and Siberia. The present collection constitutes a first record from Xinjiang. Genus Messor, Fore1 1890
Heatwole et al. (1992) described the foraging behavior and habi- tats of Messor species in the Junggar Basin. Ants of this genus are seed gatherers in semi-arid to arid habitats. Consequently, where large colonies are established, these ants may affect seed-dispersal and may regulate the size of the seed-pool of particular plants. In the area stud- ied, however, members of this genus only occurred at 8% of collection sites.




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Wu and Wang (1995) list only one species of Messor from China (M. aciculatus).
Messor aralocaspius Ruzsky 1902
Locality: Fukang 8.viii.91,25.viii.91, 27.viii.91. This species may develop large crater nests in favorable situations. It is distributed through southern Russia to the Arabian highlands and also has been recorded from Mongolia and from near Kandahar, Afghanistan at 950 m (12.1.53, leg. H. Klapperich). Messor orientalis Emery 1898
Localities: Fukang 17.viii.91, 19.viii.91, 21 .viii.91, 23.viii.91; Sangong riverside 25.viii.91, 27.viii.91; Urumqi 28.viii.9 1. This species occurred in a variety of habitats and was collected at 13 sites incuding dune slopes, salt desert, loess desert, steppe, river- side, and city precincts. It has a wide range through Central Asia, southern Russia, Cypms, Turkey and the Middle East. Messor perantennatus Arnoldi 1969
Locality: Fukang 20.viii.91, 21 .viii.91. This species was described from Turkmenistan and southern Rus- sia.
Messor subgracilinodis Arnoldi 1969
Localities: Muosuowan 2.viii.9 1 ; Fukang 8.viii.91. This species has a similar range to that of M. perantennatus. Genus Cardiocondyla Emery 1869
Cardiocondyla species are very small ants that live by general scavenging. Often, surface-active workers are seen only when the ground is damp after irrigation or rain. Both of the following species were generally abundant in sandy areas in the neighborhood of trees, shrubs and buildings. Some individuals also were recovered from the stomachs of lizards and small mammals.
Wu and Wang (1995) listed two species of Cardiocondyla from China, not including either of the two species below. Cardiocondyla elegans Emery 1869
Localities: Muosuowan 2.viii.9 1 ; Tien-Chi 6.viii.9 1 ; Fukang 9.viii.91, 13.viii.91, 14.viii.91, 17.viii.91. This is a wide-ranging southern European species, also common on the Mediterranean coast of the Near East.



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Cardiocondyla stambuloffi Fore1 1892
Localities: Fukang 5.viii.91, 9.viii.91, 10.viii.91, 12.viii.91, 13.viii.91, 17.viii.91.
This species was known previously from southwestern Russia, the southeastern Balkans, Turkey and Mongolia. Genus Monomorium Mayr 1855
Monomorium lindbergi Pisarski 1967
Locality: Fukang 21 .viii.91.
Two workers were among Solenopsis specimens collected in a salt desert habitat. This is a brown, large-eyed species (ornrnatidia in the longest dimension 0.36 x head width). The specimens nearly corre- sponded in size, color and general appearance to Pisarski's (1967) orig- inal description of specimens from Tang Sayed, northern Afghanistan, but the eyes were larger and the head more rectangular than appears to be true from Pisarski's description (although he gave no measure- ments). A simple pair of hairs are present on the pronotum and on the first gastric tergite. This species was previously known only from Afghanistan.
Genus Solenopsis Westwood 1840
Wu and Wang (1995) recorded two species of Solenopsis from China, but not S. orientalis.
Solenopsis orientalis Ruzsky 1905
Locality: Fukang 10.viii.91, 11 .viii.91. This species was collected on soil in a poplar windbreak. Like all boreo-palearctic Solenopsis, this is a small but aggressive and preda- tory ant, feeding on arthropods and on the brood of other (usually larger) ant species. S. orientalis also has been recorded from Siberia, Mongolia and Turkestan. The present paper represents the first Chinese record of this species under its own name. However, it is likely that Wheeler's (1930-3 1) record of the European species S. fugax Latreille from Foochow, China, was actually based on S. orientalis which has more diffuse sculpture and fewer body hairs. Genus Crematogaster Mayr 1861
Crematogaster subdentata Mayr 1877
Locality: Fukang 5.viii.91, 9.viii.91, 23.viii.91. This was the only species collected of this large world-wide genus. The ants were taken in the neighborhood of poplars and on rocky



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slopes fringing loess desert. C. subdentata nests in the soil but like most species of Crematogaster is very dependent on homopteran hon- eydew on trees and shrubs. This ant prevously was known from Turkestan, Afghanistan, the Caucasus, Iran and in China from the Tienshan foothills. Wu and Wang (1995), however, did not include it on their list of 11 species of Chinese Crematogster. Genus Tetramorium Mayr 1855
All of the following Tetramorium species have similar habits. They are general scavengers but also frequently collect and store small seeds. Members of this genus outnumbered all other Myrrnicinae and were present at 33% of the collecting sites. Tetramorium caespitum (Linnaeus 1758)
Localities: Muosuowan 2.viii.9 1 ; Fukang 8 .viii.9 1, 9.viii. 9 1, 10.viii.9 1, 17.viii.9 1 ; Sangong riverside 27.viii.9 1. This species was collected mainly in irrigated pasture and along river banks. This is a common and abundant Eurasian ant. Wheeler (1930-31) and Wu and Wang (1995) distinguished the Chinese form of this species as T. jacoti and Pisarski (1969) used that name for Tetramor- ium collected in many places in Mongolia. However, this is a wide- ranging and variable species and the complex of forms has not been sufficiently characterized.
Wu and Wang (1995) included Tetramorium jiangxiense, recorded from eastern China, in the synonomy of T. caespitum. Tetramorium diomedaeum Fore1 1920
Localities: Fukang 9.viii.9 1, 21 .viii.9 1, 23.viii.9 1 ; Sangong river- side 27 .viii.9 1.
This is a pale, relatively unsculptured ant with a wide petiole node. It was previously known from the eastern Mediterranean and southern Balkans and on the Maltese Islands.
Tetramorium ferox Ruzsky 1903
Locality: Fukang 21 .viii.9 1.
This coarsely sculptured species with strong propodeal spines was described from southern Russia and also has been recorded from Afghanistan and Mongolia (Pisarski 1967, 1969). Tetramorium inerme Mayr 1877
Locality: Fukang 2 1 .viii.91
In this species the propodeal spines are reduced to very short den- ticles. It was described from Turkestan and also occurs in Mongolia.



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Tetramorium nitidissimum Emery 1925
Locality: Fukang 17.viii.91.
This species was previously known from the Caspian region. Tetramorium sahlbergi Finzi 1936
Localities: Muosuowan 3.viii.91; Fukang 9.viii.91, 10.viii.91, 12.viii.91, 13.viii.91, 17.viii.91, 19.viii.91, 21.viii.91, 23.viii.91. This small black Tetramorium was often among the stomach con- tents of gerbils and lizards. The species is known from Syria and Pales- tine and these records from the Junggar Basin considerably extend its range northeastward.
Tetramorium sarkissiani Fore1 19 1 1
Localities: Fukang 9.viii.9 1, 12.viii.9 1, 20.viii.9 1, 23.viii.9 1 ; San- gong riverside 25.viii.9 1.
These ants occurred mainly in open sites in loess desert and on clay interdune plains. During the present study ants of this species were observed in intraspecific battles. Hostile workers tug at each other, often in chains, but seem to do little damage. Such engagements may continue for several days. T. sarkissiani is one of the less well known species. It is distinguished by its smooth dorsal profile and strong sculpture. This species was described from Turkey and also was known previous to this study from the Balkans (leg. B. Poldi and C. A. Collingwood).
Tetramorium striativentre Mayr 1877
Localities: Muosuowan 2.viii.91; Fukang 8.viii.91, 12.viii.91, 13.viii.91, 17.viii.91; Sangong riverside 27.viii.91. Tetramorium striativentre is a characteristic denizen of warm step- pic environments and previously was known from Turkestan and parts of Afghanistan and Iran. It is slightly larger than its congeners and also is distinguished by the finely striated first tergite of the gaster. Tetramorium taurocaucasicum Arnoldi 1967 Localities: Fukang 10.viii.91, 11 .viii.91, 12.viii.91, 14.viii.91, 23 .viii.9 1 ; Sangong riverside 27.viii.9 1. This species was collected mostly in steppic environments near Fukang and above the Sangong River. It was described from southern Russia and also occurs in Iran. It resembles a somewhat larger T. cae- spitum, which it evidently replaces in drier sites. Tetramorium tauro- caucasicum has weak reticulate sculpture on the first gastric tergite, and the petiole and postpetiole are more sculptured than in T. caespi-



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turn. T taurocaucasicum is sometimes considered a junior synomym of T. forte Fore1 1904 (Bolton 1995).
Genus Plagiolepis Mayr 1861
Plagiolepis rnanczshurica Ruzsky 1905
Localities: Muosuowan 2.viii.91, 3.viii.91; Fukang 8.viii.9 1, 10.viii.91, 12.viii.91, 13.viii.91, 14.viii.91, 23.viii.91. This is a small, somewhat cryptic species nesting in rock crevices and in leaf litter where there is moisture and some shade. It was mostly collected at sites around field stations and among poplar trees. Species of this genus mostly prey upon small arthropods in the soil. P. manczshurica was described from Manchuria and occurs throughout Mongolia, and widely in central China and in areas around Beijing (Wheeler 1930-31) and in North Korea (Collingwood 1976). Wu and Wang (1995) report this species from Xinjiang. Plagiolepis sp. indet.
Locality: Fukang 22.viii.91.
This species was collected from loess desert as a single pale speci- men that could not be identified.
Genus Lepisiota Santschi 1926 (Acantholepis Mayr actt.) Ants of this genus are surface-active and fast-moving, usually found in open areas near water and along sea coasts. Lepisiota sp. indet.
Locality: Fukang 14.viii.91.
A single damaged specimen recovered among the stomach con- tents of an agamid lizard at Fukang could not be identified to species. At least four species of Lepisiota are known from Afghanistan and Turkestan. The occurrence of this genus was previously recorded for China by Wu and Wang (1995), who reported L. rothneyi (as Plagi- olepis rothneyi) from southern China.
Genus Lasius Fabricius 1804
Both of the following are small robust species mainly dependent on honeydew from root-feeding and foliage aphids. These ants are common holarctic species but their taxonomy and nomenclature in Central Asia are subject to revision.




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Lasius niger (Linnaeus 1758)
Locality: Sangong riverside 27.viii.91.
Lasius obscuratus Stitz 1930
Localities: Sangong riverside 25.viii.9 1, 27.viii.9 1 ; Tien-Chi 26.viii.91. This species was known previously from the Caucasus. Genus Camponotus Mayr 1861
Camponotus species depend mostly on arboreal aphid honeydew and do not usually occur far from trees or shrubs. Many of the larger species are nocturnal and more likely to be caught in pitfall traps than seen in full daylight.
Camponotus fedtschenkoi Mayr 1877
Localities: Muosuowan 3 .viii.9 1 ; Fukang 13 .viii.9 1, 17.viii.91. Small pale workers of this species were collected from pitfall traps. Its previously known range was Turkestan and Afghanistan. Camponotus kurdistanicus Emery 1898
Localities: Fukang 17.viii.9 1; Sangong riverside 27.viii.9 1. This is a reddish species related to C. fallax of western Europe. Its distribution is Central Asia, including Afghanistan. Camponotus sachalinensis Fore1 1904
Locallity: Tien-Chi 26.viii.9 1
This is a black relative of C. herculeanus (Linnaeus) which it replaces in the spruce forests of northeastern Asia. All present speci- mens were taken among spruce.
Camponotus semirufus Emery 1925
Locality: Fukang 5 .viii.9 1.
Camponotus semirufus was collected among poplar trees. It was described from Turkestan and is also present in Afghanistan, taken at Scham Shir Kandahar 22.v.52 (leg. H. Klapperich), as a new record for that country. It is related to C. lateralis (Olivier) of southern Europe. Camponotus turkestanicus Emery 1887
Locality: Fukang 9.viii.9 1.
This is another Central Asian species and was taken in sandy ground among poplar trees. It has an unsually strong development of long curved subcephalic hairs characteristic of several desert-adapted species in diverse genera. This species occurs in Russian Turkestan, Afghanistan and in Mongolia (Pisarski 1969).


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