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Table of Content
    Volume 20 Issue 4
    20 July 2012

    Mongolian wild ass or Khulan (Equus hemionus hemionus) is a nationally protected animal species and has been listed as Endangered (EN) by the IUCN Red List of Threatened Species. Khulan is a typical desert dweller whose largest population in China is found in Kalamaili Mountain Nature Reserve, northern Xinjiang. Using GIS and habitat evaluation model, Lin et al. (for details see pages 411–419 of this issue) assessed the suitability of Khulan’s habitat in

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    Editorial
    Original Papers
    Habitat suitability assessment of Equus hemionus hemionus in Kalamaili Mountain Nature Reserve
    Jie Lin, Wenxuan Xu, Weikang Yang, Canjun Xia, Wei Liu
    Biodiv Sci. 2012, 20 (4):  411-419.  doi:10.3724/SP.J.1003.2012.09219
    Abstract ( 1908 )   PDF (475KB) ( 2535 )   Save
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    To improve habitat conservation and management of the Khulan (Equus hemiomus hemionus), we assessed the habitat suitability of this species in the Kalamaili Mountain Nature Reserve. Distance from water point, slope, vegetation type and human activities were selected as factors for habitat suitability assessment, and the habitat evaluation model for the assessment was conducted with GIS. In the absence of human activities, the total area of potential suitable habitat was 9,784 km2, 8,090 km2, 9,379 km2 and 10,856 km2 during the four seasons, accounting for 54.4%, 44.9%, 52.1% and 60.3% of the total reserve area, respectively. However, the reserve was negatively affected by a national highway, mine exploration, and grazing livestock. As a result, a large amount of suitable habitats were lost in the reserve. Especially in winter, nearly 6,629 km2 of suitable habitats were lost in the reserve, which accounts for 61.1% of suitable habitat during these months. Based on these results, the following measures were put forward to protect the population of Khulan: (1) strengthen protection of water resources; (2) limit the population size of livestock in the reserve during winter months; (3) regulate and/or restrict human activities in the reserve; and (4) remove border fences and rebuild the population migration and connection routes for the Khulan between China and Mongolia.
    The quantitative assessing of trail impacts on giant panda activity based on field track points and GIS
    Minghao Gong, Meng Hou, Chen Lin, Yanling Song, Zhiyun Ouyang
    Biodiv Sci. 2012, 20 (4):  420-426.  doi:10.3724/SP.J.1003.2012.07231
    Abstract ( 2025 )   PDF (456KB) ( 2288 )   Save
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    Trails have been shown to have impacts on wildlife, which may include injury, death, habitat hindrance, and then cause small population or population isolation and increase species extinction risk. Road effect zone was widely used to examine the wildlife responses to roads and trails, but had shortages in quantifying the changes of causative factors, and meeting the data requirements for habitat evaluation. In this research, we evaluated the impacts of trails on giant panda activity based on giant panda tracking data. The study sites were centered in Foping and Changqing natural reserves, and 1,042 giant panda tracking points were collected in association with three major trails. By aids of GIS, the distance of every track point to the trails was obtained, and then the amount and frequencies of track points every 20 m from trail were calculated. To identify the impact threshold of trails on giant panda activity, we established the testing points every 100 m from the trail for analysis of the activity frequency pattern. The results showed that within 1,000 m from the trails, the giant panda activity frequencies increased with the distance. The tracking point frequency showed significant changes at 500 meters and 1,000 meters from the trails, which represent impact threshold of trails on giant panda activity. The method based on track points and impact threshold can provide a more feasible and a quantitative evaluation for disturbances of trails and other infrastructure on wildlife activity.
    Seed predation and removal of Quercus wutaishanica, Prunus salicina and Pinus armandii by rodents in the Liupan Mountains
    Xingfu Yan, Yangchun Yu, Libiao Zhou, Yunfeng Zhou
    Biodiv Sci. 2012, 20 (4):  427-436.  doi:10.3724/SP.J.1003.2012.10193
    Abstract ( 2170 )   PDF (411KB) ( 2391 )   Save
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    We established a field experiment to explore effects of different releasing methods and coverage treatments on predation and removal of Quercus wutaishanica, Prunus salicina and Pinus armandii seeds by rodents. We used three release (separately releasing, assembly releasing, and blend releasing) and cover (control, litter coverage, and soil coverage) treatments in young Q. wutaishanica stands in the Liupan Mountains of Ningxia in spring 2010. We detected significant effects of seeds of different species, release methods, and coverage treatments on the in situ seed predation rate. Seed predation and hoarding rates after removal by rodents were influenced by seed size, thickness and hardness of seed testa (endocarp), release methods, and coverage treatments. Although the average distance of seed movement during predation and hoarding events was within 5 m, the hoarding distance was longer than the predation distance for all three species. Specifically, we found that the predation distance of P. armandii seeds was significantly longer than those of Q. wutaishanica and P. salicina seeds, whereas the hoarding distance of the former was significantly shorter than the other two species. Sites that rodents used as predation and cache sites tended to contain only a single seed at a majority of the detected cache and predation sites. Soil burial was the primary strategy that rodents used to cache seeds. In summary, seed physical characteristics, distributional patterns, and ground coverage in forest communities may affect seed predation and hoarding behaviors of rodents by extending seed handling time and increasing predation risk of estimation by rodents and as a result, determined seed fates.
    Reproductive and pollination characteristics of conspecific fig wasps in two geographic types of Ficus auriculata
    Pei Yang, Zongbo Li, Yanqiong Peng, Darong Yang
    Biodiv Sci. 2012, 20 (4):  437-442.  doi:10.3724/SP.J.1003.2012.06022
    Abstract ( 1988 )   PDF (320KB) ( 1971 )   Save
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    Each Ficus species is generally pollinated by one host-specific fig wasp species, which completely depends on the fig tree for reproductive success. However, there are some exceptions in which more than one host species, subspecies, variety, or geographic type shares the same pollinator species. Studies of the reproduction and pollination traits of conspecific fig wasps in different hosts could provide an important understanding of the stabilizing mechanism of the non-specific fig–fig wasp mutualisms. This study investigated the reproduction and pollination traits of Ceratosolen emarginatus, the pollinator of Ficus auriculata in its different geographic types, auriculata-form and oligodon-form. We found a significant increase in the number of offspring and seeds as the number of foundresses increased. However, average numbers of offspring or seeds per foundress were not different in the same treatment. These results indicate that when there are fewer foundresses, interference competition is limited as to maximize available female flowers. Results also showed that irrespective of host differences, seeds were more numerous than offspring in the same treatment. This indicates that reproductive interests of figs dominate fig–wasp mutualisms, and also demonstrates that female flowers regulate the overall fig reproductive cycle. Finally, our results revealed that although the number of wasp offspring in different hosts was similar, there were more seeds in oligodon-form than in auriculata-form for equivalent foundress numbers. This suggests that the two forms may have differentiated in reproductive capacity, which implies that different host types do not affect the reproduction of fig wasps but control seed production in this species.
    On the validity of the species Phenacoccus solenopsis based on morphological and mitochondrial COI data, with the description of a new body color variety
    Zhe Chen, Jiang Zhang, Hangfei Fu, Zhengzheng Xu, Kunzheng Deng, Jiayong Zhang
    Biodiv Sci. 2012, 20 (4):  443-450.  doi:10.3724/SP.J.1003.2012.08202
    Abstract ( 2428 )   PDF (409KB) ( 2362 )   Save
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    Phenacoccus solenopsis, an exotic invasive species, was firstly reported in 2008 in Guangdong, China. Since it’s discovery, P. solenopsis has been observed in Zhejiang, Guangxi and Yunnan provinces. To discuss whether two cryptic evolutionary lineages or the P. solenopsis complex species existed, we surveyed and sequenced P. solenopsis in Zhejiang Province. During surveys for P. solenopsis in Zhejiang Province, we found P. solenopsis individuals with yellowish color on the body and three pairs of yellow spots on the back. To evaluate potential genetic divergence among these phenotypes, we sequenced mtDNA COI gene sequences (694 bp) of 25 individuals from seven host plants in six locations. We analyzed these sequences and the known sequences of P. solenopsis from GenBank and discovered three haplotypes. Additionally, we calculated intra-species genetic distance of P. solenopsis and inter-species genetic distance of the genus Phenacoccus and constructed phylogenetic trees of P. solenopsis. We found that genetic divergence of P. solenopsis was 0–1.0% compared to samples from Chinese provinces (i.e., Zhejiang, Hainan, Guangdong), the United States (i.e., California), and Pakistan, and varied from 3–3.6% to samples collected from other areas of the United States (i.e., Florida). Further, intra-species genetic distance was obviously smaller than inter-species genetic distance in Phenacoccus(13.0–17.2%). Based on the morphological characters and mt COI gene sequence analysis, these individuals with phenotypic differences are likely true P. solenopsis. However, two distinct evolutionary lineages appear to exist in P. solenopsis, and further evidence is necessary to draw reliable conclusions on the existence of a P. solenopsis complex species.
    Genetic variation in rusty-necklaced partridge (Alectoris magna) detected by mitochondrial DNA
    Rong Zhou, Jiaqi Li, You Li, Naifa Liu, Fengjie Fang, Limin Shi, Ying Wang
    Biodiv Sci. 2012, 20 (4):  451-459.  doi:10.3724/SP.J.1003.2012.09221
    Abstract ( 2482 )   PDF (534KB) ( 1961 )   Save
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    The rusty-necklaced partridge (Alectoris magna) is a bird species endemic to China that has been threatened by habitat fragmentation and hunting. To examine the conservation status of this species, we examined the intraspecific variation in the mitochondrial DNA (mtDNA) control region and cytochrome b 234 individuals from 12 representative populations throughout the species’ range. Forty-four haplotypes were defined by 34 variable sites with the mean haplotype diversity of 0.916 ± 0.011 and nucleotide diversity of 0.00449 ± 0.00242 in the 1,127 bp mtDNA control region. Haplotype diversity was maximized in the Jingyuan and Minhe populations (0.894 ± 0.063) and was minimized in the Haiyuan population (0.476 ± 0.155). Fourteen haplotypes were defined by 14 variable sites with a mean haplotype diversity of 0.738 ± 0.024 and a nucleotide diversity of 0.00216 ± 0.00009 in mtDNA cytochrome b.Haplotype diversity was the highest in the Gonghe population (0.763 ± 0.059) and did not differ in the Delingha population. Correlation between genetic diversity indices and sample size did not result in statistically significant differences. We found that neighbour-joining methods revealed limited phylogenetic distinction between geographically distant populations of the rusty-necklaced partridge, and provided tentative support for a single evolutionarily significant unit. AMOVA revealed that the most genetic variation was distributed within populations; However, the genetic variation among groups and populations was also statistically significant, indicating that the appropriate short-term management unit for the rusty-necklaced partridge is a the level of the local population. Therefore, edge populations with lower genetic diversity, including Delingha, Dulan, Lixian, Zhangjiachuan and Haiyuan, have higher conservation value. We recommend that strengthening protection of populations with lower genetic diversity, including Gonghe, Guide and Jingyuan, is advantageous to retain the general level of genetic diversity in this species.
    Genetic diversity and fine-scale spatial genetic structure of different lifehistory stages in a small, isolated population of Sinojackia huangmeiensis (Styracaceae)
    Yongmei Ruan, Jinju Zhang, Xiaohong Yao, Qigang Ye
    Biodiv Sci. 2012, 20 (4):  460-469.  doi:10.3724/SP.J.1003.2012.10011
    Abstract ( 2746 )   PDF (511KB) ( 2690 )   Save
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    Knowledge of genetic diversity and fine-scale spatial genetic structure (SGS) at different age stages of small isolated populations is important for understanding population dynamics and developing effective conservation measures for fragmented populations. In this study, we used a small, isolated population of Sinojackia huangmeiensi as a case study to investigate the change in the levels of genetic diversity and SGS at different age stages. We mapped and genotyped 60 adults,175 saplings, 198 seedlings using eight microsatellite markers to detect the genetic diversity, SGS and pollen and seed dispersal patterns in a 80 m × 160 m transect located in an original secondary forest surrounded by farmlands. No significant differences in genetic diversity were found among the three life stages, and a significant heterozygote deficiency in the population may result from substantial biparental inbreeding. We found significant fine-scale spatial structure at different age stages within 10 m, suggesting that seed dispersal mainly occurred near a mother tree. Seed dispersal distance and pollen dispersal distance were 9.07±13.38 and 23.81±23.60 m, respectively, and ‘L’ shaped curves were observed in both pollen dispersal and seed dispersal patterns. The spatial distribution of the different age stages is most likely the result of little overlap in seed rain, self-thinning, biparental inbreeding and limited gene flow. Our results have important implications for conservation of extant population of S. huangmeiensis. Measures for promoting pollen flow and increasing survival rate of seedlings should be considered for in situ conservation. The presence of SGS in this fragmented population implies that seeds for ex situ conservation should be collected from trees at least 10 m apart to reduce genetic similarity between neighboring individuals.
    Species composition and community structure of a typical mixed broadleaved-Korean pine (Pinus koraiensis) forest plot in Liangshui Nature Reserve, Northeast China
    Lina Xu, Guangze Jin
    Biodiv Sci. 2012, 20 (4):  470-481.  doi:10.3724/SP.J.1003.2012.12233
    Abstract ( 2391 )   PDF (1028KB) ( 2935 )   Save
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    The mixed broadleaved-Korean pine (Pinus koraiensis) forest (MBKF) represents the climax vegetation type of the eastern mountainous area of Northeast China. It is divided into three categories according to community structure and species composition, including southern MBKF, typical MBKF, and northern MBKF. To monitor long-term dynamics of the typical MBKF, we established a 9-ha plot following the field protocol of the 50 ha plot in Panama (Barro Colorado Island, BCI) in 2005 in Liangshui National Nature Reserve of Heilongjiang Province. All free-standing plant species with DBH (diameter at breast height) ≥1 cm were mapped, tagged, and identified to species in the plot. In the 2010 census, we documented 48 woody species with 21,355 individuals, belonging to 34 genera and 20 families. Most of the species belong to the Changbai Mountain plant flora, with a minor subtropical plant species component. The DBH distribution of all individuals showed a reversed “J” type, indicating well regenerated for the community. The DBH distribution for the canopy, sub-canopy, and middle layers and the understory layer showed the reversed "J” and “L” distributions, respectively. The DBH pattern of the nine dominant species can be classified into three categories: approximate normal, reversed “J”, and “L” distribution. Spatial pattern analysis showed that the major tree species were aggregated, and aggregation intensity decreased as DBH increased. Further analysis showed that spatial distribution was closely related to topography and varied with DBH classes. For example, the distribution of Pinus koraiensis and Tilia amurensis was significantly affected by topography (P < 0.05); however, the distribution of Abies nephrolepis, Acer ukurunduense, and Ulmus laciniata correlated significantly with topography at diameter classes of I (DBH < 10 cm) and II (10 cm ≤ DBH < 30 cm). The distribution of Betula costata and Acer tegmentosum was also significantly correlated with topography at a diameter class I (P < 0.05). Overall, the impact of the topographic gradient on species distribution decreased significantly as DBH class increased.
    Disjunct distribution of vascular plants between southwestern area and Taiwan area in China
    Jianfei Ye, Zhiduan Chen, Bing Liu, Haining Qin, Yong Yang
    Biodiv Sci. 2012, 20 (4):  482-494.  doi:10.3724/SP.J.1003.2012.13056
    Abstract ( 2800 )   PDF (642KB) ( 3454 )   Save
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    Disjunct distributional patterns and the formative mechanisms resulting in distinct flora has been an important focus in the field of phytogeography. In this study, we compared vascular plant checklists of the Chinese mainland and Taiwan, and analyzed the formation mechanisms of the discontinuous taxa between southwestern area and Taiwan area in China. We documented 198 total species (including infraspecies) in 129 genera and 56 families disjunctly distributed in southwestern area and Taiwan area in China, among which 22 genera in 15 families have species-pairs with a disjunct distribution between the two regions, including six genera of ferns, one of gymnosperm, seven dicotyledons, and eight monocotyledons. The disjunct taxa are mainly herbaceous, most of which are ferns and orchids. These disjunct taxa are mainly distributed in the center-northeastern Taiwan area, and three regions of the mainland China, including eastern Sichuan–western Hubei, western Sichuan–northwestern Yunnan–southeastern Xizang, and southeastern Yunnan–western Guangxi–southwestern Guizhou. These taxa are generally distributed along an altitudinal gradient ranging from 1,550 m to 2,350 m. We believe that these disjunct patterns may have been derived from three possible origins including a northern temperate origin, southwestern China origin, and/or tropical Asia origin.
    Population diversity and phylogeny of halophiles in the Qinghai Lake
    Derui Zhu, Jian Liu, Rui Han, Guoping Shen, Fang Yang, Qifu Long, Deli Liu
    Biodiv Sci. 2012, 20 (4):  495-504.  doi:10.3724/SP.J.1003.2012.10224
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    Qinghai Lake is the largest inland saline lake in China, but the diversity of halophiles within the lake remains unknown. We isolated 35 bacterial strains from different water samples in Qinghai Lake using an Oesterhelt-Stoeckenius medium (OSM). The majority of halophiles in this lake were moderate halophiles (grew at 0.4 to 3.5 M NaCl), accounting for about 62.9% (22 strains) of total bacteria, followed by the slight halophiles (22.9%, 8 strains) which could grow at 0 to 1.0 M NaCl, whereas the halotolerant (grew at 0 to more than 1.0 M NaCl) and nonhalophile (grew at 0 to 1.0 M NaCl) represented 11.4% (4 strains) and 2.9% (1 strain) of total bacteria, respectively. We investigated and analyzed the population diversity and evolutionary relationship based on the 16S rDNA sequences. A total of 35 bacterial strains were sequenced, and homology analysis showed that a majority was affiliated with γ-Proteobacteria and Bacilli, which accounted for 68.6% (24 strains) and 17.1% (6 strains) of total bacteria, respectively, whereas Actinobacteridae (3 strains), α-Proteobacteria (1 strain) and Eurotiomycetidae (1 strain) represented a small portion of total bacteria. Based on phylogenetics, 35 strains belonged to 14 different genera. Ten strains of Halomonas in the Oceanospirillales were the most dominant species, whereas 4 strains of Marinomonas represented a minor species component. Overall, Halomonas was the dominant group of moderate halophile, which can likely be attributed to its ability to adapt to relatively low salinity environments.
    Forums
    Attitude to the traditional knowledge of international governmental organizations and suggested strategies of China
    Gong Cheng, Cheng Wang, Dayuan Xue
    Biodiv Sci. 2012, 20 (4):  505-511.  doi:10.3724/SP.J.1003.2012.07020
    Abstract ( 2123 )   PDF (323KB) ( 2532 )   Save
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    Traditional knowledge is an important issue for international governmental organizations (IGOs). A number of international conferences have been held to discuss this topic, and a number of relevant reports and resolutions have been produced. These resolutions reflect different interests at stake and express a different understanding and position towards traditional knowledge. IGOs led by the Convention on Biological Diversity focus more on environment and biodiversity, whereas IGOs including the World Intellectual Property Organization and World Trade Organization focus on economic development and international trade. Other IGOs such as the United Nations Declaration on the Rights of Indigenous Peoples focus on society and human rights. In consideration of China’s commitment to IGOs and rich history, China should attach more importance to the protection of traditional knowledge, and consider the initiation of legislation that provides the benefits of information sharing as described in the Nagoya Protocol. This will establish a benefit sharing system to protect traditional knowledge as a way to ensure harmonious development among the environment, economy, and society.
    Holdings of type specimens of plants in herbaria of China
    Yong Yang
    Biodiv Sci. 2012, 20 (4):  512-516.  doi:10.3724/SP.J.1003.2012.05076
    Abstract ( 1965 )   PDF (320KB) ( 3424 )   Save
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    Species diversity is one of the central topics of biodiversity. Type specimens are the permanent vouchers of taxonomic names, and are irreplaceable as a means to safeguard the stability of the nomenclatural system. Holdings of common specimens and type specimens are indices reflecting historical accumulation of botanical studies for a certain country or region. This paper reviews holdings of common specimens and type specimens in herbaria worldwide and those native to China. Overall, we conclude that type collections in herbaria of China are relatively less than the world on average, because plants from China were studied by western botanists for over 400 years before Chinese botanists started to collect specimens and study plants in the early twentieth century. Based on digitization of type specimens housed in native herbaria, it is now urgent to digitize not only type specimens of Chinese herbaria, but also those held in herbaria abroad. In addition, the number of collections per species in Chinese herbaria is much less than the world average. For Chinese botanical researchers, it is important to conduct investigations in poorly studied areas in China and overseas, and to enhance the overall number and representation of specimens for each species.
    Bioinventory
    The pteridophyte diversity of the Danxia landform in Hunan Province, China
    Yuehong Yan, Zuxia He, Qixia Ma, Xile Zhou, Qiyong Lu
    Biodiv Sci. 2012, 20 (4):  517-521.  doi:10.3724/SP.J.1003.2012.05215
    Abstract ( 2309 )   PDF (320KB) ( 3083 )   Save
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    The Danxia landform is a unique landscape type with a high level of floral endemism. Herein, we report the fern flora of Danxia landform in Hunan Province. We complete comprehensive surveys of pteridophytes in the Danxia landform areas of Hunan between 2007 and 2010. We report an annotated species checklist in this paper, including 183 species and 66 genera in 31 families. Nine species, including Hypolepis polypodioides, Notholaena hirsuta, Adiantum juxtapositum, Allantodia baishanzuensis, A. succulenta, Asplenium ritoense, Polystichum deltodon var. henryi, P. gymnocarpium, P. simplicipinnum and Ctenitis dingnanensi, are reported as new records to Hunan. The families with more than 10 species include the Dryopteridaceae, Athyriaceae, Polypodiaceae, Thelypteridaceae, Selaginellaceae and Aspleniaceae, and the genera with more than 8 species are Dryopteris, Selaginella, Asplenium, Allantodia and Pteris. This list not only includes calciphilous ferns, but also acidophilous and endemic ferns restricted to the Danxia landform (e.g., A. juxtapositum and P. gymnocarpium). It is hypothesized that the fern flora of the Danxia landform may be an intergradation type between Karst landform and granite landform. Lastly, we provide recommendations for the longterm protection of the floral diversity of the Danxia landform.
    The bryophyte diversity of the Danxia landform in Hunan, China
    Zuxia He, Yuehong Yan, Qixia Ma, Qiyong Lu
    Biodiv Sci. 2012, 20 (4):  522-526.  doi:10.3724/SP.J.1003.2012.05216
    Abstract ( 2145 )   PDF (381KB) ( 2616 )   Save
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    The Danxia landform is a special red bed landform with distinct vegetation and flora. Although bryophytes play an important role in succession of vegetation, their diversity and distribution throughout the Danxia landform is unknown. Based on the systematic field surveys and microscopic identification of 710 specimens, we found 174 species in 96 genera from 53 families in the Danxia landform in Hunan, of which 36 species in 24 genera from 21 families were liverworts and 138 species in 72 genera from 32 families were mosses. Among these, 6 species were endemic to China, including Campylopus taiwanensis, Trichostomum zanderi, Schlotheimia pungens, Lindergia sinensis, Jungermannia laxifolia, and Saccogynidium irregularispinosum. Thirty species are newly recorded in Hunan, such as Gymnostomum aeruginosum, Riccia huebeneriana, and Riccardia multifida. Based on the size of ecological range, 87 species were classified as rare, such as Bartramia pomiformis, Schlotheimia pungens, and Pilotrichopsis dentate, while 52 species were common or very common, such as Marchantia polymorpha. Herein, we provide a detailed checklist of bryophytes of the Danxia landform. We also put forward suggestions to increase conservation efforts in the Danxia landform.
    A checklist of plants associated with the rare and endangered plant, Parakmeria lotungensis (Magnoliaceae)
    Hongfeng Chen, Jinsong Zhou, Rongjing Zhang, Meina Wang, Qingwen Zeng, Fuwu Xing
    Biodiv Sci. 2012, 20 (4):  527-531.  doi:10.3724/SP.J.1003.2012.05013
    Abstract ( 2162 )   PDF (323KB) ( 2410 )   Save
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    Parakmeria lotungensis is a rare and endangered plant endemic to China. Based on intensive field surveys, we chose 10 plant communities where relatively more individuals of P. lotungensis were found. Based on these surveys, we developed a checklist of 477 plant species, belonging to 237 genera and 110 families. Dominant tree layer species belonged to the Fagaceae, Lauraceae, Theaceae, and Mognoliaceae plant families, whereas dominant shrub layer species belonged to the Ericaceae, Myrsinaceae, Euphorbiaceae, Symplocaceae, and Aquifoliaceae plant families. A majority of herbaceous layer species belonged to fern families, such as Dicranopteris linearis, Lindsaea chienii, Woodwardia japonica, and Blechnum orientale. We found that most species occurred in only one or relatively few communities and there are no obligate plants which associate with P. lotungensis.

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