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Research progress on karst tiankeng ecosystems.

Introduction

Karst terrain accounts for a significant portion of the global land surface. It is a natural complex formed by the interaction of water and soluble rock with special functions (Luo et al. 2014; Zhu and Chen 2005). In China, karst areas cover about one-third of the country, making them the world's largest contiguous karst areas (Luo et al. 2014; Ravbar and Sebcla 2015). Unlike other geomorphologic features, karst geomorphology has many unique natural landscapes, such as fengcong (a karst landscape with closely packed conical hills), fenglin (tower-and-cone karst), cave systems, and tiankeng (Waele et al. 2011; Waltham 2008; Zhu and Chen 2005). Among them, karst tiankeng--which can be loosely described as a giant cylindrical opening that develops due to a variety of mechanisms and subsequently undergoes a specific set of geologic and biologic processes--has enjoyed increasing attention owing to its high-value for conservation and tourism in recent years.

As a kind of negative landform geological wonder, karst tiankengs were discovered and first named at the end of the twentieth century in China. It has long been disputed whether karst tiankengs can be significantly distinguished from normal dolines (Gunn 2005; Kranjc 2009; Shui et al. 2015), however, there is growing recognition of their unique model of geophysical and ecological development in landscapes all over the world (Chen 2011; Fan 2014; Feng 2015; Jiang 2002; Lin 2005; Palmer and Palmer 2005; Shui et al. 2015; Waltham 2005a, b, c; Zhu 2001; Zhu et al. 2003a, b; Zhu and Chen 2005; Zhu and Waltham 2005). Karst tiankeng ecosystems (Fig. 1) are important natural resources and are rich in biodiversity (Fan 2014; Feng 2015; Shui et al. 2015). In karst regions, they are one of the most important living environments for many wild animals and plants, including endangered species. Since the 1990s, studies on karst tiankengs in China commenced with the discovery of Dayanwan and Xiaoyanwan Tiankengs in Xingwen county, southwest China, by Chinese-led joint expedition of international researchers (Zhu and Chen 2005). Since then, more karst tiankengs were found and explored, such as Xiaozhai Tiankeng in 1994, Dashiwei Tiankeng groups in 2000 and Wulong Tiankeng in 2001 (Zhu et al. 2003a, b; Zhu and Chen 2005). In the first 10 years of the twenty-first century, the word 'tiankeng' entered the karst lexicon and received endorsement from some karst researchers in domestic and international academic circles (Zhu 2001; Zhu et al. 2003a, b; Zhu and Chen 2005). Research on karst tiankengs experienced a progression from basic investigation to long-term experiment study and from morphology to geomorphic evolution, developmental conditions and ecological environment (Shui et al. 2015). In the past 20 years, research on karst tiankengs in China has gradually formed its own characteristics and has progressed. In the present work, we review studies related to karst tiankengs in China and analyzed the current problems. Finally, we suggest future directions for research on karst tiankeng ecosystems.

The Concept, Classification and Distribution of Karst Tiankengs

The word "tiankeng" is a transliteration from two Chinese characters, which roughly mean "sky hole" or "heaven pit" or some similar variation on that double theme. In 2001, Zhu gave the first definition of tiankeng. This was later revised to "a large, steep-walled, pit-like, negative, karst landform that opened from beneath towards the surface, with both its depth and diameter more than 100 m, developed in a great thickness of continuous soluble rocks within a deep vadose zone of the aquifer and connecting with an active cave river at its foot" (Zhu and Chen 2005). Following the above description of tiankeng, Palmer and Palmer (2005) provided a simplified definition of tiankengs as "large karst depressions with nearly vertical walls and a mean depth and diameter typically more than 100 m each". However, not all authors agreed that tiankengs should be included into the karst geomorphology lexicon. For example, Gunn (2005) stated that tiankengs lay at one extreme of the collapse doline spectrum, and a key question was whether there was a distinctive "tiankeng process" or whether the distinction was purely morphological. In addition, tiankengs have been recorded as giant dolines outside China, including Crveno jezero in Croatia, Minye in Papua New Guinea, Garden of Eden in Malaysia, and Sotano in Mexico (Zhu and Chen 2005).

Early studies mainly focused on researching cone karsts (fengcong) and tower karsts (fenglin) and there was no research on tiankengs as an integrated whole. Until recently, tiankengs were known due to their discovery and exploration for tourism development, with some being listed as World Heritage List sites. In addition to differences in purpose, methods and regions in tiankeng research, there is no accepted tiankeng classification system. For example, in a review of the literature, Shui et al. (2015) summarized the research methods into four types: morphological, genetic, morphogenetic and evolutionary approaches.

Many researchers have followed a morphological approach, which mainly stressed morphological and morphometric criteria in delineating a tiankeng as a type of giant doline, and they distinguished collapse and erosional tiankengs as sub-types (Klimchouk 2005; Shui et al. 2015). For example, based on their depth and width, tiankengs were divided into three size groups: very large, large and normal tiankengs (Zhu and Chen 2005). Klimchouk (2005) believed that this approach would lead to the blending of features of different origins.

Differing from above researchers, the other researchers followed genetic or morphogenetic approaches, believing that tiankengs were one extreme of the collapse doline spectrum (Gunn 2005) or one type of closed depressions not uniquely different from other collapse features in a consideration of the size distribution (White and White 2005). Based on their geomorphic evolution, tiankengs have been divided into three types: original, mature and degraded tiankengs (Waltham 2005c). However, it is often difficult to distinguish an original tiankeng from a doline by an evolutionary approach.

There are more than 80 recorded tiankengs worldwide, including more than 50 in China (Kong 2012; Liu et al. 2012; Shui et al. 2015; Zhu et al. 2003a, b; Zhu and Chen 2005). Karst tiankengs in China are characterized by their number, large scale, concentrated distribution, great scientific value and high aesthetic value. In the world, the three largest tiankengs all located in China, including Xiaozhai Tiankeng, Dashiwei Tiankeng and Haolong Tiankeng (Shui et al. 2015), and 9 of the 10 largest tiankengs are in China (Shui et al. 2015). These tiankengs occur mainly in southwest China, notably in Guangxi, Chongqing, Sichuan and Guizhou Provinces; followed by Hunan, Hubei and Yunnan Provinces (Fig. 2). Available data also suggested that there were only 26 known tiankengs outside China (Waltham 2005a, b, c). These tiankengs are distributed in Nakanai of New Britain, south-east Asian islands, southern Slovenia, Mexico, Brazil, Puerto Rico, Sarawak, Madagascar and Italy (Waltham 2005a, b, c; Zhu et al. 2003a, b).

Formation and Evolution Mechanism of Karst Tiankengs

Compared with other typical negative karst landforms,--such as dolines, depressions and vertical shafts, tiankengs are relatively rare and are characterized by a special set of ecological processes. Karst tiankengs are formed by the combination of various natural factors, and can be roughly divided into two types: collapse and erosion tiankengs (Shui et al. 2015; Zhu and Chen 2005). However, different researchers have afforded different views and experiences on the formation and evolutionary mechanism of karst tiankengs.

Researchers from outside of China have focused on the following hypotheses to infer possible formation and evolution mechanism of karst tiankeng: cave un-roofing (Klimchouk 2005), hydraulic processes (Palmer and Palmer 2005), evolutionary theory of tiankengs (Waltham 2005c), and mechanism theory of depressions (White and White 2005). However, these hypotheses cannot explain all aspects of tiankengs' development patterns; most explain only parts of them. For example, Waltham (2005a, b, c) placed three different tiankengs--Zhucaojing, Xiaoyawan and Dayawan Tiankengs--into an evolutionary sequence and suggested that the three adjacent sites represent three successive stages in the same process of tiankengs' development, overlooking a key characteristic of tiankengs: their changes over time and space under natural conditions (Shui et al. 2015). Similarly, Klimchouk (2005) holds that tiankeng is a youthful stage of open collapse-doline development, suggesting that the formation of tiankengs may involve various breakdown mechanisms and processes, including upward stoping by slab and block breakdown, lateral expansion of an initial opening through face retreat, and un-rooftng of a cavity by subaerial weathering. However, only few examples of tiankeng, such as the Sanqiao (Three Bridges) site (Klimchouk 2005), supported gorge formation by cave un-roofmg. Palmer and Palmer (2005) believed that tiankengs were formed most commonly by the collapse of bedrock into underlying caves that contain active rivers. However, some examples from the tiankengs of South China and the giant dolines of the Nakanai and the Muller indicated that rock collapse (including tiankengs) was not associated only with vadose drainage (Eavis 2005). Many Chinese researchers agree with the opinions of Zhu and Chen (2005), who explored the formation and evolution mechanism of karst tiankengs from their formation conditions and developmental stages and divided them into two types: erosional and collapse tiankengs. They believed that collapse and erosional tiankengs are formed by underground river flow and surface river flow, respectively, and both are directly related to large underground conduit flows in the aquifer (Zhu and Chen 2005). Since many karst tiankengs were found to be of collapse type, they mainly focused on the development of collapse tiankengs and divided these into three evolutionary stages: a cave river passage that expands to a large cave chamber, and from there becomes a tiankeng open to the surface (Wei et al. 2011; Zhu et al. 2003a, b; Zhu and Chen 2005; Zhu and Waltham 2005). Zheng et al. (2009) quantitatively analyzed various factors in a hierarchical process and calculated the relative importance of various factors. This forecast expands understanding of the developmental model of karst tiankengs by combing both internal and external factors that affect development of karst tiankengs.

Karst Tiankeng Ecosystems

Differing from other terrestrial ecosystems, karst tiankeng ecosystems have unique development models and paths, and form a unique ecological community structure, which is closely related to development of the karst tiankeng (Chen 2011; Zhu and Chen 2005). Due to the broad variability in different parts of the world, karst tiankeng ecosystems are vastly different from those of mainland and island communities. Despite their uniqueness and rarity, previous studies mainly focused on the status, definition, geomorphology and evolution of tiankengs (Shui et al. 2015) and only a few focused on biodiversity and evolution of karst tiankeng plants (Fan 2014; Feng 2015; He et al. 2004; Huang et al. 2004; Huang and Su 2015; Jiang 2002; Lin 2005; Liu et al. 2009; Su 2012; Su and Ma 2011; Su and Zhang 2012).

Plant Succession, Plant and Animal Biodiversity in Karst Tiankeng Ecosystems

Plant Succession

Similar to islands, the establishment of tiankeng plant communities was believed to go through species invading, settling and migrating. Based on the ratios of top area/bottom area (Zhu et al. 2003b), the Dashiwei Tiankeng group was divided into three stages: origin, competition and stable stages (Fan 2014; Feng 2015). They found that, in the origin stage of tiankeng development, the top area was smaller than the bottom, the sunlight was dimmer and air humidity higher than for the outside surface. The plant community mainly comprised shade-tolerant species, such as bryophytes, ferns, herbs and shrub species (Fan 2014; Feng 2015). The first stage of flora begins as bryophytes settle and ends as plants adapt to temperature and moisture levels in the environment that were higher than that of the surrounding landscape (Feng 2015). Although the temperate genus proportion is much higher than at the other tiankeng stages, the importance of these genera and species is very low (Feng 2015). In the competition stage of tiankeng development, with the increase in sunlight, humidity and soil thickness, more shade-tolerant tree species occur in the plant community (Fan 2014; Feng 2015). In the stable stage of tiankeng development, the plant community is mainly composed of heliophilous species owing to its adequate sunlight, drought bottom and the flat terrain (Fan 2014; Feng 2015).

Plant Biodiversity

Plant diversity of karst tiankengs plays an important role in maintaining biodiversity, ecosystem function and resilience in producing landscapes in karst regions, and is one of the most important parts of a karst tiankeng ecosystem. Plant biodiversity varies with the degree of maturity and the size of karst tiankengs, and is possibly related to habitat heterogeneity and diversity. Recently, some studies indicated positively correlation between plant richness and area of karst tiankengs, similar to those found on island systems (Fan 2014; Feng 2015). Compared with the closely related surrounding karst positive landforms, the karst tiankeng ecosystem possessed more vegetation cover, bearing capability and endemic species because of its relatively closed micro-environment. The main prominent feature of karst tiankengs' plant communities was their higher plant diversity which varied with different habitat environments, lower similarity and poor reproducibility. The vertical band spectrum of vegetation also showed its own unique variation law.

In recent years, some researchers have studied plant species composition, structure, succession tendency and plant reproduction in different tiankengs and outside of them (Borrero et al. 2016; Fan et al. 2014; Feng et al. 2015; He et al. 2004; Huang et al. 2011; Huang and Su 2015; Jiang 2002; Lin et al. 2005; Liu et al. 2009; Su et al. 2011; Su et al. 2012). They found that plant diversity in the karst tiankeng region was characterized by rich species diversity, ancient origin and the characteristics of secondary vegetation. For example, in Dashiwei Tiankeng groups and outside of the surface of them (Fig.3), there were more than 874 plants that included 227 pteridophyta, 7 gymnosperms and 640 angiosperms recorded (Fan et al. 2014; Feng et al. 2015; He et al. 2004; Huang and Su 2015; Su et al. 2011). Among these, there were 784 species having been identified, which were collected from Dashiwei Tiankeng groups (including Dashiwei Tiankeng, Baidong Tiankeng, Banyuedong Tiankeng, Mafengdong Tiankeng, Meiguidating Tainkeng, Shengmu Tiankeng, Sujia Tiankeng, Luojia Tiankeng, Chuandong Tiankeng, Dacao Tiankeng, Dengjiatuo Tiankeng, Liuxing Tiankeng) in Guangxi, China (Table 1). Because of its special ecological environments, there were 73 species which were found only lived in the twelve tiankengs but not found outside of these tiankengs (Table 1). In the Tiankeng-Difeng regions (Fig.4), 2107 higher plants of 176 families and vegetation including forests, bushland, grassland and farmland, cover about 80% of the land area (Jiang 2002). Among these species, there are 32 species of rare, threatened and endemic plants (Jiang 2002).

Based on the vegetation investigation and the data from sampling survey to nine tiankengs from Dashiwei Tiankeng groups (Fig. 3), Lin (2004) found that these plant species can be mainly classified into 5 types, including the evergreen-deciduous broadleaf-mixed forest, the lower subtropical evergreen broadleaf forest, the warm coniferous forest, the deciduous broadleaf forest and the open forest. He also found that the species diversity was always ranked the highest level in the lower subtropical evergreen broadleaf forest. Furthermore, differed from the vertical zonation of the vegetation in a typical mid-subtropical zone, these Tiankeng forests presented its own unique variation law. For example, the vertical zonation of the vegetation in the Tiankeng forests is, from the top to down, the warm coniferous forest (1400-1300 m altitude), the evergreen-deciduous broadleaf-mixed forest (1360-1120 m altitude), the lower subtropical evergreen broadleaf forest (1240-900 m altitude), or the deciduous broadleaf forest (1150-1045 m altitude), or the open forest (1155-1095 m altitude). In addition, he also found that the topography characteristics, human disturbance and moisture were the most important factors which effected on the community diversity. It is a pity that the researches don't analysis the differences between different tiankengs since the diversity index shows a great difference in the different forest communities.

On the basis of Lin's research, Fan (2014) and Feng (2015) investigated and analysised the the composition and structure of plant species from the twelve tiankengs of Dashiwei Tiankeng groups (Fig.3). They found that the vegetation in the origin stage Tainkengs was mainly composed by herb species (105 species belonging to 80 genera and 50 families) and the life form spectrum of the herb constituted up to 58%. The vegetation in the competition stage (246 species belonging to 185 genera and 92 families) and the stable stage (206 species belonging to 155 genera and 78 families) are mainly composed by tree species, and tree as a life form constituted up to 44% and 38% respectively. They also found that both slop position and aspect can significantly affect the plant community in these Tiankengs (Fan 2014; Feng 2015). Furthermore, in the the competition stage, the biodiversity of the plant species is the greatest (Fan 2014).

In addition, Feng (2015) has analysed the georgraphical distribution of plant species in these tiankengs. The results showed that the genera (from Shenmu Tianken, Sujia Tiankeng and Luojia Tiankeng) of tropical distribution account for 70%, which is 10% higher than for the surface ground flora, and the temperate genus proportion (from Meigui Tiankeng, Banyuedong Tiankeng and Baidong Tiankeng) is much higher than the other Tiankeng flora stages, about 50%in all of genera. Although these researches have investigated the effects of topography characteristics, human disturbance, moisture, slop position and aspect on the biodiversity of plant species in tiankengs, it does not take into account the effects of soil characteristics, such as, soil physical and chemical properties, and soil microorganism, on the biodiversity of plant species despite the strong relation between soil and plant community (Peay et al. 2013; Sheng et al. 2015).

Animal Diversity

Animals are an active and appealing component in ecosystems and animal resources are rich in karst tiankeng regions, given the unique environments with numerous mountain streams, karst caves and richness of plant species. For example, 449 vertebrates were found living in the Tiankeng-Difeng region (Fig. 4), including 135 fishes, 24 amphibians, 34 reptiles, 186 birds and 60 mammal species (Jiang 2002). Among these, there are 60 species of endangered animals in the Tiankeng-Difeng region (Jiang 2002). There are 59 animals species distributed in the Dashiwei Tiankeng group and the surface ground of tiankengs (Deuve 2002; Huang et al. 2004; Tong and Li 2007; Zeng and Su 2012, 2013; see also Fig. 3), and this includes 17 amphibian (Zeng and Su 2013) and 42 reptile species (Zeng and Su 2012). Furthermore, more than 14 animal species was only found living in the tiankengs, including two new species, Belisana zhangi sp. nov. (Tong and Li 2007) and Giraffaphaenops clarkei Deuve (Deuve 2002), and 12 rare, threatened or endangered species (Table 2).

Landscape Ecology of Karst Tiankengs

Landscape Value

With their peculiar collapse landform, diversity of surrounding landscapes, uniqueness of ecological environment and exhibition of successive stages of geological development, karst tiankengs are valuable natural resources (Huang et al. 2004; Zhu and Chen 2005). Up to now, two karst landscapes containing tiankengs have been listed in world geological parks: Xingwen World Geological Park and Leye-Fengshan Geopark (Deng et al. 2012; Wang 2012; Xu et al. 2009; Yu et al. 2009). The karst landscapes featured including high fengcong, large caves, large karst gorges, large tiankeng group, large natural bridges and subterranean rivers (Peng et al. 2006; Waltham 2005a, b; Zhu and Cheng 2005). These karst landscapes are recognized as tourist resource fairly recently, but they are remarkable because they make a strong impression upon the viewers, and are already considered a high quality tourism resource (Huang et al. 2004; Yue et al. 2008). Using field surveys, aerial photography, GIS technology, and other analytical tools some researchers have attempted to estimate the monetary and non-monetary values of tiankeng landscapes for science and tourism (Bai et al. 2010; Chen et al. 2009; Huang et al. 2004; Peng et al. 2006; Zhu 2001).

Recent Human Exploitation of the Tiankengs

The discovery and research of karst tiankengs has promoted their exploitation as tourism resources. For example, using the theory of organizational ecology, some research has proposed a model of integrated development in karst tiankeng tourism, producing both small-group exploration-based tourism and more generalized, mass tours (Wei et al. 2011). Besides physical exploration of the tiankengs, building 3D virtual reality as a supplemental tourist activity was also suggested by some (Huang et al. 2004; Wang 2012). Finally, developing ecotourism is also recommended (Liu 2006; Sun 1998). At the same time, owing to the non-renewable nature and vulnerability of karst landscapes, sustainable exploitation of karst landscapes is bound to become an important research topic. However, few studies have focused on sustainable exploitation and notably, studies on the potential impacts of the tourism development on endangered species reside in tiankengs are lacking.

Effects of Environmental Changes on Karst Tiankeng Ecosystems

Karst ecosystems are restrained by the karst environment, including how karst environments affect life and the reaction of life to the karst environment (Yuan 2001). For thousands of years, unique natural microclimates formed in karst tiankeng environments, and the wildlife of these environments cannot be recovered once lost (Fan 2014; Feng 2015; Shui et al. 2015). Therefore, tourism development of karst tiankengs must pay attention to ecological carrying capacity and environmental sensitivity. For example, tourists have strong impacts on the carbon dioxide concentrations in lateral unventilated caves (Shui and Su 2011). Although Tian (2005) and Wang (2012) tested carrying capacity of some karst tiankengs, knowledge of ecological carrying capacity in karst tiankengs is limited.

In addition to the carrying capacity of karst tiankengs, changes in the external environmental, such as global warming, acid rain and nitrogen deposition cannot be ignored. Previous studies indicated that as a consequence of their unique geomorphological structure, karst tiankengs can have 'cold trap effects', which enrich organic pollutants in soil at the bottom of karst tiankengs (Kong 2012; Kong et al. 2011; 2012; 2013; Oramah et al. 2008; Wang et al. 2009). Persistent organic pollution can be transported among some tiankengs, as found by Kong (2012) from a study of source, distribution, environmental behavior and influencing factors of persistent organic pollution in the Dashiwei Tiankeng group, through the geological and hydrological system. Although there have been many studies on the effects of organic pollutants on karst tiankengs, little is known about the effects of contaminants on organisms in karst tiankeng ecosystems.

Major Problems

Lack of a Consensus on a Basic Theory in Tiankeng Geological Evolution

In the past 10 years, karst tiankengs especially in China have received more attention from geomorphologists and geologists due to their unique value for scientific research, and tourism, as well as designation of some tiankengs as world natural heritage sites and global geological heritage (Klimchouk 2005; Palmer and Palmer 2005; Shui et al. 2015; Waltham 2005a, b, c; White and White 2005; Zhu et al. 2003b; Shui and Su 2011). Unfortunately, these studies mainly focused on the macro qualitative, empirical analysis, descriptive induction and hypothesis inference. The scientific evidence of the geological evolution process and formation of karst tiankengs is still insufficient, which prevents a good understanding in their evolutionary mechanisms (Shui et al. 2015). This in turn hinders our understanding in the formation and evolution of the tiankeng ecosystem as a whole.

Although there have been many studies on karst tiankeng ecosystems, they have mainly focused on investigating animal and plant resources (Fan 2014; Feng 2015; Huang et al. 2004; Huang et al., 2011; Jiang 2002; Lin 2005; Liu et al. 2009; Shui et al. 2015; Su et al. 2011; Su et al. 2012; Tong and Li 2007; Zeng and Su 2012, 2013). In these unique ecosystems, the distribution and composition of plants and animals in karst tiankengs are quite different from adjacent external environments. Correspondingly, microorganism communities in karst tiankeng ecosystems may greatly differ from the related external environments since soil microorganisms and plants are dependent on each other. Microorganisms play important roles in material and energy flows of ecosystems and consequently it is unfortunate that little is known about soil microorganisms in karst tiankeng ecosystems.

Lack of Systematic Research

There are two significant concerns regarding the current state of research on karst tiankengs. First, karst tiankeng studies are scattered and have limited integration with one another. Each discipline focuses on its professional area, so studies lack collaboration and innovation. For example, geologists are more concerned with geological structure, lithology and structural characteristics of karst tiankengs; geomorphologists are mainly concerned with the rock gravity collapse process, hydraulic erosion and erosion; and biologists focus on the rich species and biodiversity. There are no studies that seek to integrate these different foci.

Second, most research has focused on individual case and there is lack of systematic and co-ordinated studies of tiankengs in China and elsewhere. For example, foreign scholars tend to have a profound understanding of tiankengs in their research region, but lack good investigations and understanding of those in China. Similarly, much research from China has focused on karst tiankengs found in China. As a result, systematic research on karst tiankengs is limited, which has prevented its development.

Research Prospects

We see the need to strengthen multi-disciplinary systematic study. Multi-subject participation and integration in studies of karst tiankengs can provide better conditions for the use of new methods and technologies. Previous studies have failed to combine research on geomorphologic changes, vegetation succession and environmental changes related to human activity despite their possible interactions. In addition, comparative studies among karst tiankengs from different geographic areas of the world may be important to understand their formation and evolution mechanism since the formation of different karst tiankengs may be highly dependent on the surrounding environment.

Karst tiankengs are fragile features, which are easily disturbed by natural disasters, climate change and human activities. Once a karst tiankeng is destroyed by human activites, it can't be restored, and its loss will lead to a decline in biodiversity and produce accumulative negative environmental effects on the whole karst tiankeng ecosystem. To this end, it is important to explore the effects of the coupling of natural and human activities on karst tiankeng ecosystems. Especially, in recent decades, global climatic change (e.g., warming, chemical pollutants, nitrogen deposition and acid rain) caused by human activities has been a major concern around the world. These factors have substantial impacts on ecosystem processes, especially carbon and nitrogen cycles in terrestrial ecosystems. However, little is known about their effects on karst tiankeng ecosystems despite that some organic pollutants distribution in soil, water and air of Dashiwei tiankeng have been well knowned. This is especially so concerning soil microorganisms, which are the foundation of the earth's biosphere, and play important roles in the cycling of carbon, nitrogen, sulfur and phosphorus, as well as of various metals. Furthermore, owing to their high degree of selectivity and adaptability to environmental factors, soil microorganisms can adapt to environmental changes by adjusting their physiological structure or changing the community structure. Unfortunately, little is known about their composition, diversity and contribution concerning karst tiankeng ecosystems.

Finally, it is essential that we achieve ecosystem protection and sustainable use and management of the tiankengs. To do this, we must identify their distribution and classification, and assess their environmental quality. It is also highly desirable to establish an information management system for karst tiankeng ecosystems, and systematic monitoring of biodiversity changes in the karst tiankeng ecosystems

DOI: 10.1007/s12229-017-9179-0

Acknowledgements This study was supported by the Natural Science Foundation of Guangxi Province (2015GXNSFEA139001; 2015GXNSFAA139072), Guangxi Scientific and Technological Projcct (Guikezhongl 598014-3) and the Fund of Guangxi Key Laboratory of Plant Conservation and Restoration Ecology in Karst Terrain (GKB15-A-33), the Natural Science Foundation of Shandong Province (ZR2015CL014) and the Fundamental Research Funds for the Guangxi Institue of Botany (Guizhiye, 15004). We thank Dr. Hong Liu. International Center for Tropical Botany, Florida International University, U. S. A., for her valuable comments and sugestions during manuscript preparation. We also want to thank three anonymous reviewers and Dr. Elizabeth Hamblin for the comments and suggestions made on earlier versions of the manuscript.

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Gaozhong Pu (1,3) * Yanna Lv (2) * Guangping Xu (1) * Danjuan Zeng (1) * Yuqing Huang (1)

(1) Guangxi Key Laboratory of Plant Conservation and Restoration Ecology in Karst Terrain,Guangxi Institue of Botany, Guangxi Zhuang Autonomous Region and Chinese Academy of Sciences, No.85, Yanshan Street, Yanshan District, Guilin City, Guangxi Province 541006, China

(2) School of Pharmacy and Biological Sciences, Weifang Medical University, No.7166, Baotang street (west), Weicheng District, Shandong Province, Weifang City 261053, China

(3) Author for Correspondence; e-mail: pukouchy@ 163.com

Published online: 2 March 2017

Caption: Fig. 1 Plant community of Dashiwei Tiankeng in Guangxi, China

Caption: Fig. 2 The distribution of karst tiankeng in China

Caption: Fig. 3 Map of Leye county showing the study site of Dashiwei Tiankeng groups in Guangxi (Cite from Zhu et al. 2003b)

Caption: Fig. 4 Map of Fengjie county showing the study site of Tiankeng-Difeng regions in Chongqin (Cited from Chen et al. 2004 and Jiang 2002)
Table 1 A list of plant spccies in the twelve tiankengs from Dashiwei
Tiankeng groups in Guangxi, China

Family             Species                                      Areal-
                                                                types
                                                                  of
                                                                genera

Pteridophyta       Asplenium nidus L.                             1
Aspleniaceae       A. sarelii Hook.
                   A. saxicola Roscnstock
                   A. tenuifolium D. Don
                   A. varians Wallich ex Hooker & Greville
Athyriaceac        Anisocampium niponicum (Mettenius)             7
                     Yea C. Liu, W. L.Chiou & M. Kato
                   Athyrium vidalii (Franchet & Savatier)         1
                   Deparia boryana (Willdcnow) M. Kato            4
                   D. lancea (Thunbcrg) Frascr-Jenkins
                   D. okuboana (Makino) M. Kato
                   Diplazium donianum (Mettenius) Tardieu         3
                   D. megaphyllum (Baker) Christ
                   D. prolixum Rosenstock
                   D. viridissimum Christ
Blechnaceae        Blechnum orientale L.                          2
                   Woodwardia japonica (L. f.) Smith              1
                   W. unigemmata (Makino) Nakai
Cyatheaeeae        Alsophila spinulosa (Wallich ex                2
                     Hooker) R. M. Tryon
Davalliaceac       Araiostegia perdurans (Christ) Copeland        14
                   Humata griffithiana
                     (Hooker) C. Christensen
Dcnnstaedtiaccae   Microlepia hancei Prantl                       4
                   M. marginata (Panzer) C. Christensen
                   M. marginata var. villosa
                     (C. Presl) Y. C. Wu
                   M trichocarpa Hayata
Dryopteridaeeae    Arachniodes amabilis (Blume) Tindale           2
                   A. assamica (Kuhn) Ohwi
                   A. chinensis (Rosenstock) Ching
                   * Ctenitis subglandulosa (Hance) Ching         2
                   Cyrtomium nephrolepioides                      6
                     (Christ) Copeland
                   C. tsinglingense Ching & K. H. Shing
                   C. caryotideum (Wallich ex Hooker &
                     Greville) C. Presl
                   Dryopteris assamensis (C. Hope)                1
                     C. Christensen &Ching
                   D. atrata (Wallich ex Kunzc) Ching
                   D. bodinieri (Christ) C. Chr istensen
                   D. scottii (Beddomc) Ching
                     ex C. Christensen
                   Polystichum balansae Christ
                   P. craspedosorum (Maximowicz) Diels
                   P. deltodon (Baker) Diels
                   P. excellens Ching
                   P. fimbriatum Christ
                   P. makinoi (Tagawa) Tagawa
                   P. tenuius (Ching) Li Bing Zhang
Hymenophyllaceae   * Hymenophyllum badium Hooker & Greville
                   * H. barbatum (Bosch) Baker
Hypodematiaceae    Hypodematium crenatum                          6
                     (Forsskal) Kuhn & Decken
Lindsaeaccae       Lindsaea orbiculata                            2
                     (Lamarck) Mettenius ex Kuhn
                   Odontosoria chinensis (L.) J. Smith
Lycopodiaceae      Lycopodium cernuum L.                          1
                   Phlegmariurus phlegmaria (L.) Holub            2
Lygodiaceae        Lygodium japonicum (Thunberg) Swartz
                   L. micmphyllum (Cavanilles) R. Brown
Polypodiaceae      Arthromeris lungtauensis Ching                 14
                   Drynaria roosii Nakaike                        5
                   Lemmaphyllum carnosum                          14
                     (Wallich ex J. Smith) C. Presl
                   L. drymoglossoides (Baker) Ching
                   L. micmphyllum var. micmphyllum
                   Lepidomicrosorium buergerianum (Miquel)
                     Ching & K.H. Shing ex Xu
                   Lepisorus contortus (Christ) Chmg              6
                   L. thunbergianus (Kaulfuss) Ching
                   Leptochilus decurrens Blume                    7
                   * L. henryi (Baker) X. C. Zhang
                   L ellipticus (Thunbeig) Nooteboom
                   L. ellipticus var. longipes
                     (Ching) Nooteboom
                   Loxogramme chinensis Ching                     1
                   Micmsorum membranaceum (D. Don) Ching          12
                   Neolepisorus ovalus                            6
                     (Wallich ex Beddome) Ching
                   N.fortunei (T. Moore) Li Wang
                   Phymatosorus cuspidatus                        4
                     (D. Don) Pichi Sermolli
                   Polypodiodes amoena (Wallich ex Mettenius)     14
                   Pyrrosia lingua (Thunberg) Farwell             1
                   P. tonkinensis (Giesenhagen) Ching
Psilotaceae        * Psilotum nudum (L.) P. Beauvois              2
Pteridaceac        Aleuritopteris argentea (S. G. Gmelin) Fee     1
                   Adiantum. caudatum L.
                   A. flabellulatum L.
                   Antmphyum obovatum Baker                       6
                   Cheilanthes nitidula Wallich ex Hooker         2
                   Coniogramme itermedia var. intermedia
                   Haplopteris flexuosa (Fee) E. H. Crane
                   Pteris cretica var. laeta (Wallich
                     ex Ettingshausen) Christensen & Tardieu
                   P. decrescens Christ
                   P. deltodon Baker
                   P. ensiformis var. merrillii
                     (C. Christensen ex Ching)S. H. Wu
                   P. esquirolii Christ
                   P. henryi Christ
                   P. insignis Mettenius ex Kuhn
                   P. semipinnata L.
                   P. splendida var splendida
                   P. vittata L.
Selaginellaceae    Selaginella delicatula (Desv.) Alston          1
                   S. doederleinii Hieron.
                   S. helferi Warb.
                   S. moellendorffii Hieron.
                   S. uncinata (Desvaux ex Poiret) Spring
Tectariaceae       Tectaria devexa (Kunzc) Copeland               2
                   T. phaeocaulis (Rosenstock) C. Christensen
                   T. simonsii (Baker) Ching
                   T. subtriphylla (Hooker & Amott) Copeland
Thelypteridaceae   Cyclosorus acuminatus (Houttuyn) Nakai
                   C. aridus (D. Don) Ching
                   C. parasiticus (L.) Farwell
                   Parathelypteris glanduligera (Kunze) Ching
                   Phegopteris decursive-pinnata (Hall) Fee       14
                   Pronephrium penangianum (Hook.) Holtt.         7
                   P. gymnopteridifrons (Hayata) Holttum
Spemaiophyla
Cephalotaxaccae    Cephalotaxus fortunei Hooker                   14
Cupressaccae       Fokienia hodginsii (Dunn) A. Henry             7
                     & H. H. Thomas
Pinaceae           Pinus kwangtungensis Chun & Tsiang             8
                   P. massoniana Lambert
                   Pseudotsuga brevifolia W. C.                   9
                     Cheng & L. K. Fu
Taxaceae           * Taxus wallichiana var. mairei                8
                     (Lemee & H. Leveille) L. K. Fu & Nan Li
Taxodiaceae        Cunninghamia lanceolata (Lambert) Hooker       15
Schisandraccae     Schisandra henryi subsp. henryi                9
Aristolochiaceae   * Asarum caudigerum Hance                      8
Piperaceae         Pepemmia blanda (Jacquin) Kunth                2
                   * Piper betle L.
                   * P. bonii C. de Candolle
                   * P. sintenense Hatusima
                   * P. wallichii (Miquel) Handel-Mazzetti
Saururaceae        Gymnotheca chinensis Decaisnc                  15
                   Houttuynia cordata Thunberg                    7
Annonaceae         * Fissistigma glaueescens (Hance) Merrill      4
                   * F. retusum (H. Leveille) Rehder
                   * F. wallichii (Hooker & Thomson) Merrill
                   * Miliusa sinensis Finet & Gagnepain           5
Magnoliaceae       * Manglietia aromatica Dandy                   7
                   * Michelia martini (H. Leveille) Finet
                     & Gagnepain ex H. Leveille
Illiciaceac        Illicium difengpi K. I. B.                     9
                     et K. I. M. ex B. N. Chang
Lauraceac          * Actinodaphne kweichowensis                   7
                     Yen C. Yang & E. H. Huang
                   Beilschmiedia kweichowensis W. C. Cheng        2
                   Cinnamomum burmannii                           3
                     (Nees & T. Nees) Blume
                   C. pingbienense H. W. Li
                   C. saxatile H. W. Li
                   Cryptocarya concinna Hance                     7
                   Lindera erythrocarpa Makino                    9
                   L. glauca (Siebold & Zuccarini) Blume
                   L. megaphylla Hemsleyey
                   L. metcalflana var. dictyophylla
                     (C. K. Allen) H. P.Tsui
                   L. pulcherrima var
                     hemsleyana (Diels) H. P. Tsui
                   Litsea elongata (Nees) J. D. Hooker            3
                   L. mollis Hemsley
                   Machilus austroguizhouensis                    7
                     S. K. Lee & F. N. Wei
                   M. calcicola C. J. Qi
                   M. glaucifolia S. K. Lee et F. N. Wei
                   M. multinervia H. Liu
                   M. oreophila Hance
                   Neolitsea cambodiana Lecomte
                   N. confertifolia (Hemsley) Merrill
                   N. homilantha C. K. Allen
                   Phoebe neurantha (Hemsley) Gamble
Chloranthaccae     *Chloranthus multistachys Pei
                   *C. sessilifolius K. F. Wu
                   Sarcandra glabra (Thunberg) Nakai
Araceae            Amorphophallus dunnii Tutcher                  4
                   A. variabilis Blume
                   Arisaema decipiens Schott                      8
                   A. franchetianum Engler
                   A. heterophyllum Blume
                   Pinellia ternata (Thunberg)                    14
                   Tenore ex Breitenbach
                   * Pothos chinensis (Rafincsque) Merrill        5
Burmanniaceae      * Burmannia disticha L.                        2
Dioscoreaceae      Dioscorea collettii var hypoglauca
                     (Palibin) C. T. Ting et al.
                   D. exalata C. T. Ting & M. C. Chang
                   D. fordii Prain & Burkill
                   D. hemslevi Prain & Burkill
                   D. velutipes Prain & Burkill
Stemonaceae        Stemona tuberosa Loureiro                      5
Liliaceae          Aletris spicata (Thunberg) Franchet            9
                   Allium macrostemon Bungc                       8
                   Asparagus cochinchinensis                      4
                     (Loureiro) Merrill
                   A. lycopodineus (Baker) F. T. Wang & Tang
                   Campylandra wattiiC. B. Clarke                 14
                   * Disporopsis aspersa (Hua) Engler             7
                   Disporum sessile D. Don                        14
                   Lilium brownii F. E. Brown ex Miellez          8
                   L. nepalense D. Don
                   Liriope graminifolia (L.) Baker                14
                   L. spicata (Thunberg) Loureiro
                   Ophiopogon chingii Wang & Tang
                   O. sparsiflorus F. T. Wang & L. K Dai
                   Polygonatum cyrtonema Hua                      8
                   P. kingianum Collett & Hemsley
                   * Reineckea carnea (Andrews) Kunth             14
                   * Paris cmnquistii (Takhtajan) H. Li           10
                   * P. fargesii Franchet
                   Smilax basilata F. T. Wang & Tang              2
                   S. chapaensis Gagnepain
                   S. ocreata A. de Candolle
Orchidaccae        Cymbidium faberi Rolfe                         5
                   C. jloribundum Lindley
                   C. lancifolium Hooker
                   C. mastersii Griffith ex Lindley
                   C. seiratum Schlechter
                   C. tracyanum L. Castle
                   Cvpripedium henryi Rolfe.                      8
                   Epipactis helleborine (L.) Crantz
                   Liparis chapaensis Gagnepain                   1
                   L. distans C. B. Clarke
                   L. nervosa (Thunberg) Lindley
                   L. giganlea C. L. Tso
                   L. pauliana Handel-Mazzetti
                   L. viridiflora (Blume) Lindley
                   * Odontochilus elwesii C. B.                   7
                     Clarke ex J. D. Hooker
                   Panisea cavaleriei chlechtcr                   14
                   Paphiopedilum hirsutissimum                    7
                     (Lindley ex Hooker) Stein
                   P. micranthum Tang & F. T. Wang
                   Peristylus goodveroides (D. Don) Lindley       5
                   Pholidota yunnanensis Rolfe.
                   Tropidia angidosa (Lindley) Blume              3
                   * Zeuxine gengrnanensis                        4
                     (K. Y. Lang) Ormerod
Arecaceae          Guihaia argyrata (S. K. Lee & F. N. Wei)       7
                     S. K. Lee. N. Wei & J. Dransfield
Musaceac           Musa acuminata Colla                           5
                   Musella lasiocarpa                             15
                     (Franchet)C. Y. Wu ex H. W. Li
Zingibcraceae      Alpinia brevis T. L. Wu & S. J. Chen           5
                   A.japonica (Thunberg) Miquel
Cypcraccac         Care.x aita Boott                              1
                   C. baccans Nees
                   C. brunnea Thunberg
                   C. cruciata Wahlenberg
                   C. fargesii Franchet
                   C. hattoriana Nakai ex Tuyama
                   C. ischnostachya Steudel
                   C. ligulata Nees
                   C. perakensis C. B. Clarke
                   C. pruinosa Boott
                   C. stipitinux C. B. Clarke ex Franchet
                   C. tristachya var. pocillifonnis
                     (Boott) Kukenthal
                   Cyperus iria L.
                   C. pilosus Vahl
                   C. rotundus L.
                   Eriophorum comosum Nees                        8
                   Fimbristylis bisumbellata                      2
                     (Forsskal) Bubani
                   F. complanata vai exalata (T. Koyama)
                     Y. C. Tang ex S. R. Zhang & T. Koyama
                   F. disticha Boeckelcr
                   F. littoralis var. littoralis
                   F. thomsonii Bocckeler
                   Fuirena umbellata Rottboll
                   Kyllinga brevifolia Rottboll
                   K. brevifolia vat leiolepis
                     (Franchet & Savatier) H. hara
                   Rhynchospora rubra (Loureiro) Makino           1
                   Scleria levis Retzius                          2
                   S. parvula Steudcl
Poaccae            Arthraxon hispidus (Thtmberg) Makino           4
                   Arundinella decempedalis (Kuntze) Janowski     2
                   A. setosa Trinius
                   Bothriochloa bladhii (Retzius) S. T. Blake
                   Chimonobambusa quadrangularis                  14
                     (Franceschi) Makino
                   Cynodon dactvlon (L.) Persoon                  2
                   Cyrtococcum patens var.                        4
                     latifolium (Honda) Ohwi
                   Digitaria ischaemum (Schrcbcr) Muhlenberg      2
                   D. longiflora (Retzius) Persoon
                   D. ternata (Hochstetter ex
                     A. Richard) Stapf
                   Eragrostis japonica (Thunberg) Trinius
                   E. multicaulis Stsudel
                   E. pilosa (L.) P. Bcauvois
                   Eulalia pallens (Hackel) Kuntze                4
                   Festuca elata Keng ex E. B. Alexeev            1
                   F. parvigluma Steudel
                   Hackelochloa granulans (L.) Kuntze             2
                   Imperata cylindrica (L.) Racuschcl
                   Ischaemum barbatum Retzius
                   I. ciliare Retzius
                   Miscanthus floridulus (Labillardicre)          6
                     Warburg ex K. Schumann & Lauterbach
                   Paspalum thunbergii Kunth ex steudel           2
                   Phyllostachys nidularia Munro                  14
                   Pogonatherum crinitum (Thunberg) Kunth
                   P. paniceum (Lamarck) Hackel
                   Themeda quadrivalvis (L.) Kuntze               4
Berberidaceac      *Dysosma versipellis (Hance.)                  14
                     M. Cheng ex T. S. Ying
                   Epimedium acuminatum Franchet                  10
                   E. wushanense T. S. Ying
                   Mahonia bealei (Fortune) Carricrc              9
Lardizabalaccac    Akebia trifoliata (Thunberg) Koidzumi          14
                   Stauntonia cavalerieana Gagnepain
Menispermaceac     Cyclea sutchuenensis Gagnepain
                   Stephania kwangsiensis H. S. Lo                4
                   Tinospora sagittata (Oliv.) Gagnepain
Papaveraccac       Corydalis balansae Prain                       8
                   C. esquirolii H. Leveille
                   C. saxicola Bunting
Ranunculaceac      Anemone hupehensis (Lemoine) Lemoine           1
                   Clematis apiifolia de Candolle
                   C. chingii W. T. Wang
                   C. henryi Oliver
                   C. meyeniana Walp.
                   C. uncinata Champion ex Bentham
                   C. urophylla Franchet
                   Delphinium anthriscifolium Hance               8
                   Ranunculus cantoniensis de Candolle            1
                   Thalictrum ichangense Lecoyer ex Oliver        8
Sabiaccac          Meliosma fordii Hemsley                        3
                   Sabia dielsii H. Leveille                      7
                   S. japonica Maximowicz
                   S. pariflora Wallich
                   S. swinhoei Hemsley
Buxaccae           * Pachvsandra axillaris Franchet               9
                   * Sarcococca ruscifolia Stapf                  7
Crassulaccac       Sedum bulbiferum Makino                        8
                   S. stellariifolium Franchet
                   * Sinocrassula indica (Decaisne) A. Berger     14
Daphniphyllaceae   Daphniphyllum paxianum K. Rosenthal            7
Saxifragaccae      * Chrysosplenium delavayi Franchet             9
                   * C. hydrocotylifolium
                     H. Leveille & Vaniot
                   * Deutzia setchuenensis Franchet
                   * Saxifraga stolonifera Curtis                 8
Vitaceac           Parthenocissus dalzielii Gagnepain             9
                   P. tricuspidata (Siebold
                     & Zuccarini) Planchon
                   Tetrastigma obtectum (Wallich ex M.            5
                     A. Lawson) Planchon ex Franchet
                   Vitis hevneana Roomer & Schultcs               8
Fabaceae           Bauhinia touranensis Gagnepain                 2
                   Callerya nitida var nitida                     5
                   Campylotropis trigonoclada                     11
                     (Franchet) Schindler
                   Dalbergia assamica Bentham                     2
                   Derris cavaleriei Gagnepain
                   Desmodium multiflorum Candolle                 3
                   * Dumasia villosa Candolle                     4
                   Hylodesmum podocarpum subsp. oxyphyllum        9
                     (Candolle) H. Ohashi & R. R. Mill
                   Lespedeza thunbergii subsp. formosa
                     (Vogel) H. Ohashi
                   Pueraria montana var. lobata                   7
                     (Willdenow) Maesen & S.M. Almeida
                     ex Sanjappa & Prcdeep
Polygalaceae       Polygala arillata                              1
                     Buchanan-Hamilton ex D. Don
                   P. caudata Rehder & E. H. Wilson
                   Salomonia cantoniensis Loureiro                5
Elaeagnaceac       Elaeagnus bockii Diels                         8
                   E. difficilis Servettaz
                   E. glabra Thunberg
Moraceae           Broussonetia papvrifera(L.)                    7
                     L. Heritier ex Ventenat
                   Ficus fistulosa Reinwardt ex Blume             2
                   F. tikoua Bureau
                   Maclura cochinchinensis (Loureiro) Corner      1
                   Moms mongolica (Bureau) C. K. Schneider        8
Rhamnaceac         Berchemia edgeworthii M. A. Lawson             9
                   B. lineata (L.) Candolle
                   Rhamnus crenata Siebold & Zuccarini            8
                   R. esquirolii H. Levcille
                   R. Julvotincta F. P. Metcalf
                   R. henryi C. K. Schneider
                   R. lamprophylla C. K. Schneider
                   R. longipes Merrill & Chun.
                   R. napalensis (Wallich) M. A. Lawson
                   R. rosthornii E. Pritzel
                   R. wilsonii C. K. Schneider
                   R. wumingensis Y. L. Chen & P. K. Chou
Rosaceac           Cerasus campanulata
                     (Maxim owicz) A. N. Vassiljeva
                   C. conradinae (Kochne) T.
                     T. Yii et C. L Li
                   C. cyclamina (Kochne) T. T. Yu & C. L. Li
                   C. dielsiana (C. K. Schneider) T. T. Yii
                     & C. L. Li
                   C. yunnanensis (Franchet) T. T.
                     Yu & C. L. Li
                   Cotoneaster glaucophyllus Franchet             9
                   * Laurocerasus undulam (Buchanan-Hamilton      2
                     ex D. Don) M. Roemer
                   Pyrus callervana Decaisne                      10
                   P. serrulata Rehdcr
                   Photinia bodinieri H. Leveille                 9
                   Rubus amphidasys Focke                         1
                   R. adenophorus Rolfe
                   R. tsangorum Handel-Mazzetti
                   Spiraea kwangsiensis T. T. Yu
Ulmaceac           Celtis biondii Pampanini                       2
                   C. cerasifera C. K. Schneider
Urticaceae         Boehmeria clidemioides Miquel
                   B. macrophylla Homemann
                   B. nivea (L.) Gaudichaud-Beaupre
                   Debregeasia longifolia                         6
                     (N. L. Burman) Weddell
                   Elatostema albopilosum W T. Wang               4
                   E. balansae Gagnepain
                   E. brachyodontum W. T. Wang
                   E. coriaceifolium W. T. Wang
                   E. cyrtandrifolium (Zollinger
                     & Moritzi) Miquel
                   E. dissectum Weddell
                   E. ficoides Weddell
                   E. macintyrei Dunn
                   E. nanchuanense W. T. Wang
                   E. polystachyoides W. T. Wang
                   Gonostegia hirta (Blume ex                     5
                     Hasskarl) Miquel
                   Nanocnide lobata Weddell                       14
                   Oreocnide frutescens (Thunberg) Miquel         7
                   Pilea angulata (Blume)Blumc                    2
                   P. boniana Gagnepain
                   P. cavaleriei H. Leveille
                   P. elliptilimba C. J. Chen
                   P. hilliana Handel-Mazzetti
                   P. plataniflora C. H. Wright
                   P. pumila (L.) A. Gray
                   P. semisessilis Handel-Mazzetti
                   * Procris crenata C. B. Robinson               6
                   Urtica fissa E. Pritzel                        1
Betulaceae         Alnus trabeculosa Handel-Mazzetti              8
                   Betula alnoides
                     Buchanan-Hamilton ex D. Don
                   Carpinus tschonoskii Maximowicz
Fagaceae           Castanea mollissima Blume
                   Castanopsis fabri Hance                        9
                   C. hystrix J. D. Hooker
                     & Thomson ex A. de Candolle
                   Cyclobalanopsis austroglauca Y. T. Chang       7
                     ex Y. C. Hsu & H. W. Jen
                   C. glauca (Thunberg) Oersted
                   C.jenseniana (Handel-Mazzetti) W. C.
                     Cheng & T. Hong ex Q. F. Zheng
                   Lithocarpus corneus (Loureiro) Rehder          9
                   L. hancei (Bentham) Rehder
                   Quercus acutissima Carruthers                  8
                   Q. fabri Hance
                   Q. phillvreoides A. Gray
Juglandaccac       Cyclocarya paliurus (Batalin) lljinskaya       15
                   Engelhardia roxburghiana Wallich               7
                   Juglans regia L.                               8
                   Platycarya strobilacea Siebold & Zuccarini     14
                   Pterocarya stenoptera C. de Candolle           6
Begoniaccac        * Begonia parvula H. Leveille & Vaniot         2
Coriariaceac       Coriaria nepalensis Wallich                    8
Cucurbitaceac      Zehneria japonica (Thunberg) H. Y. Liu         4
                   Trichosanthes cucumeroides var.                5
                     dicaelosperma (Clarke) S. K. Chen
Celastraccac       Celastrus aculeatus Merrill.                   2
                   C. gemmatus Loosener.
                   C. monospermus Roxburgh
                   Euonymus euscaphis Handel-Mazzetti             1
                   E. echinatus Wallich in Roxburgh
                   E.fortunei (Turczaninow) Handel-Mazzetti
                   E. laxiflorus Champion ex Bentham
                   E. myrianthus Hemsley
Elaeocarpaccae     * Elaeocarpus japonicus                        5
                     Siebold & Zuccarini
Oxalidacoae        Oxalis corniculata L.                          1
                   0. griffithii Edgeworth & J. D. Hooker
Clusiaccac         Hypericum acmosepalum N. Robson
Euphorbiaccac      Antidesma bunius (L.) Spreng                   4
                   Glochidion daltonii                            2
                     (Miiller Argovicnsis) Kurz
                   G. hirsutum (Roxb urgh) Voigt
                   * Leptopus fangdingianus (P. T. Li)            5
                     Vorontsova & Petra Hoffmann
                   Mallotus apelta (Loureiro)                     4
                    Miiller Argoviensis
                   M. barbatus Miiller Argoviensis
                   M.japonicus (L. f.)Miiller Argoviensis
                   M. nepalensis Miiller Argovicnsis
                   M. repandus (Willdenow)
                     Miiller Argoviensis
                   * Mercurialis leiocarpa Sicbold                9
                     & Zuccarini
                   * Sauropus androgynus (L.) Merrill             5
Linaceac           Tirpitzia ovoidea Chun & F. C                  7
                     How ex W. L. Sha
Flacourtiaceae     Xylosma controversa Clos                       2
Passifloraceae     * Passiflora papilio H. L. Li
Thymelacaceae      Daphne papyracea Wallich ex Steudel            8
                   Wikstroemia indica (L.) C. A. Meyer            5
Violaceac          Viola kosanensis Hayata V. pilosa Blume        1
Geraniaccae        Geranium franchetii R. Knuth
                   G. nepalense Sweet
Mclastomataccae    Sarcopyramis bodinieri H. Leveille             7
                     & Vaniot
Myrtaccac          Syzygium buxifolium Hooker & Arnott            4
Stachyuraccac      * Stachyurus himalaicus J. D. Hooker           14
                     & Thomson ex Bentham
                   * S. yunnanensis Franchet
Staphylcaceae      Turpinia montana (Blume) Kurz                  3
Aceraccac          Acer cordatum var. cordatum                    8
                   A. davidii Franchet
                   A. tonkinense Lecomte
Anacardiaccae      Colinus coggvgria var.
                     glaucophylla C. Y. Wu
                   Pistacia chinensis Bunge                       12
                   Rhus chinensis var. chinensis                  8
                     (C. B. Clarke) Ridley
                   Toxicodendron succedaneum (L.) Kuntze          9
Hippocastanaceae   Handeliodendron bodinieri                      15
                     (H. Leveille) Rehder
Meliaccae          Cipadessa baccifera (Roth) Miquel              7
                   Melia azedarach L.                             4
                   Munronia pinnata (Wallich) W. Theobald         7
                   Toona ciliata M. Roemcr                        5
Rutaccac           Clausena dunniana H. Leveille                  4
                   Murraya euchrestifolia Hayata                  7
                   Skimmia reevesiana (Fortune) Fortune           14
                   Tetradium ruticarpum
                     (A. Jussieu) T. G. Hartley
                   Zanthoxylum echinocarpum Hemslcy               2
                   Z. esquirolii H. Leveille
                   Z. ovalifolium Wight
Simaroubaceae      Ailanthus vilmoriniana Dode                    5
                   Picrasma chinensis P. Y. Chen                  3
Sterculiaccae      * Reevesia pubescens Masters                   7
Brassicaceae       Capsella bursa-pastoris (L.) Medikus           1
                   Cardamine hirsuta L.
                   C. impatiens L
                   Descurainia Sophia (L.) Webb cx Prantl         8
                   * Neomartinella violifolia                     15
                     (H. Leveille) Pilger
                   Rorippa indica (L.) Hiem                       1
Loranthaceae       Taxillus levinei (Merrill) H. S. Kiu           7
Olacaccac          Erythropalum scandens Blume
                   * Malania oleifera Chun et S. K. Lee.          15
                   * Schoepfia jasminodora Siebold                3
                     & Zuccarini
Amaranthaceac      Achyranthes aspera L.                          2
                   Amaranthus tricolor L.                         1
Caryophyllaceae    Brachystemma calycinum D. Don                  14
                   Mvosoton aquaticum (L.) Moench                 10
                   Silene baccifera (L.) Roth                     8
                   Stellaria alsine Grimm                         1
                   S. media (L.) Villars
Phytolaccaceae     Phytolacca acinosa Roxburgh                    2
Polygonaccac       Fagopyrum esculentum Moench                    9
                   F. tataricum (L.) Gaertncr
                   Fallopia multiflora (Thunberg) Haraldson       8
                   Polygonum capitatum                            1
                     Buchanan-Hamilton ex D. Don
                   P. chinense L
                   P. longisetum Bruijn
                   P. molle D. Don
                   P. perfoliatum L.
                   P. plebeium R. Brown
                   P. posumbu Buchanan-Hamilton ex D. Don
                   Rumex japonicus Houttuyn
Alangiaceae        Alangium chinense (Loureiro) Harms             4
Comaceae           Cornus controversa Hemsley                     8
                   C. hongkongensis subsp. tonkinensis
                     (W. P. Fang) Q. Y. Xiang
                   * Helwingia japonica                           14
                     (Thunberg) F. Dietrich
Actinidiaccae      Actinidia callosa Lindley
                   A. fasciculoides C. F. Liang
                   A. henryi Dunn
                   A. callosa var. discolor C. F. Liang
                   A. nibricaulis Dunn
                   A. rufotricha C. Y. Wu
Balsaminaccac      * Impatiens chlorosepala Handcl-Mazzetti       2
                   * I. mengtszeana J. D. Hooker
Ebenaccac          Diospyros dumetorum W. W. Smith
                   D. kaki var. silvestris Makino
                   D. lotus L.
                   D. reticulinervis C. Y Wu D.
                     vunnanensis Rehder & E. H.Wilson
Ericaceae          Lyonia ovalifolia (Wallich) Drudc              9
                   Rhododendron mariesii Hemsley                  8
                     & E. H. Wilson
                   Vaccinium cuspidifolium C. Y-Wu
                     & R. C. Fang
Myrsinaceac        Ardisia alyxiifolia Tsiang ex C. Chen          2
                   A. caudata Hemsley
                   A. chinensis Bentham
                   A. ensifolia E. Walker
                   A. gigantifolia Stapf
                   A. pusilla A. de Candolle
                   Maesa hupehensis Rehdcr                        4
                   M. insignis Chun
                   Myrsine faberi (Mez) Pipoly & C. Chen          6
                   M. stolonifera (Koidz umi) E. Walker
Primulaccac        Androsace runcinata Handel-Mazzctti            8
                     A. umbellata (Loureiro.) Merrill
                   Lysimachia fooningensis C. Y Wu                14
                   L. insignis Hemsley
                   L. lobelioides Wallich
                   L. microcarpa Handcl-Mazzetti ex C. Y. Wu
                   L. navillei (H. Leveille) Handcl-Mazzetti
                   L. paridiformis Franchet
Sapotaccac         Sinosideroxylon pedunculatum                   7
                     war. pedunculatum
Styracaccae        Alniphyllum fortunei (Hemsley) Makino
                     Huodendron biaristatum var. biaristatum
Symplocaccae       Symplocos racemosa Roxburgh S. sumuntia        2
                     Buchanan-Hamilton ex D. Don
Thcaccac           Camellia sinensis (L.) Kuntze                  7
                   Eurya chinensis R. Brown                       3
                   E. saxicola Hung T. Chang
Apocynaceae        Alstonia yunnanensis Diels                     2
                   Alyxia schlechteri H. Leveille                 5
                   * Rauvolfia verticillata (Loureiro)            2
                     Baillon
                   Trachelospermum jasminoides (Lindley)          9
                     Lemaire
Asclcpiadaceae     Adelostemma gracillimum (Wallich ex            7
                     Wight) J. D. Hooker
                   Cynanchum atratum Bungc                        10
                   Dregea volubilis (L. f.) Bentham               6
                     ex J. D. Hooker
                   Hoya jungii Merrill                            7
                   H. lyi H. Leveille
                   Periploca forrestii Schlcchter                 6
                   * Tylophora silvestris Tsiang                  4
Aucubaccac         Aucuba himalaica J. D. Hooker & Thomson        14
Gcntianaceae       Gentiana rhodantha Franchet                    1
Loganiaccac        Buddleja asiatica Loureiro                     2
                   B. davidii Franchet
                   B. officinalis Maximowicz
                   * Gardneria multiflora Makino                  7
                   * Mitreola pedicellata Bentham                 2
Rubiaceac          Aidia pyenantha (Drake) Tirvengadum            4
                   Lasianthus formosensis Matsumura               2
                   L. japonicus Miquel
                   * Myrioneuron faberi Hemsley                   7
                   Ophiorrhiza cantonensis Hance
                   O.japonica Blume
                   Paederia foetida Linnaeus
                   * Pavetta hongkongensis Bremekamp              4
                   Rubia oncotricha Handel-Mazzetti               8
Boraginaccae       Bolhriospermum zeylanicum                      14
                     (J. Jacquin) Druce
                   Cynoglossum lanceolatum ForsskiH               8
                   Symphytum officinale L.                        10
                   Trigonotis funingensis H. Chuang               9
                   T. leyeensis W. T. Wang
Convolvulaceac     Cuscuta chinensis Lamarck                      2
                   Dinetus racemosus (Wallich) Sweet              7
                   Ipomoea purpurea (L.) Roth                     2
Solanaceae         Atropa belladonna L.                           12
                   Lycianthes bijlora (Loureiro) Bitter           3
                   Solanum americanum Miller                      1
Acanthaceac        Justicia austrosinensis H. S. Lo               2
                   J. latiflora Hemsley
                   Strobilanthes cusia (Nees) Kuntze              6
Bignoniaccae       Catalpa fargesii Bureau                        9
Gesneriaccae       * Briggsia longipes (Hemsley                   7
                     ex Oliver) Craib
                   Calcareoboea coccinea C. Y. Wu ex H.W. Li      15
                   * Chirita leyesienesis F. Wen                  7
                   * Didymocarpus glandulosus var. minor          6
                     (W. T. Wang) W. T. Wang
                   Hemiboea cavaleriei H. Leveille                14
                   H. cavaleriei var. paucinervis W. T. Wang
                     & Z.Y. Li ex Z. Y. Li
                   Lysionotus pauciflorus Maximowicz
                   Paraboea glutinosa (Handel-Mazzetti)           7
                     K. Y. Pan
                   P. rufescens (Franchct) B. L. Burtt
                   P. sinensis (Oliver) B. L. Burtt
                   * Petrocosmea martini (H. Leveille)
                     H. Leveille
                   * P. minor Hemsley
                   * Rhynchoglossum obliquum Blume                3
Lamiaccac          Clinopodium gracile (Bentham) Matsumura        8
                   Elsholtzia blanda (Bentham) Bentham            10
                   Hanceola sinensis (Hemsley) Kudo               15
                   Leucas ciliata Bentham                         4
                   * Paraphlomis gracilis var                     7
                     lutienensis (Sun) C. Y. Wu
                   Prunella vulgaris L.                           8
                   Salvia chinensis Bentham                       1
Oleaccac           Jasminum lanceolaria Roxburgh                  2
                   Ligustrum lucidum W. T. Aiton                  10
                   L. sinense Louteiro
                   Osmanthus Jragrans Loureiro                    9
Phrymaceac         Phryma leptostachya subsp.
                     asiatica (H. Hara) Kitamura
Plantaginaceac     Plantago asiatica L.                           1
Scrophulariaccae   Alectra avensis (Bentham) Merrill              6
                   Brandisia hancei J. D. Hooker                  7
                   Ellisiophyllum pinnatum (Wallich
                     ex Bentham) Makino
                   Mazus pulchellus Hemsley                       5
Vcrbenaccac        Callicarpa bodinieri H. Leveille               2
                   C. cathayana Chang
                   C. erythrosticta Merrill & Chun
                   C. giraldii Hesse ex Rehdcr
                   C. longissima (Hemsley) Merrill
                   C. longifolia var. longifolia
                   C. rubella var rubella
                   Clerodendrum chinense (Osbeck) Mabberley
                   C. fortunatum L.
                   C. mandarinorum Diels
                   C. serratum var. amplexifolium Mold-enke
                   Premna puberula var. bodinieri                 4
                     (H. Leveille) C. Y. Wu & S. Y. Pao
                   Vitex negundo var. cannabifolia (Siebold       2
                     & Zuccarini)
                   Handel-Mazzetti
Aquifoliaceac      Ilex macrocarpa Oliver
                   I. pentagona S. K. Chen et al.
Asteraceae         Ainsliaea ramosa Hemsley                       14
                   Blumea axillaris (Lamarck) Candolle            4
                   Carpesium abrotanoides L.                      10
                   Chrysanthemum indicum L.                       8
                   Cirsium racemiforme Y. Ling & C. Shih
                   Cissampelopsis spelaeicola (Vaniot)            7
                     C. Jeffrey & Y. L. Chen
                   Dichmcephala benthamii C. B. Clarke            4
                   Gynura bicolor (Roxburgh ex Willdenow)
                     Candolle
                   G. divaricata (L.) Candolle
                   G. procumbens (Loureiro.) Merrill
                   Senecio scandens Buchanan-Hamilton             1
                     ex D. Don
                   * Sinosenecio bodinieri (Vaniot)               14
                     B. Nordenstam
                   * S. oldhamianus (Maximowicz)
                     B. Nordenstam
Campanulaccac      Asyneuma chinense D. Y. Hong                   10
                   Campanula pallida Wallich                      8
                   Campanumoea javanica Blume                     7
                   Cyclocodon lancifolius (Roxbuigh) Kun
                   Lobelia heyneana Schults                       2
                   L. nummularia Lamarck
                   Wahlenbergia marginata (Thunbeig)
                     A. Candolle
Escalloniaccac     Itea chinensis Hooker                          9
                     & Arnott f. chniensis
                   I. glutinosa Handel-Mazzetti
                   I. indochinensis Merrill
                   I. yunnanensis Franchet
Adoxaceae          Sambucus javanica Blume                        8
                   Viburnum atrocyaneum C. B. Clarke
                   V. cinnamomifolium Rehder
                   V. corymbiflorum P. S. Hsu et S. C. Hsu
                   V. cylindricum Buchanan-Hamilton ex D. Don
                   V. foetidum var.
                     rectangulatum (Graebner)Rehder
Caprifoliaccae     Lonicera hvpoglauca Miquel
                   L. ligustrina Wallich
                   L. ligustrina var. pileata
                     (Oliver) Franchet
                   L. macrantha (D. Don) Sprengcl
Dipsacaceae        Dipsacus japonicus Miquel.                     10
Linnaeaceae        Abelia uniflora R. Brown                       9
Valerianaceae      Patrinia monandra C. B. Clarke                 10
                   P. scabiosifolia Link
                   P. villosa (Thunberg) Dufrcsnc
                   * Valeriana hardwickii Wallich                 8
                   * V. jatamansi W. Jones
Apiaccac           Hydrocotyle nepalensis Hooker                  2
                   Pternopetalum trichomanifolium                 14
                     (Franchet) Handel-Mazzetti
                   P. vulgare (Dunn) Handel -Mazzetti
                   Sanicula orthacantha S. Moore                  1
Araliaccae         Hedera nepalensis var. sinensis                12
                     (Tobler) Rehder
                   Macropanax rosthoniii (Harms) C.               7
                     Y. Wu ex G. Hoo
                   Metapanax davidii (Franchet) J. Wen
                     & Frodin
                   Schefflera bodinieri (H. Leveille) Rchdcr      2
                   S. bre\'ipedicellata Harms
                   S. elliptica (Blume) Harms
                   S. minutislellata Merrill ex H. L. Li
                   S. pauciflora R. Viguier
                   * Tetrapanax papyrifer (Hooker) K. Koch.       15
Pittosporaceac     Pittosporum brevicalyx (Oliver) Gagnepain      4
                   P. kwangsiense H. T. Chang & S. Z. Yan
                   P. pauciflorum Hooker & Amott
                   P. pulchrum Gagnepain
                   P. tonkinense Gagnepain

Data citied from Feng (2015) and Fan (2014), which were collected from
Dashiwci Tiankeng, Baidong Tiankeng, Banyuedong Tiankeng, Mafengdong
Tiankeng, Meiguidating Tainkeng, Shengmu Tiankeng, Sujia Tiankeng,
Luojia Tiankeng, Chuandong Tiankeng, Dacao Tiankeng, Dengjiatuo
Tiankeng and Liuxing Tiankeng. Asterisk (*) indicated plant species
were only found in these tiankengs but not found on the outside of
them. The plant species' name is following Flora of China and the
family is ordering the Angiosperm Phylogeny Group IV (APG IV 2016).
The areal-typcs of genera of seed plants is following Wu et al. (2006,
2011), and fern plants following Zang (1998) and Chen ct al. (2013).
Areal-types legends: 1. Cosmopolitan; 2. Pantropic; 3. Tropical Asia &
Tropical America disjunced: 4. Old world tropics; 5. Tropical Asia &
tropical Australasia; 6. Tropical Asia & tropical Africa; 7. Tropical
Asia; 8. North temperate; 9. Eest Asia & North America disjunced; 10.
Old world temperate; 11. Temperate Asia; 12. Mediterranea, west Asia
to central Asia; 13. Central Asia; 14. East Asia; 15. Endemic to China

Table 2 A complete list of the rare, threatened or endangered species
of vertebrates and invertebrates lives in the tiankengs of Dashiwei
Tiankeng groups

Family         Species

Goneplacidae   Typhlocarcinus vittosus Stimpson
Pholcidae      Belisana zhangi sp. nov.
Trechinae      Giraffaphaenops clarkei Deuve
Ranidae        Hoplobatrachus rugulosus Wiegmann.
Accipitridae   Accipiter nisus L.
Phasianidae    Chrysolophus pictus L.
               Lophura nycthemera L.
               Syrmaticus humiae Hume.
               Tragopan caboti Gould
Cuculidae      Cuculus canorus L.
Oriolidea      Oriolus chinensis diffusus Sharpe
Sciuridae      Petaurista petaurista
Viverridae     Prionodon pardicolor Hodgson
               Viverra zibetha L.

Data citied from Deuve (2002), Huang et al. (2004), Tong and Li
(2007), Leye Dashiwei Tiankeng Group National Geopark in Guangxi
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Author:Pu, Gaozhong; Lv, Yanna; Xu, Guangping; Zeng, Danjuan; Huang, Yuqing
Publication:The Botanical Review
Article Type:Report
Geographic Code:9CHIN
Date:Mar 1, 2017
Words:10402
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