Records of Odonata from Phong Nha-Ke Bang National Park and Its Buffer Zone, Central Vietnam

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Records of Odonata from Phong Nha-Ke Bang National Park 37

Entomologie heute 24 (2012): 37-49

Records of Odonata from Phong Nha-Ke Bang National Park and its Buffer Zone, Central Vietnam

Untersuchungen der Libellenfauna des Phong Nha-Ke Bang Nationalparks und seiner Pufferzone, Zentral-Vietnam

PHILIP STEINHOFF

Summary: From September 2010 until August 2011, the dragonfl y fauna of several habitats in the Phong Nha-Ke Bang National Park and its buffer zone in Central Vietnam was investigated. It is the fi rst time that research on dragonfl ies was undertaken in that area. The habitats surveyed in the buffer zone were rivers, water-fi lled bomb craters, paddy fi elds and forest streams, while the habitats where research on dragonfl ies was carried out inside the national park were forest rivers and forest streams. In total, 61 species were determined at 16 different sites, while the identifi cation of 15 more species was possible just at genus level; an identifi cation of the species requires further research. The species were identifi ed either in the fi eld, with the help of photographs or (when a specimen was collected) in the laboratory. The differences of the habitats surveyed and the species found at each location are discussed and compared with the knowledge of the dragonfl y fauna of Vietnam. Furthermore, an outlook on further research in the future is given.

Keywords: Dragonﬂ ies, distribution, Vietnam, Phong Nha-Ke Bang National Park, buffer zone

Zusammenfassung: Von September 2010 bis August 2011 wurden qualitative Untersuchungen der Libellenfauna im Phong Nha-Ke Bang Nationalpark und der zugehörigen Pufferzone in Zentral-Vietnam durchgeführt. Das sind die ersten Erfassungen von Libellen im Phong Nha-Ke Bang Nationalpark. In der Pufferzone wurden Libellen an Flüssen, wassergefüllten Bombenkratern, nassen Reisfeldern und Waldbächen kartiert, im Nationalpark wurden Waldﬂ üsse und Waldbäche untersucht. Insgesamt wurden an 16 verschiedenen Lokalitäten 61 Libellenarten festgestellt. 15 weitere Arten konnten bislang nur bis zur Gattung bestimmt werden. Die Libellenarten wurden entweder im Feld oder später anhand der Fotobelege bestimmt, in Ausnahmefällen wurden auch Tiere gesammelt, diese wurden dann mithilfe eines Mikroskops im Labor bestimmt. Die Unterschiede der untersuchten Libellenbiotope werden diskutiert und mit den bekannten Libellenvorkommen in Vietnam verglichen. Darüber hinaus wird ein Ausblick auf mögliche Ergebnisse von weitergehenden Untersuchungen in der Region gegeben.

Schlüsselwörter: Libellen, Verbreitung, Vietnam, Phong Nha-Ke Bang Nationalpark, Pufferzone

1. Introduction Quang Binh belongs to northern Central Vietnam and thus is part of the subtropical The Phong Nha-Ke Bang National Park climate zone (STERLING et al. 2007). (PNKB NP) is located in Quang Binh, Cen- The Phong Nha-Ke Bang area was de clared tral Vietnam, bordering Laos to the west and a national park in 2001. Two years later in approximately 500 kilometers south of the ca- 2003, the national park was declared a World pital city, Hanoi (Figs. 1, 2). Due to its length, Natural Heritage site by UNESCO (ZIEGLER Vietnam stretches across the subtropical 2004). The PNKB NP covers an area of and the tropical climate zone. The Province 85,000 hectares and is known worldwide;

Entomologie heute 24 (2012) 38 PHILIP STEINHOFF in Vietnam it is even famous for its special of animals and plants (ZIEGLER et al. 2004), karst landscape and the caves typical of including a high number of endemics, such that landscape (CLOUGH 2008). The biggest as the Phong Nha-Ke Bang Gecko, Cyrtod- cave in Vietnam is located in the PNKB NP actylus phongnhakebangensis Ziegler et al., 2003 (HERRMANN & PAGEL 2000). (HERRMANN & PAGEL 2000). The term “karst” refers to the weakly soluble Although a lot of research has been done in carbonate rock mountains, which form the recent years, the Odonata fauna of Vietnam topography of the central Vietnamese high- is still poorly known. The main focus of lands. Here, distinctive surface features like odonatologists is still on taxonomy (e.g. DO caves and underground rivers result from the 2011), while just very few publications add weathering of the limestone (LESER 2009). to the knowledge of Vietnamese odonate Together with the Hin Namno biodiversity diversity (DO 2007). protection area in Laos, the PNKB NP is Consequently, almost no published infor- one of the biggest karst forest areas in all of mation about the distribution of dragonﬂ ies Indochina that still boasts a high biodiversity in Central Vietnam is available, and the Odonata fauna of the PNKB NP is basically unknown. In April 2011 the discovery of Rhinagrion hainanense Wilson and Reels, 2001 in the PNKB NP during my investigation of the last year, was mentioned by KAWASHIMA et al. (2011).

Fig. 2: Position of the Phong Nha-Ke Bang National Park in Quang Binh, Central Vietnam Fig. 1: The Province Quang Binh in Central Viet- (from: Wikimedia commons 2011, changed). nam (from Wikimedia commons 2011, changed). Abb. 2: Lage des Phong Nha-Ke Bang Natio- Abb. 1: Die Provinz Quang Binh in Zentralviet- nalparks in Quang Binh, Zentralvietnam (aus: nam (aus Wikimedia commons 2011, verändert). Wikimedia commons 2011, stark verändert). Records of Odonata from Phong Nha-Ke Bang National Park 39

Thus, the present investigation contributes sites could not be visited with the same to the knowledge of distribution of dragon- frequency of visits in every location, thus fl ies in Vietnam. In addition, the importance qualitative research methods were used. of nature protection and the necessity of In total, Odonata were recorded during 74 protected areas for Odonata and the confi eld trips to 16 different locations (see Fig. 3). servation of unique nature areas such as the Every location was visited at least two times, karst forests of the PNKB NP is pointed except water body No. 13, where just one out. Furthermore, this investigation expands survey was possible. The surveys were carried the knowledge of the dragonfl y species in out regularly when the weather was fi ne. In rural areas of Vietnam and in primary karst Central Vietnam, the year can be divided into forest regions such as Phong Nha-Ke Bang. two main seasons: a wet and cold winter, This investigation and its results should be and a dry and hot summer. The rainy season seen as a starting point for more detailed is between September and December, and research in the future. January and February are usually the coldest months (STERLING et al. 2007). 2. Research area and methods In 2011, the cold weather lasted till March and the summer months were unusually wet. Odonata were investigated from September Thus, from November to March only seven 2010 to August 2011. excursions were possible due to the rainy Because of several unpredictable circum- season and the long winter. stances, such as fl oods and other unfortunate For each location the geo-coordinates were meteorological conditions, the investigation determined with a Garmin “GPS Map60SX”.

Fig. 3: The PNKB NP with extension area in the north-west and the research area in the east. The 16 localities where Odonata were surveyed are marked with black dots. Adapted from: FZS and CZ (unpubl., modiﬁ ed). Abb. 3: Der Phong Nha-Ke Bang Nationalpark mit Erweiterungsgebiet im Nordwesten und dem Untersuchungsgebiet im Osten. Die 16 untersuchten Habitate sind durch schwarze Punkte markiert. Adaptiert von: FZS und CZ (unveröff., stark verändert).

Entomologie heute 24 (2012) 40 PHILIP STEINHOFF

Tab. 1: Overview about the different habitat types where dragonﬂ ies were investigated. The GPS coordinates are given in brackets, the characteristics of size, bottom structure and vegetation of each waterbody are summarized. Sizes are based on estimations and Google Earth measuring (see also Figs. 4a, b, e, f). Tab. 1: Die unterschiedlichen Habitattypen, an denen Libellen kartiert wurden. Die GPS-Koordinaten sind in Klammern angegeben, charakteristische Merkmale der Größe, Vegetation und des Unter- grundes der Gewässer sind zusammengefasst. Die Größenangaben beruhen auf Schätzungen und Messungen mit Google Earth (s. Abb. 4a, b, e, f). Records of Odonata from Phong Nha-Ke Bang National Park 41

At each location adult dragonflies were habitats into lentic and lotic ecosystems and sought along the shore, over the water surface distin guished between the river systems of and in the surroundings. Each survey lasted the north, the south and the central part of between one and six hours and most surveys Vietnam. For the PNKB area, I differentiate were carried out between 11 a.m. and 2 p.m. between the habitats of the national park and Most species were identifi ed in the fi eld or those of its buffer zone, due to the differen- later by photographs with the use of various ces in structure and degree of pollution in books and papers (e.g. LAIDLAW 1950; HÄ- the two types of habitats. Furthermore, the MALÄINEN 1991; ORR et al. 2004; BEDJANIC lentic waters are divided into rice fi elds, bomb et al. 2007). In some cases specimens were craters and temporary standing waters or collected for a detailed examination later in bayous. Bayou is used here as a term to de- the laboratory, but it was not possible to get scribe the mostly sluggish, sometimes even a permit for collecting dragonfl ies. lentic tributaries of the forest rivers that can be found particularly in the dry season. 2.1. Localities The lotic waters are divided into rivers and streams by means of their different channel 16 reference sites were investigated for morphology. Inside the national park, such Odonata, eight in the buffer zone and eight a classifi cation is only a helpful construct as inside the national park boundary (Fig. 3, water systems in a primary landscape, espe- Tab. 1; see also Figs 4a, b, e, f). cially a karst area such as PNKB NP, change The sampling sites were chosen by two cri- their conditions very often according to the teria: First, the area had to be within reach, high dynamics in the landscape (e.g., fl oods). so that a fi eld-survey including the travel to the site and back was possible on a single 2.2.1. Water bodies inside the national park day; and second, the localities were chosen to cover a typical spectrum of Odonata The forest streams inside the Phong Nha- habitats present in the PNKB NP. Ke Bang National Park are characterized Location No. 4 lies outside the border of by a high density of shore vegetation and the national park but belongs to the area of sudden changes of the watercourse, which the park because it was bought by the park might include sections of subterranean for carrying out the reintroduction of en- fl ow, especially in the dry summer months. dangered animals. Rapids and waterfalls are frequent, as is typical of running waters in a karst landscape 2.2. Habitat types (LESER 2009). The stream beds are mostly rocky, regularly interrupted by clay or sand For the Odonata water bodies are the most sediments. Depending on the season and important habitats, because dragonfl ies spend weather, forest streams inside the national the longest part of their life as larvae in these park vary in width approximately between waters. Regional exceptions may be traced 50 cm and 5 m. within the genus Gynacantha which shortly Forest rivers inside the PNKB NP are much was found to develop in temporal water broader, but their water levels also change sig- bodies in very short times (see DE MARMELS & nifi cantly during the course of the year. The NEISS 2011 for a Brazilian representative of rivers receive all the water from the catchment this genus). In addition, the waterbodies serve area completely situated within the borders as reproductive sites for the adults. of the national park, and transport it out I follow the classifi cation of water types of it. When the water level drops, the rivers by DO (2007) who divided the Odonata become ephemeral and numerous bayous

Entomologie heute 24 (2012) 42 PHILIP STEINHOFF and temporary standing waters come into and Heliocypha biforata, are typical of a single existence in the river bed. These temporary habitat type. standing waters are among the only lentic Because of this and the diffi culties of defi n- waters inside the national park, and they ing the habitat types in an always changing attract a high number of dragonfl y species. landscape, I propose a classification of the different habitat types with the use of 2.2.2 Water bodies in the buffer zone of Odonata coenoses, as they are found in the the national park PNKB NP and its buffer zone. This classifi cation is preliminary and should Beside rare signs of human presence (also be seen as a fi rst step to a more precise de- sometimes even inside the national park) fi nition in the future. such as broken branches or small paths along The forest streams inside the national park the water, forest streams in the buffer zone as well as the forest streams in the buffer resemble those inside the national park. zone can be characterized by a high diversi- Rivers in the buffer zone are no longer typi- ty of species belonging to the superfamily cal forest rivers, because they fl ow through Calopterygoidea. Especially the species Neu- a rural landscape, especially rice fi elds and robasis chinensis (Fig. 4d), Heliocypha biforata small villages. They are characterized by the and Euphaea masoni, can be seen as typical nearly complete absence of riverine vegeta- species of these water types. tion and increasing pollution, with nutrients At the forest rivers in the national park, fe- from agriculture and waste waters from wer species occur than at the forest streams. households and livestock farming. Also, more eurytopic species can be found, The standing waters of the buffer zone are probably because of the capacity of the bomb craters, paddy fi elds or buffalo ponds. river to create standing waters in the dry The term buffalo pond is used to describe months. Core species could be Zygonyx iris water bodies that are used frequently by and Heliocypha perforata. buffalos and do not serve any other pur- The temporary standing waters inside the pose. The vegetation varies, depending on PNKB NP are inhabited mostly by libellu- the use of the waterbody. Some, like the rice lids such as Orthetrum glaucum and Orthetrum fi elds, dry out in summer; others are used chrysis, but also species of the family Coena- for irrigation agriculture. If the number and grionidae are present, for example Ischnura frequency of buffalos visiting the waters is senegalensis. high, these waterbodies become turbid and The rivers in the buffer zone can be clearly polluted, and consequently the number of defi ned by the absence of the stenotopic dragonfl ies present is reduced. Calopterygoidea species that occur at every clear river and stream inside the national 3. Results park. Instead, species such as Trithemis pallidinervis and Pseudagrion microcephalum are During the one year of investigation 76 common. species were found in total, 61 of them were The standing waters of the buffer zone can identifi ed to the species level. The species be characterized by species that are euryto- belong to 50 genera in 13 families. pic. Here most of the libellulids occur, such As shown in table 2 (see also Fig. 4), certain as Brachydiplax farinosa, Diplacodes nebulosa and species, e.g. Orthetrum pruinosum and Pantala Brachythemis contaminata. fl avescens, are eurytopic and can be found About 90 % of the species recorded from at nearly every waterbody, regardless of inside the Phong Nha-Ke Bang National Park habitat type. Others, such as Mnais mneme are found along forest streams. The forest Records of Odonata from Phong Nha-Ke Bang National Park 43 List of species recorded; indication of of recorded; study sites and number the site where they were species found. Liste der nachgewiesenen Arten mit Fundorten; Anzahl der Arten je Fundort. Tab. 2: Tab. 2: Tab.

Entomologie heute 24 (2012) 44 PHILIP STEINHOFF Continued. Fortsetzung. Tab. 2: Tab. 2: Tab. Records of Odonata from Phong Nha-Ke Bang National Park 45 Continued. Fortsetzung. Tab. 2: Tab. 2: Tab.

Entomologie heute 24 (2012) 46 PHILIP STEINHOFF

a b

e f

g h

Fig. 4: Karstforest in the Phong Nha-Ke Bang Nationalpark (a); forest-stream and waterfall inside the nationalpark (b); Aristocypha fenestrella, female (c); Neurobasis chinensis, male (d); Song San River in the buffer zone of the national park (e); Ecotrail River inside the national park (f); Trithemis aurora, male and the secondary genitalia of Idionyx victor (g); male (collected specimen, light microscopic picture) (h). Records of Odonata from Phong Nha-Ke Bang National Park 47 stream No 3 (see Tab. 2) is an exception, inside the PNKB NP fl ew at forest streams, because it was completely dry during the time whereas 66.6% of the species that occurred of investigation. Forest rivers and standing only in the buffer zone, were found only waters show a signifi cant smaller quantity at standing waters and rivers and 33.3 % of species, with 36 % and 25 % respectively. only at forest streams. This shows, that in In the buffer zone, the proportions of spe- general the species that occur only inside the cies occurrence at the different sampling national park are stenoecious, whereas most sites are more equally distributed. The forest of the species that were found only in the streams show the greatest diversity, with buffer zone are euryoecious. about 69 % of the species occurring in the For 15 other species, a reliable identifi cation at buffer zone. But the differences from lentic the species level could not be made (Tab. 3). waters (50 %) and rivers (approximately 35 %) are less than in the national park. 4. Discussion Species found inside the national park are about 73 % of the total number. This To identify the species listed in table 3, amount is essentially the same in the buffer further research, including the collecting zone, where 77 % of the total species occur. of specimens, is required. For example, Thus, all of the species that occurred only Macrogomphus lankanensis is known only as

Tab. 3: Species that could not be surely identiﬁ ed and the site at which they were found. Tab. 3: Arten die nicht absolut sicher bestimmt werden konnten mit Fundorten.

Abb. 4: Karstwald im Phong Nha-Ke Bang-Nationalpark (a); Bergbach mit Wasserfall im National- park (b); Aristocypha fenestrella, Weibchen (c); Neurobasis chinensis, Männchen (d); Song San-Fluss in der Pufferzone des Nationalparks (e); Ecotrail-Fluss im Nationalpark (f); Trithemis aurora, Männchen (g); die zur Bestimmung notwendige Aufnahme der sekundären Geschlechtsorgane von Idionyx victor, Männchen (Präparat, lichtmikroskopische Aufnahme) (h).

Entomologie heute 24 (2012) 48 PHILIP STEINHOFF endemic to Sri Lanka (BEDJANIC et al. 2007). 27 belong to the family Libellulidae. This To confi rm that it also occurs in Vietnam, remarkable fact was also shown in earlier a photographic proof is not enough, but a studies in other countries (TANG et al. 2010), detailed comparison of genital structures because the family Libellulidae consists of must be done with a specimen collected. Ad- many eurytopic species which are quite fl ex- ditionally, the two Coeliccia species that were ible and can tolerate pollution and changes found during this investigation cannot be of their habitat (ORR et al. 2004). Especially identifi ed, even though they were collected: in the buffer zone, the number of libellulid Coeliccia sp. is a new species, and the species species increased, whereas the more sten- Coeliccia scutellum and Coeliccia tomokunii can- oecious species, living only in clear forest not be distinguished (DO 2008). streams (ORR et al. 2004), could not be found According to the present study, it is possible in the buffer zone. This was to be expec- to state that the species which were recorded ted, because the anthropogenic impact on for the Phong Nha-Ke Bang National Park the buffer zone is much higher than in the and its buffer zone during this investigation national park itself. The rivers and ponds represent the typical species diversity of are used by humans and livestock (e.g. cattle) Odonata in central Vietnam (DO pers. comm.). and are therefore much polluted. Whereas Without doubt, especially in the primary forest rivers and ponds can thus be easily distingu- regions of the Phong Nha-Ke Bang National ished by means of their dragonfl y species, Park, more species could be found if more forest streams inside the national park and intensive research was carried out. This has in the buffer zone show little difference in already become clear through this year of species diversity: 11 of these species were research, during which the limitations beca- found only in national park streams, and me obvious. There have always been some 6 only at buffer zone streams. In general, species that could not be identifi ed or caught. the same typical species were found at The weather was often bad, so during entire forest streams both inside the national park months no investigations were possible. As a and at the border of the national park in result, conditions for fi eld trips were frequently the buffer zone. This interesting fact mainly unsuitable, especially after fl oods. Additionally, results from the further fact that most of it will be possible to check more different sites the species found are generally common at and water bodies if more researchers could be clear forest streams. But it is also due to the integrated in the study of dragonfl ies in this presence of forest in the buffer zone, alt- area. It is dangerous to walk through the karst hough it is restricted in extent. Many people forest around Phong Nha, especially alone, pass through the forest and some even bring taking into account the karst topography with their cattle for grazing, but the streams are steep and sharp ridges, holes in the ground and mostly unaffected. The forest is not used for unstable limestone formations (LESER 2009). agriculture, and people are not allowed to cut It is also necessary to collect more specimens wood, thus especially close to the border of so that correct determinations of additional the national park, the forest of the buffer species can be done. Many species have to be zone is of similar value to dragonfl ies as the identifi ed by the use of the original descrip- secondary forest inside the PNKB NP. tion, because there is no other information Although this might be the case in Phong (and certainly not fi eld keys) available. Iden- Nha-Ke Bang, it should not be forgotten that tifi cation of many tropical species is nearly in other regions the buffer zone forest is victim impossible without a specimen on hand. to logging, which makes protected areas such Among the 61 species found in and around as the Phong Nha-Ke Bang National Park even the Phong Nha-Ke Bang National Park, more important for conservation (DO 2001). Records of Odonata from Phong Nha-Ke Bang National Park 49

Acknowledgements HÄMÄLÄINEN, M. (1991): Idionyx victor spec. nov. (An- isoptera: Corduliidae) and some other odonata I am grateful to PHAN QUOC TOAN and from Hong Kong. Odonatologica 20(3): 343-347. NGUYEN THANH HAI for help with fi eldwork HERRMANN, H.-W., & PAGEL, T. (2000): Phong Nha-Ke Bang-Regenwaldschutzprojekt des in the forest. I thank MATTI HÄMÄLÄI NEN and Kölner Zoos in Vietnam. Zeitschrift des especially DO MANH CUONG for help with the Kölner Zoo, Heft 2: 1-13. identifi cation of some species and manifold in- KAWASHIMA, I., SASAMOTO, A., PHAN, Q. T., & DO, formations about dragonfl ies and odonatology M. C. (2011): First discovery and description of in Vietnam. Beside this, I would like to thank female and larva of Rhinagrion hainanense Wilson MARTIN SCHORR for tireless help with literature. and Reels, 2001 (= R. yokoii Sasamoto, 2003) GABRIELE UHL granted me access to a light mi- (Zygoptera: Megapodagrionidae) from Viet- croscope and MELANIE WITTHUHN teached me nam. Acta odonatologica Japonica 53: 93-99. the handling. Furthermore, I thank KATALIN LAIDLAW, F.F. (1950): A survey of the Chlorocy- phidae (Odonata: Zygoptera), with diagnoses SCHEIDLER, TABEA LUTZKER, MARTIN SCHORR of proposed new genera, and description of and especially JÜRGEN OTT for numerous a new geographical subspecies. Transactions explanatory notes on the manuscript. DENNIS Royal Entomological Society London 101 (8): PAULSON gave my English the fi nal touch. 257-280, fi gs. 1-8. LESER, H. (2009): Geomorphologie. Westermann; References Braunschweig. ORR, A.G., BUTLER, S.G., HÄMÄLÄINEN, M., & BEDJANIC, M., CONNIFF, K., & DE SILVA WIJEYE- KEMP, G.R. (2004): Insecta: Odonata. Pp. RATNE, G.(2007): Gehan’s Photo Guide. A 409-442 in: Freshwater Invertebrates of the Photographic Guide to the Dragonfl ies of Malaysian Region (YULE, C.M., & YONG, H.S., Sri Lanka. Jetwing Eco Holidays: Colombo. eds.). Academy of Sciences; Malaysia. CLOUGH, L.D. (2008): UNEP “Phong Nha-Ke Bang STERLING, E.J., HURLEY, M.M., & LE, D.M. (2007): National Park, Vietnam”. In: Encyclopedia of Vietnam: a natural history. Yale University Earth (CUTLER, J. C., ed.). Washington, D.C.: Press; Yale. Environmental Information Coalition, National TANG, H.B., WANG, L.K., & HÄMÄLÄINEN, M. Council for Science and the Environment). First (2010): A photographic guide to the dra- published in the Encyclopedia of Earth August gonfl ies of Singapore. Raffl es Museum of 22, 2008. http://www.eoearth.org/article/ Biodiversity Research; Singapore. Phong_Nha-Ke_Bang_National_Park,_Vietnam VOGT, M., FORSTER, B., PAGEL, T., & ZIEGLER, DE MARMELS, J., & U.G. NEISS (2011): Description of T. (2006): Neues vom Naturschutzprojekt the larva of Gynacantha auricularis Martin, 1909 des Kölner Zoos in Vietnam. Zeitschrift des (Odonata: Aeshnidae). Zootaxa 3137: 64-68. Kölner Zoo, Heft 1: 35-49. DO, M.C. (2007): Checklist of dragonfl y from ZIEGLER, T. (2004): Erforschung der Artenvielfalt Vietnam.Vietnam National University Pub- im Vietnamprojekt des Kölner Zoos: Die Am- lisher; Hanoi. phibien und Reptilien von Phong Nha-Ke Bang. DO, M.C. (2008): Coeliccia hoanglienensis spec. nov., Zeitschrift des Kölner Zoo, Heft 4: 147-171. a new platycnemid damselfl y from hoang lien ZIEGLER, T., HERRMANN, H.-W., VU N.T., LE K.Q., mountains, the north of Vietnam (Zygoptera: NGUYEN T.H., CAO X.C., LUU M.T., & DINH, Platycnemididae). International Journal of H.T. (2004): The amphibians and reptiles Odonatology, 11: 21-26. of the Phong Nha-Ke Bang National Park, DO, M.C. (2011): Burmagomphus schneideri sp. nov., Quang Binh Province, Vietnam. Hamadryad new dragonfl y from the south of Vietnam Vol. 28, Nos. 1 & 2: 19-42. (Odonata: Gomphidae). International Journal of Odonatology, 14: 223-231. Philip Steinhoff DO, T. (2001): Characteristics of Karst eco systems Trelleborger Weg 1 of Vietnam and their vulnerability to human D-17493 Greifswald impact. Acta Geologica Sinica, 75: 325-329. E-Mail: [email protected]

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