1* Abstract Kon Ka Kinh National Park belongs to Kon Tum Plateau and has high value of diversity in Forestry. Since the first survey in 1999, the in Kon Ka Kinh is little known. We conducted the investigation to study their population and factors affect butterfly distribution in a mountainous forest. Pollard walk method with a hand-net method was used for observation. 323 species were recorded, including 235 species in this survey. There were three new records for Vietnam and three new records in Central Vietnam. is the dominating family. Our result indicated the impact of seasons, water resource, elevation and forest type on butterfly diversity. Their diversity is higher in the rainy season, where above ground water resource is highly available. The diversity indexes also are higher at a lower elevation and disturbed forest area.

1

I. INTRODUCTION ...... 1 1.1. Background ...... 1 1.2. Aims, scope, research question ...... 5 II. LITERATURE REVIEW ...... 6 2.1. Systematics...... 6 2.2. Ecology and behaviors...... 7 2.3. Butterflies as an indicator of environment ...... 8 2.4. Factors influencing butterfly diversity ...... 10 2.5. Butterfly conservation: ...... 12 III. MATERIAL AND METHODS ...... 14 3.1. Study area...... 14 3.2. Transect selection: ...... 17 3.3. Data collection: ...... 18 3.4. Data analysis ...... 19 IV. RESULTS ...... 21 4.1. Butterflies in Kon Ka Kinh National Park ...... 21 4.1.1. The butterfly fauna...... 21 4.1.2. Ecological complexes of tropical butterflies in Kon Ka Kinh N.P...... 23 4.1.3. Kon Ka Kinh butterfly in conservation ...... 24 4.1.4. New distribution records: ...... 25 4.1.5. Biogeographical features ...... 25 4.2. Butterfly indexes in Kon Ka Kinh National Park ...... 26 4.3. Factors influence to butterfly population distribution ...... 28 V. DISCUSSION ...... 33 5.1. Butterfly fauna in Kon Ka Kinh National Park...... 33 5.2. Butterflies diversity along the transect ...... 35 5.3. Factors impact to butterfly diversity: ...... 36 VI. CONCLUSION ...... 38 VII. ACKNOWLEDGMENT ...... 40 VIII. APPENDIX ...... 41 IX. REFERENCES ...... 53

LIST OF FIGURES Figure 1:Kon Ka Kinh national park map ...... 15 Figure 2: Average Monthly Temperature and Rainfall...... 16 Figure 3: Percentage of butterfly species in each family recorded in Kon Ka Kinh N.P. (with and without consideration of species abundance) and entirely in Vietnam ...... 22 Figure 4: Taxonomical compositions of butterflies in Kon Ka Kinh N.P...... 23 Figure 5: Percentage of common and forest butterfly species recorded ...... 24 Figure 6: Correspondence analysis ...... 29 Figure 7: Butterfly rarefaction curve for seasons ...... 30 Figure 8: Butterfly rarefaction curve based on water condition ...... 31 Figure 9: Rarefaction curves by forest types and altitude factors ...... 31

LIST OF TABLE

Table 1: List of endemic species in Kon Ka Kinh National Park ...... 26 Table 2: Butterfly diversity indices ...... 28 Table 3: Diversity permutation test ...... 28

I. INTRODUCTION 1.1. Background Among invertebrate conservation group, the butterflies are among one of the most popular focal groups. They are well known as indicators of tropical forest ecosystems and their ecological condition. (Oostermeijer & Van Swaay, 1998). Butterflies are a good group of for studying the effects of human activity. Under pressure from population growth, the forest area is destroying to increase the fragmentation and biodiversity loss. A butterfly is a sensitive group with the changing of the environment (Mecenero, 2015). In many representatives of this group, the specific characteristics of coloration, wing pattern, and genital morphology may demonstrate biogeographical information and provide useful parameters for studying the geographic variability of species. Butterflies represent a suitable object for solving the problems related to the specific features of geographic, landscape, seasonal, and climatic distribution of species (A. Monastyrskii, 2007b). Among many reasons that could be named to justify the use of this group as an object for solving the above problems, the following are most important: (1) The butterfly is well-developed (De Jong, Vane-Wright, & Ackery, 1996; Rod & Ken, 1988); (2) They are present in practically all the known tropical habitats of Vietnam (A. Monastyrskii & Holloway, 2013; A. L. Monastyrskii & Devyatkin, 2015); (3) Their life cycle is often associated with specific host plants and some other groups of insects and vertebrate . The changes in the vegetation are always accompanied by the changes in the taxonomic composition of primary consumers, including butterflies (Dover, 1996; Ehrlich & Raven, 1964); (4) Butterflies can be easily observed and counted in nature, and can therefore be used for demonstrating seasonal changes in abundance (Pollard, 1975; Pyle, Hughes, & Institute, 1992); (5) The boundaries of distribution ranges of butterflies can be vertified with a satisfactory degree of precision based on museum collections and illustrated

1 guides; (6) In many representatives of this group, the specific traits of coloration, wing pattern, and genital morphology provide the biogeographical information and provide convenient parameters for studying the geographic variability of species (Benedick et al., 2007; Dover, 1996; Gilbert & Singer, 1975) During the last twenty years, butterflies have been successfully utilized by many national and international programs for conservation activity as a perfect tool supplementing the main methodology for the assessment of natural resources(Bonebrake, Ponisio, Boggs, & Ehrlich, 2010; T. New, Pyle, Thomas, Thomas, & Hammond, 1995; T. R. New, 1997). Practically nearly all conservation projects implemented in Vietnam from 1993 have included butterfly study into biodiversity research programs. These faunistic surveys showed a high level of butterfly diversity and character of species distributions (A. Monastyrskii, 2007a; A. L. Monastyrskii & Devyatkin, 2015). There were discovered a high number of species which have been described as new to science. Many of these new species are endemics to Vietnam and were found very locally (A. L. D. Monastyrskii, Alexey, 2000, 2012). The majority of endemic species are characterized by restricted ranges and are located in some topographically isolated mountain areas including mountain massifs of Vietnamese central highlands: Central Truong Son Range; Kon Tum Plateau, and for example, Kon Ka Kinh and Ngoc Linh national parks. Kon Ka Kinh is an isolated mountain massif located in Gia Lai Province of Kon Tum Plateau. It is situated within Endemic Bird Area and supports six restricted-range bird species(Le, 2000). Kon Ka Kinh is also a globally important site for the conservation of amphibian diversity. This area supports a number of amphibian endemics (Tordoff, Tran, Nguyen, & Le, 2004). The National park also promotes a high variety of insects, in particular butterflies. During the only two-months survey in March/April 1999 conducted by BirdLife International, there were recorded over 200 butterfly species. Eight species collected at that time were previously unknown to

2 science, and they have been described as new to science (A. L. Monastyrskii & Devyatkin, 2015). Unfortunately, since 1999 there were no additional organized surveys that may be extended our knowledge on faunistic, biogeographic, ecological and conservation aspects of this topographically isolated area In French colonial times (before 1945), the forests here were not exploited or impacted. Roads and vehicles were not in good conditions, so forest products could not be transported to the market. In 1964, American troops landed in the National Park area. They occupied and developed military bases on many hills. They exploited and depleted around 1000 hectares of primary forests for different purposes (Le, 2000). Following the Birdlife report (Le, 2000), in 1980, the Mang Yang forestry company was established and managed the forest in the region. During this time, the main task of the company was logging. Each year, the company harvested approximately 15,000-20,000 m3. Exploitation is carried out from this area to other areas. An activity of forest logging has gravely affected and depleted the forest resources. The majority of secondary forests today are a consequence of this period. Besides these activities, illegal logging also occurred. This contributed significantly to forest degradation in Kon Ka Kinh forest. In 2002, Kon Ka Kinh National Park was established. The logging activities have been prohibited completely. Instead of logging, forest protection, reforestation, enrichment planting, and restoration activities have been promoted. However, illegal logging has still occurred in some remote places with fewer forest rangers and larger trees. During this time, forest resources have also been affected by the activities of the local people as farming, grazing, harvesting of forest products such as honey, orchids and so on. Kon Ka Kinh forms part of a contiguous landscape of natural habitats in north- eastern Gia Lai province which supports some of the most intact faunal and floral communities in the central Truong Son. The Park maintains a range of montane habitat

3 types. The important forest area are 2,000 ha of mixed coniferous and broadleaf forest containing Fokienia hodginsii, situated above 1,300 m. The National Park is characterized by a very high diversity of animals and plants. There are 1,022 vascular plant species, belonging to 568 genera recorded in the National Park. Kon Ka Kinh also supports some globally threatened mammal species, including grey-shanked douc, yellow-cheeked crested gibbon and tiger. Kon Ka Kinh is located within the Kon Tum Plateau Endemic Bird Area, and supports some amphibian species endemic to the Annamite Mountains, including four species assessed as globally threatened (Le, 2000). The buffer zone of Kon Ka Kinh National Park has belonged to seven communities. There are 71% Ba Na ethnic and 27% Kinh in total 27,200 people. Economic development influences forest protection. Whole 1,300 ha wet rice and 2,900 ha hill rice, the result from an investment in 1999 demonstrated that forest land is one of the most important resources, that can be cleared for agriculture. (Le, 2000) From the establishment of the Park, many new species were found: A new record of gibbon species N. annamensis in Kon Ka Kinh (Van Ngoc Thinh, Thanh, & Roos, 2010), new reptile and amphibian species (Tao, Matsui, & Eto, 2014). A new flora taxon, Kerivoula kachinensis, also was described from Kon Ka Kinh (Vislobokov, Nuraliev, Kuznetsov, & Kuznetsova, 2016). Eight new butterfly species were described from the survey in 1999 (A. L. Monastyrskii & Devyatkin, 2015; A. L. D. Monastyrskii, Alexey, 2012) The butterfly of Vietnam is the results of combing the fauna group from Holarctic, Indo – Malayan regions and Indo – Burmese (A. Monastyrskii, 2007b). They resemble the butterfly fauna of neighboring countries. The local butterfly fauna is fascinating and unique in Kon Ka Kinh National Park. According to the first survey undertaken in 1999 the local butterfly fauna includes over 200 species (Le, 2000). Thirteen species (~6.5%) are endemics to eastern Indochina, demonstrating links with

4 Sino-Himalayan, Indo-Burmese and Sundanian faunas (A. Monastyrskii & Holloway, 2013). We supposed that a new survey would bring further information on the composition of the local butterfly fauna, its endemic portion, and unknown taxa. Such monitoring comparative study is rather important because it will be carried out in time of climate change. Conducting a further study will show if there is any evidence to suggest that changes in environmental conditions, and have affected butterfly species in the region (Menéndez et al., 2007). Therefore, after 20 years, it was necessary to carry out the research on “Butterfly diversity and conservation in Kon Ka Kinh National Park.” 1.2. Aims, scope, research question

 Aim: The aim of the study is the to determine the diversity and factors impact to butterflies in Kon Ka Kinh national park

 Objectives: The specific objectives of the research are: 1. To apprise the butterfly recorded in Kon Ka Kinh National Park; 2. To compare the butterfly diversity indices in different typical habitat; 3. To determine the factors afecting to butterflies’ diversity

 Question: The study will answer the following questions: - What is butterfly population in Kon Ka Kinh N. P at present? - How is the diversity of butterfly fauna in Kon Ka N.P? - What are the factors impacting to butterfly diversity in the tropical rainforest?

5 II. LITERATURE REVIEW 2.1. Systematics. The diversification of butterflies which is believed to be at around the K–T extinction event, about 100–60 million years ago (Boggs & Dau, 2004). More than 60% of all species on the planet are insects (May, 1992), and butterflies are undoubtedly one of the best-known groups of insects. They are also relatively easy to identify and diverse, which makes them well-suited as biodiversity indicators. Butterfly (Rhopalocera) has belonged to order of Insecta class. The estimated number of the butterflies is about 13000 -20,000 species in total 160,000 Lepidoptera species (Torben Bjørn Larsen, 2005; T. R. New, 1997; Rod & Ken, 1988). This group of Lepidoptera was generally represented in three super-families which are skippers (Hesperioidea), true butterflies (Papilionoidea), and butterfly-moths (Hedyloidea) without clubbed antennae was suggested recently (Scoble, 1986). At the same time, the modern progress in the molecular analysis has allowed developing a new phylogenetic tree that strongly contradicts the traditional hypothesis (Kawahara & Breinholt, 2014) Recently, the trending of the butterfly’s systematic study is moving to molecular analysis. However, morphology still plays an essential role in butterfly phylogenetic. Butterfly morphology presents their evolution and evaluating the evolutionary scenarios of phylogenetic trees. (Simonsen, de Jong, Heikkilä, & Kaila, 2012). Butterfly species recorded in Vietnam was increasing from 455 species in 1957 (Metaye, 1957) to 992 species in a checklist in 2003, and up to 1181 species in the newest checklist in 2015(A. L. Monastyrskii & Devyatkin, 2015). Based on morphology characters, all butterfly fauna belongs to 6 families (De Jong et al., 1996; Espeland et al., 2018). That includes Papilionidae – 69; – 57; Nymphalidae – 418; Riodinidae – 31; – 318; and Hesperiidae – 288. The total number of species-level taxa is 1277 (A. L. Monastyrskii & Devyatkin, 2015).

6 Some critical taxonomical works and researches have been done relating to the butterfly of Vietnam. In 2009, Callaghan review 29 Riodinid taxa, which were found in Vietnam. It provided information about taxonomy, diagnosis, behavior habitat and range. It also presented on biogeography data of each taxon and compared with distribution in Indo-China (Callaghan, 2009) Taxonomical works reveal unknown or poorly studied butterflies researching process of which may include a description of new taxa and revision notes. The extensive study was undertaken on the general Scobura, Hidari, Celaenorrhinus, and Potanthus (Devyatkin, 2004; A. L. Monastyrskii & Devyatkin, 2015). Monastyrkii revised butterflies fauna of Vietnam on his book series “Butterflies of Vietnam”: volume 1 - Nymphalidae (), volume 2 - Papilionidae, and volume 3 – Nymphalidae (, Amathusiinae) (A. Monastyrskii, 2007a; A. L. Monastyrskii, 2005, 2011). Six species discovered in Kon Ka Kinh during the survey in 1999 were determined as new to science (A. L. D. Monastyrskii, Alexey, 2000, 2012): Delias vietnamensis (Pieridae, Pierinae); konkakini, L. melisana, Faunis bicoloratus, Aemona kontumei, A. simulatrix, Stichophthalma eamesi, (Nymphalidae, Satyrinae), Dodona katerina (Riodinidae). 2.2. Ecology and behaviors Temperate montane butterfly prefers feeding on mud, herbivore dung, and carnivore dung (Boggs & Dau, 2004), while tropical species feed on a wide variety of sources including fruit, carrion, and pollen (K. C. Hamer et al., 2006). Pollen feeding butterflies, probably are a result of selection for longer adult lifespan (Bonebrake et al., 2010). Female adults of Eurema species have a reproductive diapause phase with larval host plant conditions. They are sensitive to the change in rainfall and photoperiod (Gilbert & Singer, 1975; Jones & Rienks, 1987). A large number of Lycaenidae butterfly is a close relationship with ant species (Thomas, Elmes, Wardlaw, & Woyciechowski, 1989). The symbiosis between

7 butterflies and ant usually are mutually beneficial. However, an ant can be parasitic, and competitive. If a host ant species becomes extinct, it will bring the affiliated butterfly to go along with it (Bonebrake et al., 2010; Koh et al., 2004). The evolution of butterflies supports defense from their predators in multiple ways, such as toxins in mulciber (Wei, Lohman, Peggie, & Yen, 2017), faked head - Spindasis lohita (Torben B Larsen, Van Hao, & Van Lap, 2005), or bird dung form in caterpillars. For non-poisonous species, mimicry is the efficient way of hiding their predators. Mimicry has an essential effect on the habitat distribution of tropical butterflies (R. I. Hill, 2010). The first research on mimicry in butterfly had been studied on Heliconiines, Ithomiines, Papilionids, and Pierids (Bates, 1862; Bonebrake et al., 2010). A huge of poisonous and aposematic butterflies fly leisurely, that allows them to show their characteristic warning colors and patterns (Torben B Larsen et al., 2005) In butterfly fauna, many eggs are laid on the underside of the leaves. After a few days, small larvae will eat their eggs cover and enter into the world (Rod & Ken, 1988). The caterpillar is immature. This is feeding time, and they have several stages. When the larva reaches their full size, they will change the stage, and become a pupa. Many male specimens often push itself out of its chrysalis before the female, so that the sex ratio in population changes during the flight season (Gilbert & Singer, 1975; T. R. New, 1997) Temperature impact on butterfly activity. Therefore, a thermoregulatory behavior is complex and involves a range of postures and activities, because of the butterfly need to elevate their body temperature for flight requirement (T. R. New, 1997) 2.3. Butterflies as an indicator of environment The methodology of watching and observation for tropical butterflies represents in a number of scientific monographs and papers (e.g., (Basset et al., 2011; Pyle et al.,

8 1992)). The butterflies are less abundant or are narrowing their distribution than they used to be (T. R. New, 1997). Although butterflies cannot represent for some lifestyles and habitats, example, few butterflies have predacious or parasitic larvae; no species is aquatic; and very few feeds on decaying wood (T. New et al., 1995), their caterpillars are depended on a hostplants, they are coevolution, so sensitive group reflect the changing of environment (Ehrlich & Raven, 1964; Mukherjee, Banerjee, Saha, Basu, & Aditya, 2015; Rusman, Atmowidi, & Peggie, 2016). Especially, diversity of forested species population can present for disturbing habitat (Ghazoul, 2002). Therefore, studying on butterfly pattern population can be used as the indicator of forest disturbance (J. Hill, Hamer, Lace, & Banham, 1995). Butterfly species diversity was uncorrelated with disturbance of forest in general. However, there is a strong relationship between rare species diversity and environmental quality (Hayes, Mann, Monastyrskii, & Lewis, 2009; Rod & Ken, 1988). According to butterfly ecology and behavior, fruit-baited trapping method denote butterfly fauna is still well remained in the small isolated primary forest rather than disturbed forest (Daily & Ehrlich, 1995) Long-term studies of butterfly communities at different sites in Vietnam have defined a group of the common butterfly species. This group includes about 105-110 diurnal butterfly species belonging to six families (A. L. Monastyrskii & Devyatkin, 2002). All of these species are very widely distributed in Vietnam and common in adjacent countries (A. Monastyrskii, 2010). They comprise the most typical inhabitants of degraded forest habitats, secondary vegetation, agricultural lands and urban areas at low elevations (V. L. Vu & Yuan, 2003). Flight periods typically occur throughout the year. The group includes a number of eurytopic, opportunistic and migratory species, which also often happen in ruderal areas and near the primary forest. Their larvae usually feed on a variety of plant species, most of which are also common. As a whole, species within the group are characterized by high levels of abundance and ease of

9 recording. For these reasons, species in the group are considered suitable indicator species for assessing levels of disturbance in natural habitats. Results of butterfly surveys carried out at sites throughout Vietnam indicate that the proportion of common butterfly species of the butterfly fauna in primary and virgin tropical forest habitats rarely exceeds 20-25% (A. Monastyrskii, 2010). In lightly disturbed and secondary forest habitats, percentages of common butterflies may vary between 25-45% (Spitzer, Jaros, Havelka, & Leps, 1997; Spitzer, Novotny, Tonner, & Leps, 1993). In the poor quality forest, bamboo and mixed secondary forest, levels of common butterflies may vary between 45-50% and are in many cases higher. Highest proportions of common butterfly species are usually found in secondary vegetation in the vicinity of roads and cultivated areas and can exceed 60% of the entire butterfly fauna in these habitats (Fahrig & Rytwinski, 2009; A. Monastyrskii, 2010) 2.4. Factors influencing butterfly diversity There are many factors affecting butterfly diversity (Menéndez et al., 2007; Spitzer et al., 1997). Non- flora factor, including shelter, isolation, the width of hedgebank or grass verge, and uncultivated habitats effect on butterfly assemblages (Dover, 1996). In the scope of research, data were analyzed by the following factors: season; availability of water resources; habitat type richness, and elevation. In the tropical rainforest, temperature and rainfall, and day-length vary only within very narrow margin throughout the year. Butterfly community is represented by an enormous range of species (Rod & Ken, 1988). Many studies demonstrate the relationship between seasons and butterfly diversity. The most significant differentiating factor between temperate and tropical zone is winter temperature. During the winter, tropical butterflies do not face to very cold and harsh temperature as in temperate zone. However, in the tropical season, the humidity is different. Therefore, some butterflies have two forms: a dry season and a wet season form (Bonebrake et al., 2010).

10 The butterfly diversity indexes are higher in pristine forest areas (J. Hill et al., 1995). In the same forest type, canopy has higher butterfly species richness than understory, example in Ecuadorian rainforest (DeVries, Murray, & Lande, 1997) or in Bornean rain forest (Schulze, Linsenmair, & Fiedler, 2001). There are significant differences between diversity indices calculated for butterfly assemblages occurring at lower and higher elevations. The values of diversity indices in butterfly communities at lower elevations are usually higher than at high mountain areas (V. L. Vu & Yuan, 2003). Groundwater supply influence on butterfly distribution (Oostermeijer & Van Swaay, 1998). The proportion of forest species in dense forest is higher than the community in steam trails. In the opposite trend, the common species is higher in stream areas. Besides, the stream sides are the higher number of individual number, while the disturbed forest contains the greatest species number (Hayes et al., 2009; L. V. Vu & Quang Vu, 2011) The proportion of the rare butterfly species often decreases from the natural primary forest to the stream sides, while the percentage of common species increase from the natural forest to the stream sides. The stream sides have the greatest individual number, while the disturbed forest contains the greatest species number (L. V. Vu & Quang Vu, 2011). The Vietnamese butterflies also are influenced by geographical factors such as isolation, continuity of habitat (A. Monastyrskii, 2010; A. Monastyrskii & Holloway, 2013; Spitzer et al., 1993). The forest preferred species that have a smaller range of distribution of endemic type, whereas common species are wide distribution (Spitzer et al., 1993). Butterflies in lowland forest areas are higher diversity than mountainous forest. They are also similar to higher diversity in dry and transitional season, lower in the wet season (A. Monastyrskii, 2007b). However, richness in species is not reflected in abundance in numbers of individual, for a characteristic of these rainforests in the

11 relatively small sample of each species which can be seen on the wing in any single period, so that no one species appears conspicuously abundant or dominant (Rod & Ken, 1988). The result from Spitzer pointed out that temporal change in forest butterfly species of Vietnam is determined by relatively higher seasonality of species with biogeographical affinities to the seasonal `East Himalayan' region (Spitzer et al., 1993). 2.5. Butterfly conservation: and butterfly extinction is less well known, but butterflies are over half of the described species in the world. Its studies have been used as models of tropical insect diversity (DeVries et al., 1997; Groombridge, 1992). However, at present, lacking knowledge of metapopulational theory in tropical butterfly population studies or conservation efforts lead to a paucity of population studies in the tropics (Bonebrake et al., 2010). Only 4 butterfly species are known to have gone extinct over the past 150 years(IUCN, 2018), despite over 80% destruction of the forest in the area, but many risks are threatening to butterfly (Bonebrake et al., 2010). Threats to tropical butterfly fauna include habitat loss, global climate change, invasive species, pollution, and exploitation and harvest activities. Because of shifting usage purpose, forest land become agriculture land, plantation forest or rubble plantation that destroy the habitat (J. K. Hill, 1999; T. R. New, 1997; Sodhi & Brook, 2006) In 2006, in IUCN Red List database, there are 43 are butterflies (CR: 1, EN: 14, VU: 28) from South East (Koh, 2007). All of them belonged to only two families Papilionidae and Nymphalidae. In 2018, 65 species were listed in categories: EW, RE, CR, EN, VU, LR/cd, NT or LR/nt. The total 253 butterflies in the world were listed in the above categories (IUCN, 2018). In Vietnam, there are ten species were listed in Vietnam Data Red book: Stichophthalma uemurai, Zeuxidia masoni, albofasciata, Baysa crassipess,

12 Papilio elephenor, Papilio noblei, Teinopalpus aureus, Teinopalpus imperialis, Troides helena, and Troides aeacus (Ban et al., 2007).

13 III. MATERIAL AND METHODS 3.1. Study area Kon Ka Kinh National Park was formed by the name of the highest peak, Kon Ka Kinh Mount (1750m). The park is located in two districts: Mang Yang and KBang. It contains parts of five communes: Ha Dong, Ayun, Kroong, Kon Pne, and Dak Roong. It is bounded by the coordinates 14º03' to 14º36’N and 108º12' to 108º32’E. The altitude range is from 570m (at Ba river) to 1750m (Kon Ka Kinh peak). The geology of Kon Ka Kinh is determined by metamorphic rocks, principally gneiss characteristics (Le, 2000) Hydrology: There are two main watersheds belonged to two rivers: Ba and Dak Pne River. The stream and river in Park are often shot, narrow, and fast flowing. Because of stream influence to butterfly distribution (Spitzer et al., 1993), so in this research, the survey was conducted in both two watersheds: - Ba River: it contains two transects (number 4 and 5). The Ba River has a highly seasonal flow during three months October, November, and December - Dak Pne River: watershed cover three transects (number 1, 2 and 3)

14

Figure 1:Kon Ka Kinh national park map Meteorology: There are two regular seasons in Kon Ka Kinh National Park: the dry season, from December to April, and the wet season from May to November (Le, 2000). In the dry season, the average temperature in the hottest month (April) is 23.6oC. The total rainfall in the dry season is about 25% - 30% total annual rainfall. In the wet season, the average temperature of the hottest month (August) is 25.1oC. The highest rainfall is 427.44mm in October (World Bank Group, 2016) see on Figure 2

15

Figure 2: Average Monthly Temperature and Rainfall at location (14.23,108.37) from 1991-2015(World Bank Group, 2016) Social and economic condition: Timber extraction was banned from April 1999. Although it was not caused by shifting cultivation, it increased the forest degradation in the National Park. Harvesting none-timber forest product is focusing on honey, rattans, Litsea bark, and Amomum fruit (Le, 2000). However, the price of honey was ten time increased (compare with price in 1999), up to VND 300000 per liter (in 2018). Tourism: The beauty of nature attracts tourists in Kon Ka Kinh National Park. Surrounding of magnificent of Kon Ka Kinh top, there are several waterfalls and peace of landscapes. Six tourist routes are developing in the Park (Thang Long et al., 2016). Tourist development influence to transect 1, transect 2, and transect 3, but do not affect to transect 4 and 5.

16 3.2. Transect selection: Choosing transect was based on the main features characterized by a particular ecosystem. According to result from the previous study on altitude and forest types in Kon Ka Kinh National Park (Le, 2000). this research was surveyed at five transects. It represented for 5 different habitats in the Park: - Transect 1: flat area in valley, riverine and disturbed forest. The surveys were conducted following the tourist trail. It started from the point (14.20773N; 108.317198E) to Frankfurt station (14.220347N; 108.317172E). The transect is regularly influenced by economic development. During the survey in 2018, we met both Vietnamese tourists and local people. The range of elevation from 800 – 1000m. The transect is 3-5m width and 1.5km length. In this trail, because survey cross streams, the groundwater is a factor influence butterfly distribution. - Transect 2: is defined as logging trail. Since 1980, Mang Yang forestry company was established and managed the forest in the region. This trail was affected by forest harvesting until 2002, Kon Ka Kinh national park was established. Nowadays, this trail was closed for collection, but it is still affected by local people. In this transect, none of the streams was cross, so groundwater flow does not influence the butterfly population. The elevation variation of transect 2 is from 1000m to 1150m - Transect 3: was performed in the disturbed forest in the past, but now it was well protected since Kon Ka Kinh national park established. The altitude of the transect is from 1200m to 1500m. The typical area in this transect is defined as near primary forest in steep gradient. - Transect 4: was defined as primary forest trails. This trail crosses the many streams, so the habitat was impacted by water sources. The intermittent observation in transect 4 ensure the similar characters of primary forest and water condition. Its elevation is from 1200m to 1500m

17 Transect 5: was described the high mountainous primary forest. This area was determined as untouched area and is located in Kon Ka Kinh mountain area. The transect started from camping site (14.332232N, 108.406921E) to Kon Ka Kinh peak (14.320913N, 108.397784E)

3.3. Data collection: There are some methods for butterfly’s observation: visual encounter surveys (VES), line transects distance sampling (LTDS), and fruit-baited traps (FBT). FBT had low efficient, less suitable for comparing butterfly assemblages (Basset et al., 2011). VES is more efficient for recorded butterflies individually. LTDS provide a quick method to evaluate the butterfly density (Kral, 2018). The data on butterfly assemblages were collected using the transect method described by Pollard (Pollard, 1975). This standard protocol was modified by Spitzer (Basset et al., 2011; Spitzer et al., 1993). Transect counts provide an index of population size and therefore can be used to measure changes in abundance One full observation will include one pass. The observation will be executed out along each transect from 9 am to 3 pm at a speed of approximately 20 m per min, and all observed butterflies will be recorded. Species that require further identification or taxonomical study will be collected using a butterfly hand-net. For butterfly identification, we follow the system in the book “Butterflies of Vietnam”. This system based on newer taxonomic classification by morphological and molecular analysis (A. L. Monastyrskii & Devyatkin, 2015). During the transect, I and experts observed both two side to record butterfly abundance. The study was conducted in both dry and wet season. In the dry season, the survey was carried from 31st March to 12th April. During this survey, the rain did not happen all the time. It contrasts to the study in the wet season when the rain occurred every day after 3pm

18 3.4. Data analysis Butterfly species diversity indices (Shannon_H, Menhinick, Margalef indices, Equitability_J and Fishers α) and dominance indices (Dominance_D; Simpson_1-D; BergerParker) were calculated by the following formulas: Simpson biodiversity index

∑

Where is the number of individuals displaying one trait; and = the total number of all individuals (Simpson, 1949) Buzas and Gibson's evenness

Where H (Shannon index) = -sum((ni/n)ln(ni/n)) and S = number of taxa (Hammer, 2001). Margalef diversity index:

where S is the number of species, and N is the total number of individuals in the sample (Gamito, 2010; Margalef, 1958). Fisher’s log-series

( )

where S is the number of taxa, n is the number of individuals and α is Fisher's alpha (Fisher, Corbet, & Williams, 1943; Hammer, 2001). Individual rarefaction:

19 It will estimate how many taxa expect to find in a sample with a smaller total number of individuals (Hammer, 2001). Rarefaction curves are species distribution as a function of an individual from a sample (Checa Villafuerte, 2016) Let N be the total number of individuals in the sample, s the total number of species, and Ni, the number of individuals of species number i. The expected number of species E(Sn) in a sample of size n and the variance V(Sn) are then given by

Many programs could be used to calculate for diversity indexes. In this study, data was analyzed by Past program version 3.2 (Hammer, 2001) Data will be analyzed using Correspondence Analysis (CA) ordination to calculate the relative significance of spatial and temporal variability in their compositions. Assemblage data were calculated by Correspondence Analysis that is a direct ordination method relating the composition of samples to variables (season, transect and vertical level) (Hammer, 2001).

20 IV. RESULTS 4.1. Butterflies in Kon Ka Kinh National Park 4.1.1. The butterfly fauna A total of 323 butterfly species were recorded at Kon Ka Kinh National Park, including 235 species recently recorded during our survey in 2018 (Appendix 1). These species belong to six butterfly families. Nymphalidae is the dominant family during the observation (Figure 3). Percentage of number of species recorded in each family included Nymphalidae - 41.8%; Lycaenidae - 21.1%; Hesperiidae - 19.0%; Pieridae - 8.4%; Papilionidae - 5.9%; and Riodinidae - 3.8%. Percentage of the butterfly species will slightly change if we consider the species abundances recorded. The ranking of butterfly families based on a percentage of species number and their abundance looks similar to preceding analysis: Nymphalidae, Lycaenidae, Pieridae, Hesperiidae, Papilionidae, and Riodinidae (Figure 3)

21 60.0%

50.0%

40.0%

30.0%

20.0%

10.0%

0.0% Nymphalidae Lycaenidae Hesperiidae Pieridae Papilionidae Riodinidae

without within Vietnam

Figure 3: Percentage of butterfly species in each family recorded in Kon Ka Kinh N.P. (with and without consideration of species abundance) and entirely in Vietnam

During the transect (see Figure 4), the number of the recorded taxon is highest in transect 1 (163 species) and lowest in transect 3 (41 species). Flat valley and riverine forest (transect 1) are characterized by highest species richness (163 spp.) whereas the lowest species richness was observed at previously disturbed forest area (41 spp.) (Figure 4). Nymphalidae is the dominant family in all five transects. The highest percentage of this family occurred in transect 5, with 53.45%

22 120.00%

100.00% 12.07% 16.82% 19.51% 18.75% 24.54% 5.17% 80.00% 3.74% 2.08% 4.88% 4.17% 8.62% 3.07% 10.28% 1.72% 12.50% 9.82% 5.61% 17.07% 60.00% 6.13% 7.32% 53.45% 40.00% 44.86% 39.88% 47.92% 39.02%

20.00%

18.69% 18.97% 16.56% 12.20% 14.58% 0.00% Trans_1 Trans_2 Trans_3 Trans_4 Trans_5

Hesperiidae Nymphalidae Papilionidae Pieridae Riodinidae Lycaenidae

Figure 4: Taxonomical compositions of butterflies in Kon Ka Kinh N.P.

4.1.2. Ecological complexes of tropical butterflies in Kon Ka Kinh N.P. Common and forested butterflies in Kon Ka Kinh N.P.: In total, 48 (20.3%) common, and 82 (34.3%) forest butterfly species were found in Kon Ka Kinh national park (Appendix). Percentage of common and forest species recorded in each transect is shown in Figure 5. The number of forest species is higher than common butterflies in all transect. Besides, the range of common species is 10.4% to 25.2%, and for forest species is 29.4% to 47.9%.

23 120.0%

100.0%

80.0% 41.3% 43.1% 45.4% 51.4% 50.0%

60.0%

40.0% 29.4% 30.8% 36.1% 52.2% 44.8% 20.0% 25.2% 17.8% 13.9% 12.1% 0.0% 6.5% Trans_1 Trans_2 Trans_3 Trans_4 Trans_5

% common % forest % other

Figure 5: Percentage of common and forest butterfly species recorded in transects

- In transect 1, the survey counted 163 species, 443 records. Total 41 common species were recorded in 112 (25.28%) records. The number forested butterfly is 48 species in 136 records. - Transect 2: 107 butterfly species were counted in 261 records. There are 20 common species and 32 forested species - Transect 3: 41 species were found in 81 records. In there, only five species are the common butterfly, and 14 species are the forested butterfly. - Transect 4: along the transect, we found 48 species in 112 records. There were five the common butterflies and 23 forested species - Transect 5: 58 species distributed in this transect. Among them, there were seven common butterfly and 27 forest species 4.1.3. Kon Ka Kinh butterfly in conservation The butterfly is the most traded order; even they are the most focus of conservation group in an invertebrate group. These survey recorded three observations

24 in total ten butterfly species in Vietnam Data Red book (Ban et al., 2007). All of them was classified as Vulnerable in Vietnam Red book: Stichophthalma uemurai, Troides helena, Teinopalpus imperialis. Two of them were recorded at below 1100 elevation: St. uemurai, and T. helena. The only one species was recorded at Kon Ka Kinh peak, T. imperialis, which also belong to NT criteria in IUCN Red List (Collins & Morris, 1985; Gimenez Dixon, 1996). Many wild animals hunted from protected areas are sold to foreign markets, and this extends to butterflies and other insects as well. For instance, T. helena and T. imperialis was listed in CITES Appendix II (UNEP-WCMC (Comps.), 2014) 4.1.4. New distribution records: In this study, some new information on butterfly ecology were found: Three new records for the butterfly of Vietnam: Taraka mahanetra Doherty, 1890; Logania watsoniana; and Callenya lenya Evans,1932. Furthermore, the new information on butterfly niche for Fanis genus Beside of new records for Vietnam, there were three first record for Central of Vietnam: Coladenia indrani; Logania sp.; and Ethope diademoides. 4.1.5. Biogeographical features Endemic species of Vietnam and Indochina found in Kon Ka Kinh National Park During previous and current surveys some species characterized by restricted distribution were observed in Kon Ka Kinh National Park. Some species are the endemics of Central Vietnam which have not been seen in other parts of the country and outside Vietnam. For example, Aemona simulatrix, Aemona kontumei, Stichophthalma eamesi During the survey, eight endemic species of Vietnam were discovered in Kon Ka Kinh National Park (Table 1). Among them, six species were recorded in the wet season and four species in dry season. There are two species recorded in both dry and wet season

25 F. bicoloratus and D. vietnamensis. The distribution of F. bicoloratus did not change during two seasons, while the distribution of D. vietnamensis is altered. In the first survey, dry season, D. vietnamensis was recorded three species in Kon Ka Kinh top (transect 5). While in the wet season, we found only one specimen in transect 3 and did not record in transect 5. Table 1: List of endemic species in Kon Ka Kinh National Park No. Dry Wet Family Subfamily Species season season 1 Nymphalidae Satyrinae Aemona simulatrix x 2 Nymphalidae Satyrinae Faunis bicoloratus x x 3 Nymphalidae Satyrinae Stichophthalma eamesi x 4 Nymphalidae Satyrinae Stichophthalma uemurai x 5 Nymphalidae Satyrinae Enispe cycnus x 6 Nymphalidae Satyrinae Lethe melisana x 7 Pieridae Pierinae Delias vietnamensis x x 8 Riodinidae Riodininae Taxila dora x

4.2. Butterfly indexes in Kon Ka Kinh National Park Diversity indices The diversity indices are calculated for the butterfly fauna based on their distribution along the transects. In this report, data were analyzed both in theory information (Simpson biodiversity index) and statistic information (Fisher’s alpha and Margalef). In Table 2, the data was calculated by Past version 3.0. The result demonstrates that Kon Ka Kinh is a high diversity value region of butterfly fauna (Simpson’s value: 0.9834) The Simpson biodiversity index in all transect is higher than 0.920. It means based on theory; in Kon Ka Kinh National Park, all five surveyed transect is high butterfly diversity value. The butterfly community richness is concentrated in transect

26 1 while it’s value (0.9859) is slightly higher than the Simpson’s value of total butterfly fauna (0.9834).

27

Table 2: Butterfly diversity indices Trans_1 Trans_2 Trans_3 Trans_4 Trans_5 Total Taxa_S 163 106 37 45 58 235 Individuals 443 258 77 109 104 995 Simpson_1-D 0.9859 0.9741 0.9314 0.9359 0.9577 0.9834 Margalef 26.59 18.91 8.288 9.379 12.27 34.19 Fisher_alpha 93.12 67.25 27.98 28.69 54.06 98.43 The statistic information from Margalef and Fisher’s alpha demonstrate the same result; transect 1 is highest butterfly’s diversity. The diversity model of statistic is homologous to the result from theory. However, the values of Margalef and Fisher’s alpha will be higher if data is calculated for all five transects. It means the species component in Kon Ka Kinh is significantly different (Table 3) Table 3: Diversity permutation test

Perm Perm Perm Perm Perm Perm Perm Perm Perm Perm p(1_2) p(1_3) p(1_4) p(1_5) p(2_3) p(2_4) p(2_5) p(3_4) p(3_5) p(4_5) Simpson 0.0001 0.0001 0.0001 0.0002 0.0007 0.0002 0.028 0.7883 0.0239 0.0756 Margalef 0.0105 0.0001 0.0002 0.2159 0.004 0.0049 0.4874 0.6975 0.0279 0.0299 Fisher alpha 0.0189 0.009 0.0027 0.0783 0.0053 0.0083 0.4511 0.9478 0.0615 0.0472 Comparing diversities in two samples of abundance data in Table 3 demonstrate the similarity of abundance in five transects. Butterfly abundance in transect 3 and transect 4 is similar (p_simpson = 0.7883; p_margalef = 0.6975; p_fisher = 0.9478). Other pairs of permutation test imply that they are different. 4.3. Factors influence to butterfly population distribution Many research demonstrated that many factors affect tropical butterfly distribution, such as floral variables, stream, elevation, forest cover (K. C. Hamer et al., 2006; J. K. Hill, 1999; A. Monastyrskii, 2007b; L. V. Vu & Quang Vu, 2011; V. L. Vu & Yuan, 2003). In this study, to determine the factors affect butterfly diversity in Kon Ka Kinh National Park, we divided data by the following factors: season; water

28 condition; forest types and altitude. We use Simpson t-test for comparing the butterfly abundance to determine the factor impact to butterfly fauna.

Figure 6 illustrated the correlation between butterfly species in Kon Ka Kinh National Park. The butterfly concentrated in the following characters: wet season, low elevation, available water, and disturbance forest.

Figure 6: Correspondence analysis The data will be compared in separate factors to understand how factors influence the butterfly diversity. Seasonality: Testing on Renyi index in Past program prove the diversities are comparable. The results of the butterfly population in the dry and rainy season were shown in Figure 77. It demonstrates the higher butterfly richness in the wet season. In the dry season, the butterfly is less diverse both in a number species and number an individual. Comparison of the diversities in two samples, we used a t-test, which described by Poole in the Past program. It indicated two butterfly populations in dry season and wet

29 season are significantly different (t = 5.872, df= 410.11; p= 8.9037E-09). Therefore, it illustrates the impact of seasons on butterfly fauna.

Figure 7: Butterfly rarefaction curve for seasons Water resource factor: Transect 1 and transect 4 are similar in term of water resources condition. Streams are present in both of them. There is no stream or above groundwater sources in transect 2 and 5. So to determine the influences of groundwater on butterfly distribution, butterfly data will be compared in a pair of group 1_4 and group 2_5 on Figure 8Figure 8: Butterfly rarefaction curve based on water condition The result from diversity t-test for Simpson (t = -2.372, df =615.09, p = 0.017994) performs the butterfly abundance in Kon Ka Kinh is significantly different between two conditions of the water source. Rarefaction curve confirmed the higher butterfly richness when water is available. The number of observation for butterfly in there is also higher than in none water forest areas.

30

Figure 8: Butterfly rarefaction curve based on water condition Forest types and altitude: Both factors, forest types, and elevation have the same influence to butterfly distribution in pair transect 1, 2; transect 4, 5 on Figure 9

Figure 9: Rarefaction curves by forest types and altitude factors

31 We tested the sampling conditions in both transect 1_2 and transect 4_5. Using diversity t-test for Simpson, the result (t= -4.5305, df = 229.04, and p = 8.89E-06) demonstrated the significant difference of butterfly diversity between two conditions in transect 1_2 and transect 4_5. It confirms the effect of the forest types and elevation on butterflies’ distribution. The sample size was standardized through rarefaction curves. Butterfly species richness was higher in transect 1_2. It means the butterfly diversity in disturbed sites below 1200m is higher than their distribution in high elevation and primary forest. The number of observation in low-land disturbance area is higher than in the primary. Butterfly rarefaction curve in transect 4_5 is under the rarefaction curve of transect 1_2. However, the trending in both two communities are similar within less than 200 observations

32 V. DISCUSSION 5.1. Butterfly fauna in Kon Ka Kinh National Park In this study, butterflies were collected by hand-nets, and the observation followed the Pollard walk method for data collection (Pollard, 1975). This method was similar to what was used in the survey in 1999 (Le, 2000). Furthermore, the butterfly fauna sampling from observation with hand-nets is more variation in species diversity in the tropical forest (Caldas & Robbins, 2003). Data from Pollard walk is more representative monitoring of rainforest butterfly assemblages (Basset et al., 2011). In this survey, 235 species were recorded in Kon Ka Kinh. Compared to the previous survey in 1999, 89 species did not record in this survey. However, it contributed 124 new records for the Park. The reason for population distinction may be the periods of study (Basset et al., 2011; Spitzer et al., 1993). In the previous study, the research was carried only in dry season (Le, 2000), but in this report, research periods consider both dry and wet season. It is difficult to compare the data come from two short periods in different years (Spitzer et al., 1993). Total butterfly species in Kon Ka Kinh is 323 recorded species. Because of environmental impact to butterfly diversity (Ackery & Vane-Wright, 1989; Checa Villafuerte, 2016), the similarity of natural condition leads the homogeneous on butterfly fauna between Kon Ka Kinh N.P and Ngoc Linh N.P, Kon Tum. Based on biogeographical zonality, the specific biogeographical features of the Vietnamese fauna of butterflies demonstrated by the example of classification (A. Monastyrskii, 2007b). Nine distributional biogeographical classes were recognized from most restricted to the most widespread: 1. Endemics of Indochina: Vietnam, , Cambodia and E. ; 2. Species range: E. Himalaya and S. China; 3. Indo- Burmese fauna (from to Indochina); 4. Indo-Malayan distribution; 5. Indo- Malayan and Australasian species; 6. Species distributed mainly from the Palaearctic extending into the Indo-Malayan region; 7. Old-World tropics species; 8. Species mostly

33 distributed in the Holarctic extending into the Indo-Malayan region; 9. Cosmopolitans. During the survey, we found some new butterfly distribution in Vietnam. Three new records for Vietnam: Taraka mahanetra; Callenya lenya; and Logania watsoniana. Besides the new record for Vietnam, there were three new record of Central Vietnam: Coladenia indrani; Logania sp.; and Ethope diademoides. The further information of their distribution discussed below: There are two species in Taraka genera in Indo-China region: T. hamada, and T. mahanetra (Inayoshi, 2018). The species T. Mahanetra distributed in Peninsula Thailand, the Malay Peninsula and Borneo (Corbet, 1938; Marshall & de Nicéville, 1890). In Vietnam, during the research, this species and C. lenya were found in transect 4 in primary forest. In Indo-China region, C.lenya is the broader distribution in forested habitat in southern , Peninsula Thailand, Malay Peninsula, Borneo (Inayoshi, 2018). For another new record, L. watsoniana species, the previous its distribution was reported in Manipur, Myanmar, Thailand, Laos (Inayoshi, 2018) In Vietnam, C. indrani species was found in the South of Vietnam at Binh Thuan province. During the present survey, this species was found in the rainy season. The Logania sp. also was encountered in this season. This survey was the first time recorded Logania genera in Central Vietnam (A. L. Monastyrskii & Devyatkin, 2015). We need to do more taxonomical work to understand this specimen in the future. For another species, E. diademoides was found in Lam Dong province in previous data(A. L. Monastyrskii & Devyatkin, 2015) In Kon Ka Kinh, this survey recorded three species in Faunis genera: F. bicoloratus, F. canens, and F. eumeus. In there, F. eumeus is a common butterfly and wide distribute in Vietnam (A. L. Monastyrskii, 2011), e.g., Cuc Phuong N.P (Torben B Larsen et al., 2005), Kon Ka Kinh N.P (Le, 2000). However, this is the first time F. canens was collected in the same area with F. bicoloratus. The species F. canens niche in the previous study have been being distributed in North Vietnam up to Quang Tri

34 province (A. L. Monastyrskii, 2011; A. L. Monastyrskii & Devyatkin, 2015), while species F. bicoloratus was recorded from Quang Ngai province to the South of Vietnam (Inayoshi, 2018; A. L. Monastyrskii, 2011). 5.2. Butterflies diversity along the transect In Vietnam, about 1200 butterfly species were described. In there, the proportion of six families in turn Papilionidae (5.8%), Pieridae (4.8%), Nymphalidae (35.4%), Riodinidae (2.6%), Lycaenidae (26.9%), and Hesperiidae (24.4%) (A. L. Monastyrskii & Devyatkin, 2015). In our results, although the order of Pieridae, Papilionidae, and Riodinidae differ from the ratio of number species in Vietnam, the ranking of remaining families is similar to their order in butterflies of Vietnam. Generally, the study on correlations among taxa in the richness, or the relative importance of rare and common species is poor (Pearman & Weber, 2007). Common butterflies show the apparent response to changes in soil nutrient status, acidity or moisture (Oostermeijer & Van Swaay, 1998). In Vietnam, there are 105 common species (A. L. Monastyrskii & Devyatkin, 2002). The disturbed and riverine forest are area has a higher proportion of common species (L. V. Vu & Quang Vu, 2011). The most common species occurs in low and middle altitudes (A. Monastyrskii, 2007b). It is similar to our results in Kon Ka Kinh National Park in the present study. Monitoring of butterfly species richness may provide information on rare butterflies (Pearman & Weber, 2007). The endemic species occur mainly in isolated montane areas, such as Kon Tum and Da Lat plateaus. In Kon Tum plateau, there was a high diversity of endemic butterflies (A. Monastyrskii & Holloway, 2013). The present study found six endemic species in both dry and wet season. The different factors in every transect led to differences in butterfly diversity, richness (J. Hill et al., 1995; Rod & Ken, 1988). Each transect represented for specific characters of the forest. They could be the causes of differences in transect diversity

35 indices. The Simpson index is higher in better forest condition (K. Hamer et al., 2005). It contrasted to his previous result in 2003 (K. Hamer et al., 2003). 5.3. Factors impact to butterfly diversity: Season factor: Butterfly species in one season is not necessarily ecologically equivalent to that butterfly species in another season. The seasonal dynamic of butterfly fauna increase in the wet season, and a decrease in the dry season (Bonebrake et al., 2010). Butterfly species richness is most closely associated with annual temperature sum because of butterflies less active in a cold season (Torben B Larsen et al., 2005; Menéndez et al., 2007). Butterfly populations increased significantly during the period of high temperature and low precipitation (Boonvanno, Watanasit, & Permkam, 2000). The statistic information proves the significant difference of butterfly fauna richness between rainy and dry season. The causes of different on butterfly fauna not only by plant phenology, climate generally but also biogeography and fauna history evolution (Spitzer et al., 1993). The more diverse of butterfly community in the rainy season similar to other researches, such as Cuc Phuong N.P. (Torben B Larsen et al., 2005), Pu Mat N.P. (A. L. Monastyrskii & Pham, 2016), Tam Dao N.P. (Spitzer et al., 1993) Water condition Butterflies are sensitive to the water source (Oostermeijer & Van Swaay, 1998). The living environment condition attracts more butterflies (L. V. Vu & Quang Vu, 2011). Because the streamside provide mud- patches or water-seepages, ingesting salts, minerals, and water, butterflies concentrate along the rivers (Torben B Larsen et al., 2005). In our results, butterfly abundant is significantly higher in streamsides. It differs from the study in Czech - the water body is unimportant to butterflies (Storch, Konvicka, Benes, Martinková, & Gaston, 2003). It could be explained by butterfly species in Vietnam is strong differ from Czech.

36 Habitat and geographic factors: Vietnam is facing the incredible rate of habitat loss and degradation in the humid tropics. Many endemic butterflies are meeting global extinction current levels of deforestation to continue (Koh, 2007; Koh et al., 2004). Our result indicates the influence of habitat and geographic to butterflies distribution. In the reviews of butterfly fauna in South East Asia, Koh pointed out 16 studies showed the effect of the environment to butterfly richness (Koh, 2007), similar to our conclusion of the third testing. The positive impact of primary forest to butterfly distribution was reported in seven papers, while nine reports showed the negative results in undisturbed forest (Koh, 2007). The geographic location may be a driving factor influencing the effectiveness of butterfly indicators(Grealey, 2011). Furthermore, in Thailand, species with the smallest geographic ranges tend to be the least abundant and occurred most frequently in the undisturbed site (Ghazoul, 2002). Besides, the impact of altitude on butterfly also verified in India, though the geographical classification is different with the present study (Sreekumar & Balakrishnan, 2001). However, the geographic do not relate to genetic distance (Benedick et al., 2007)

37 VI. CONCLUSION After 19 years, the survey repeated the Pollard walk for butterfly survey. The present study was conducted in both dry and wet season in Kon Ka Kinh N.P, Gia Lai. Total 323 butterfly species were recorded in Kon Ka Kinh National Park, including 235 species recorded in 2018. Compare to previous data in 1999, this survey added 124 new records to the Park, while 89 species were not recorded

All six butterfly families in Vietnam were recorded in Kon Ka Kinh N.P. The proportion of butterfly families in the community decrease from Nymphalidae, Lycaenidae, Hesperiidae, Pieridae, Papilionidae, end at Riodinidae. Along the transects, Nymphalidae also was the dominant family.

There were 20.3% common butterflies, and 34.3% forested species. The number of forest species is higher than common butterflies in all transect. It illustrates the Kon Ka Kinh N.P is a well-protected forest. It plays an important role in butterfly conservation in Vietnam. Furthermore, the proportions between common species and forested butterflies along the transect 4 and 5 demonstrate the high diversity in these areas. The forest status among them is the best for butterfly conservation. In addition, eight endemic butterfly species were recorded during the research. Most of them were found in the wet season. There were three species in the present survey belonged to the Vulnerable group in Vietnam Data Red book

During the research, the study contributed three new records for Vietnam: Taraka mahanetra Doherty; Logania watsoniana de Nicéville; and Callenya lenya Evans. Both T. mahanetra and C. lenya were found in transect 4, where is the highest number of forested species. The L. watsoniana was found in transect 1, where is highest butterfly diversity. There were three new records for Central Vietnam: Coladenia indrani; Logania sp.; and Ethope diademoides. We also verified the

38 overlap of two species in Faunis genera, F. bicoloratus, F. canens. This is the first time F. canens was recorded in Kon Ka Kinh N.P.

All of the transects is high diversity, the Simpson value of total recorded species in the present survey is 0.9834. Transect 1 is highest butterfly diversity. Diversity permutation test indicates high homogeneous on butterfly diversity between transect 3 and 4, while others are significant differences.

The present data proved the impact of environmental factors, season; water condition; forest types and altitude, to butterfly distribution. In the wet season, the butterfly is significant higher diversity and richness. The butterfly is more concentrated along the riverine forest or somewhere exist above groundwater. The result of the last factor, forest types, and altitude concluded the butterfly is more diverse in the disturbed area at lower altitude forest.

39 VII. ACKNOWLEDGMENT Sincere gratitude to my advisors, Alexander L. Monastyrskii and Le Bao Thanh, for their guidance, and butterflies’ identification during the thesis. Thank you Vu Tien Thinh for all your help with my data analysis. I would also like to thank David Lohman, Bui Manh Hung, and Hoang Van Lam for taking the time to review and edit the proposal and lending your various areas of expertise

The research presented in this paper would not have been possible without support from the study site, including Nguyen Van Hoan, and all staffs of Kon Ka Kinh National Park. A special thanks to To Van Quang and Nguyen Anh Tuan who assisted with field surveys

I would also like to thank the foundations, Rufford and National Geographic Young Explorer, for providing funding for carrying out fieldworks. I would also like to thank my employer, Vietnam Forestry Museum, for their flexibility and support through this process. Thank the Vietnam National University of Forestry for their support for my study.

Lastly, I especially thank my parents and sister for their constant loving support; thanks to my friends for their encouragement.

40 VIII. APPENDIX

Camping in nature forest Transect 1

Transect 2 Transect 3

Transect 4 Transect 5 © All pic. Tan Pham - 2018

41

Delias belladonna Aemona simulatrix

Taraka mahanetra Lexias dirtea

Stichophthalma eamesi Faunis eumeus © All pic. Tan Pham - 2018

42 1 - endemic species of Vietnam; 2 - endemic species of Indochina; 3 - Endemic subspecies of Vietnam and Indochina; 4- the first time in Vietnam; 5 - the first time in Central Vietnam * common butterfly ** forested butterfly

Dry Dry Wet Family/ n/n Genus/Species season season season Subfamily 1999 (2018) (2018) Hesperiidae

1 Coeliadinae Choaspes subcaudatus - v - 2 Coeliadinae Choaspes stigmatus** v - v 3 Coeliadinae Choaspes furcatus** v - - 4 Coeliadinae Hasora vitta* v v - 5 Coeliadinae Hasora chromus v - - 6 Coeliadinae Hasora schoenherr** v - - 7 Coeliadinae Hasora malayana v - - 8 Coeliadinae Hasora taminatus v - - 9 Coeliadinae Badamia exclamationis* v - - 10 Pyrginae Capila lidderdali** - - v 11 Pyrginae Capila pauripunetata** 3 - - v 12 Pyrginae Capila penicillatum** 3 - - v 13 Pyrginae Celaenorrhinus asmara** v - - 14 Pyrginae Celaenorrhinus aspersa** - - v Celaenorrhinus 15 Pyrginae - - v leucocera** 16 Pyrginae Celaenorrhinus patula** - - v 17 Pyrginae Celaenorrhinus putra** v v v 18 Pyrginae Celaenorrhinus sp. - - v 19 Pyrginae Coladenia agni - - v 20 Pyrginae Coladenia indrani 5 - - v 21 Pyrginae Darpa striata v - - 22 Pyrginae Darpa pteria v - v 23 Pyrginae Gerosis tristis** - - v 24 Pyrginae Pseudocoladenia dan* - - v 25 Pyrginae Mooreana trichoneura - - v 26 Pyrginae Seseria sambara 3 v - - 27 Pyrginae Satarupa gopala** - - v

43 28 Pyrginae Tagiades cohaerens - - v 29 Pyrginae Tagiades gana* - - v 30 Pyrginae Tagiades japetus - v - 31 Pyrginae Tagiades litigiosa* - - v 32 Pyrginae Tagiades menaka - - v 33 Hesperiinae Ancistroides nigrita* - v v 34 Hesperiinae Astictopterus jama* v v v 35 Hesperiinae Arnetta atkinsoni* v - - 36 Hesperiinae Baoris penicillata - v - 37 Hesperiinae Borbo cinnara v - - 38 Hesperiinae Caltoris sirius v - - 39 Hesperiinae Caltoris tenuis v - - 40 Hesperiinae Caltoris cormasa v - - 41 Hesperiinae Caltoris confusa v - - 42 Hesperiinae Halpe sp. 1 - - v 43 Hesperiinae Halpe sp. 2 - - v 44 Hesperiinae Halpe pelethronix v - - 45 Hesperiinae Hyarotis adrastus v - - 46 Hesperiinae Iambrix salsala* v - - 47 Hesperiinae Iton semamora - - v 48 Hesperiinae Koruthaialos rubecula ** - - v 49 Hesperiinae Koruthaialos sindu** - v - 50 Hesperiinae Lotongus calathus v - - 51 Hesperiinae Matapa sasivarna** - - v 52 Hesperiinae Notocrypta clavata - v v 53 Hesperiinae Notocrypta curvifascia** - - v 54 Hesperiinae Notocrypta feisthamelii** v v v 55 Hesperiinae Notocrypta paralysos* - v v 56 Hesperiinae Oriens goloides - - v 57 Hesperiinae Parnara sp. - v - 58 Hesperiinae Parnara bada v - - 59 Hesperiinae Polytremis eltola - v v 60 Hesperiinae Polytremis lubricans* v v - 61 Hesperiinae Potanthus sp. v - v 62 Hesperiinae Pyroneura margherita v - - 63 Hesperiinae Sebastonyma dolopia** - v - 64 Hesperiinae Suada swerga** v - -

44 65 Hesperiinae Telicota ohara - v - 66 Hesperiinae Thoressa sp. - - v Lycaenidae 67 Curetinae Curetis bulis* v - v 68 Lycaeninae Heliophorus epicles - - v 69 Lycaeninae Heliophorus ila v v v 70 Lycaeninae Heliophorus sp. - v - 71 Miletinae Allotinus drumila** v v v 72 Miletinae Logania sp.** 5 - - v 73 Miletinae Logania watsoniana** 4 - - v 74 Miletinae Miletus chinensis v - v 75 Miletinae Miletus croton - - v 76 Miletinae Miletus mallus v v v 77 Miletinae Taraka hamada - v - 78 Miletinae Taraka mahanetra** 4 - v - 79 Acytolepis puspa v v v 80 Polyommatinae Caleta elna - v v 81 Polyommatinae Caleta roxus v v v 82 Polyommatinae Callenya lenya ?** 4 - v - 83 Polyommatinae Callenya melaena** v v v 84 Polyommatinae Castalius rosimon* - v - 85 Polyommatinae Catochrysops panormus* - - v 86 Polyommatinae Celastrina lavendularis - - v 87 Polyommatinae Celatoxia marginata v v v 88 Polyommatinae Discolampa ethion v - v 89 Polyommatinae Ionolyce helicon - - v 90 Polyommatinae Jamides alecto** - - v 91 Polyommatinae Jamides bochus - - v 92 Polyommatinae Jamides caeruleus** - v - 93 Polyommatinae Jamides celeno* v v - 94 Polyommatinae Jamides pura v - - 95 Polyommatinae Leptotes plinius - - v 96 Polyommatinae Lycaenopsis haraldus - v v 97 Polyommatinae Megisba malaya v - v 98 Polyommatinae Nacaduba beroe - v - 99 Polyommatinae Nacaduba sp. v - v 100 Polyommatinae Niphanda tesselata - v -

45 101 Polyommatinae Pithecops corvus v v v 102 Polyommatinae Prosotas bhutea** - v v 103 Polyommatinae Prosotas sp. v v - 104 Polyommatinae Udara akasa** - v - 105 Polyommatinae Udara albocaerulea** v - v 106 Polyommatinae Udara dilecta v v v 107 Polyommatinae Udara placidula** v v v 108 Polyommatinae Zizina otis v - - 109 Polyommatinae Zizeeria maha* - - v Ravenna nivea 110 3 v - - miyagawai** 111 Theclinae Amblypodia anita* - v v 112 Theclinae atosia** - - v 113 Theclinae Arhopala aurelia** - - v 114 Theclinae Arhopala birmana** v - - 115 Theclinae Arhopala eumolphus** v - - 116 Theclinae Bindahara phocides - v - 117 Theclinae Hypolycaena amasa* - - v 118 Theclinae Hypolycaena erylus* v - v 119 Theclinae Rapala rhoecus - - v 120 Theclinae Rapala manea v - - 121 Theclinae Rapala pheretima v - - 122 Theclinae Rapala nissa v - - 123 Theclinae Surendra quercetorum v - v 124 Theclinae tripunctata v v v Nymphalidae 125 Apaturinae Eulaceura osteria - v v 126 Apaturinae Hestinalis nama - v - Rohana nakula 127 Apaturinae 3 - - v bernardii** 128 Apaturinae Rohana parisatis** - v v 129 Biblidinae Ariadne merione v v v 130 Biblidinae Chersonesia risa v - v 131 Biblidinae Chersonesia intermedia** - v - 132 Biblidinae Cyrestis thyodamas* v v v 133 Biblidinae Cyrestis themire** v - - 134 Biblidinae Laringa horsfieldi** v - -

46 Dichorragia 135 Biblidinae v v - nesimachus** 136 Biblidinae Stibochiona nicea** v - v Calinaga sudassana 137 Calinaginae 3 - v - distans 138 Charaxinae Charaxes kahruba** - v - 139 Charaxinae Polyura athamas* - - v 140 Danainae Danaus genutia* v - - 141 Danainae Ideopsis vulgaris* v - - 142 Danainae Euploea algea v v v 143 Danainae Euploea doubledayi v v v 144 Danainae Euploea eunice - v - 145 Danainae Euploea mulciber* v v v 146 Danainae Euploea radamanthus* v v v 147 Danainae Euploea core v - - 148 Danainae Euploea tulliolus* v - - 149 Danainae Parantica aglea* v v v 150 Danainae Parantica melaneus v v v 151 Danainae Parantica sita v - v 152 Danainae Parantica swinhoei - v - 153 Heliconiinae Argyreus hyperbius* v - - 154 Heliconiinae Phalanta phalantha v - - 155 Heliconiinae Cirrochroa chione** 2 v - v 156 Heliconiinae Cirrochroa tyche v v v 157 Heliconiinae Paduca fasciata** v - - 158 Heliconiinae Cupha erymanthis* v v v Terinos clarissa 159 Heliconiinae 3 v - v militum** 160 Heliconiinae Vagrans egista* v - v 161 Heliconiinae Vindula erota* - - v 162 Heliconiinae Cethosia biblis v - - 163 Limentidinae Euthalia monina - - v 164 Limentidinae Euthalia bunzoi ssp.** 3 - - v 165 Limentidinae asura - - v 166 Limentidinae Athyma cama** - - v 167 Limentidinae Athyma kanwa** - v v 168 Limentidinae Athyma nefte** - - v 169 Limentidinae v - v

47 170 Limentidinae Bhagadatta austenia** - - v 171 Limentidinae Cynitia lepidea* v v - 172 Limentidinae Lebadea martha** v - - 173 Limentidinae Lexias dirtea** v v v 174 Limentidinae Moduza procris* - - v 175 Limentidinae Neptis ananta** v v v 176 Limentidinae Neptis clinia* - v v 177 Limentidinae Neptis hylas* v v v 178 Limentidinae Neptis leucoporos** v v - 179 Limentidinae Neptis miah v - v 180 Limentidinae Neptis namba - - v 181 Limentidinae Neptis nata - - v 182 Limentidinae Neptis soma** - - v 183 Limentidinae Neptis zaida** 3 v v v 184 Limentidinae Neptis yerburii v - - 185 Limentidinae Neptis sankara v - - 186 Limentidinae Neptis cartica** 3 v - - 187 Limentidinae Neptis radha** v - - 188 Limentidinae Neurosigma siva** v v v 189 Limentidinae Pantoporia aurelia** v - v 190 Limentidinae Pantoporia sandaka - v - 191 Limentidinae Phaedyma columella v - v 192 Limentidinae Sumalia daraxa** v - - 193 Limentidinae Tanaecia julii* v v v 194 Libytheinae Libythea myrrha v - v 195 Libytheinae Libythea geoffroyi v - - 196 Doleschallia bisaltide* - - v 197 Nymphalinae Symbrenthia hypselis - - v 198 Nymphalinae Symbrenthia lilaea* v v - 199 Nymphalinae Hypolimnas bolina* v - - 200 Nymphalinae Junonia almana* v - - 201 Nymphalinae Junonia atlites* v - - 202 Nymphalinae Junonia iphita* - - v 203 Nymphalinae Junonia lemonias v v v 204 Nymphalinae Kallima alicia** 3 v v - 205 Nymphalinae Kallima incognita** - - v 206 Nymphalinae Kaniska canace* v v -

48 207 Satyrinae Aemona simulatrix** 1 v - v 208 Satyrinae Aemona kontumei** 1 v v v 209 Satyrinae Aemona tonkinensis** 3 - v v 210 Satyrinae Callerebia narasingha** v v - 211 Satyrinae Coelites nothis** v - v 212 Satyrinae Elymnias patna** v - - 213 Satyrinae Enispe cycnus** v v - 214 Satyrinae Erites falcipennis** v v v 215 Satyrinae Ethope diademoides** 5 - v - 216 Satyrinae Faunis bicoloratus** 1 - v v 217 Satyrinae Faunis canens** - v v 218 Satyrinae Faunis eumeus* v v v 219 Satyrinae Lethe dura - - v 220 Satyrinae Lethe insana v v v 221 Satyrinae Lethe latiaris v - v 222 Satyrinae Lethe mekara* - v - 223 Satyrinae Lethe melisana 1 - - v 224 Satyrinae Lethe naga - - v 225 Satyrinae Lethe verma v v v 226 Satyrinae Lethe vindhya v v v 227 Satyrinae Lethe confusa* v - - 228 Satyrinae Lethe distans** v - - 229 Satyrinae Lethe kansa v - - 230 Satyrinae Lethe rohria v - - 231 Satyrinae Lethe sinorix** v - - 232 Satyrinae Lethe konkakini** 1 v - - 233 Satyrinae Mandarinia regalis** - v - 234 Satyrinae Melanitis leda* v - v 235 Satyrinae Melanitis phedima** - - v 236 Satyrinae Melanitis zitenius** v - - 237 Satyrinae Mycalesis anaxias** v v - 238 Satyrinae Mycalesis francisca** v v v 239 Satyrinae Mycalesis mineus* v v v 240 Satyrinae Mycalesis perseoides v - v 241 Satyrinae Mycalesis sangaica - v v 242 Satyrinae Mycalesis mucianus v - - 243 Satyrinae Mycalesis mnasicles** v - -

49 244 Satyrinae Neope bhadra** - - v 245 Satyrinae Penthema darlisa** v - - 246 Satyrinae Ragadia crisilda** - v v 247 Satyrinae Ragadia critias** v - - 248 Satyrinae Stichophthalma eamesi** 2 v - v Stichophthalma 249 Satyrinae 1 - - v mathilda** Stichophthalma 250 Satyrinae 1 - - v uemurai** 251 Satyrinae Thaumantis diores** v v v 252 Satyrinae Ypthima huebneri* v - - 253 Satyrinae Ypthima tappana** v - - 254 Satyrinae Ypthima baldus* v v v 255 Satyrinae Ypthima dohertyi** v v - 256 Satyrinae Ypthima nebulosa - - v 257 Satyrinae Ypthima savara** v v - Papilionidae 258 Papilioninae Atrophaneura varuna** - - v 259 Papilioninae Byasa dasarada** v - - 260 Papilioninae Graphium macareus - v - 261 Papilioninae Graphium agamemnon* v v - 262 Papilioninae Graphium antiphates* - - v 263 Papilioninae Graphium agetes - v - 264 Papilioninae Graphium eurypylus v - - 265 Papilioninae Graphium doson* v - - 266 Papilioninae Lamproptera curius* - - v 267 Papilioninae Lamproptera meges* - - v 268 Papilioninae Meandrusa lachinus - v - 269 Papilioninae Papilio arcturus** - v - 270 Papilioninae Papilio protenor v - - 271 Papilioninae Papilio alcmenor** v - - 272 Papilioninae Papilio polytes* v - - 273 Papilioninae Papilio demoleus* v - - 274 Papilioninae Papilio helenus* v v v 275 Papilioninae Papilio memnon* v v v 276 Papilioninae Papilio nephelus** v - v Teinopalpus imperialis 277 Papilioninae 3 - v - gillesi**

50 278 Papilioninae Troides aeacus v - - 279 Papilioninae Troides helena v v v Pieridae 280 Coliadinae Catopsilia pomona* v v v 281 Coliadinae Catopsilia scylla v - - 282 Coliadinae Catopsilia pyranthe v - - 283 Coliadinae Dercas verhuelli** v v v 284 Coliadinae Eurema andersoni** v v v 285 Coliadinae * v v v 286 Coliadinae Eurema hecabe* v v v 287 Coliadinae Eurema ada indosinica** 3 v - - 288 Coliadinae Eurema brigitta v - - 289 Coliadinae Eurema novapallida** 2 - v v 290 Coliadinae Eurema simulatrix - - v 291 Pierinae Ixias pyrene* v - - 292 Pierinae Appias albina* v - v 293 Pierinae Appias indra v v v 294 Pierinae Appias lalassis - v v 295 Pierinae Appias pandione** v v v 296 Pierinae Appias nero v - - 297 Pierinae Appias lyncida* v - - 298 Pierinae Appias lalage v - - 299 Pierinae Appias olferna v - - 300 Pierinae nadina* - - v 301 Pierinae Prioneris philonome v - - 302 Pierinae Prioneris thestylis* v - v 303 Pierinae Delias vietnamensis** 2 v v v 304 Pierinae Delias belladonna** v v - Delias agostina 305 Pierinae 3 v - - annamitica** 306 Pierinae Delias pasithoe* v - - 307 Pierinae Hebomoia glaucippe* v - v 308 Pierinae Leptosia nina v v v 309 Pierinae Pareronia anais v - v 310 Pierinae Pieris canidia* v - - 311 Pierinae Talbotia naganum** 3 v v v Riodinidae

51 312 Riodinidae Abisara attenuata** v - v 313 Riodinidae Abisara burnii** - v v 314 Riodinidae Abisara fylla** v - v 315 Riodinidae Abisara neophron** v - v 316 Riodinidae Abisara echerius* v - - 317 Riodinidae Dodona ouida** v - v 318 Riodinidae Dodona egeon v - - 319 Riodinidae Dodona katerina** 1 v - - 320 Riodinidae Stiboges elodinia ** v - v 321 Riodinidae Taxila dora** 2 v - v 322 Riodinidae Taxila hainana** - v v 323 Riodinidae Zemeros flegyas* v v v

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