INTRODUCTION
There are a great number of fish inhabiting Indochina in East Asia and it is reported that around 1,000 species of freshwater fish inhabit the region including the Mekong River (Kottelat, 2001; Kottelat and Whitten, 1996; Kottelat et al., 1993; Rainboth, 1996). As there are a number of tributaries in the upstream of Mekong River where the vast part is a mountainous area, several hydroelectric power plants are currently in operation and under construction to produce electricity.
The Bolaven Plateau is located in a volcano that erupted millions of years ago, in southern Laos and is crossed by two main rivers including many tributaries and has many big waterfalls. Elevation of this pleateau ranges approximately from 800 to 1,300m.
In addition, Xe-Pian Xe-Namnoy Hydropower Project has been established to construct a hydroelectric power plant in the Bolaven Plateau, where there are two rivers flowing in the Xe Kong, one of the tributaries of the Mekong River. Dams will be built in these two rivers, the Xe Pian and the Xe Nammoy as part of the Xe-Pian Xe-Namnoy Hydropower Project in the Bolaven Plateau. Although, both rivers flow in Xe Kong, they are two river systems divided by the waterfall.
In previous years, the fish inhabiting these two rivers of the Bolaven Plateau have been studied by Kottelat (2001), and the study on the impact of Xe-Pian Xe-Namnoy Hydropower Plant Project on fish has been conducted by Kottelat (2013).
The fish fauna on the Bolaven Plateau is characterized by the high endemic rate (Kottelat, 2013). However, some species is doubted to validity of species owing to intraspecific variation (Kottelat, 2001, 2013; Reberts, 1998). The present study aimed to provide the fish fauna and distribution data on the Bolaven Plateau, and discuss intraspecific trophic morphs in Poropuntius bolovenensis.
MATERIALS AND METHODS
1.Survey Periods and Sites
The survey was conducted in the mainstream and tributaries of Xe Pian and Xe Namnoy in order to investigate fish fauna during the period of 11th March to 1st April 2014. A total of twenty sites were selected in two river systems on the Bolaven Plateau (Figure 1). The survey was conducted in the upstream, main stream and tributaries of Xe Pian and Xe Namnoy. The survey sites and the name of rivers are listed in Table 1 as below.
2.Study Methods
A rangefinder and direct observation were used to investigate the river width, the water width, the structure of river bed, and other features of the surrounding environment. Additionally, a portable electronic temperature gauge was used to measure the water temperature. Fishes were captured by using the gill net (10×10 mm), casting net (6×6 mm), and skimming net (5×5 mm). And the collected fishes were preserved in 10% formalin with photos, counts and measurements undertaken.
In the majority of survey areas, observation via skin diving was conducted underwater to investigate behavior of fishes and the underwater environment factors while taking photos and filming using an underwater digital camera. In each survey site, over an hour was spent in an approximately 20-50 meter radius in the pool and rapid of the river to film and observe fish.
The identification of fish was determined based on the study of Rainboth (1996), Roberts (1998), Kottelat (2001; 2013), and the scientific name and the classification were determined by Kottelat (2001, 2013) and Nelson (2006). A dissecting microscope was used to count and vernier calipers were used for measuring the collected samples.
RESULTS AND DISCUSSION
1.Physical Environment in Each Survey Area
The physical environment in each survey site describes Table 2 as below. The river bed structure of all sites is composed of bedrock was caused by volcano. However, the edge and flat water areas displayed boulders, pebbles, sand and mud.
2.Fish Fauna
A total of 8 families and 28 species of fish were identified in the mainstream and tributaries of Xe Pian and Xe Namnoy on the Bolaven Plateau (Table 3). In a further study to classify the fish species it was found that the most common taxon was Cyprinidae and Balitoridae as they had ten species respectively (35.7%) in the area, Channidae and Sistoridae followed by of two species respectively (7.1%), and lastly, Clariidae, Synbrachidae, Mastacembelidae and Anabantidae of one species (3.6%) respectively. Meanwhile, the seven species, Garra cyrano, Balitora annamitica, Schistura sp.1, Schistura sp.3, Schistura sp.4, Monopterus albus and Channa gachua were found only in Xe Pian river system, and the ten species such as Garra cambodgiensis, Mystacoleucus atridorsalis, Onychostoma meridionale, Opsarius pulchellus, Scaphiodonichthys acanthopterus, Schistura tizardi, Glyptothorax macromaculatus, Macronathus sp., Channa striata, and Anabas testudineus were found only in Xe Namnoy river system.
Among the total of twenty eight species of fishes found in the survey sites, Devario salmonatus, Poropuntius bolovenensis, Annamia sp., Sewellia elongata, Schistura bolavenensis, Schistura sp.1, Schistura sp.2, Schistura sp.3, Schistura sp.4, and Macronathus sp. were identified as the endemic species in the Bolaven Plateau, revealing a rate of endemic species of 35.7% in the area. Glyptothorax macromaculatus and Macronathus sp. were first collected on the Bolaven Plateau.
Meanwhile, Kottelat (2013) have reported 63 fish species on the Bolaven Plateau and downstream under the plateau. Among them, 37 species have been collected on the Bolaven Plateau. In this study, we collected 28 species on the Bolaven Plateau and such findings show a few difference when compared to the study in 2013, which reported 37 species. It is caused by difference of collecting methods. Kottelat (2013) used more diverse, as batterypowered electric shocker and ichthyocides.
The fish species affected by the Xe Pian-Xe Namnoy Hydroelectric project in the impact areas (Dam sites) and non-impact areas of Xe Pian and Xe Namnoy, consisted of fifteen fish species that were found in three survey sites (St. 4, 5, 6) in the impact area of Xe Pian. In Xe Namnoy, fourteen species were found in four survey sites (St. 10, 13, 17) in the impact area (Table 4).
Kottelat (2013) argued Devario salmonatus, Poropuntius bolovenensis, Schistura sp.2, Schistura sp.3, Sewellia elongata are rated endangered fish species because of their small distribution ranges and most of which will be either flooded by the reservoirs. On the one hand, endemic fish species, including 5 species in the Bolaven Plateau were discovered in impact area of Xe Pian and Xe Namnoy in this study. However, they were also found in the non-impact area of Xe Pina and Xe Namnoy as there are a large number of tributaries.
After the dam construction, the places where the water fills will become reservoir and the existing lotic system will become lentic system. Within the dam, fish species that live in flat waters will be prosperous. Contrariwise, fishes in flow waters will be almost disappeared. Thus, continuous surveys will be needed to view the changes of fish fauna after the dam construction completion.
3.Trophic Polymorphism of Poropuntius bolovenensis
Poropuntius species on the Bolaven Plateau have two mouth shapes (Figure 2). One is broad mouth, and the other is narrow mouth. Thus, Poropuntius species were captured to investigate their morphological characteristics in this species. However, as the major characteristics of the Poropuntius spp. were shared between the species, we could not distinguish between each (Table 5).
Of the Poropuntius species, P. bolovenensis was first discovered in Xe Namnoy and reported as a new species by Roberts (1998). He insisted that P. bolovenensis was divided into 3 subspecies, despite they inhabit the same site without barriers. However, his claim was later challenged by Kottlelat (2000), who argued that the claim is violated by the International Code of Zoological Nomenclature; and thus the three species should be regarded as main species rather than subspecies. The three species were named as P. bolovenensis, P. lobocheiloides, P. consternans (Kottelat, 2000). Later, Kottlelat (2000) discovered one more species in Houay Makchan, the tributary in Xe Namnoy and was named as P. solitus. According to Kottelat (2013), there are five species of Poropuntius (P. bolovenensis, P. consternans, P. lobocheiloides, P. solitus, and P. sp.) inhabiting the Bolaven Plateau. In his later survey conducted on Hydroelectric Power Project Environmental Impact Assessment, however, he described that P. consternans is a variation of Poropuntius bolovenensis. In conclusion, Kottelat (2013) reported that there are three species of fish, P. bolovenensis, P. lobocheiloides, P. solitus in Xe Namnoy, and two species of fish, P. solitus and P. sp. in Xe Pian.
Meanwhile, Kottelat (2000, 2013) argued that Poropuntius species that inhabits the Bolaven Plateau can be clearly distinguished by their characteristics such as mouth shapes, lateral line scales, branched soft rays of dorsal fin, predorsal scales (Table 6).
During the survey it was found that Poropuntius species cannot be distinguished by lateral line scales, dorsal fin rays, predorsal scales and scales below lateral line due to the overlaps of these (Table 5). All of the young individuals less than 70 mm (SL) of Poropuntius species have narrow mouth shape of the generalized type on the Bolaven Plateau. However, the mouth shape of adult individuals was divided to two phenotype (Roberts, 1998; Kottelat, 2013; Present study). Thus, we interpret Poropuntius lobocheiloides, P. solitus and P. sp. are the same species as P. bolovenensis and the phenotypic variation of mouth parts is the intraspecific trophic polymorphism.
It seems that the difference of their mouth shapes are trophic morph types caused by their unique feeding habit (Kornfield and Taylor, 1983; Robinson and Parsons, 2002; Swanson et al., 2003; Whiteley, 2007). The feeding behavior of P. bolovenensis was observed under the water by skin diving during the dry season, April. The individuals of broad mouth type always graze on benthic algae on the surface of bedrock. On the other hand, the individuals of narrow mouth type mainly graze on algae but sometimes suck on drift organic particles in rapids (personal observation). Thus, trophic polymorphism relates to common resource use in same habit and is thought to be a possible step in speciation.